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>J / k? f U, I I ^o J

Harvard College
Library

FROM THE BEQUEST OF

FRANCIS BROWN HAYES

CUm of 1839

OP LIZINGTON, MASSACHUSITTS

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THE

CAMBRIDGE NATURAL HISTORY

EDITED BY

S. F. HARMER, ScD., F.R.S., Fellow of King*s College, Cambridge ;
Superintendent of the University Museum of Zoology

A. E. SHIPLEY, M.A., Fellow of Christ's College, Cambridge;
University Lecturer on the Morphology of Invertebrates

VOLUME VIII

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:S&(Sm

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AMPHIBIA AND REPTILES

By Hans Gadow, M.A. (Cantab.), Ph.D. (Jena), F.R.S., Strick-
land Curator and Lecturer on Advanced Morphology of
Vertebrata in the University of Cambridge.

ILontion
MACMILLAN AND CO.. Limited

NEW YORK: THE MACMILLAN COMPANY
I 90 I

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S 7670.ll^(x)j

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PREFACE

LiNNA£US h£id but a poor opinion of the Amphibia and their
deacribera, or he would not have called the former " peseima
tetraque animalia," nor would he have dismissed the latter with
the terse remark : " Amphibiologi omnium paucissimi sunt
nullique veri." That was, however, nearly 150 years ago; and
at the present time there are fewer difficulties in writing a book
on Amphibia and Beptiles. Those who care for the study of
Amphibia and Beptiles — the Herpetologists, to give them their
scientific title — have never been numerous ; but most of them
have been serious students. One reason for the fact that this
branch of Natural History is not very popular, is a prejudice
against creatures some of which are clammy and cold to the
touch, and some of which may be poisonous. People who
delight in keeping Newts or Frogs, Tortoises or Snakes, are, as
a rule, considered eccentric. But in reality these cold-blooded
creatures are of fascinating interest provided they are studied
properly. The structure of animals is intimately connected with
their life-habits ; and this correlation is perhaps more apparent in
Amphibia and Beptiles than in any other class. The anatomist
who studies internal and external structure is as much struck
with the almost endless variety in details as he who takes the
trouble to observe the living animal in its native haunts, or at
least under conditions not too unnatural. He will agree with
V. von Scheffel's Toad " that those above seem to have no

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VI PREFACE

notion of the beauties of the swamp" — brilliantly coloured
Newts engaged in amorous play, concert-giving Frogs, and meta-
morphosing Tadpoles. The motto assigned to the Eeptiles seems
singularly appropriate when we consider that poisonous snakes
have been developed from harmless forms, and that many kinds
of reptiles have lost limbs, teeth, and eyesight in the process
of evolution.

The present work is intended to appeal to two kinds of
readers — to the field-naturalist, who, while interested in life-
histories, habits, and geographical distribution, beauty or strange-
ness of forms, is indifferent to the homologies of the metastemum
or similar questions ; — and to the morphologist, who in his turn
is liable to forget that his specimens were once alive.

A great portion of the book is anatomical and systematic.
It was necessary to treat anatomy, especially that of the skeleton,
somewhat fully, since it has long been recognised that it is
impossible to base a scientific classification upon external
characters. The reader familiar with Vertebrate anatomy has a
right to expect that questions of special moq^hological interest
will be dwelt upon at length. Those who have no anatomical
foundation must be referred to one of the now numerous intro-
ductory manuals on the subject.

The account of the .Vmphiljia is more complete than that of
the Rei)tilia. It was possible to diagnose practically all the
recent genera ; and this has lx*en especially done in the Anm'a,
in order to show how in an otherwise very liomogeneous group
almost any part of the body, internal or external, can l)e nuxlified
in kaleidoscopic variety. The same could not be done with the
Reptilia. Their principal groups, — called sub-classes in the
present work, in order to emphasise their taxonomic importance
in comparison with the main groups of Riixls and Maunnals, —
differ so much from each other tlmt it was decided to refrain

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PREFACE vii

from attempting a general account of them. Moreover, the
number of species of recent lizards and snakes is so l:>ewilder-
ing, the genera of many families being but tedious variations of
the same theme, that only those forms have been described
which are the most important, the most striking, or which the
traveller is most likely to come across. The student who wishes
to go farther into systematic details must consult the seven
volumes of the Catalogue of Reptiles in the British Museuw
(London, 1889-1896). Mr. G. A. Boulenger, the author of this
magnificent series, has rendered the systematic treatment of
recent Amphibia and Reptiles an easy task. During many
years of the most friendly intercourse I have profited on count-
less occasions by his ever -ready advice. Although he has
kindly read the proofs of the part dealing with the Amphibia
it would be unfair to associate him with any of its short-
comings or with contestable opinions, for which I alone am
responsible.

Cope's large work on the Crocodilians, Lizards, and Snakes
of North America {Rep. UX Nat. Mus. for 1898 (1900)) has
unfortunately appeared too • late to be used in the present
work.

The drawings on wood were, with few exceptions, made by
Miss M. E. Durham, mostly from living specimens — a procedure
which has to a great extent determined the selection of tlie
illustrations.

Since both the metric and the English systems of measure-
ments have been employed, it may be well to state for the
convenience of the reader that the length of a line of the text is
four inches or approximately ten centimeters.

I have frequently and freely quoted accounts of previous
authors instead of paraphrasing them. Especial thanks are due
to Messrs. Longmans, Green, and Co., and to Messrs. Murray,

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vni PREFACE

for their courteous permission to make several long quotations
from Sir J. E. Tennent's Cei/lon, and from H. W. Bates' Naturalist
on the River Amazons.

Lastly, a remark about my Editors. Instead of being a
source of annoyance they have rendered me the greatest help.

H. GADOW.

Cambridge, December 19, 1900.

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CONTENTS

PAOB

Preface v

Scheme of the Classification adopted in this Book . • . . xi

PART I. AMPHIBIA

CHAPTER I

Characteks and Definition— Position of the Class Amphibia in the
Phylum Vertebrata— Historical Account of the Classification of
Amphibia 3

CHAPTER II

Skeleton of Urodela and Anura — Skin — Colour-changing Mechanism—
Poison-glands — Spinal Nerves — Respiratory Organs — Suppression
of Lungs— Urino-genital Organs — Fecundation— Nursing Habits
— Development and Metamorphosis .11

CHAPTER III
Nkoteny— Regeneration— Temperature — Geographical Distribution . 63

CHAPTER IV
Stegocephali or Labyrinthodonts— Lissamphibia— Apoda ... 78

CHAPTER V
Lissamphibia {continvbd) — Urodela 94

CHAPTER VI
Lissamphibia (co.vt/a'fjkd)— Anura 138

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CONTENTS

PART II. REPTILIA
CHAPTER VII

PAGE

Definition and CHARArrERs— Position of the Class Reptilia in the

Phylum Vertebrata— Classification— Skull and Vertebrae . 277

CHAPTER VIII
Proreptilia — Prosauria— Theromorpha 285

CHAPTER IX
Chelonia— Atheoae— Thecophora 312

CHAPTER X

DiNOSAURIA — CrOCODILIA 412

CHAPTER XI
Plesiosauria — Ichthtosauria— Pterosauria— Pythonomorpha . . 473

CHAPTER XII

SaURIA— AUTOSAURI OR LaCERTILIA— LlZAUDS 491

CHAPTER XIII
Sauria (COAT/iVT^D)— 0 phi di a— Snakes 581

INDEX 651

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SCHEME OF THE CLASSIFICATION ADOPTED
IX THIS BOOK

CLASS AMPHIBIA.

Sub-Clasa.

8TEQOCE-
PHALI (p. 78)

Order.
Stogoco-

Sab-Order.
C Branch 10-

Family.

Sub-Family.

phali Lepo- 1 sauki (p. 80).

spondyli
(p. 80) I

Btogooe-
phali Temso-
spondyli
(p. 81)
Btegoce-
phaU
Stereo-
■pondyli
^(p.83)

' (p. 84)

Urodela

(p. 94)

AlSTOPODES

(p. 81).

PHIBIA(p.84)

( AOLOSSA

(p. 143).

Annra

(p. 138)

Phanero-

OLOSSA

(p. 152)

CuE(:iLIIDAE(p.89).

[ Amphiumidae
(p. 97).

Salamandridae I
(p. 102) 1

Pkoteiiiae
(p. 132).
, SiRENIDAE (p. 136).

( DiSCOGLOSSIDAE

(p. 152).

Pelobatidae

(p. 160).

BUFONIDAE
(p. 166).

UYLID.A.E (p. 185)-!

Cystignathidae I
(p. 209) 1

Kngystomatidae I

(p. 22:0

[ Ran I dak ([>. 237)

Desmognathinae (p. 102).
Plethodontinae (p. 103).
Amblystomatinae (p. 109).
Salamandrinae (p. 115).

Amphignathodontinae

(j). 188).
Hylinae (p. 189).
Uemiphi-actiuae (p. 210).
Cystigiiathinae (p. 211).
De ndroph ry nisei nae

(p. 224).
Engystomatinae (p. 225).
Dyscophinae (p. 235).
Genyophryninae (p. 236).
Ceratooatrachinae (p. 237).
Raiiinae (p. 238).
Deudrobatinae (272).

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Xll

SCHEME OF CLASSIFICATION

CLASS BEPTILIA (p. 277).

PBOBEPTILIA (p. 285). Eryops (p. 286). Cricotus (p. 287).

Sub-Class.

PBOSAUBIA

(p. 288)

THEBO-
MOBPHA

(p. 300)

CHELONIA

(p. 312)

Order. Sub-Order

( nicrosauri (p. 288).
\ Prosanri f Frotorosauri (p. 290).
[ (p. 290) \ Rhynchocephali (p. 292).

[ Fareiasaori (p. 304).
I Theziodontia (p. 30C).
I Anomodontia (p. 809).
I Placodontla (p. 311).

fAtheca(p. 383)

Family.

CBOCODIUA

(p. 431)

FLESIO-
SAUBIA

(p. 473)

ICHTHYO-
SAUBIA

(p. 478)

PTEBO
SAUBIA

(p. 484)

FYTHONO-
MOBPHA

(p. 487)

Thecophora

(p. 838)

DINOBAUBIA

(p. 412)

^Stegosavri
425).
Ornithopoda
426)

1 Ornit
I (p. 4

Sauropoda

(p. 418).
Theropoda

(p. 420).

Orthopoda

(p. 424)

Ceratopsia

(p. 430).

Psendosnchia (p. 432).
Parasnchia (p. 433).

Ea8nchia(p. 434)

' Nothosauri

(p. 476)

Plesiosauri

(p. 477)

Ichthyosauri

(p. 483).

Pterosaur!

(p. 486)

' Dolichosauri

(p. 489).
Mosasauri

, (p. 489).

f Cryptodiua
(p. 388)

Pleurodira I

(p. 388) ]

Trionychoidea
(p. 404)

Sphargidae (p. 838).
Chelydridae (p. 838).
Dermatemydidae (p. 341).

CiNOSTERNIDAE (p. 342).

Platysternidae (p. 345).
Testudinidae (p. 345).
Chelonidae (p. 378).
Pelomedusidae (p. 390).
Cuelydidae (p. 399).
Carettochelydidae (p. 404).
Trionychidae
(p. 404).

'Tei.eosauridae (p. 450).
Metriorhynchidae (p. 451).
Mackorhyxchidae (p. 451).
Gavialidae (p. 451).
Atofosauridae (p. 453).

GONIOPHOLIDAE (p. 453).
^CUOCODILIDAE (p. 454).

/ Mekosafridae (p. 476).
\ XOTHOSAVniDAE (p. 477).
f PLIOiSAUKIDAE (p. 477).

4 Plesio.sai'ridae (p. 478).
( Elasmosauridae (p. 478).

' Pterodactyli
(p. 486).
Pteranodontes

. (p. 487).

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SCHEME OF CLASSIFICATION

Xill

:3ut><:ias«.

Order.

Sub-Order.

r GeCKONES
(p. 502)

aAURIA

fp. 491)

LacerUlia

(p. 491)

•i Lacertae
(p. 513)

Chamaeleon-
tes (p. 567)

1 OpUdia
(p. 581)

Family. Sub-Family.

( Geckoninae (p. 507).
Geckokidae I Eublepharinae
(p. 507J ^1 (p. 512j.

I Uroplatinae (p. 512).
Agamidae (p. 515).
Iguamdae (p. 528).
Xenosauridae (p. 536).

ZON'URIDAE (p. 536).
ANGUIDAE(p. 537).

Helodeumatidae (p. 540).
Lanthanotidae (p. 541).
Varan ID ae (p. 542).
Xantuhiidae (p. 547).
Tejidae (p. 547).
Lacertiiiak (p. 549).
Gerrhosauridae (p. 559).
SriNciDAE (p. 559;.
Anelytkopidae (p. fj64).

DiBAMIDAE (p. 564).

Aniellidak (p. 564).
Amphksbaexidae (p. 565).
.Pyoopodidae (p. 567).

JChamaeleontidae (p. 573).

Typhlopidae (p. 693).
GLArCONIIDAE (p. 594).
Ilysiidae (p. 594).
Uropeltidak (p. 595).

BoiDAvrn 6Q6U^ Pythoninae (p. 5981.
UOIDAE (p. 596)^ g^.j^^^ ^j^ gQj^

Xenopeltidae (p. 605).

Acrooliordiuae

Aglypha
(p. 606)

Opistho-
glypha
(p. 623)

Protero-
glvjiha
(p. 625)

A.MDLY(E-

phalidae
(p. 637).

VlPEKIIlAE

(p. 6.37)

(p. 606).
Colli hriuae (p. 607).
Kliachiodoutinae
, (p. 622).
( Dij)sadomorphinat'

(p. 623).
I Elachistodontinac
^1 (l». 625).
I Homalo])sinae

I (p. 625;.
i Elaiiiiiac fp. 626 j.
-! Ilyarophinae.
\ (p. 63r.).

J Viperiiiae (]>. 638).
I Crotaliiiae (p. 644,.

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PART I
AMPHIBIA

VOL. VIII * B

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' 's scheint, dass die hier obeii keine
Ahnung haben von dem Sampf nnd
Seiner Pracht."

The "plattgedriickteKrote,"

Scheffel's Trompeter von SdkkingfM.

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CHAPTER I

AMPHIBIA

CHARACTERS AND DEFINITION POSITION OF THE CLASS AMPHIBIA

IN THE PHYLUM VERTEBRATA HISTORICAL ACCOUNT OF THE

CLASSIFICATION OF AMPHIBIA

A Bird is known by its feathers, a Beast by its hairs, a Fish by
its fins, but there is no such obvious feature which characterises
the Amphibia and the Eeptilea In fact, they are neither fish,
flesh, nor fowL This ill-defined position is indicated by the
want of vernacular names for these two classes, a deficiency which
applies not only to the English language. All the creatures
in question are backboned, creeping animals. Those which are
covered with horny scales, and which from their birth breathe
by lungs only, as Crocodiles, Tortoises, Lizards, and Snakes, are
the Eeptilea. The rest, for instance, Newts or Efts, Frogs and
Toads, are the Amphibia. Their skin is mostly smooth and
clammy and devoid of scales ; the young are different from the
adult in so far as they breathe by gills and live in the water,
before they are transformed into entirely lung-breathing, terres-
trial creature& But there are many exceptions. Proteus and
Siren the mud-eel, always retain their gills ; while not a few
frogs undergo their metamorphosis within the egg, and never
breathe by gilk. If we add the tropical limbless, burrowing
Coecilians, and last, not least, the Labyrinthodonts and other
fossil forms, the proper definition of the class Amphibia, — in
other words, the reasons for grouping them together into one
class, separated from the other backboned animals, — requires the
examination of many other characters.

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AMPHIBIA CHAP.

So far as numbers of living species are concerned, the
Amphibia are the least numerous of the Vertebrata. There are
about 40 limbless, burrowing Apoda ; 100 Urodela or tailed
two- or four-footed newts, and about 900 Anura, or tailless,
four-footed frogs and toads ; in all some 1000 different species.
Few, indeed, in comparison with the 2700 Mammals, 3500
Eeptiles, nearly 8000 Fishes, and almost 10,000 Birds. But
we shall see that the Amphibia have not only " had their day,"
having flourished in bygone ages when they divided the world, so
far as Vertebrata were concerned, between themselves and the
Fishes, but that they never attained a dominant position. Inter-
mediate between the aquatic Fishes and the gradually rising terres-
trial Eeptiles they had to fight, so to speak, with a double front
during the struggle of evolution, until by now most of them have
become extinct. The rest persist literally in nooks and comers
of the teeming world, and only the Frogs and Toads, the more
recent branch of the Amphibian tree, have spread over the whole
globe, exhibiting almost endless variations of the same narrow,
much specialised plan. The greatest charm of the Anura lies in
their marvellous adaptation to prevailing circumstances ; and
the nursing habits of some kinds read almost like fairy-tales.

Characters of the Amphibia.^

1. The vertebrae are (a) acentrous, (6) pseudocentrous, or (c) notoceiitroiis.
•2. The skull articulates with the atlas by two condyles which are formed
by the lateral occipitals. For exceptions see p. 78.

3. There is an auditory columellar apparatus, fitting into the fenestra o\^lis.

4. The limbs are of the tetrapodous, pentadactyle type.

ft. The red blood -corpuscles are nucleated, biconvex, and oval.

6. The heart is (a) divided into two atria and one ventricle, and (h) it has

a conus provided with valves.

7. The aortic arches are strictly symmetrical.

8. Gills are present at least during some early stages of development.

9. The kidneys are provided with persistent nephrostomea

10. Lateral sense-organs are present at least during the larval stage.

1 1. The vagus is the last ci*anial nerve.

12. The metiian fins, where present, are not supporte<l by spinal skeletal raya

13. Sternal ribs and a costal or true sternum are absent

14. There i.** no paired or unpaired medio- ventral, copulatory apparatus.
If). Development takes place without amnion and allantois.

None of these characters is absolutely diagnostic, except 1 (r), and this
applies only to the Anura and most of the Stegocephali.

References to explanations of the terms used below will be found in the index.

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I CHARACTERS AND POSITION 5

Numbers 1 (6), 1 (c)y 2, 3, 4 and 12 separate the Amphibia froiii the Fishes.

Numbers 1, 6 (6), 7, 8, 9, 11, 13, 15 separate them from the Reptiles,
Birds, and Mammals.

Number 2 separates them from the Fishes, Reptiles, and Birds.

Number 5 separates them from the Mammals.

Number 6 (a) separates them from the Fishes (excL Dipnoi), Birds ami
Mammals,

We can, therefore, very easily define all the Amphibia, both
recent and extinct, by a combination of the characters enumerated
above. For instance, by the combination of numbers 2, 3 or 4
with either 7, 8, 9, 11, 13 or 15.

Ampkicondylotbs Attavmia would be an absolutely cx)rrect and
all-sufficient diagnosis, but it would be of little use in the deter-
mination of adult specimens ; and the tetrapodous character is of
no avail for Apoda. Amphicondylovs animals without an intra-
cranial hypoglossal nerve is a more practical diagnosis.

In the case of living Urodela and Anura the absence of any
scales in the skin affords a more popular character ; it is unfor-
tunately not applicable to the Apoda, many of which possess
dermal scales, although these are hidden in the imbricating
transverse rings of the epidermis ; and the frequent occurrence of
typical scales of both ecto- and meso-dermal composition in many
of the Stegocephali forces us to discard the scales, or rather their
absence, as a diagnostic character of the class Amphibia. The
same applies to the mostly soft, moist, or clammy, and very glan-
dular nature of the skin.

The position of the class Amphibia in the Phylum Verte-
brata. — There is no doubt that the Amphibia have sprung from
fish-like ancestors, and that they in turn have given rise to the
Eeptilia. The Amphibia consequently hold a very important
intermediate position. It was perhaps not a fortunate innova-
tion when Huxley brigaded them with the Fishes as Ichthyopsida,
thereby separating them more from the Sauropsida ( = Eeptilia
and Aves), than is justifiable, — perhaps more than he himself
intended. The connecting-link, in any case, is formed by the
Stegocephali ; all the recent Orders, the Apoda, Urodela, and
Anura, are far too specialised to have any claims to the direct
ancestral connection& The line leading from Stegocephali to
fossil Eeptiles, notably to such Proreptilia as Uryops and
Cricotus, and even to the Lepospondylous Prosauria, is extremely
gradual, and the steps are almost imperceptible. Naturally,

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AMPHIBIA

assuming evolution to be true, there must have lived countless
creatures which were a " rudis indigestaque moles," neither
Amphibia nor Keptilia, in the present intensified sense of the
systematist. The same consideration applies equally to the line
which leiids downwards to the Fishes. But the great gulf
within the Vertebrata lies between Fishes and Amphibia, between
absolutely aquatic creatures with internal gills and " fins," and
terrestrial, tetrapodous creatures, with lungs and fingers and
toes. On the side of the fishes only the Dipnoi and the Crosso-
pterygii come into consideration.

The piscine descent of the Amphibia is still proclaimed by
the following features. — (1) The possession by the heart of a
long conus arteriosus, provided with, in many cases, numerous
valves, or at least (Anura) one series at the base, another at the
beginning of the truiicus where the arterial arches branch off;
(2) the strictly symmetrical arrangement of these arches; (3)
the trilocular heart is still like that of the Lung-fishes or Dipnoi ;
(4) the occurrence of as many as four or even five branchial
skeletal arches in the larval stage; (5) the glottis is supported
by cartilages which themselves are derivatives of posterior visceral
arches; (G) the development of the vertebrae (Stegocephali and
Urodela) from four paii*s of arcualia, and the formation of the
intervertebral joints by a split across the intervertebral ring of
cartilage : this feature is unknown in Keptilia, but it occurs also
in LepidosteuSy most probably also in Polypterus ; (7) the hypo-
glossixl still retains the character of a post-cranial or cervical
spinal nerve; (8) the presence of lateral sense-organs; (9) the
possession of external gills is of somewhat doubtful phylogenetic
value, although such gills occur amongst fishes only in Dipnoi
and Crossopterygii. It is not unlikely that in the Amphibia
these organs owe their origin to entirely larval requirements,
while the suctorial mouth of the larvae of the Anura and many
fishes has certainly no ancestral meaning, but is a case of con-
vergent development.

The usual diagnoses of the Amphibia contain the statement
that they, or most of them, undergo a metamorphosis, or pass
through a larval stage. The same applies to various fishes ;
while, on the other hand, the larval (not ancestral) stage has
become permanent in the Proteidae and Sirenidae ; and lastly, we
cannot well speak of larvae in the viviparous Salamandra atru.

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CLASSIFICATION

The evolntion of an adequate dassiflcation of the Amphibia

has been a long process. Even their recognition as a class,
separate from, and of equal rank with that of, the Eeptilia, was
bj no means generally accepted until comparatively recent times.
A historical sketch of the laborious, often painful, striving for
light, in France and Germany, then in England, and lastly in
America, is not without interest.

The term Amphibia was invented by Linnaeus for the third class of animals
in his fiunous " Systema Naturae." It comprises a very queer assembly,
which, even in the 13th edition (1767), stands as follows : —

1. Rbftiles pedatt, with the four "genera" Testudo, Dracoy Lacerta^
and Eana. Lctcerta includes Crocodiles, Lizards, and Newts !

2. SeRPENTES APODE8.

3. Nantes pinnati. Elasmobranchs, Sturgeons, Laiujireys, and various
other fishes.

Laurenti, 1768, in a dissertation entitled "Specimen medicum, exhibens
Synopein Reptilium . . .," uses Brisson's term. Reptiles, and divides
them into : —

Beptilia balientia, these arc the Anura.
Gradtentia, namely the Urodela and Lizards.
Serpbntia, the SntJces and the Apoda.

Brongniart, 1800, " Essay d'une classification naturelle des Reptiles,"^ dis-
tinguishes : — '

Chelonu, Sacrii, Ophidii, Batrachii ; the last for the Frogs, Toads,
and Newts.

Latreille, 1804, "Nouveau Diet. Hiftt. Nat." xxiv.,- accepts the four Orders of
Brongniart's " Reptiles," but clearly separates the fourth Order,
"Batrachii," from the rest by the following, now time-honoured,
diagnoBiB: Doigis des pattes n^ayant pas d^ongles; des hranchies, du
fiwins pendant un temps; des melamorphoses. But there is not one
word about ** Amphibia " in opposition to " Reptilia."

Dumeril, 1806, " Zoologie analytique " (p. 90), and "^l^mens de lldstoire
naturelle," 1807, divides the " Reptiles batraciens," or " Batracii," into
EcAUDATi and Caudati ; he also introduces the terms " Amoures "
and " Urod^les " as their equivalents ; but since these terms appear in
the French form purists do not admit their having any claim to
recognition !

Oppel, 1811, "Die Ordnungen, Familien und Gattungen der Reptilien,"
establishes the term Apoda for the Coeciliae, and recognises their
affinity to the Ecaudata and Oaudata l)y removing them from the
Snakes.

De Blainville, 1816, "Prodrome d'une nouvelh* distribution du regne
animal " ^ —
Ahphibiens squamif^reb. [The Reptilia.]

„ NUDIPELLIF^RES ft. Ichthyoidos. [The Amphibia.]

* Bull, Soe. Fhilom. ii. p. 81. * Tableaux viithodiques, p. 61.

» BulL Soc. Philom, p. 11-3.

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AMPHIBIA cHAr.

GRADIENTIA

DiPNOA. [The Amphibia]

Merrem, 1820, " Tentameu systeniatis Amphibiorum."
Pholtdota. [The Reptilia.]
Batrachia: APODA.

SALIENTIA.

'Mutabilia [with metaniorphoBiB, e.g.

Newta.]
Amphipneusta [Perennibranchiate Uro-
deles.]

F. S. Leuckart, 1821, "Einiges ueber die fischartigeii Amphibien."^
MoNOPNOA. [The Reptilia.]

Twith temporary gills: Eiaudata + Caudata

pt.
with permanent gills : " Proteidae," Meno-
pama and Amphiuma.
Latreille, 1825, " Families naturelles du regne animal" The Vertebrata are
divided into Haematherma and Haemacryma. Tliese terms for wami-
and cold-blooded creatures were later on amended by Owen to Haemato-
therma and Haematocrya. The latter are divided by Latreille as
follows : —

Reptilia. Still including the Coeciliae amongst the Snakes.
. rCaducibranchiata.

\ Perenn ibranchiata.

PiSCBS.

Wagler, 1830, "Systema Amphibionun."

Testudines, Crocodili, Lacertae, Serpentes, Argues, Coeciuae,

RaNAE, ICHTHTODI.

Ranae I. AGLOSSA.

II. FHANEROGLOSSA : 1. Cauda nulla, [The Anura.]
„ „ 2. Cauda distincta. [The Sala-

mandridae.]
IcHTHYODi 1. ABRANCHIALES. Menopoma [Cryjftohranchus]

and Amphiuma.
II. BRANCHIALES, [The Perennibranchiate Urodela.]
J. Miiller, 1831, "Beitriige zur Anatomic . . . der Amphibien."^

Gymnophiona, Dbrotremata, Proteidae, Salamakdrina, Bat-
rachia.
J. Bell, 1836, Todd's " Cycloj)aedia of Anatomy and Physiology," Art.
" Amphibia."

Amphipneusta, the Perennibranchiate UrodeW ; Anoura, Urodela ;
Abranghia, Menop<yma. and Amphiuma : Apoda.
Stannius, 1856, " Handbucli der Zootomie : Anatomie der Wirbelthiei-e.*'
(2nd ed.)

Amphibia Monopnoa. The Reptilia.
Amphibia Dipncv. 1. Urodela. PERENNIBRANCHIATA.

DEROTREMATA: Amphinma and

Menopoma.
MYCTODERA}'

^ Jsin, lf*21. - TreviranuH' ZvUschr, /. Physiol. 1831, p. 190.

•' 5^/y»7, neck ; yui'w, close.

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CLASSIFICATION

2. Batrachia. A0L08SA.

PHANER0GL08SA : Systomata =
Engystomatidae.
Buibninae. Without manubrium

' stemi.
Raninae. With manubrium.
Hyloidea. With adhesive finger-
discs,
GYMNOPHIONA.
Qegenbaur, 1859, "Grundziige der vergleichenden Anatomie."

Amphibia as a separate claas, equivalent to that of the Reptilia, are
divided into the four Orders: PERENNIBRANGHIATA, SALA-
MANDBINAy BATRAGHIA, and GYMNOPHIGNA. In the
second edition of the "Grundzuge" (1870) they are divided into
URGDELA, ANURA, and GYMNOPHIGNA,
Huxley, 1864, "The Elements of Comparative Anatomy."
Mammals.

Sauboids, subsequently changed into Sauropsida = Reptilia + Aves.
IcHTHYOiDS, „ „ loHTHYOPSiDA = Amphibia +

Pisces.
Haeckel, 1866, "Generelle Morphologie."

Amphibia. A. Phractamphibia s. Ganocephala = Labyrinthodonta +
Peromela [Apoda}
B. LissAMPHiBiA s. Sozobranchia = Sozura [Urodela] +
Anui-a.
Cope, 1869.1

Steqocephali, Gymnophidia, Urodela, Proteidea, Trachy«tomata,
Anura.
Huxley, 1871, "A Manual of the Anatomy of Vertebrated Animals."
Amphibia I. Saurobatrachia [v.d. Hoeven's term] 8. Urodela
1. Proteidea.
•1. Salamandridae.
II. Labyrinthodonta.

III. Gymnophiona.

IV. Batrachia s. Andra.

Boulenger, 1882, ** Catalogue of the Batrachia Gradientia s. Caudata and
Batrachia Apod a," divides the Caudata simply into : SAI^AMAND-
RIDAE, AMPHIUMIDAE, PROTEIDAE, and SIRENIDAE.
1882, "Cat. Batrachia Salientia s. Ecaudata," see p. 140.
Cope, 1890, " Synopsis of the Families of Vertebrata." -'
Class Batrachia.

Sub-Class I. Stegocephali.

Order 1. Ganocephali : TrimerorJuichis, Archegosaunts.

2. Rhachitomi : Eryops

3. Embolomeii : Cricotus.

4. Microsauri : Branchiosaurus^ HylonomuSj etc.

' Proc. Ac. PMlad, p. 209. * Amertc. Natural, .\xiii. ji. 849.

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I O AMPHIBIA

Siil)-Cla68 II. Urodela.

Order 1. Proteidae : Proteus.

2. Pseudosauria. [All the rest of the Urodela -H

Coeciliidae.]

3. Trachystomata : SireiiidaeL
III. Salientia.

and F. Barasiu, 1890, "Zur Entwickluiigsgeschichte der Ceylonesificheu
Blindwiihle, Ichthyophis glutinosa." *

Sub-Class I. Archaeobatraghi s. Stkgocephali.
II. Nbobatrachi.
Order I. URODELA.

a. Salaniandroidea. [The Urodela.]
6. Coeciloidea = Amphiumidae + Coeciliidae.
2. ANURA.

The classification adopted in this volume is as follows : —

Class Amphibia.

Sub-Cla«»8 I. Phractamphibia.

Onler I. Stkgocephali Lepospondyli.
Sub-Older 1. Branchiosauri.
Sub-oi-der 2. Aistopod&s.
Older II. Stegocephali Temnosiwndyli.
Onler III. Stegocephali Stereospondyli.
Sub-Class II. Lissamphibia.
Onler I. Apoda.
Onler II. Urodela.
Order III. Anura.

Sub-order 1. Aglossa.
Sub-order 2. Phaneroglosc?fi.

Sarasins' Ergebnisse . . . Ceylon, 1887-1890.

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CHAPTER II

SKELETON OF URODKLA AND ANURA SKIN COLOUR -CHANGING

MECHANISM POISON - GLANDS SPINAL NERVES RESPIRA-
TORY ORGANS SUPPRESSION OF LUNGS URINO -GENITAL

ORGANS FECUNDATION NURSING HABITS DEVELOPMENT

AND METAMORPHOSIS

Skeleton of the Urodbla

The vertebral column. — The number of vertebrae in smallest in
the terrestrial, greatest in the entirely aquatic forms, and is excep-
tionally large in the eel-shaped Amphiuma. In the following
table the sacral vertebra is included in those of the trunk.

Trunk.

Tail.

Siren lacertina

. 22

35 + '

Nedturus maculatus

Proteus anguinus

. .30

28 +

Cryptohranchus alleghauiei

isis 20 or 21

24 +

C. scheuehzeri

. 21

C. japonicus

. 22

22 to 26

Amphiuma means

. G.3

.35 +

Amhlystoma tigrinum .

. 17 or 10

.32 +

^lamandra maculosa .

. 17

Triton cristatus .

. 17

Triton taeniatus .

14 or 1")

36 +

Triton palmatvs .

23 to 25

Salamandrina perspicilUtU

< . 15

32 to 42

Spelerpes fnscvs .

. 16

The vertebrae of the Urodela and those of the Apoda differ
from those of all the other Tetrapoda ^ by possessing no special
centra or bodies. That part which should correspond with the
centrum is formed either by the meeting and subsequent complete
co-ossification of the two chief dorsiil and ventral pairs of arcualia

' Crediier's term for all Vertebrates higher than fishes.

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URODELA

(tail-vertebrae), or entirely by the pair of chief dorsal arcualia.
There is consequently no neuro-central suture. Moreover, the
central region of each vertebra is strongly pinched in laterally,
widening towards the ends. Another feature of the vertebral
column of the Urodela is the possession of a considerable amount
of intervertebral cartilage, by which the successive vertebi-ae are
held together. This cartilage does not ossify, and it either remains
continuous, serving in its entirety and owing to its flexibility as
a joint, or it becomes more or less imperfectly separated into a cup
and ball portion, the cup belonging to the posterior end of the
vertebra. Such joints are called opisthocoelous, and occur in the
Desmognathinae and Salamandrinae. In the adult the cup and
ball frequently calcify, and the chorda dorsalis or notochord is
completely destroyed. Those vertebrae between which the inter-
vertebral cartilage remains unbroken, are called amphicoelous,
since in them, most obviously in macerated or dried skeletons,
the vertebrae appear hollowed out at either end. In such
amphicoelous vertebrae a considerable amount of the chorda
always remains, running in an unbroken string through the
whole length of the vertebral column. Towards adult life
the chorda becomes constricted, and is ultimately squeezed out
or destroyed, in the middle of the vertebra, by the invasion of
cartilage from the cliief arcualia. This intravertebrally situated
ciirtilage has Ijeen described erroneously as chordal cartilage.

The development of the vertebrae proceeds as follows. First
appear a pair of basidorsalia and a pair of basiventralia (Fig. 1, i,
B.D, B.V), blocks of cartilage, imbedded in and resting upon the
thin sheath of the chorda dorsalis. Next appears a pair of inter-
dorsal blocks, immediately behind the basidorsals ; and somewhat
later appears a pair of interventral blocks. These four pairs of
cartilages or " arcualia " each meet, above or below the chorda, and
form semi-rings, which again by extending upwards or downwards
fuse into complete rings, in such a way that the interdorsal and
interventral elements form the intervertebral mass spoken of above.
The basidorsals fuse with the basiventrals, and form the body of
the vertebra, the fusion being effected chieiiy by the calcification
and ossification of the lateral connecting portion of the skeleto-
genous layer. The basidorsalia form the neural arches witli
their unpaired short spinous or neural, and the paired anterior
and posterior zygapophysial processes. Concerning the basi-

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VERTEBRAL COLUMN

ventralia we have to distinguish between the trunk and the tail.
In the latter they produce a pair of ventral outgrowths or
haemapophyses, which ultimately enclose the caudal blood-vessels.
In the trunk the basiventral blocks of cartilage are suppressed ;
they appear in the early larvae, but disappear during or even
before metamorphosis.

Towards the end of the tail the vertebrae diminish in size,
and their constituent cartilages assume a more and more

Fio. 1. — 1-6, Five successive stages
of the development of a caudal
vertebra of a newt ; 6-7, the
second and the first cervical ver-
tebra of Chryptobranchus ; 8-9,
side view of the constituent
cartilaginous blocks of a caudal
vertebra (8) and a trunk -ver-
tebra (9) of Archegosaunts as
typical examples of Temno-
spondylous quadripartite and
tripartite vertebrae. The cross-
hatched parts indicate the artic-
ular facets for the ribs. The
anterior end of all the vertebrae
looks towards the right side.
a/y In 7, articulating facet for
the occipital condyle ; B.Djhasi-
dorsal piece or neural arch ;
Ji.Vj basiventral piece or ven-
tral arcli ; Ch, chorda dorsalis,
or notochord ; 7. Z>, interdorsal
piece ; /. T, interventral piece :
I.V.Ly intervertebral ligament ;
A\ spinal nerve — these are num-
bered I, II, III in 6 and 7 ; R,
rib ; T, in 7, rib-like tubercle
on the first vertebra.

indifferent shape, until they become confluent into a continuous
rod of cartilage, resembling in this respect the Dipnoi and
Holocephali. A periodical revival of this rod, at least of its
connective tissue, appears in the tail-filament of the male Triton
jxilnuitiis during the breeding-season.

The first vertebra, called the atlas, because it carries the
head, is remarkable for the possession of an odontoid process.
The latter is formed by a pair of cartilages and represents part
of a vertebra, the dorsal portion of which seems to have been
added to the occipital part of the cranium.

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14 URODELA CHAP.

All the trunk -vertebrae, with the exception of the atlas,
carry ribs, at least vestiges thereof. Owing to the early dis-
appearance of the basiventral cartilages the capitular portions of

the ribs are much reduced, and
are mostly represented by strands
of connective tissue only. The
ribs develop therefore occasion-
ally at some distance from the
vertebral column, and that por-
tion of the rib which in the
metamorphosed young newt looks
like the capitulum is to a great
extent really its tuberculum.

Fig. 2.— Transverse section through a Witness the position of the ver-

trunk- vertebra of a larva of &/a- ^j^^^al artery, which Still indi-
inandra macidoaa^ enlarged. The nght "^

side shows the actually existing state, cates the true foramen trsus-
whUe on the left side the rib and its yersarium. The homologies of

attachments are restored to their pro- '^* "*•**»*"*• -^"^ **v*x*v vgx^>« v

bable original condition. A, Verte- these parts are Still more ob-

bral artery within the true transverse „^„^^j "u„ ±.\^^ x* ^4. 4.1,^4. ^ •^^^r

canal ;^.r, remnant of the basi-ventral SCUred by the fact that a neW

cartilage; Ch, chorda dorsalis; Sp,c, procesS growS OUt from the rib,

Hpinal canal ; *, the false transverse , i'-Li.i_ ^ j.*.

canal. t)y which the latter gams a new

support upon a knob of the
neural arch. Thus an additional foramen is formed, sometimes
confounded with the true transverse canaL The meaning which
underlies all these modifications is the broadening of the body,
the ribs shifting their originally more ventral support towards
the dorsal side. The whole process is intensified in the Anura ;
it is an initial stage of the notocentrous type of vertebrae. The
transverse ossified processes of the adult are often much longer
than the vestiges of the ribs themselves, and are somewhat com-
plicated structures. They are composed first of the rib-bearing
cartilaginous outgrowths of the neural arches ; secondly, of a broad
string of connective tissue which extends from the ventro-lateral
corner of the perichordal skeletogenous layer to the ribs.

The shoulder-girdle is extremely simple. It remains almost
entirely cartilaginous, and the three constituent elements are not
separated by sutures. Ossification is restricted to the base of the
shaft of the scapula, and may extend thence over the glenoid
cavity. The coracoids are broad, loosely overlap each other, and
are " tenon and mortised " into the triangular or lozenge-shaped

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LIMB-GIRDLES I S

cartilaginous sternum, which latter has no connection with the
ribs. The precoracoid is a large, flat process, directed forwards,
not meeting it& fellow ; it is absent in Siren.

The humerus articulates with both radius and ulna, and these
two bones of the forearm remain separate. The elements which
compose the wrist and hand exhibit an almost ideally simple
arrangement, slightly varied by the frequent fusion of two or
more neighbouring carpalia into one, and by the reduction of the
number of fingers. Most frequently the intermedium and the
ulnar carpal element fuse together, and there is more often one
centrale instead of two. The wrist and hand of the Urodela
represent, however, no longer the entirely primitive pentadactyle
type, owing to the loss of one finger together with its metacarpal
and carpal element. Comparison with the Anura makes it
probable that the Urodela have lost the poUex, their four fingers
being consequently the 2nd, 3rd, 4th, and 5th. Siren has four
or three fingers ; Protetts has only three fingers and three large
compound carpal cartilages. In Amphiuma, with either three or
two fingers, the ulnare, intermedium, and carpale are fused
together, the radiale w^th the neighbouring carpale. The number
of phalanges in the four-fingered species is generally 2, 3, 3, 2
respectively.

The pelvic girdle. — The ilium stands vertically to the vertebral
axis, slanting slightly forwards and downwards. It is attached
by means of a rib to only one vertebra, and this ilio-sacral
connection is acetabular in its position, i.e. it lies in the same
transverse plane with the acetabulum, in other words vertically
above it. The ventral portion of the pelvis is formed by one
large continuous mass, the united pubo-ischia, the anterior or
pubic portion of which extends forwards in the shape of a broad
triangle {Nectiirus) or as a slender, stalked, Y-shaped cartilage,
the epipubis, which is often movably jointed at its basa The
lateral portion of the pubic cartilage is always perforated by
the nervus obturatorius. Ossification is restricted to the ischium
and to the middle of the shaft of the ilium. The acetabular fossa
for the femur is closed. The tibia and fibula remain separate.
The foot is still more primitive than the anterior extremity, as
the majority of Urodela possess the full complement of five
toes, with 2, 2, 3, 3, 2 phalanges respectively. Concrescence of
the tarsalia applies most frequently to the fourth and fifth distal

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1 6 URODELA

CHAP.

and to the two centralia ; exceptional, for instance, in Crypto-
hranchns Japonicus, are as many as three centralia, but this is an
individual, even a one-sided variation, as shown for instance by
a spedmen in the Cambridge Museum. Loss of the fifth toe
occurs sporadically in genera of different groups, namely, in
ScUamandrella, Batrachyperiis, ScUamandrina, Necturus, Manculus,
Batrachoseps. In Amphiuma the number is reduced to three
or two ; in Proteus to two ; and in Siren the hind limbs,
with their girdle, are altogether absent. Lastly, in some species
of Spelerpcs and Batrachoseps both fore and hind limbs have
become so small as to be practically without function, parallel
cases being found among various Scincidae and other Lizards.

Tlie hyoid apparatus is still very primitive in many,
especially in larval, Urodela. Besides the hyoid there are as many
as four pairs of branchial arches, which, however, decrease in size
and completeness, so that the last two have lost their connection
with the median copular piece, and become attached in various
ways to the second branchial arch. This is the arrangement
apparently in all larvae, but four pairs of branchials persist in
the adult Siren, Amphiuma, and Cryptohranchus alleghaniensis.
The whole branchial apparatus is reduced to three pairs of
arches in Nectvrus and Protens, to two in the adult Crypto-
h^anc-Jius japonicus and in the Salamandridac. Of considerable
interest is the vestige of a fifth pair of arches in the larvae of
Triton and Salamandra, in the 8hai>e of a pair of tiny cartilages,
which lie in front and on each aide of the opening of the
trachea, and give T\m to tlie formation of the laryngeal cartilages,
better developed in the higher Vertebrata.

The following are noteworthy characters of the skull of
Urodela. The articulation of tlie skull with the vertebral column
is not always effected entirely by the two condyles of the lateral
occipital bones, but the median basal cartilage often }X)8sesse8 a
pair of facets for the odontoid-like process of the first vertebra ;
such additional facets are perhaps best developed in Crypto-
hranchus and in the Salamandrinae.

The middle portion of the primitive, cranium, from the exit
of the optic ner\^e to the ethmoid cartilage, is formed by a pair
of separate bones, the orbito- sphenoids. The parietal and
frontal bones remain separate. One or more periotic bones
exist, besides the prootic, in the aquatic families.

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SKULL

A pair of prefrontal bones is present in most Salamandridae,
e.g. Salamandra, Triton, Amhlystoma, especially in the larva, and
in CryptohranchiLS ; these bones are absent in Amphluma, Necturus,
Proteus, and Siren.

The lacrymalia are still separate in some Amblystomatiuae,

Fio. 8. — Skulls of various
Urodela. 1, Salaniandra ma-
culosa, ventral view, and 2,
dorsal view ; 3, Axolotl stage of
Amblystofna ; 4, adult stage of
Avibly stoma ; 5, Salamandrina
perspicillata (after Wieders-
heim) ; 6, Salamandra ma-
culosaf dorsal view of the lower
jaw. A, Articulare ; C„ 63,
outer and inner occipital con-
dyles ; Chf choana or posterior
nasal opening ; d, dentary ; JC,
ethmoid ; F^ frontal ; ZO,
lateral occipital ; if, maxillary ;
iV, nasal ; ^'b^nostril ; OS^ orbito-
sphenoid ; P, parietal ; /y, pre-
frontal ; Pit palatine ; Pm^ pre-
maxillary ; Po^ prootic ; PS,
paraspbenoid ; /*<, pterygoid ; Q,
quadrate ; S, angulo-splenial ;
Sq, squamosal ; St, stapes ; Vo,
vomer ; II, VII, X, exits of the
optic, facial, and glosso-vagun
nerves.

e.g. Ranidens and Hynohius. A pair of nasalia are generally
present, but are absent in Necturus, Proteus, and Siren. The
l^irasphenoid is furnished with teeth in the Plethodontinae and
Desmognathinae.

Separate palatine bones exist in Necturus and Proteus, and in
the larva of AmUystoma, but in the adult form they fuse with
the vomers, producing the vomero-palatines characteristic of the
majority of Urodela.

VOL. vni c

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1 8 ANURA CHAP.

The pterygoid bones are most fully developed, so as to reach
the vomero-palatines, in the Amblystomatinae, in Nectums, and
in Proteus ; they are reduced, so as to leave a gap, in Crypto-
branchus, and still more in the Salaraandrinae ; they are absent
in Amphiuma and in Siren.

The quadrates are directed forwards in NectvruSy Proteus, and
Siren, while in the other Urodela they extend transversely and
almost horizontally. The hyomandibular remnant, the so-called
operculum, is small, and forms a plate which fits into the fenestra
ovalis, extending as a ligamentous process upon the quadrate.

The quadrato-jugal elements are reduced to ligaments. In
many Salamandrinae the large orbito-temporal space is divided
into an orbital and a temporal fossa by an arch which is formed
by the meeting of two corresponding processes from the
squamosal and frontal bones respectively. This bridge is rarely
bony {Salarnandrina, Triton), mostly ligamentous : — apparently
a reminiscence of the Stegocephalous condition. The two pre-
maxillary bones are liable to fuse into one, for instance in •
Cryptohratichus, generally in adult Tritons. They are most
reduced, and are toothless, in Siren,

The two maxillary bones are absent only in Nectiirus, Proieus,
Typhloviolgey and Siren. Their posterior end is frequently free,
loosely connected by ligaments with the pterygoid in Crypto-
Itranchus ; or with the distal portion of the quadrate, and in this
case .either just touching it {Tylototriton), or forming a broad
junction (Pachytriton).

Each half of the lower jaw consists of a dentary, articular
and angulo - splenial. The splenial remains as a separate
element in Siren ; in others only during the larval period. There
are no mento-Meckelian elements.

Skeleton of the Anura

The vertebral column. — The distinctive peculiarities of
the vertebrae of the Anura are that they are notocentrous, and
that about a dozen of them are modified and fused into an os
coccygeum. The whole column is the most specialised found in
the Vertebrata ; and various stages are rapidly hurried through
and obscured caenogenetically during the embryonic development-
Paired cartilages appear on the dorsal side of the thin chordal
sheath, and whilst tending to enclose the spinal cord in a

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11 THE DEVELOPING VERTEBRAL COLUMN 19

canal, their bases grow head- and tail-wards into what will
ultimately become the intervertebral region. This extension of
cartilage leads to a fusion with that of the next following pair
of arches, so that the axial column at this early stage consists of
a right and left longitudinal ridge of cartilage which sends off
dorsal processes, neural arches, in metameric succession. Next,
the intervertebral cartilage increases in such a way as to
constrict the chorda either laterally (Ranu) or obliquely from
above downwards and inwards (Bufo, Hyla), We recognise in this
cartilage the interdorsalia. Ventral arcualia are late and much
obscured. There is scarcely any cartilage which could represent
the interventralia, the intervertebral cartilage being almost
entirely made up of the interdorsalia. These fuse together and
form a disc or nodule, which later fuses either with the
vertebra in front, and in this case fits into a cup carried by the
vertebra next behind (precocious vertebrae), or the knob is added
to the front end of the vertebra, fitting into a cup formed by
the tail end of the vertebra next in front (opisthocoeloiis
vertebrae). Much later than the two longitudinal dorsal bands
there appears on the ventral side an unpaired band in which
appear metamerically repeated swellings of cartilage, likewise
impaired. These swellings become confluent, in a way similar
to that which produced the dorsal bands, and form the unpaired
ventral band of cartilage, the hypochordal cartilage of some
authors. The swellings in this band, equivalent to the basi-
ventralia, become semilunar in a transverse view, their horns
tending upwards towards the basidorsal cartilages, but there is
no actual meeting. Both dorsal and ventral elements are,
however, joined together and form the chief portion of the vc r-
tebrae, owing to the rapidly proceeding calcification and latt r
ossification of the all-surrounding " membrana reuniens " or
skeletogenous layer so far as that is not cartilaginous.

Procoelous vertebrae exist in the overwhelming majority of
Anura ; opisthocoelous are those of the Aglossa, the Discoglossidae,
and of some Pelobatidae. The systematic value of this pro-
or opistho-coelous character has been much exaggerated. A^'e
have seen that the centra of the vertebrae of the Anura are
formed entirely by the interdorsal elements, hence the term
"notocentrous," and these centra sometimes remain in adult
specimens of Pddbates as separately ossified and calcified pieces,

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20 ANURA

not fused with the rest of the vertebrae. This important dis-
covery has been made by Boulenger, but Stannius had previously
mentioned a specimen of Felohates in which the second and
fourth vertebrae are biconvex, the third, sixth, and eighth bicon-
cave. Moreover, since the sacral vertebra, generally the ninth,
in all the Anura is invariably biconvex, the eighth being
biconcave in the procoelous families, opisthocoelous like the
remaining seven vertebrae in the other families, it is not
difiScult to imagine that in the Anura the production of pro- or
opistho-coelous vertebrae depends simply upon the centra or
articulating knobs happening to fuse either with the hind or the
front end of the vertebrae. This must of course ultimately be
determined by a mechanical problem of motion.

A second type of the vertebrae amongst the Anura is the
epichordal type, an exaggeration in degree of the notocentrous
tendencies of the more usual perichordal arrangement. It shows,
namely, the almost complete suppression of all the ventral
cartilaginous elements, so that the chorda remains for a long
time on the ventral surface of the axial column in the shape of
a flattened longitudinal band. These two types are not un-
connected. The suppression of the ventral elements applies
most typically to the trunk region, while hypochordal cartilage
exists in the anterior cervical vertebrae, and above all in the
coccyx. Typically epichordal are the vertebrae of Fipa, Xeno-
piis, Bomhiiiato)\ Pelohntes, Discoglossvs and Alytes. It is
significant that the epichordal often coincide with opisthocoelous
vertebrae, and still more suggestive is the fact that Bomhina.i(yr
is eminently aquatic, Pij)a and Xenoims entirely so, having lost
the tympanum, at least externally. The epichordal feature is
not necessarily indicative of relationship. It has probably been
developed independently in various groups, in correlation with a
resumption of aquatic life. Various genera of Pelobatidae and
most likely some Cystignathidae, e.g. rseudis, will not improbably
connect the two types and their several correlated features, for
instance, the frequent reduction of the tympanic cavity.

Theoscoccygeum has retained rather primitive features in
so far as much dorsal and ventral cartilage is developed ; but this
has almost entirely lost its metameric arrangement, and the
posterior half of the coccyx is formed chiefly by the ventral mass
of cartilage, while the dorsal elements are more or less reduced.

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VERTEBRAL COLUxMN

Only two vertebrae, generally the tenth and eleventh of the
whole column, are clearly visible, each being composed of a pair
of dorsal and a pair of ventral cartilaginous blocks. The sacral
vertebra articulates with the coccyx by one or two convexities,
but in the Aglossa, in some Pelobatidae, and a few others, the
coccyx is fused with the sacral vertebra. Beyond the first
and second component vertebrae of the embryonic coccyx, the
cartilage is continued in the shape of two dorsal, and one ventral,
bands, which soon fuse with each other. Dorsally this cartilage
surrounds the spinal cord ; the latter degenerates towards the
end of the tadpole-stage, leaving, however, the empty spinal
canal. The chorda, completely surrounded by cartilage, persists
into the post -larval stage, but is destroyed long before the
creature attains maturity. Ultimately the whole coccyx ossifies.

The tail proper, namely that portion which is absorbed
during the metamorphosis, remains throughout its existence in
an apparently primitive condition. The chorda dorsalis and the
spinal cord extend through its whole length, surrounded ])y
continuous connective tissue without any cartilage ; in fact it
represents a piece of typical vertebral column before the appear-
ance of cartilage. The reduction of this swimming organ begins
at the hind end.

The vertebral column of the adult. — The first vertebra
(we will call it the atlas since it carries the skull) is not, as in
the Urodela, provided with an odontoid process. It articulates
by two cups with the condyles of the occiput. In some Anura it
co-ossifies, rather incompletely, with the second vertebra, regularly
in the fossil Palaeohatrcichus, often in Ceratophrys, Breviceps,
and occasionally in Pelohates, BvfOy Eana, and Xenopus,
This is, however, no justification for looking upon the first
vertebra as a complex of two vertebrae, although the atlas is
frequently very thick and broad, and even carries, in the
Aglossa, considerable lateral wings or diapophyses. Those of the
trmik-vertebrae are often very long, acting thereby as substitutes
for ribs which are absent, except on the second, third, and fourtli
vertebrae of the Discoglossidae, and on the second and third of
the Aglossii. In the adult Aglossa these ribs fuse with the
processes which carry them.

The diapophyses of the sacral vertebra carry no ribs, the
ilia being attached to them directly. They are either cylindri(.al

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ANURA

as ill the Eanidae and Cystignathidae, or they are more or less
dilated as in all the other families, most strongly in the
Pelobatidae and the Aglossa. In some members of the large
sub -family of the Cystignathidae the otherwise cylindrical
diapophyses are slightly dilated.

The sacrum is formed by the ninth vertebra, but there are
a few interesting exceptions. Frlohates, Pipa, and Hymenochir%bs
possess two sacral vertebrae ; and, neglecting individual abnor-
malities, these three genera form the only exception amongst
recent Amphibia. In the three genera the coccyx is fused with
the second sacral vertebra, and such a fusion occurs elsewhere
normally only in Bombinator with its single^ sacral vertebra.
The morphologically oldest condition is normally represented by
Pelohatcs, the sacral vertebrae being the tenth and ninth. One

Fkj. 4. — Dorsal view of the sacral or ninth vertebra (9), with the attachment of the ilium,
of (I) Rana teinpomria, (2) Jiu/o vtt/gariSf showing the whole coccyx and pelvis, (3)
l*elobates fu^ciis, as examples of cylindrical and of dilated sacral diapophyses.
(About nat. size.) a. Acetabulum ; c, coccyx ; i, ilium ; z, anterior zygapopbyses.

cast* has been recorded by Boulenger of Bombinator jiachi/p^cs
'' with eleven segments,** the last carrying the ilium. Individual
lop-sided abnormalities have been described in Bombinator and
Alytes, where the right ilium articulated with the tenth, the left
ilium with the ninth vertebra. This shifting forwards of the
ilium to the extent of one metamere has been continued further
in Pipa, in which the sacrum is formed by the ninth and eighth
vertebrae, their diapophyses fusing on either side into extra broad
wing-like expansions. In old specimens oi Palaeohatrachiisfritschi
the seventh vertebra is in a transitional condition, the ilium
being carried by the ninth and eighth, and sli^jlitly also by the
diapophyses of the seventh vertebra ; and in P. (Jiluviaims the

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VERTEBRAL COLUMN 23

diapophyses of all these vertebrae are united iuto one broad plate
to which the ilia are attached. Lastly, in Hymenochirus the
first sacral is the sixth vertebra, and this creature has thereby
n^duced the pre-sacral vertebrae to the smallest number known.

This shifting forwards of the iliac attachment implies the
conversion of original trunk into sacral vertebrae, and the
(»riginal sacral vertebra itself becomes ultimately added to the
urostyle. The second sacral, the tenth of Pelobates, the ninth
of Fipa, and the tenth on the right side of the abnormal
BombinatoTy are still in a transitional stage of conversion. In
Discoglossidae the tenth is already a typical post-sacral vertebra,
and is added to the coccyx, but it still retains distinct, though
short, diapophyses. In the majority of the Anura the tenth
vertebra has lost these processes, and its once separate nature is
visible in young specimens only. In Bomhinator even the
eleventh vertebra is free during the larval stage. In fact the
whole coccyx is the result of the fusion of about twelve or more
vertebrae, which from l)ehind forwards have lost their in-
dividuality. We conclude that originally, in the early Anura,
there was no coccyx, and that the ilium was attached much
farther back ; and this condition, and the gradual shifting for-
wards, supply an intelligible cause of the formation of an os
coccygeum. The fact that the sacral vertebrae of the Anura
possess no traces of ribs as carriers of the ilia, is also very
suggestive. The ilia have shifted into a region, the vertebrae
of which had already lost their ribs. By reconstructing the
vertebral column of the Anura, by dissolving the coccyx into
about a dozen vertebrae, so that originally, say the twenty-first
vertebra carried the ilia, we bridge over the enormous gap which
exists between the Anura and Urodela. That whole portion of
the axial continuation behind the coccyx, more or less coinciding
with the position of the vent, is the transitional tail.

The disappearance of both notochord and spinal cord, and
the conversion of the cartilaginous elements into a continuous
rod in the case of the os coccygeum, find an analogy in the
hinder portion of the tail of Dipnoi and Crossopterygii, and in
the tail-end of most Urodela, portions which are not homologous
with the OS coccygeum. The term urostyle should be restricted
to such and similar modifications of the tail-end, and this latter
liappens to be lost by the Anura during metamorphosis.

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24 ANURA CHAP.

Strictly speaking, or rather in anatomical parlance, the
Vertebrate tail begins with the first post-sacral vertebra. In
the Anura that portion of the whole tail has retained most
cartilage, and has become the coccygeum, which is required as
a " backbone " for the often enormous belly. This require-
ment is an outcome of the great shortening of the trunk proper
(if the trunk be defined as ending with the pelvic region), and this
shortening of the trunk is again intimately connected with the
jumping mechanism, enlargement of the hind-limbs, elongatiou
of the ilia, and throwing the fulcral attachment forwards as
much as possible. The pre-acetabular ilio-sacral connection is
carried to the extreme in the Anura.

The shoulder-girdle and " sternum " are more complete than
in the Urodela, there being also a pair of clavicles, fused with
the precoracoidal bars. The whole apparatus presents two types.
In the arciferous type tlie coracoids and precoracoids retain
a great amount of cartilage in their distal portions, and these
cartilages (the epicoracoids of some authors) overlap each other
movably on one another, the right usually lying ventrally upon
the left. The epicoracoidal cartilage of each side, by connecting
the distal end of the coracoid with the precoracoid of the same side,
forms an arc, hence "arciferous." In the firmisternal type
the epicoracoidal cartilages are much reduced, and, instead of
overlapping, meet in the middle line and often fuse with each
other, forming thereby a firm median bar, which connects tlie
ventral ends of the precoracoids with those of the coracoids.
This type is morphologically the higher and more recent, and
passes in the larval stage through the arciferous condition.
It is restricted to the Eanidae, Engystomatinae, and Aglossji.
Although these two types afford an excellent distinctive char-
acter for the main divisions of the Anura, they are to a certain
extent connected by intermediate forms in such a way, that, for
instance, in Bufo and among Cystignathidae in Ceratophrys, the
two opposite epicoracoidal cartilages begin to unite at the
anterior end.

In many Engystomatinae the precoracoids together with the
clavicles are much reduced, sometimes to thin ligaments, beint*-
in this case mostly curved back and lying closely against the
coracoids ; or they may be lost completely. Very rarely the
precoracoidal bars are actually much stronger than the coracoids.

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SHOULDER-GIRDLE

and the median symphysial bar of cartilage is lost ; this is the
case in ffemistts.

The scapula is always large and curved into transverse,
dorsally broadening blades, the dorsal greater portion of which,
the so-called supra-scapula, does not ossify but calcifies.

It is very doubtful if the Anura possess a true sternum, if
by sternum we understand a medio -ventral apparatus which
owes its origin to the ventral portions of ribs. The so-called

Fto. 5. — Ventral views of the shoulder-girdles of various Anura. (Slightly enlarged.) 1,
Hombinalor igneus, and 2, Bu/o vulgaris, as examples of the arciferous type ; 3,
adnlt, 4, metamorphosing Rana temporarioy showing change from the arciferous
into the firmistemal type ; 5, Hemisus guttatum ; 6, Breviceps gibbosus ; 7, Cauxpus
systoma, {5, 6, 7, after Boulenger. ) Cartilaginous parts are dotted ; ossified part«
are left white. O?, Clavicle ; Co, coracoid ; JS, epicoracoidal cartilage ; //, humerus ;
J/, metastemum ; 0, omostemum ; P, precoracoid ; Sc, scapula ; S.S, supra-
scapula.

sternal apparatus of the Anura consists of two pieces. One,
anterior, variously named episternum, presternum, or omosternum,
rests upon the united precoracoida and extends headwards, being
either styliform or broadened out. Sometimes it is partly ossified,
with a distinct suture at its base ; this is the case especially in
the FirmiBtemia ; in many Arcifera the omosternum remains
cartilaginous and is continuous, without a sutural break, with
the cartilage of the precoracoids, indicating thereby its genetic
relation to the shoulder -girdle. Hence omosternum is the

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26 ANURA

preferable name. It is frequently much reduced, even absent,
for instance in most Bufonidae and in the Engystomatinae.
The posterior so-called sternal part may be termed metasfernum.
It forms the posterior counterpart of the omosternum. It
is attached behind to the epicoracoidal cartilages, or fusing
with them forms their posterior continuation. It appears
mostly in the shape of a style, which is frequently ossified, and
broadens out behind into a cartilaginous, partly calcified blade.
In the Discoglossidae only it diverges backwards into two horns,
assuming a striking resemblance to the typical xiphisternum of
the Amniota. In yoimg Anura the metasternal cartilage is
intimately connected with the i)ericardium, an indication of its
being derived not from ribs but from the shoulder-girdle.

The glenoid cavity is always formed by the coraooids and by
the scapula, but the precoracoid often takes part in its forma-
tion, for instance in Bufonidae, Hylidae, and Discoglossidae.

In the fore-limb the humerus has a crest, stronger in the males
than in the females ; it assumes extraordinary strength in some
Cystignathidae, notably in the male Leptodactyhis. Eadius and
ulna are fused into one bone. The carpalia are originally nine
in number : radiale, ulnare, two centralia, and five carpalia distalia,
tlie fifth of which is reduced to a tiny nodule or to a ligamentous
vestige. The primitive condition still prevails in the Disco-
glossidae. In most of the other Anura the fourth and third
distal carpalia, in any case very small, fuse with the enlarged
ulnar centrale ; the radial centrale comes, in the Bufonidae and
Pelobatidae, into contact with the radius, so that the forearm
articulates with three elements as in the Urodela, but with this
difference, that the intermedium of the Urodela has been lost by
the Anura. There are five metacarpalia and five fingers, but
the elements of the first or thumb are nearly vestigial, so that
the poUux is reduced to one or two nodules, scarcely visible
externally. The normal number of the phalanges of the second
to fifth fingers is 2, 2, 3, 3. The distal phalanges are generally
straight, either pointed or expanded or with Y or T-shaped ends ;
hut in the Hylidae, in Hylamhatefi amongst the Eanidae, and in
Ceratohyla, one of the Hemiphractinae, the terminal phalanges
are produced into curved claws which support the adhesive
finger-discs. There are, however, many genera of different
families, which possess finger-discs and have no claw-shaped

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PELVIC GIRDLE 2/

phalanges. The Hylidae, and many of the climbing members of
the Banidae with adhesive discs, possess an extra skeletal piece
intercalated between the last and last but one phalanges of the
fingers and toes. This piece, a mere interarticular cartilage in
Hyla, is in the following Baninae developed into an additional
phalanx, so that their numbera are 3, 3, 4, 4 in the hand and
3, 3, 4, 5, 4 in the foot : GassiTta, Hylnmhates, Rappia, Mega-
lixalus, RharophoTus, Chiromantis, Ixalus, and Nyctixaliis, All
the other Eanidae are without this additional phalanx, irrespec-
tive of the presence or absence or size of digital expansions.^

The pelvic girdle looks like a pair of tongs (see Fig. 4,
1>. 22). The ilium is enormously elongated and is movably
attached to the sacral diapophyses. This connection is always
pre-acetabular in position. The ilium and ischium co-ossify com-
pletely, and make up nearly the whole of the pelvis ; the pubis
is very small, and remains cartilaginous unless it calcifies. It
rarely possesses a centre of ossification, for instance in Felohates,
where the osseous nodule is excluded from the acetabulum,
recalling certain Labyrinthodonta, whose ossa pubis likewise
do not reach that cavity. The latter is open or perforated in
young Anura and remains so in the Discoglossidae, but in the
others it becomes closed up as in the Urodela. The ventral
halves of the pelvis, besides forming a symphysis, closely approacli
e.ach other, just leaving room for the passage of the rectum and
the urino-genital ducts.

The hind-limbs are in all cases longer than the fore-limbs.
The femur is slender, the tibia and fibula are fused into one bone.
The tarsus is much modified by the great elongation of the two
proximal tarsalia (there being no intermedium) into an astragalus
and a calcaneum, both of which* fuse together distally and
proximally, or completely as in Pelodytes ; in the latter case the
limb assumes a unique appearance, since it consists of three
successive and apparently single bars of nearly equal lengtli.
The other tarsal elements, especially the luore lateral ones, are
practically reduced to pads. The Anura have thereby acquired
two well-marked joints, one cruro - tarsal, the other tarso-
metatarsal; this shows a high stage of specialisation in coni-
])arison with the TJrodelous and Stegocephalous type of still
xmdefined joints.

^ Boulenger, P.Z.S. 1888, p 201.

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2 8 ANURA CHAP.

The Anura possesses five well-developed toes with normally
2, 2, 3, 4, and 3 phalanges, and the rudiments of a sixth digit,
the so-called prehallux, which consists of from two to four
pieces, including the one which represents its metatarsal. This
prehallux, as a vestige of a once better developed digit, is
exactly like the elements on the radial side of the wrist, which,
we are certain, are the remnants of a once complete finger, namely
the poUex. The only weighty diflBculty against its interpreta-
tion as a prehallux lies in the fact that hitherto no six-toed
Stegocephali have been found ; but the fact that there are no
Stegocephali known witli more than four fingers coidd be used as
an argument against there being a poUex- vestige in recent
Anura with just as little reason.

The skull of the Anura differs from that of the other recent
Amphibia in the following features : —

The orliital region of the primitive cranium remains carti-
laginous, but further forward the cranial cavity is closed by the
unpaired si)henethmoid, which forms a ring round the anterior
portion of the brain -cavity, hence called " os en ceinture " by
some anatomists. The frontals and parietals fuse into one pair
of fronto-parietal bones, and these again can fuse together in the
middle line ; as in Aglossa and relohates. The palatal portion of
the palato-quadrate cartilage is complete, reaching forwards to the
sides of the ethmoid region. The curved arch, formed by this
cartilage, is covered by the following bones: (1) the quadrato-
jugal, reduced to a thin splint which connects the quadrate and
squamosal with the posterior end of the maxilla ; (2) the ptery-
goid, always strong, extending from the distal inner corner of
the quadrate to the maxilla, sometimes also to the palatine, and
with a broad, median process to the parasphenoid, this process
covering ventrally most of the otic region ; (3) the palatinen,
which vary considerably in shape and size ; they are placed
transversely and meet in the middle line; in Bomhinator and
Pelodytes they are absent.

The quadrates are directed transversely and backwards, in
conformity with the wide gape of the mouth. Tlie squamosal is
always well developed, covering the wliole of the quadrate on its
outer side ; it lias a forwardly directed process which ends freely
in RanUy meets a corresponding process of the maxilla and forms
a bony arch with it in Discoglossus, Pf^lohatni, and others, or

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II SKULL 29

is scarcely developed at all, for instance in Bvfo. In Felolatea
cidtripes the squamosal is very wide and forms a junction with
the fronto-parietals, thus producing a broad bridge across the
temporal fossa.

The nasal bones are large and meet in the middle line.
Frequently they leave a space between them and the diverging
anterior portion of the fronto-parietals, through which gap
appears part of the dorsal surface of the ethmoid cartilage.
A fontanelle between the frontals occurs in most Hylidae, many
(•ystignathidae, some few Bufonidae, in PeJodytes amongst the
Pelobatidae, and in the Discoglossidae.

The tympanic cavity is bordered in front, above, and below
by the squamosal and quadrate, behind by the musculus depressor
mandibulae, internally by the otic capsule, and by the cartilage
of the cranium between this and the lateral occipital bone.
The cavity communicates, however, by the wide and short
Eustachian tube with the mouth, the passage being bordered
anteriorly by the pterygoid, posteriorly by soft parts. Partly
imbedded in these soft tissues is the styloid process or stylohyal,
which is attached to the cranium, mostly behind the otic region,
and is continued downwards into the anterior horn of the hyoid.
The whole partly cartilaginous, ligamentous, and osseous string
is, in fact, the entire ventral half of the hyoid arch, while the
dorsal half or hyomandibular portion of tliis, the second visceral
arch, is modified into the coliunellar or auditory chain. The
inner end of this chain, the stapes, is inserted into and around
the fenestra ovalis of the otic capsule, while the outer end is
somewhat T-shaped, and is loosely attached to or near the upper
rim of the tympanic ring and to the middle of the tympanic
disc. In many Anura this terminal bar can be seen from the
outside. The middle portion of the columellar chain is ossified,
the rest remains cartilaginous. But the whole chain exhibits
various modifications in different genera, especially in the
number and the extent of the processes sent out by the outer
cartilaginous portion ; these are attached in various ways to
the tympanum and its rims. The tympanic disc is carried
by a cartilaginous ring, which rests against a special process
sent out by the quadrate, and is probably itself a differentiation
of this element.

In some very aquatic genera the whole tympanic cavity is

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30 ANURA CHAP.

much reduced, for instance in Felobates, Bomiinator, Liopdma.
In Batrachophrynus not only the cavity, but also the Eustachian
tubes are suppressed. In the Aglossa only the two tubes an*
united into one short but wide median canal, opening at the level
of the pterygoids on the roof of the mouth.

The lower jaw is remarkable for the possession of mento-
Meckelian cartilages, absent only in the Aglossa and Diaco-
glossidae. At first they are much longer than the rest of the
jaw ; during the larval life they indeed form the functional
jaw, and they are now covered with homy sheatlis instead of
teeth. Owing to the absence of teeth on them, these mento-
Meckelian cartilages are later not invested by bone, although in
many Anura they ultimately ossify, either retaining their sepa-
rate nature or fusing partly with the dentary bones. The bulk
of the lower jaw, the Meckelian cartilage, becomes invested by
the dentary, a small articulare, and an inner angulare, while a
splenial element is absent. The dentary itself is mostly reduced
to a small dentigerous splint, while the angulare forms by far
the greater part of the bony jaw.

Teeth are more restricted in their occurrence than in the
Urodela. On the jaws they always stand in one row. With
the exception of the Hemiphractinae, Amphignathodontinae,
Ceratobatrachinae, and Genyophryninae, no recent Anura carry
teeth on the lower jaw, and even in these genera they are mostly
much reduced in size and firmness, having all the appearance of
vanishing structures. The premaxillae and maxillae are frequently
furnished with teeth, except in the Dendrobatinae, Genyophry-
ninae, Engystomatinae, Dendrophryniscinae, Bufonidae, Pipa^ and
Hj/menochii'f/s. The vomers mostly carry a series of teeth on
their posterior border ; when these teeth are absent, as in many
species of Bufo, a kind of substitute sometimes occurs on the
palatines in the shape of a row of tuberosities. The palatines
carry teeth in Hemiphractinae. The parasphenoids are toothed
in Triprioji and Biaglena, and occasionally in Pelohates cvltripes.

A few Anura possess peculiar substitutes for teeth in tlie
anterior portion of the lower jaw, namely, a pair of conical bonv
processes, sometimes rather long, but always covered by the
dense gums, or investment of the jaws ; eg. LepidohcUrach%ts,
several Bana, ejj. B. (irfsperaft, B. Ihasiamt^ B. knhli, and Crypiotiji
brer is.

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II SKIN 3 1

Cranial dermal ossifications are developed in some species of
BufOy still more in the Hemiphractinae, and above all in Peh)-
hates cidtripes and in the Cystignathoid genus Cnh/ptocephalns.

The hyoid apparatus of the Anura is complicated. It is
originally composed of the hyoidean and four branchial arches,
with one median, copular piece. The branchial arches form in
the early life of the tadpole the elaborate framework of the
filtering apparatus mentioned on p. 44. During metamorphosis
the whole filter disappears, owing to resorption of the greater
part of the branchial arches ; only their median portions remain,
and fuse with the enlarged copular piece and the hyoidean
arches into a broad shield -shaped cartilage {corpus linguae),
whence several lateral processes sprout out, the posterior pair of
which are generally called thyrohyals or thyroid horns. The
true hyoid horns give up their larval lean-to articulation with
the quadrate, become greatly elongated, and gain a new attach-
ment on the otic region of the cranium. The transformation of
the whole apparatus has been studied minutely by Kidewood, in
Pelodytes punctatus}

Skin

The epidermis of the young larvae of Amphibia is furnished
with cUia, which later on are suppressed by the development of
a thin hyaline layer or cuticula, but clusters of such cilia
remain, at least during the larval life and during the periodical
aquatic life of the adult, in the epidermal sense-organs. In the
frog, currents are set up by the ciliary action at an earlier stage,
and are maintained to a later stage than in the newt. In the
latter the tail loses its ciliation, whereas in the frog it remains
active almost up to the time of the metamorphosis. In tadpoles
of 3-10 mm. nearly the whole surface is ciliated (Assheton).^
The cilia work from head to tail, causing the little animal, when
perfectly quiet, to move forwards slowly in the water. Beneath
the cuticula, in the Perennibranchiata and the larvae of the
other Urodela, lies a somewhat thicker layer of vertically striated
cells, the so-called pseudo-cuticula, which disappears with the
transformation of the upper layers of the Malpighian cells into
the stratum corneum. The latter is very thin, consists of one or
two layers of flattened cells, and is shed periodically by all
' P.Z.S. 1897, p. 577. * Q,j,]if,s. xxxviii. 1896, ].. 465.

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32 AMPHIBIA

Amphibia in one piece. In the Urodela it generally breaks
loose around the mouth, and the animal slips out of the delicate,
transparent, colourless "shirt," which during this process of
ecdysis or moulting becomes inverted. In the Anura it mostly
breaks along the middle line of the back, the creature struggles
out of it, pokes it into its mouth, and swallows it. Urodela
also eat this skin. As a rule the first ecdysis takes place
towards the end of the metamorphosis, preparatory to terrestrial
life. So long as the animal grows rapidly, the skin has to be
shed frequently, since this corneous layer is practically dead and
unyielding. Adult terrestrial Urodela do not seem to moult
often, mostly only when they take to the water in the breeding
season. Anura, on the other hand, moult often on land, at least
every few months. The surface of the new skin is then quite
moist and slimy, but it soon dries and hardens.

The Malpighian stratum consists of several layers, thickest
in the Perennibranchiata ; in them it contains mucous cells
throughout life, in others sucli slime-cells are restricted to larval
life. Later, regular slime -glands are developed, which open
on tlie. surface. They are very numerous, and more evenly
distributed, over most parts of the body, than the specific or
poison-glands, which are restricted to certain parts, often form-
ing large clusters, especially on the sides of the body. They
reach their grciitest development in the " parotoid glands " of
the Anura. Both kinds of glands are furnished with smooth
muscle-fibres, which are said to arise from the basal membrane
underlying and forming part of the Malpighian layer ; these
muscle-cells extend later downwards into the corium. For the
action of the poison, see p. 37.

The stratum corneum is mostly thin, but on many parts of
the body, especially in Anura, the epidermal cells proliferate and
form hard spikes or other rugosities, generally stained dark brown.
With these may be grouped the nuptial excrescences so frequent in
tlie Anura, especially on the rudiment of the thimib, and on the
under surface of the joints of the Sogers and toes. In many
Anura, less frequently in the I^rodela, the tips of the fingers and
toes are encased in thicker horny sheaths, producing claws or
nails. They are best developed among newts in Onychodactylus,
among the Anura in Xenojnis and Hymniochirus. The horny
covering of the metatarsal tubercles reaches its greatest size in

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II SKIN 33

the digging spur or spade of Pelohates, In most of these eases
the cutis is elevated into more or less wart-like papillae, covered,
of course, by the proliferated and cornified epidermis. In the
female of Rana temporaria nearly the whole surface of the body
becomes covered with rosy papillae during the breeding season.
Similar nuptial excrescences are common, and are most note-
worthy in the male of the Indian Bana liebigi.

The epidermis also contains sense-organs. They attain
their highest development in the larvae ; later on they undergo a
retrogressive change. Each of these sense-organs is a little
cup-shaped papilla, visible to the naked eye. It is composed of
elongated cells which form a mantle around some central cells,
each of which ends in a stiff cilium perforating a thin, hyaline
membrane which lines the bottom of the cup, and is perhaps the
representation of the cuticula. These ciliated cells are connected
with sensory fibres, the nerve entering at the bottom of the
whole organ. The cilia are in direct contact with the water,
but the outer rim of the whole apparatus is protected by a short
tube of hyaline cuticula-like secretion. These sense-organs are,
in the larvae, scattered over the head, especially near the mouth
and around the eyes, whence they extend backwards on to the
tail, mostly in three pairs of longitudinal rows, one near the
vertebral column, the others lateral. They are supplied by
the lateral branch of the vagus nerve. They disappear during
the metamorphosis, at least in the Anura, with the exception of
Xeiiopiis, in which they form conspicuous white objects. The
white colour is caused by the tubes becoming choked with
the debris of cells or coagulating mucous matter, so that it is
doubtful if these organs, which moreover have sunk deeper into
the skin, are still functional. In the terrestrial Urodela these
organs undergo a periodical process of retrogression and rejuven-
escence. During the life on land they shrink and withdraw
from the surface, and their nerves likewise diminish, but in the
breeding season, when the newts take again to aquatic life, they
revive, are rebuilt and become prominent on the surface. They
are an inheritance from the fishes, in which such lateral line
organs are universally present.

The cutis of most Amphibia is very rich in lymph -spaces,
which, especially in the Anura, assume enormous proportions,
since the so-called subcutaneous connective tissue forms com-

VOL. VIII I)

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34 AMPHIBIA

paratively few vertical septa by which the upper and denser
layers, the corium proper, are connected with the underlying
muscles. The spaces are filled with lymph, and into some of them
the abnormally expanded vocal sacs extend, notably in Paliidicola,
Leptoddctylm, and other Cystignathidae, and in Rhinoderma.

The cutis frequently forms papillae and prominent folds,
sometimes regular longitudinal keels on the sides of the back ; but
dermal, more or less calcified or ossified scales are restricted to
the Stegocephali and to the Apoda, q,v., pp. 79, 87. We con-
clude that the Urodela and Anura have entirely lost these organs.
Dermal ossifications, besides those which now form an integral
part of the skeleton, like many of the cranial membrane-bones, are
rare, and are restricted to the Anura. They are least infrec^ueut
on the head, where the skin is more or less involved in the
ossification of the underlying membrane-bones, for instance iu
Triprion, Calyptocephalus, Hemiphractus and PeJohates, The
thick ossifications in the skin of the back of several species of
Ceratophrys are very exceptional. In Brachycephalus ephippiina
these dermal bones enter into connection with the vertebrae ;
small plates fuse with the dorsal processes of the first to third
vertebrae, while one large and thick plate fuses with the rest
of the dorsal vertebrae. Simple calcareous deposits in the
cutis are less uncommon, for instance, in old specimens of Bufo
vulgaris. AVe are scarcely justified in looking upon these various
calcifications and even ossifications as reminiscences of Stego-
cephalous conditions.

The skin contains pigment. This is either diffuse or granular.
Diffuse pigment, mostly dark brown or yellow, occurs frequently
in the epidermis, even in the stratum corneum. The granular
pigment is stored up in cells, tlie cliromatophores, which send
out amaiboid processes, and are restricted to the cutis, mostly
to its upper stratum, wliere they make their first appearance.
Contraction of the chromatophores withdraws the pigment
from the surface, expansion distributes it more or less equallv.
The usual colours of the pigment are black, brown, yellow, and
red. Green and blue are merely subjective colours, due to
interference. A peculiar kind of colouring matter is the white
pigment, which probably consists of guanine, and is likewise
deposited within cells ; cf the description of the white spots in
the skin of Hyla voerulea.

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II CHANGES OF COLOUR 35

Most Amphibia are capable of changing colour, the TJrodela,
however, far less than the Anura, some of which exhibit an
extraordinary range and adaptability in their changes.

The mechanism by which the change of colom* is produced in
frogs has been recently studied by Biedermann.^ If we examine
the green skin of the common Tree-frog, Hyla arborectj under a
low power and direct light, we see a mosaic of green, polygonal
areas, separated by dark lines and interrupted by the openings of
the skin -glands. Seen from below the skin appears black.
Under a stronger power the black layer is seen to be composed of
anastomosing and ramified black pigment-cells. Where the light
shines through, the skin appears yellow. The epidermis itself is
quite colourless. The mosaic layer is composed of polygonal
interference-cells, each of which consists of a basal half which is
granular and colourless, while the upper half is made up of yellow
drops. Sometimes the tree-frog appears blackish, and if then
the black pigment-cells are induced to contract, for instance, by
warming the frog, it appears silver-grey ; in this case the pig-
ment in the yellow drops is no longer diffuse, but is concentrated
into a round lump lodged between the interstices of the gi-anular
portions; the black pigment -cells are likewise balled together.
These black chromatophores send out numerous fine branches,
which occasionally stretch between and round the polygonal
cells. When each of these is quite surrounded and covered by
the black processes, the frog appears black. On the other hand,
when the black pigment-cells withdraw their processes, shrink up,
and, so to speak, retire, then the light which passes through the
yellow drops is, by interference, broken into green.

Stoppage of the circulation of the blood in the skin causes
the black chromatophores to contract. Carbon dioxide paralyses
them and causes them to dilate. This is direct influence
without the action of nerves. But stimulation of the central
nerve - centres makes the skin turn pale. Low temperature
causes expansion, high temperature contraction, of the chrom-
atophoras. Hence hibernating frogs are much darker than they
are in the summer. Frogs kept in dry moss, or such as have
escaped into the room and dry up, turn pale, regardless of light
or darkness, probably owing to a central, reflex, nerve-stimulus.

Tree-frogs turn green as the result of the contact with leaves.
^ Arch. (jes. Fkysiol. li. 1892, p. 455.

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36 AMPHIBIA

Dark frogs will turn green when put into an absolutely dark
vessel in which there are leaves. This is reflex action, and
blinded specimens do the same. The principal centres of the
nerves which control the chromatophores, lie in the corpora
bigemina and in the optic thalami of the brain. When these
centres ai*e destroyed, the frog no longer changes colour when
put upon leaves, but if a nerve, for instance the sciatic, be
stimulated, the corresponding portion of the body, in this case
tlie leg, turns green. Kough surfaces cause a sensation which
makes the frog turn dark. Rana seems to depend chiefly upon
temperature and the amount of moisture in the air, so far as its
changes of colour are concerned. Biedermann concludes that the
" chromatic function of frogs in general depends chiefly upon the
sensory impressions received by the skin, while that of fishes
depends upon the eye."

All this sounds very well, but the observations and experi-
ments are such as are usual in physiological laboratories, and the
frogs, when observed in their native haunts, or even when kept
under proper conditions, do not always behave as the physiologist
thinks they should. There is no doubt that in many cases the
changes of colour are not voluntary, but reflex actions. It is
quite conceivable that the sensation of sitting on a rough
surface starts a whole train of processes : roughness means bark,
bark is brown, change into brown ; but one and the same tree-
frog does not always assume the colour of the bark when it
rests, or even sleeps upon, such a piece. He will, if it suits
him, remain grass -green upon a yellow stone, or on a white
window - frame. I purposely describe such conditions, changes,
CO incidences, and discrepancies in various species, notably in
Hyla arhorea, If. coerulea, Rana teinporaria, Bufo vii-idis, to show
that in many cases the creature knows what it is about, and
that tlie eye plays a very important part in the decision of what
colour is to be produced. The sensory impression received
through the skin of tlie belly is the same, no matter if the board
be painted white, black, or green, and how does it then come to
pass that the frog adjusts its colour to a nicety to the general
hue or tone of its surroundings ?

Boulenger ^ has given us a summary of the action of the
poison of Amphibia :

^ Xat. ScL i. 1892, p. 185.

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POISON 37

It is well known to all who have handled freshly -caught
newts, and certain toads, especially Bombinator, that their secre-
tion acts as a sternutatory, and causes irritation of th6 nose and
eyes, the effects produced on us by Bombinator being comparable
to the early stages of a cold in the head. Many collectors of
Batrachians have learned, to their discomfiture, how the intro-
duction of examples of certain species into the bag containing the
sport of their excursion may cause the death of the other
prisoners ; for although the poison has no effect on the skin of
individuals of the same species, different species, however closely
allied, may poison each other by mere contact. But when
inoculated the poison acts even on the same individual.

Miss Ormerod, to personally test the effect, pressed part of the
back and tail of a live Crested Newt between the teeth. " The
first effect was a bitter astringent feeling in the mouth, with
irritation of the upper part of the throat, numbing of the teeth
more immediately holding the animal, and in about a minute
from the first touch of the newt a strong flow of saliva. This
was accompanied by much foam and violent spasmodic action,
approaching convulsions, but entirely confined to the mouth
itself. The experiment was inmiediately followed by headache
lasting for some hours, general discomfort of the system, and half
an hour after by slight shivering fits."

Numerous experiments have shown that the poison of toads,
salamanders, and newts is capable, when injected, of killing
manmials, birds, reptiles, and even fishes, provided, of course, that
the dose be proportionate to the size of the animal Small
birds and lizards succumb as a rule in a few minutes ; guinea-
pigs, rabbits, and dogs in less than £m hour.

This poison of Amphibia is not septic, but acts upon the
heart and the central nervous system. That of the common
toad has been compared, in its effects, to that of Digitalis and
Erythrophlaeum. Some authorities hold that the poison is an
acid, others regard it as an alkaloid.

Phisalix^ has come to the conclusion that toads and sala-
manders are possessed of two kinds of glands, different both
anatomically and physiologically. These are, first the mucous
glands, spread over the greater part of the body, with an alkaloid
secretion, which acts as a narcotic ; secondly, specific glands, as

' C. JR. Ac, SH, cix. 1889, pp. 405, 482.

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38 AMPHIBIA

the parotoids and larger dorsal glands, the secretion of which
is acid, and acts as a convulsive.

The Indians of Colombia are said to employ the secretion of
Dendrohates tiiictomus for poisoning their arrows. The poison is
obtained by exposing the frog to a fire, and after being scraped
off the back is sufficient for poisoning fifty arrows. It acts on
the central nervous system, and is used especially for shooting
monkeys. Concerning the use of this poison for "dyeing"
parrots, see p. 272.

The milky secretion of toads protects them against many
enemies, although not always against the grass-snake. A dog
which has once been induced to bite a toad, suffers so severely
that it will not easily repeat the experiment. The handling of
tree-frogs also irritates both nose and eyes. The hind limbs of
the Water-frog, Rana esculenta, have a very bitter, acrid taste.
In short, most, if not all. Amphibia are more or less poisonous,
and it is significant that many of the most poisonous, e.g.
Sidamandra maculosa, Bomhimitor, Dendrohates, exhibit that very
conspicuous combination of yellow or orange upon a dark ground,
which is so widespread a sign of poison. Other instances of
such warning colours, protective in a defensive sense, are the
Wasps and Helodermay the only poisonous lizard.

Nerves

Spinal nerves. — Each spinal nerve issues originally immedi-
ately behind the neural arch of the vertebral segment to which it
belongs. This intra-vertebral position is ultimately modified into
a more inter- vertebral one, owing to the predominant share of the
neural arches, basidorsalia, in the composition of the whole vertebra.
Consequently the nerves issue behind their corresponding vertebrae.

The first spinal nerve, or N. suboccipitalis, is exceptional in
several respects. It develops a dorsal and a ventral root like
a typical spinal nerve, but the dorsal root soon degenerates in
all Amphibia, while in the Phaneroglo8sal Anura the whole nerve
disappears. Tlie first spinal nerve reduced to its ventral half
persists therefore only in the Apoda, Urodela, and the Aglossal
Anura. It issues originally between the occiput and the atlas,
but in the adult it is partly imbedded in the anterior portion of
the atlas. Its own vertebra is lost, having probably been added
to the cranium.

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NERVES 39

In the Urodela the first spinal nerve either remains separate,
or it joins the second spinal, forming with it and with a branch
from the third nerve the cervical plexus, which supplies the
muscles of the cervical region. The third, fourth, and fifth
nerves, and sometimes also the sixth, form the brachial plexus.

In the Aglossal Anura N. spinalis I. mostly sends a fine
thread to the second spinal nerve, the rest supplies chiefly the
M. levator scapulae, in Piiya the abdominal muscles also. In all
the other Anura this N. spinalis I. is lost ; occasional vestiges
have been reported in Bvfo vulgaris and Rana cateshiana, and
remnants of it may possibly be found in Pelobatidae and Discoglos-
sidae. The first actually persisting nerve of the Phaneroglossa is
conse(j[uently N. spinalis II.

The brachial plexus is composed as follows: — Fipa, N.
spinalis II. and III.; Jienojyus and Phaneroglossa, K spinalis
III. and IV., with a small branch from the second ; the next
following three nerves, numbers V., YL, and VII., behave like
ordinary trunk nerves.

The pelvic plexus of the Phaneroglossa is formed in Bana by
the VIII. + IX. + X. + Xlth nerves, the tenth issuing between
the sacral vertebra and the coccyx. In Bufo and Ifyla the
plexus is composed of five nerves, the seventh spinal sending
a branch to it. Occasionally the twelfth nerve contributes a
small branch to the posterior portion of the plexus. This and
the eleventh nerve leave the coccyx by separate holes, thereby
indicating its composition. The rest of the spinal cord gives oft'
no more recognisable nerves, owing to its reduction during the
later stages of metamorphosis ; its terminal filament passes out
of the posterior end of the coccygeal canal.

Concerning the cranial nerves it is necessary to draw atten-
tion to one point only. The last nerve which leaves the
cranium of the Amphibia is the vagus or tenth cranial nerve.
There is consequently no eleventh, and no twelfth or hypoglossal,
pair of cranial nerves. Their homologues would be the first and
second spinal nerves, but the whole tongue of the Amphibia,
with its muscles, is supplied by the glossopharyngeal, or ninth
cranial pair, and is morphologically not homologous with the
tongue of the Amniota.

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40 AMPHIBIA CHAP.

Respiratory Organs

A very important and characteristic feature of the Amphibia
is the development of two sets of respiratory organs : Gills and
Lungs. It is as well to give definitions of these organs. Lutujs
are hollow evaginations from the ventral wall of the pharynx,
and their thin, vascularised walls enable the blood to exchange,
by osmosis, carbon dioxide for oxygen from the air which enters
the lungs by the mouth or the nostrils, and the windpipe. The
latter is unpaired, the lungs themselves are paired. Gills ai*e
highly vascularised, more or less ramified excrescences, covered
by a thin epithelium of ecto- or endo-dermal origin, which permits
of the exchange of carbon dioxide for oxygen from the air which is
suspended in the surrounding water. It is obvious that this
definition applies to all sorts of well-vascularised organs whose
thin surface comes into contact with the water. Various recesses
of the pharyngeal cavity, the dorsal and ventral folds of the
tail-fin, nay, even any part of the skin of the body can, and does
occasionally, assume additional respiratory functions. The proper
definition of gills, in Vertebrates, requires, therefore, the restric-
tion that they must be developed upon and carried by visceral
arches.

The general statement that the Amphibia breathe by lungs,
and, at least during some stage of their life, also by gills, requires
various restrictions. As a rule the majority of Amphibia first
develop gills, later on also lungs, whereupon, during the meta-
morphosis, the gills are gradually suppressed, so that the perfect
animal breathes ])y lungs only (see p. 61). But a number of
Urodela retain tlieir gills throughout life, although the lungs are
also functional. These are the Perennibranchiata, not a natural
group, but a lieterogeuous assembly, Proteidae and Sirenidae.
Some species of Amhly stoma remain individually Perenni-
branchiate (cf Axolotl, p. 112). On the other hand, in some
Anura the gills are almost or entirely suppressed, or restricte<l
to the embryonic jx^riod only. Lastly, a considerable numbt^r of
Salamandridae liave lost their lungs; they breathe by gills until
their metamorphosis, but have in the adult state to resort to
respiration by the skin (cf. p. 46).

The general plan of the development of the branchial re-
spiratory apparatus is as follows: — The six visceral arches.

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II GILLS 4 1

namely, the mandibular, the hyoidean, and the four branchial
arches, correspond, long before they are cartilaginous, with four
main arterial arches of the trimcus arteriosus. The first, the
arteria hyo-mandibularis, belongs to the hyoidean and mandibular
segments, the second to the first branchial, the third to the second
branchial, while the fourth soon splits in two for the third and
fourth or last branchial arch. On the dorsal side these branchial
arterial arches combine to form the radix of the dorsal aorta.
These arches, especially the three branchials, appear in newts,
less clearly in frogs, as transverse ridges on the sides of the
future neck. Between the arches the pharynx gradually bulges
out in the shape of five lateral gill-pouches; the first between
the mandibular and the hyoidean arch, the second between the
hyoidean and the first branchial arch, etc. These pouches soon
break through to the outside and become gill-clefts, except the
first pouch in l^rodela. Before the breaking through of the
clefts there appeare upon the outside of the middle of the rim of
each arch a little knob, which soon ramifies and forms an external
gill. The knob owes its origin to the development of a blood-
vessel which buds from the arterial arch, ramifies and breaks up
into capillaries, and returns a little further dorsalwards into the
arch. A secondary loop to the outside of the primary arterial
arch is thus fonned ; and whilst this outer loop sprouts out
further, driving before it the likewise proliferating skin, and thus
producing the gill, the middle portion of the primary arch
remains in the Urodela as a short cut, but in the Anura it
partly obliterates, and henceforth acts as the internal efferent
vessel of the gilL When, during metamorphosis, the gills dis-
appear, their intrinsic afferent and efferent vessels vanish likewise,
and the short cut completes the circuit. In order to do this
they have, in the Anura, to form new connections with the trunks
of the afferent vessels.

The arterial arches themselves are modified as follows : — The
first pair become the carotids, the second form the right and left
aortic arches, while the third and fourth unite and are trans-
formed into the pulmonary arteries and " ductus Botalli," the last
arterial arch having previously sent a branch into the developing
lungs. In the Anura the third arch obliterates.

The gills and clefts present various modifications. The
Urodela possess three pairs of j^jills, one eacli upon the dorsjil

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42 AMPHIBIA CHAP.

half of the three branchial arches, just near the upper corners of
the clefts ; and the skin of the body is continued upon the stem
of each gill, pigmented like the rest of the surface of the body.
Such a gill is more or less like a blade, standing vertically, and
is composed of a stem of connective tissue, thick at the base, and,
as a rule, carrying two series of fine lamellae, which, however, do
not form two opposite series, but hang downwards, being, so to
speak, folded down, so that the upper surface of the stem is bare,
and carries the lamellae on its under side. In the Axolotl some
of these lamellae are further subdivided. In Xectwriis they are
enormously increased in numbers, but are rather short, and they
stand no longer in two rows, but are crowded into one. Those
of Proteus form two rows of dendritic filaments : those of Siren
are likewise much ramified.

The larvae of the Urodela have four clefts. In the adult
Siren these are reduced to three, the first, namely, that between
the hyoid and the first branchial arch, being closed up. In
Nee turns y Proteus, and Typhlomolge the clefts are further reduced
to two, owing to the closing up of the first and last, only those
between the first, second, and third arches remaining. Arnphiuma,
and usually Cryjptohraiuhus alleghaniensis, possess only one pair
of clefts, while in C. japonicvs and in the Salamandridae all the
clefts are abolished.

The gills of the Urodela are always uncovered, although a
short operculum is formed from the posterior margin of the
hyoidean arch ; the halves of this fold meet below the throat,
and persist in various terrestrial and aquatic species as the " gular
fold." It reaches its greatest size just before metamorphosis, but
scarcely ever produces a proper outer gill-chaml)er, and it does
not cover the gills owing to their rather pronounced dorsal
position. It is perhaps best developed in Typhlomolge, and even
there its dorsal portion is continued upon the first of the three
broad vertical and short-fringed blades which form the gills.

A d('Scription of the gills of the Apoda will be found in the
systematic part.

In the Auura the gills are complicated, owing to the develop-
ment of the so-called internal gills. Fii^st appear, exactly in the
same way as in the Urodela, the external gills, one upon each of
the first three branchial arches. In the larva of Bana escideiitay
5 mm. in length, a little protuberance appears upon the first.

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II GILLS 43

and then upon the second arch. In the 6 mm. larva the first
gill shows four knobs, the second two, the third one knob. They
are always- delicate and thin, although sometimes pigmented, long,
and much -ramified structures. The first pair is always the
largest; well developed and persisting a long time in Sana
temporaria ; smaller in B, esculenta and Biifo vulgaris ; very
short, scarcely forked, in B. viridis and Hyla arborea. They are
relatively largest in Alytes, while still in the egg. Numerous
descriptions of these gills will be found in the systematic part.

CJreat changes take place about the time when the fourth or
last branchial arch and the pulmonary arteries are developed.
This occurs in B. esculenta when the larva is about 9 mm. long.
The sprouting of the gills extends gradually downwards along the
arches upon their ventral halves, and these new gill-filaments or
loops transform themselves into numerous dendritic bundles,
resting in several thickset rows upon the hinder margin of the
first to the third arch, one row only on the fourth arch, which
carries no external gill. These "internal gills" look like red
l>olster8 or thick and short-tasselled bimches. Whilst they are
developing the dorsal, older gills become arrested in their growth
and disappear, and at the same time a right and left opercular
fold grows out from the head and covers these new gills, shutting
them up in an outer branchial chamber, just like that of Teleostei
and other Tectobranch fishes. This is the reason w^hy these new-
gills have been called internal, aud the mistaken notion has
sprung up that they are comparable with the true internal gills
of fishes. In reality Amphibia have only external gills. They
are always covered by ectoderm, are restricted to the outside of the
branchial arches, and are developed before the formation of the
clefts. These gills are in many cases directly continuous with
the more dorsally and more superficially placed earlier external
gills; but although nearly every one who has studied their
development has observed this agreement, the old error still
prevaila They are morphologically as little internal as the true
internal gills of Elasmobranch embryos are external gills, because
these have become so elongated that they protrude out of the
gill-clefts.

The fact that the Amphibia possess only external gills throws
important light upon their phylogeny. Xot only do the Apoda,
Urodela, and Anura agree much more with each other than

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44 AMPHIBIA CHAP.

would be the case if the Anura possessed both internal and
external gills, but the Amphibia reveal themselves also in this
point as connected with the Crossopterygii and the Dipnoi, some
of which fishes also possess external gills. It is of course quite
possible that tlie Amphibia have developed these organs in-
dependently, but we understand now that the latter are accessory,
and not the primitive respiratory organs; they are developed
in adaptation to embryonic conditions and to prolonged larval,
occasionally perennibranchiate, aquatic life (cf. the chapter on
Neoteny, p. 63).

There is no valid reason for supposing that the Stegocephali
had true internal gills. We know their branchial skeleton, and
we can discern even gill-rakers on the arches. Such gill-rakers
occur also, although but feebly developed, in Urodela. The
whole branchial framework of the Urodela and Apoda undergoes
simple reductions during metamorphosis (see p. 86), but in the
Anura these arches are in early tadpole life transformed into a
most complicated basket-work which acts as a straining apparatus
or filter, to prevent any particle of food or other foreign matter
from finding its way into the delicate gills, the current of water
passing from the mouth through the filter, past the gills and out
of the clefts. During metamorphosis this whole elal)orate
apparatus is again transformed, almost beyond recognition, into
the hyoidean apparatus for the sux:)port of the generally very
movable and much-specialised tongue. The fact that the hyoid
apparatus of the Aglossa, especially that of Xenopus, is con-
structed upon the same lines, is a strong indication that these
creatures have arrived at their tongueless condition through the
loss of this organ, and this is intelligible in correlation with
their absolutely aquatic life.

The opercular folds assume great dimensions in all tadpoles.
They cover the whole gill-region, thereby producing on either
side an outer gill-chamber. The posterior margins of the folds
gradually l)ecome continuous with the rest of the surface of the
body. Each gill -chamber o]jens at first by one lateral canal,
usually called the spiracle. This condition prevails in the tadpolas
of the Aglossa. In the Discoglossidae the two canals gradually
converge and combine into one median opening on the middle of
the belly. In all the other Anura the right opening becomes
closed, or rather its canal passes over to and joins that of the

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II GILLS 45

left side, both opening by one short tube laterally on the left
side, at a variable distance between the eye and the vent. Hence
the elegant terms of Amphi-, Medio-, and Laevo-gyrinidae {yvplvo^
being the Greek for tadpole).

The external gills lead to a further consideration. Protopterus
possesses a vestigial external gill on the shoulder-girdle. Lepi-
dosiren has them on the gill-arches, besides true piscine internal
gills, and Polypterus has a large biserially fringed external gill (in
some cases not disappearing until the fish is adult), which starts
from the mandibular arch, at the level of the spiracle or first
visceral cleft, and overlaps the operculum externally. The axis
of this peculiar organ is possibly based upon the homologues of
the spiracular cartilages, which themselves are the branchiostegal
rays of the dorsal half of the quadrato-mandibular arch. The
branchiostegal rays of the hyoideiin arch, at least their material,
have given rise to the elaborate opercular apparatus ; and, in con-
formity herewith, the hyomandibular itself is not known to carry
a gilL Quite possibly the large external gill of Polypterus is not
serially homologous with other external gills — it may not be a
true gill at all, it has perhaps quite a different function — but it
seems to throw light upon a mysterious pair of organs which are
common in larval and young Urodela, in the larval Aglossa and
in the Apoda. These are the " balancers."

In Triton tasniatus, before hatching, there appears a little
protuberance behind and below the eye ; it rests upon the angle
of the mandibular arch, and is separated from the first trans-
verse, externally visible ridge of the first branchial arch by the
beginnings of the hyoidean arch. A few days later the arteria
hyomandibularis sends a vessel into this knob, forms a vascular
coil, and leaves it as a vein which, instead of returning into the
arterial arch, passes into the veins of the body. Its epithelium
is not covered with flat, but with cubical cells ; and sensory cells
have not been found in it. These organs attain some size, and are
shaped like rods, with thickened ends ; they are movable, and are
used by the larvae as " balancers," keeping the head from sinking
into the slime at the bottom. But they may have other functions
besides, and it is not unlikely that they develop into sensory
organs like feelers. They occur in many Salamandridae, and are
not reduced until, or even after, the metamorphosis, and during this
time they shift their place with relation to the eye and the mouth.

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46 AMPHIBIA

The same kind of organs occur in Amhlystoma} They appear,
previous to the breaking open of the gill-clefts, as protrusions of
epiblast, long before any of the external gills on the branchial
arches. When the clefts have broken open, the quadrate sends
out laterally a tiny crescent-shaped process a little above the jaw-
joint, and this process extends to the base of the balancer, but
not into it, and a bundle of muscle-cells grows into the l)alancer.
It is easy to recognise the same organ in the extremely long
thread-like structures of the larva of Xempus. In the Apoda
they are likewise present, but are retained permanently as highly
specialised, probably tentticular organs (cf. p. 86, Apoda).

One of the most unexpected features is the suppression of
the lungs in various kinds of Salamandridae. The lungs are
either reduced to useless vestiges or they are quite absent. This
occurs in aquatic and terrestrial, American and European forms,
and it is noteworthy that the reduction of the lungs does not
apply to all the species of the various genera, nor is it restricted
to one sub-family.

The following list is due to the researches of H. H. Wilder,-
L. Camerano,^ E. Lonnberg,* and G. S. Hopkins ^ : — All the
Desmognathinae and Plethodontinae ; Ambly stoma tinae, Amhly-
stoma 02)acum; Salamandrinae, Salamandrina perspiciUata. In
Tritofi and other Salamandrinae the length of the lungs varies;
in some they extend more, in others less, than half way down the
distance between head and pelvis. Hopkins remarks : " Two
questions are naturally suggested by this apparently aberrant
condition of the respiratory organs. First, what structures or
organs have taken on the function of the lungs and branchiae ;
and secondly, is there any modification in the form or structure
of the heart which in any way may l)e correlated with the above-
mentioned peculiarities of the luugless forms ? " Wilder con-
cluded that respiration was probably carried on l)y the skin, and
perhaps, to some extent, by the mucosa of the intestine. Camerano
thinks that, at least in the European forms, respiration is effected
by the bucco- pharyngeal cavity, and that the skin affords no
efficient aid. The left auricle in the luugless forms is much

* Orr, Quarf. J. Micr. Set. xxix. 1889, p. 316.

'^ "Lungenlose Salamandriden," Jnat. Anz. 1894, ji. 676 ; 1896, ji. 182.
^ " Nuove ricerche anatomo-fisiologiche iutonio ai Salamandridi nonnaluieute
aimeunioni." Torino, 1894.

•» ZooL Anz. 1896, \\. W-i ; 1899, p. 545. ^ Amcr. Xatural, xxx. 1886, p. 829.

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LUNGS — VOICE

smaller in comparison than the right, and there is no pulmonary
vein. The auricular septum has a large aperture, the eonuuuni-
cation between the auricles being larger than even in Nvcturus
(which breathes essentially by gills). The sinus venosus, instead
of opening into the right auricle only, opens more freely into
the left than into the right, and the latter communicates more
directly with the ventricle than the left, instead of about equally.
In short, the heart of these creatures appears almost bilocular,
instead of being trilocular, at least functionally.

The lungs of the Urodela are always simple, extremely thin-
walled bags. They are highly developed in the Anura, the walls
being modified into numerous air-cells, whereby tlie respiratory
surface is considerably inci-eased. The lungs are filled with air
by the pumping motion of the throat while the mouth is closed,
the nostrils being provided with muscular valves. A muscular
apparatus assists the filling of the lungs in the Anura.^

Most, if not all, Anura and some Urodela have a voice pro-
duced by the larynx, which, especially in the Anura, is provided
with a complicated cartilaginous and muscular apparatus and
with vocal cords. The voice
of the Urodela is at the best
a feeble squeak. The females
of the Anura are either mute
or they produce a mere grunt,
but that of many males is
very loud, and^ moreover, in
many species it is intensified
by vocal sacs which act as
resonators. These sacs are A B

diverticula of the lining of j,,q e.-Intemal view of the moutli of A,

the mouth-cavity, and bulge liana esaUenta, B, Bufo calamita (cf. Fig. 52,

. . , , 1 . J ^1 p. 269). Ch, Choaiia, or inner nasal oneninK ;

out the outer skm and the ^; opening of the Eustachian tube ; S, slit

muscles, chiefly the mylo- leading into the vocal sac ; T, tongue ; To,

, ., i. .1 .1 . rni patches of teeth on the vomers.

hyoid, of the throat. The

nostrils and the mouth are firmly closed during the croaking.
"The sacs are called internal when they are covered by tlie
unmodified gular integument, however much this may be dis-
tended ; external when their membrane projects through slits at

* For the mechanism of the frog's respiration, see Gaupp, Arch. Anaf. 1896,
p. 239.

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48 AMPHIBIA

the sides of the throat, as in Rana esculenta (Fig. 52, p. 269),
or when the skin is thinned and converted into a bladder-like
pouch, as in Hyla arharea." ^ These sacs exhibit many modifica-
tions. They may be impaired and median, and open by two slits
into the mouth, on either side below the tongue ; in Bufo one of
the slits or openings, either the right or the left, is obliterated.
They may be paired and symmetrical, and open one on each
side of the head, below and near the posterior angle of the jaws.
These modifications differ in closely allied species. They reach
their greatest complication in Rhinodervia and in some of the
Cystignathidae by extending far back beneath the skin into the
wide lymphatic spaces. In Rhinoderma they are put to the
unique use of nurseries for the young (see p. 228). Leptoddctyivs
typhoniits has a very distinct pair of outer vocal sacs and a
well-marked unpaired sac which extends into the belly and com-
municates with each outer sac. Several species of Faludicola,
e.g. P.fuscomacidata and F. signifera, have a similar arrangement,
in addition to an unpaired gular sac which can be inflated
independently of the rest (see Fig. 45, p. 220).

Urino-Genital Organs

The kidneys and the male generative glands are still inti-
mately connected with each other. The general plan is as
follows : —

The kidneys consist of a large number of glomeruli, produced
by the coiled segmental tubes, each of whicli is composed of a
nephrostome or funnel opening into the body-cavity, a Aial-
pighian body and an efferent canal. The latter combine to form
the segmental duct which opens into the cloaca. The testes,
composed of a large number of sperm-producing glands, are
drained by transverse canals which combine into a longitudinal
canal, and this again sends off numerous efferent canals which
open into the efferent canals of the kidney, so that the segmental
duct (Leydig's duct of many authors) conveys both sperma and
urine.

In the female the network of transverse and longitudinal
canals, which originally connect the generative glands with the
kidney's efferent canals, is reduced in so far as the connection is

1 Boiilenger, The Tailless Bat rack inns of Evroiu. Ray Soc, 1896.

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URINO-GENITAL ORGANS

Fig. 7. — Diagrammatic representation of modifications of the urino - genital ducts.
1, 2, Male and female Newt ; 3, a tubule of the kidney ; 4, male Ra)ia ; 5,
male Bufo; 6, male Bombinatm'; 7, male Discoglossus ; 8, male Alytes. a, Artery
entering, and producing a coil in, the Malpighiau body, M ; B^ Bidder's organ ;
e/.8.e, efferent segmental canal ; F.B, fat- body ; gl, glomerulus ; A", kidney ; l.c.r,
longitudinal collecting canal ; M, Malpighian body ; J/rf, Miillerian duct ; ^V,
nephrostome ; 0, ovary ; Ov, oviduct ; s.dj segmental duct ; T, testis ; (/V, ureter ;
V.d, vas deferens ; F.«, vesicnla seminalis.

VOL. VIII E

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50 AMPHIBIA

interrupted and the vestiges of the transverse canals are no longer
functional. The eggs fall into the body-cavity and are caught
up by the ostium or inner abdominal opening of a special duct,
the oviduct (Miillerian duct of many authors). Vestiges, more or
less complete, of these oviducts persist in the males of most
Amphibia.

This general scheme presents some modifications in the
various groups of Amphibia.

The Apoda retain the most primitive conditions. The kidneys
are still long and narrow, and the glomeruli are, at least in the
anterior part of the organ, still strictly segmental, agreeing in
number and position, each with a vertebral segment ; later, the
number of the glomeruli is greatly increased, and the former
agreement becomes quite disturbed. The generative glands still
retain their segmental arrangement, but they are restricted to
a much shorter region than the kidneys. In the male Apoda a
considerable portion of the cloaca can be everted by special
muscles, and acts as an intromittent organ. Both sexes possess
a .ventral urinary bladder.

In the Urodela both kidneys and testes are much concenti^ated,
the testes especially have lost all outward appearance of seg-
mentation, and their efferent canals, connecting them with the
longitudinal collecting canal, are much reduced in numbers. The
greater portion of the kidneys, at least their anterior half, has
all the appearance of a degenerating organ and is on the way to
losing its urinary function, although it still possesses Malpighiau
bodies and complete ducts ; the main function of the latter is
now the conveyance of the sperma. In the Perennibranchiata,
and in some others, e.g. Spelcrpes variegatns, the longitudinal
collecting canal, between testis and kidney, is sometimes sup-
pressed, a very simple, but pseudo- primitive arrangement. A
urinary bladder is present. The cloaca is not evei^sible.

In most Anura, e.g. Jiaiia and Bnfo (Fig. 7 ; 4, 6), the same
scheme is adhered to. The efferent canals of the testis form a
network, with a longitudinal canal, and open into the efferent
canals of the kidney, in the substance of which they are more <jr
less deeply imbedded. The ducts which lead out of the kidney
to compose Leydig*s duct, are frequently dilated, or the latter
duct is much elongated, convoluted or varicated, and this whole
portion is enclosed in a sheath of connective tissue, giving an

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II URINO-GENITAL ORGANS 5 1

appearance as if the single duct itself were dilated in the greater
part of its length ; hence the occasional name of vesicula seminaUs.
Such means of storing the sperma enable the latter to be ejected
suddenly in great quantities.

In BomMnator (6) some of the most anterior seminal canals do
not perforate the kidney, but run over it superficially and open
directly into a branch of Leydig's duct. This branch, no doubt
equivalent to a number of segmental canals which have lost
their uriniferous function, is curved round the upper end of the
permanent kidney, while its forward continuation, ending blindly,
is the remnant of its former headward extension. This arrange-
ment of BomMnator is carried further in Discoglossm (7). The
testis conveys its sperma through a wide duct directly into Leydig's
canal, without interfering with the kidney, and all the testicular
efferent network is lost. The anterior end of Leydig's duct
still extends headwards ; its middle portion acts solely as a vas
deferens, while the lower portion still behaves like a typical
segmental duct, conveying both sperma and urine. Lastly, in
Alytes (8) the functional division of the old segmental duct has been
carried to an extreme. The kidney is drained by one canal only,
now a true ureter, and this is of course produced by a consolida-
tion of the multiple exclusively uriniferous canals of the lower
half of the kidney. The whole of the segmental duct is now
in the service of the testis, and near its junction with the ureter
it forms a large diverticulum or true vesicula seminalis.

Eemnants of oviducts, or Miillerian ducts, are common in the
male Anura; they are best developed in Bufo, much reduced,
and individually absent, in Rajia, In Bomhinator each duct is
restricted to its upper or abdominal portion, and is attached to
the vestigial headward extension of Leydig's duct. Lastly in JJis-
coglossus and in Alytes aU traces of oviducts seem to have vanished,
at least in the adult males.

It is interesting to note that in the arrangement of the m^ino-
genital ducts the Discoglossidae are the most advanced of all
Amphibia, instead of showing the most primitive conditions.
This is rather unexpected, but is paralleled by the epiehordal
type of the vertebral column.

The oviducts of the Apoda and Urodela remain more or less
straight ; in the viviparous species they form uterus-like dilata-
tions. In the Anura they become greatly elongated during the

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5 2 AMPHIBIA CHAF.

breeding season and form many convolutions. As a rule each
oviduct opens separately into the cloaca, but in Hyla they have
one unpaired opening, while in Bufo and Alytes the lower parts
of both oviducts are themselves confluent.

All Amphibia possess Fat-bodies. They consist of riclily
vascularised lymphatic tissue, the meshes of which are filled with
lymph-cells, globules of fat and oil. In the Apoda these bodies
lie laterally to the generative glands, and along the posterior half
of the kidneys. In the XJrodela they accompany the anterior
half of the kidney. In the Anura they are lobate, and are
placed upon the anterior end of the testes or ovaries. Their
exact function is still doubtful, but it is intimately connected
with that of the generative glands. The old notion, that
they are simply stores of fat for the nourishment of the animal
during hibernation, is quite untenable. The fat-bodies do not
decrease during this period, on the contrary they attain their fullest
size in the spring at the time of the rapidly awaking activity of
the reproductive organs, and they enable considerable quantities
of sperma and of eggs to be produced and ripened without detri-
ment to, or utter exhaustion of, the animals, which often spawn
before they have had time or opportunity to feed. After the
spawning season the fat-bodies have dwindled down to incon-
spicuous dimensions.

l^astly, there is in some Anura, hitherto observed in Bufo only,
a mysterious organ, intercalated between the fat-body and the
testis or ovary. This is " Bidder's organ " and it seems to be a
rudimentary ovar}', or rather that upper, anterior portion of the
whole organ wliich undergoes retrogressive metamorphosis. It
disappears in old female toads, but in the males it Bometimes
assumes a size equal to, or surpassing that of the testes. The
males are in this respect hermaphrodite, and cases are known in
Avhich parts of the generative glands have developed testes and
egg-bearing ovaries.

The spermatozoa of the Apoda and Urodela have an undulat-
ing membrane along the tail, while the head-end is either pointed
or truncated. Those of Spelerpes fnscus and of Ichthyophis
(jlutiiwsa measure about 0*7 mm. in total length, those of the
other I'rodela being much smaller. A peculiarity of tlie Urodela is
that their spermatozoa are massed together in or upon spermato-
pliores, an arrangement which undoubtedly facilitates the internal

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SPERMA-EGGS 5 3

fecundation of the female without actual copulation. The female
takes up such a deposited spermatophore with the cloacal lips,
squeezes the sperma out of the capsule which remains behind, and
either conveys the former into a special receptaculum seminis,
e,g, in Salamundra atra and in Triton, or the spermatozoa wriggle
their way, thanks to the undulating tail, directly up the oviducts
to the ova.

The spermatophores are composed of a colourless, soft, gela-
tinous mass, which is probably produced by the cloacal gland.
The shell of jelly is in fact a cast of the cloacal
cavity, reproducing all its ridges, furrows and
folds, while a toad -stool -shaped papilla of
the cloaca makes the inside lumen of the cast,
e^. in Triton, Those of Salamandra maculosa
are much simpler, consisting, in conformity
with the absence of a cloacal papilla, merely of
a cone with a irlobular mass of sperma on the „„.,„,

^, n > , , . .1 Fio. 8.— A bell-sliai^ed

top. Those of Amolystoma are similar. Kpermatophore of

The spermatozoa of the Anura show con- Jj^iz^rV^^'
siderable differences in the various genera, of
which, however, only the European forms have been properly
examined. The " head " is wound like a corkscrew in Discoglossus,
FelobcUes, and Pdodytes ; spindle-shaped, more or less curved, in
Rana ttmporaria and R. agilis, Hyla, Bvfo and Bomhinator, in
the latter with an irregular membrane on one side ; cylindrical
in Rana esculenta and R, arvcdis. The tail is mostly long and
filiform, but in Bufo vulgaris and Diseoglossus it is provided with
an undulating membrane. Their size is generally veiy small,
only about 01 mm., excepting those of Diseoglossus which reach
the astonishing length of 3 mm. Tliese differences in shape,
especially that of the head, explain why species of the same
genus, e.g. Ra7ia temporaria and R, arvalis, cannot fertilise each
other.

The eggB differ much in size, colour, and numbers. They
are holoblastic, with unequal cleavage, but those species which
possess an unusual amount of food-yolk, for instance R/iacophorifs
scUegeli and the Apoda, approach the meroblastiiJ type of segmen-
tation. As a rule, the greater the amount of yolk, the snialler
is the number of eggs produced. But the number which is laid

> ZeitBchr, wUs, Zod, xlix. 1889, p. 583.

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54 AMPHIBIA CHAP.

during one season is not only difi&cult to calculate, but it varies
individually, old females laying more than young specimens.
Moreover, some kinds, e.g, the Discoglossidae, spawn several times
in one year. Alytes, Rhiiioderma, HylodeSy Hhdcophorus, Fipa,
in fact those kinds which are remarkable for special nursing
habits, lay only a few dozen eggs at a time. JETyla arhorea pro-
duces up to 1000, BaTut temporaHa about 3000, Bufo 'vulgaris
averages 5000, Bufo vi7*idis find Rana esculeiita up to 10,000
and more. T. H. Morgan ^ has observed a Bufo lentiginosus
which laid 28,000 e^s within ten hours! The nimiber of eggs
produced by the Apoda and Urodela is comparatively moderate, in
the average a few dozen, Amhly stoma alone laying about 1000.

The eggs possess a gelatinous mantle of variable thickness and
coaisistency. In Amphiuma they are strung together like the
beads of a rosary, and the envelope hardens into a kind of shelL
Many Xewts and some Anura fasten their eggs singly on to
plants and other objects in the water, with or without threads of
stiffening mucus. In many Anura, e.g. Bufonidae, they pass out
as closely-set strings of beads, one string out of each oviduct ; in
others, e.g. Ranidae, they are disconnected, and form large, lumpy
masses, especially when the gelatinous mantle swells up in the
water. The use of this mantle seems to be chiefly the protection
of the growing embryo, which in many species, when hatched out .
of the egg proper, drops into and remains for some time in the
softened jelly. Possibly the latter affords some nutriment to the
early larva.

Concerning the mode of fecundation it is to be remarked
that copulation proper takes place only in the Apoda. For the
Urodela Boulenger^ has given the following sunanary. In no
case does actual copulation take place. The male deposits the
spermatophores which it is the ofl&ce of the female to secure : —

I. No aiuplexiis, but a lengthy courtshii) in the water ; the male is more
brilliantly coloured than the female, and omainented with dorsal and
caudal crests, or other aj)pendagef5 : Triton, cf. also systematic part^
II. Aniple.\us takes place ; there are no marked sexual differences in colour
and no ornamental dermal appendages.
A. Amplexus of short duration, partly on laud, but deposition of the
s])erma in the water. No accessory sexual characters : Terrestrial
Salamanders, namely Salamandra, Chioglossa, Salamandrina, Spel-
erpes breeds in damp caves without water.

^ Amer. Natural, xxv. 1891, p. 753. « ^^^^i Jahrb. Syst, vi. 1892, p. 447.

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OVIPOSITION — NURSING 5 5

B. Aniplexus of lengthy duration and in tlie water.

a. The male, distinguished by a greater development of the fore-
limbs, which are armed with temporar}* excrescences, clasps the
female in the axillary region with the fore-limbs : Triton waltli.

b. The male, distinguished by a greater development of the liind-
limbe and a prehensile tail, clasps the female in the lumbar and
caudal regions : The Euproctus-gronii of newts : Triton asper^
T. rusconii, and T. montavus.

The act of fecundation of most of the other kinds of Urodela,
notably Cryptohramhus, Amphiuma, Proteus, has not yet been
observed.

Embracing of the two sexes is the universal rule with the
Anura, the male creeping on to the back of the female and
clasping her firmly with the arms and hands either in the
inguinal region, higher up, or under the armpits. See the
numerous statements in the systematic part. This often ex-
tremely forcible, pressing embrace seems to be necessary, although
the females can deposit the eggs without the help of the male,
but in such cases the expulsion takes place at irregular intervals
instead of at one time. When the eggs appear at last, and this
happens in many species many hours, or even some days, after
the beginning of the embrace, the male voids the contents of
its seminal vesicles over them. Fertilisation is consequently
external, with the possible exception of Pipa, q.i\ p. 152.

Deposition of the eggs and nnrsing habits. — The majority
of the Amphibia are oviparous, but some Apoda and Urodela
are viviparous. It is unnecessary to call the latter condition
ovo- viviparous, since this is really a distinction without a
difference.

Viviparous forms : — amongst Urodela ; Salamandra maculosa y
the young burst the egg-membrane in the act of being born, and
are provided with long gills ; S, atra, the young undergo their
whole development and metamorphosis within the uterus (see
p. 119); Spderpes fusc%(s, the young are likewise born in the
perfect condition : amongst Apoda ; Typhlonectes coinjyressicauda
and Dermophis thomensis.

The oviparous Apoda, at least Ichthyopthis and Hypogeophis,
and a few of the Urodela, as Desmognathus and Amphiuina, take
care of their eggs by coiling themselves aroimd them in a hole
underground.

Nursing habits are verj' common amongst the Anura.

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56 ANURA

Boulenger * has summarised the various conditions concerning the
deposition and care that is taken of the eggs, in the following
list, in which more recent diacoyeries have been interpolated.

I. The ovum i^ Hiuall, and the lan-a leaves it in a comparatively early
embryonic condition.

A. The eggB are laid in the water : —

a. Without further care or preparations : probably the majority
of Anura ; all European forms, except Alytes.

b. The eggs are laid in a specially walled -in jjart of the pond :
Hyla faber.

B. The eggs are (Ieixj»ite(l out of the water : —

a. In holes, or under grass, near the banks of pools. The larvae
are lil)erated and washed into the water by the next heavy
rain : Lepiodactylus ocellatus, L. myitadnusy Palndicola gracilis^
Pseudophryne australu and P. bibroni.

b. On leaves alx)ve the water, the larvae dropping down when
leaving the egg : Chiromantis rufescenSy Phyllomedum iheringi^
Ph. hypochondrialis,

II. The yolk is very large and the young undergoes the whole or part of
the metamorphosis within the egg ; at any rate the lar\*a does not
afwnime an independent existence until after the loss of the gills.

A. The eggs are deposited in damp situations, or on leaves. The
young escape as : -

a. Tadpoles : Arthroleptu aeyckellensis^ Rhaeophorus sehUgeliy Bh
macuUituK.

b. Perfect, air-breathing frogs : Rana opistkodon, HyMes vmr-
tinicenm, Hyla nebulom.

B. The eggs are carrieil by a jmrent.

a. By the male : —

a. Round the legs ; the young leaves the egg in the tadpole

stage : Alytes.
p. In the enlarged vocal t^acs ; the young leave in the j^rfect

state : Rhinodernia.

b. By the female : —

a. Attached to the belly : Rhaeophorus reticidatus,
p. -Attached to the back ; the young complete their metamor-
phosis within the egg : Pipa,
y. In a dorsal pouch which the young leave as tadpoles : Koto-
trema marsupiatum ; — or in the perfect state : Nototrema
testudineuviy N, comutumy N. oviferum, N. JistipeSy and Hyla
yoeldti.

The development and metamorphoBii of many species have
been described in the systematic part. The following is a short
general account of some of the more important features. Meta-
morphosis in the Apoda and Urodela is restricted chiefly to the
reduction of the gills, the closing of the clefts, and the loss of the

» Ann. Xat. Hist. (5), xvii. 1886, p. 463.

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TADPOLES 57

gill-chamber and the finny margins of the tail ; but the change
from the tadpole to the final Anurous animal implies an almost
entire reorganisation.

In the earliest condition the embryo consists of a large head
and body, while the tail is still absent. Behind the beginnings
of the future mouth appears a transverse crescentic fold, with the
convexity looking backwards, which develops into the paired or
unpaired adhesive apparatus. This consists of large complex
glands, developed in the Malpighian layer, originally covered by
the cuticula, which soon disappears, whereupon the sticky secre-
tion enables the larva to attach itself to the gelatinous mantle of
the egg, later on to weeds or other objects in the water. The
name of suckers, often applied to this apparatus, conveys a wrong

Fio. 9. — Four stages of the development of the adhesive apparatus {A) of Bvfo vulgaris ;
i/, Month ; Sp.T. spiracular tube. In 8 the gills are almost completely hidden by
the united right and left opercular folds. The small outlined figures indicate the
shape and natural size of the tadpoles. (After Thiele.)

idea, there being neither muscles nor any suctorial function. The
shape of this organ undergoes many changes during the early
life of the individual, and differs much in the various genera,
affording thereby diagnostic characters.^ At first a crescent, it
divides into a right and a left oval or disc, which either remain
asunder and behind the mouth {Rana, Bufo), or they move for-
wards to the comers of the mouth {Hyla) or further back, and
unite again more or less completely, as in Diacoglossus and
Bombinator, It is mostly of short duration, and disappears by
the time that the larva, by the proper development of the gills
and the tail and the functional mouth, changes into tlie tadpola
But in a few species these discs transform themselves into an
elaborate ventral disc. Such an organ persists throughout the
greater part of the tadpole-stage in certain Oriental species of
Hana, all of which, when adult, possess fully webbed toes and
1 J. Thiele, ZeiUchr, tf^iM, Zool, xlvi. 1888, p. 67.

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S8 ANURA

strongly dilated discs on the fingers and toes, e.g. Raim ivhite-
headi, R. iiatatriXy and R, cavitympanum of Borneo, R, jerboa of
Java (this larva was originally described and figured as that of
Rhacophoni^ reinwardti), and R, afghana of the Himalayan system.
These tadpoles, at least those of R, jerboa, are further remarkable
for having the " spiracular " opening very far back on the left
side, nearer to the base of the tail than to the snout, so as to be
well out of the way when the creature has attached itself by the
adhesive disc.

The mouth of the tadpoles of Anura is furnished with horny
armaments, substitutes for teeth. Their development and that
of the mouth in general has been well described by Gutzeit.^ In
the young larvae of Rana temporaria, one or two days after
hatching, a sliallow groove appears above the conspicuous pair of

adhesive organs. The groove
becomes rhombic in outline,
and when the mouth has
been formed in its centre,
the jaws appear in the
median corners of the
rhombus. The epidermis
then rises like a circular
wall around the jaws, and
FIG. 10. -1 Front view of the mouth of a tad- divides into an Upper and

pole of Kaiui teniporana, showing the trans- * *

verse rows of tiny homy teeth ; 2, three loWCr Up ; furrOWS appear

'^~\^^^'^y'^''^^'^''^^'^^^^^'^^^^' on them, and between

these various papillae and
comb-like transverse plates of teeth. The papillae are pos-
sibly tactile organs, but although nerves enter them, nerve-
endings of a sensory nature have not yet been discovered.
On the fourth day the jaws become black, by the tenth day
horny teeth have appeared upon all the plates of the mouth-
armature, and on the seventeenth day the mouth-apparatus has
reached the configuration typical of the tadpole, which is now
about 14 mm. long. The number of horny teeth in R. temporaria
amounts to about 640. These teeth are not cuticular products,
but cornified cells ; they are very small, and consist each of one
liorny cell, which is shaped like a nightcap, the apex of which is
curved back and serrated. The little teeth are shed continu-

^ Zeitschr. inss. Zool. xlix. 1889, p. 43.

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TADPOLES 59

ously, the renewal taking place by successive cells growing into
the bases of the older series. The shape and size differ much in
the various genera and species. The comb-like plates, composed of
those teeth which 'surround the lips, seem to be used chiefly for
the fixing or hooking of the food, while those which compose
the horny beak proper, the armature of the jaws, are used like
the radulae of snails. These beaks are likewise composed of a
great number of individual teeth, closely packed together in
several rows, but the teeth themselves are simple and not serrated.

In Hyla arbor ea there are in all about 560 teeth. The
development of the mouth does not begin before the eleventh day ;
the horny teeth break through, and the jaws get black edges, on
the eighteenth. In Pdohates fvscus the number of horny teeth
is increased to about 1100. In Borborocoetes taeniatus the horny
teeth form series of five bells, which fit into each other like the
joints of a rattlesnake's tail.

One of the most extraordinary kinds of tadpoles is that of
Metjalophrys montana} Mr. Annandale (Skeat Expedition) found
it at Bukit Besar, Malay Peninsula, from 2000 to 3000 feet
above the level of the sea. The tadpoles (Eig. 11) were found
in the beginning of the month of May 1899 in sandy streams
and in pools of rain-water ; they floated in a vertical position,
the peculiar membranous funnel-shaped expansion of the lips
acting as surface-floats. The inside of the funnel is beset with
radiating series of little horny teeth, and the whole apparatus
is possibly used for scraping the under -surface of the leaves
of water-plants in search of food. Total length of the tadpoles
1 inch.-

The gills, the formation of the operculum, and the modifica-
tions of the branchial arterial arches have been described fully
on p. 43 ; those of the hyo-branchial skeleton on p. 31. Fusion
of the opercular fold with the skin of the neck, across the
branchial region, causes the head to become confluent with the
trunk (cf. Fig. 9, 3, p. 57). The body becomes oval, more or less
globular, and the alimentary canal is greatly elongated and stowed
away in the shape of a neat, very regular spiral, shining through

^ M. Weber, Ann. Jard, Botan, Buitenzorg, Siippl. ii. 1898, p. 5.

^ For "A Synopsis of the Tadpoles of European Batrachians," see Bouleuger,
P, Z. 8, 1891, pp. 593-627, pis. xlv.-xlvii. ; also Bedriaga, "Tableaux synoptiques
ixjur servir k la determination des larves des Batracieus Urodtles," C I\. Ass.
Fran^. Sci. ii. 1891, pp. 540-546.

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ANURA

the ventral wall of the body ; the anus opens at the end of a
somewhat protruding tube, either in the median line, just in front
of the ventral fin (Discoglossidae, Pelohates, Bufo), or it assumes
an asymmetrical position by turning to the right side {Hyla,
Baria).

Although both pairs of limbs begin to bud simultaneously, or

Fig. 11.— Tadpoles of Mfi/alophri/s wontana from Bukit Besar, Malay Peninsula, x 3.

the fore-limbs even earlier, the hind-limbs are hurried on, and
appear first, long before the fore-limbs. The latter lie ready
beneath the skin of the gill-chamber, and the right always breaks
through the skin, while the left does the same in the Medio-
gyrinidae, while in the Laevogyrinidae it is generally pushed
through the left -sided spiracular opening, immediately behind
the outer gills. According to Barfurth the right limb appears,
in about 80 per cent, of Raim escidenta, from two to eight hours
before the left.

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METAMORPHOSIS 6 1

Meanwhile the lungs are being developed, and the tadpole
occasionallj rises to the surface to breathe air. The gills, which,
as has been explained elsewhere, are less ancestral than they are
larval organs, degenerate, and all the organs are modified for the
coming terrestrial life. The fins of the tail are absorbed, the
homy armature of the mouth and lips is shed in pieces and makes
room for the true teeth, the eyes receive lids, and the whole
cranium, especially the apparatus of the jaws, undergoes the final
modifications — widening and lengthening of the mouth, arresting
of the mento-Meckelian cartilages, elongation of the Meckelian
cartilages or lower jaw proper, shifting backwards of the aus-
pensorium, and lengthening of its orbital process to form the
pterygo-pjJatine bridge.

The tadpole ceases to feed, the whole intestinal canal is voided
of its contents, and by " histolysis " is thoroughly rebuilt, becoming
wider and shrinking to about one-sixth of its original length,
— undoing thereby the spiral — preparatory for the coarser food,
which consists of insects, worms, and other strictly animal, living
matter. Hitherto the tadpoles have lived on " mud," confervae,
Diatoms, rotting vegetable and animal matter. The anal tube
collapses, becomes ultimately absorbed, and a new vent is formed
at and below the root of the tail.

Barfurth ^ has made interesting observations and experiments
with regard to the absorption of the tail and other organs which
disappear during the metamorphosis. This is retarded by
low temperature; it is accelerated by rest and freedom from
mechanical disturbances, as, for instance, concussion of the water.
Hunger shortens or hurries on the last stages of metamorphosis,
the absorption of the tail taking place in four instead of five days.
Amputation of the tail has no retarding influence ; it is followed
at once by regeneration, although the tadpole may be on the
verge of reducing the tail. Whilst hungering the whole organism
draws upon its available store of material, naturally first upon
those parts which sooner or later are to become superfluous.
This applies eminently to the tail, which represents a consider-
able amount of " edible " matter, and also to that portion of the
skin which still covers the fore -limbs. The elements of the
cutis are resorbed, thereby thinning the skin ; and consequently
the limbs break through earlier in fasting than in well-fed
^ Arch. mikr. Anat. xxix. 1887, p. 1.

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62 ANURA

specimens. Natui-e herself seems to apply hunger as an acceler-
ator. Mile, von Chauvin found that the larvae of Urodela
normally fast during the transformation, and according to Barfurth
the larvae of Rana temporaria eat less after their hind-limbs
are fully developed. This is, however, also • preparatory for the
reorganisation of the gut, which has to be more or less empty
during the shortening process.

The loss of the tail is not due to a sudden dropping off of this
organ — a crude but by no means uncommon belief — but is
brought about by a very gradual process of resorbtion. When
the fore-limbs begin to break through the skin, the tip of the
tail shrinks and becomes black, owing to an increase, or rather
concentration, of the pigment cells. The reduction proceeds from
the tip forwards until on about the fifth day there remains only
a short, conical, black stump. From the beginning of this process
of reduction the tail is scarcely used for locomotion, the tadpole
rowing with its legs, or it crawls and hops about, although the
tail may still be 20 mm. long. The cells of the epidermis atrophy,
shrink, and peel off, while those of the cutis, blood-vessels, nerves,
muscles, and chorda dorsalis become disintegrated, often under-
going fatty degeneration. The leucocytes eat up the debris and
other dissolved tissue, and carry it away through the lymphatic
vessels, to be used as new building material in the rest of the
animal.

Barfurth asks very properly. Why do these tissues degenerate
and die ? Because the vasomotor nerve-fibres cease to regulate
the circulation. And why does this trophic influence of the
central nervous system stop ? Because the function of the tail
becomes superfluous through the appearance of the fore-limbs.
The tail is doomed, and degenerates like any other organ without
a function. The whole process is, of course, a recapitulation of
ancestral, phylogenetic evolution.

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CHAPTER III

NEOTENY REGENERATION TEMPERATURE GEOGRAPHICAL

DISTRIBUTION

Neoteny. — It has long been known that the larvae of the Spotted
Salamander occasionsJly attain the size of 80 mm. or about 3
inches, whilst the majority undergo metamorphosis when they
are only 40 mm. long. Again, larvae of IViton have been found,
in the months of April and May, 80 to 90 mm. long, still with
functional gills, but with the sexual organs fully developed. De
Filippi ^ found in one locality in Lombardy, besides a few normal
fully metamorphosed specimens of only 30 mm. in length, more
than forty specimens, which, although they had attained full
size, about 55 mm., and were sexually mature, still retained their
gills. According to him such gill-breathing, otherwise mature
specimens, occur constantly in a small lake in the Val Formazzo,
on the Italian slope of the Alps, in the province of Ossola. Later
Dura^ril ^ astonished the world by his account of the metamor-
phosis of the Mexican gill-breathing Axolotl into an entirely
lung-breathing and terrestrial creature, hitherto called Amhhj-
stoma, and supposed to be not only a difierent species, but to
belong to a different family from the Axolotl, which was known
as Siredon axolotl s. pisciforme, and naturally classed with the
Perennibranchiatii.

This discovery led to a series of observations and experiments,
chiefly conducted by Marie von Chauvin, instigated thereto by
Koelliker and by Camerano.^ It was then found that many,
if not most of the European Amphibia, both Urodela and Anura,

^ Arch, per zool. eper I'anat. comp.y Geneva, 1861, p. 206.
2 Ann, sei, not, (5), vii. 1876.

' Mem, Ace, Torino, xxjcv. 1883, and Atti Ace. Torino, xvii. 1883, \\ 84. See
also Wolteredorff, Zool Garten, 1896, p. 327.

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64 ' AMPHIBIA CHAi'.

occasionally postpone their metamorphosis, and also that such
Urodela sometimes become adult for all practical purposes, but
retain their gills.

This retardation, the retention of larval characters beyond the
normal period, was called Neotenie by KoUmann ^ (i/€09, young :
Teti/ft), extend, stretch). He distinguished further between : — 1.
Partial Xeoteny, namely, simple retardation of the metamor-
phosis beyond the normal period, for instance, the wintering of
tadpoles of Pelohates fuscns, Borribinator pachi/pus, Pelodytes
punctatitSy Alytes obstetricans, Hyla arhorea, Rana esculenta, H.
Umporrfvia, Biifo vulgaris, and B, viridis: II. Total Neoteny,
where the animal retains its gills, but becomes sexually mature ;
hitherto observed in Urodela only, ag. Triton vulgar is, T. alpes-
iris, T, cristatuSy T. boscai, T. walUi and Amhlystoma, Inter-
mediate stages between these two categories are not imcommon.

A satisfactory explanation of the meaning of neoteny is beset
with difficulties. Some authorities look upon the phenomenon
simply as the result of adaptation to the surroundings, which
make it advantageous for the creature to retain its larval features.
Others think that the surroundings somehow or other retard or
prevent the assumption of the adult characters. Undoubtedly
there are many cases in which larvae have been reared in water-
holes with steep walls, so that they could not change from aquatic
to terrestrial life, and it stands to reason that abnormally forced
and prolonged use of the gills and of the tail may stimulate these
organs into further growth at the expense of the limbs and
other organs whicli are intended for terrestrial life. But not
unfre([uently typical neotenie and overgrown specimens occur side
by side with others which have completed their metamorphosis,
and the same is true of larvae of newts which w^re reared, for
experimental purposes, under exactly the same conditions — for
instance, in a high- walled glass vessel.

Weisniann tried to explain neoteny as cases of reversion to
atavistic ancestral conditions, but this idea is l>ased upon an
assumption which is probably wrong. His idea necessitates
the supposition that all the Amphibia were originally gill-
bn^a thing, aquatic, and limbless animals, and that every feature
s(^en in a larva must necessarily indicate an ancestral phylo-
•ronetic sta.Lce. It is, on the contrary, much more probable that

* rcrh. (Jes. Basel, vii. 1882, p. 387.

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Ill NEOTENY 65

the external gills of the Urodela have been developed in adapta-
tion to their embryonic and larval, essentially aquatic, life. Con-
sequently the possession of such gills would be a secondary, and
not, strictly speaking, an atavistic feature. Normal loss of these
gills, exclusively pulmonary respiration, and preponderating
terrestrial life characterise the final adult Amphibian. These
casea of neoteny are therefore instances of more or less complete
retardation, or of the retention, of partially larval conditions.

The whole problem is, however, by no means simple. Sala-
maiidra atra has become viviparous, and the whole metamor-
phosis takes place within the uterus ; in fact, the young have an
embryonic, but no larval period, if by the latter we understand
the free swimming and still imperfect stage. Similarly, various
Anura — for instance, Hylodes martinicensis — pass rapidly through
their metamorphosis, and have suppressed the stage of free
swimming tadpoles. On the other hand, in many newts, the
duration of the larval period is much prolonged, find moreover
is very subject to individual variation. In the Axolotl this
larval period is continued until and after sexual maturity is
reached. The extreme condition would then be represented by
the Perennibranchiate genera. It may seem reasonable to look
upon these as the youngest members of the Urodela, and the loss of
the maxillae in the Sirenidae and Proteidae supports this idea.
But it so happens that the majority of the most neotenic genera
are more primitive in the composition of the skull and the verte-
bral column than the typically terrestrial and rapidly meta-
morphosing genera. Witness the amphicoelous vertebrae, the
completeness of the pterygoids, the separate nature of the pala-
tine bones, and the separate splenials, as mentioned in detail in
the description of their skull.

We have therefore to conclude, first, that the various Perenni-
branchiate genera do not form a natural group, but are a
heterogeneous assembly ; secondly, that they have become Perenni-
branchiate at a phylogenetically old stage — in fact, that they are
the oldest, and not the newest, members of the pi-esent Urodela.
At the same time, it would be erroneous to suppose that the
first Urodela were aquatic creatures, provided with a finny tail,
with small, ill-developed lungs, and with epidermal sense organs.
All these features are, on the contrarj', directly correlated with
aquatic life, and are larval acquisitions, not ancestral reminis-

VOL. VIII F

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66 AMPHIBIA CHAP.

cences. It would be equally wroug to allude to the absence of
lungs in many newts as a piscine and therefore ancestral feature.
The development of the typical pentadactyloid limb, the con-
nexion of the pelvic girdle with the vertebral column, the
development of the lungs, and absolute suppression of internal
gills point without doubt to terrestrial creatures. What then,
may we ask, were the first Amphibia like ? and how about
the external gills ? They were undoubtedly akin to the less
specialised Lepospondylous Stegocephali, in particular the gill-less
Microsauri, and the various stages may perhaps be reconstructed
as follows : —

(1) Terrestrial, with two pairs of pentadactyloid limbs ;
breathing by lungs only ; with a fully developed apparatus of
five pairs of gill-arches, which during the embryonic life perhaps
still carried internal gills ; with or without several pairs of gill-
clefts. Eeduction of the dermal armour and of the cutaneous
scutes had taken place.

(2) Additional respiratory organs were developed by the
embryo, in the shape of external gills ; these were at first re-
stricted to embryonic life (as in the existing Apoda), but were
gradually used also during the aquatic life of the larva. These
external gills, together with the lungs, have superseded the
internal gills, of which there are now no traces either in Urodela
or in Anura.

(3a) Some Urodeles, retaking to aquatic life, retained and
further enlarged the external gills into more or less permanent
organs (cf also Siren, p. 186).

(3&) The majority of Urodela hurried through the larval,
aquatic stage, and sQme — e.g. Salamandra atra — became abso-
lutely terrestrial. The possession of unusually long external gills
by this species and by the Apoda indicate that these organs are
essentially embryonic, not larval, features.

Begeneration. — Most Amphibia possess the faculty of re-
generating mutilated or lost limbs. This takes place the more
certainly and quickly the younger the animal The amputation
necessary to study these phenomena need not be experimental.
Axolotls and other Urodelous larvae frequently maim ecwjh other
fearfully, by biting off the gills or one or more limbs. The gills do
not even require amputation. If the larvae are kept in stagnant
water the gills often shrivel up or slough off and grow again.

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Ill REGENERATION TEMPERATURE 6j

The same applies to the larvae of viviparous species, e.g. Sola-
mandra atra, which, when cut out of the uterus and put into
water, soon cast oflf their long, tender gills and produce a stronger
set. In an Axolotl,^ two years old, a hand was cut off. After
four weeks there was a conical stuinp ; after the sixth week this
stump had two points; in the eleventh week three or four
fingers were discernible, and a week later the complete hand.
Frequently these creatures reproduce five instead of the normal
four fingers. But the more proximal the cut, the more liable is
the new limb to reproduce supernumerary fingers, or even extra
hands and feet. Complete regeneration of the limb, cut oflf in the
middle of the humerus, took place within five months.

Triton taeniatus, adult, reproduces cut fingers within five or
six weeks, and if the hand be cut above the carpus, new finger-
stumps appear in about one month. Gotte has observed that an
adult Proteus did not completely reproduce its whole leg until
after eighteen months ; and, according to Spallanzani, more than
one year elapses before the limb, bones, and cartilages of Triton
regain their normal strength.

The Anura are likewise capable of regenerating their limbs,
the more readily the younger the specimens. For instance, in
a tadpole of Bana temporaria, in which the fore -limbs were
still hidden, the hind-Umb, cut at the middle of the thigh,
reproduced nineteen days later a knee, followed by a short two-
toed stump. Ultimately the whole limb became completed. The
tail of tadpoles regenerates very quickly and completely, even
if it be cut off shortly before the final metamorphosis, when the
tail would in any case be reduced. Metamorphosed Anura have
almost entirely lost this faculty, but not absolutely. I myself
have kept two specimens of Rana te7nporaria, which, when
already adult, had each lost a hand at the wrist. First there
was only the clean-cut stump with a scar, but within a year this
changed into a four-cornered stump, and two of the protuber-
ances developed a little further, reaching a length of about 4 mm.
These specimens lived for four years without further changes.

Temperature. — Amphibia, like Fishes and Reptiles, are, as
a rule, classed as cold-blooded animals, in opposition to the warm-
blooded Birds and Mammals. This distinction is one of degree
only. The terms poikilothermous and homothermous (ttolkiXo^,
^ Barfurth, Arch. EMxoiekmech, I. 1895, p. 117.

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68 AMPHIBIA

CHAP.

variable; ofw^, equable) are based upon a sounder principle,
but are likewise liable to exceptions. Those creatures which,
like Birds and Mammals, possess a specific temperature of their
own under normal conditions, that of hibernation being excepted,
are homothermous. Cold-blooded creatures have no specific
temperature ; they more or less assume that of their surround-
ings. Frogs and newts, for instance, when living in the water,
naturally assume its temperature, which is, of course, many
degrees lower in a cold spring than in a shallow pond warmed by
the sun on a hot summer's day. The same applies to the
changes from day to night. Dark-coloured tortoises basking in
the sun are sometimes so hot that they are disagreeable to
touch, since they possess but little mechanism for regulating
their heat. The same individual cools down during a chilly
night by perhaps 40° C. Anum are, however, very susceptible
to heat ; most of them die when their temperature rises to about
40° C. Under such conditions they die quickly when in the
water, but in tlie air their moist skin counteracts the heat,
lowering it by evaporation ; otherwise it would be impossible for
a tree-frog to sit in the glaring sun in a temperature of 120" F.
Toads and others with drier skins seek the shade, hide under
stones, or bury themselves in the coolest spots available, and many
Amphibia and Reptiles aestivate in a toi-pid condition during the
dry and hot season. Many of them can endure a surprising
amount of cold, and during hibernation their temperature may
sink to freezing-point. This power of endurance does not apply
to all alike ; tropical species can stand less than those which live
in temperate and cold regions. In spite of many assertions to
the contrary, it may safely be stated that none of our European
frog's, toads, and newts sunive being frozen hard. They may
be cooled down to nearly — 1"* C, and they may be partially frozen
into the ice. Circulation of the blood is suspended in such cooled-
(lown frogs ; their limbs may become so hard that they break like
a piece of wood, but the citadel of life, the heart, must not sink
mucli below freezing-point, and must itself not be frozen, if the
animal is to have a cliance of recovering. The protoplasm resists
a long time, and so long as some of it is left unfrozen the rest
will recover. Hibernating frogs are lost if they are reached by
prolonged frost during exceptionally severe winters. Everj" frog
will be killed in an artificial pond with a clean concrete bottom,

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(iEOGRAPHICAL DISTRIBUTION 69

but if there is sufficient mud, with decaying vegetable matter, the
creatures survive, simply l^ecause they are not absolutely frozen.
A severe winter not infrequently kills off all the younger
creatures, while the older and more experienced hide themselves
more carefully and live to propagate the race.

Oeographical Distribution.

There is a very ably written chapter on the geographical
distribution of the Amphibia by Boulenger in the Catalogue of
RitrarhUi Salientiay pp. 104-118. He came to the important
conclusion that the geographical distribution of the Amphibia
agrees in general with that of the freshwater fishes. Glinther's
division into a Northern, Equatorial, and Southern zone is
modified only in so far as the last two are combined into one,
*' Tasmania and Patagonia not differing in any point regarding tlieir
Frog Fauna from Australia and South America respectively."

Boulenger recognises —

I. The Northern zone — (1) Palaearctic, (2) North American, i-egion.
11. The Equatorial Southern zone.

A. Firmistemia division = (*yprinoid division uf (lunther.

1. Indian region.

2. African region.

JL Arcifera di\'Tsion = Acyprinoid division of Giinther.
I. Tropical American region.
•2. Australian region.

In the chapter on geographical distribution in Bronn's lliienrich,
ViHjel^ Systematischer Theil, p. 296 (1893), and in my Ciassijinf'
Hon of Verfebraia (1898), due attention had been paid to the
Amphibia as well as to the other classes of Vertebrata. It
will be seen in the following pages that my arrangement is well
applicable to the Amphibia so far as fundamental principles are
concerned.

It cannot be sufficiently emphasised that any attempt to form
the various faunas of the different classes of animals into one
scheme must necessarily Ije a peiitio ^?r?'//r/y;//. The time-
honoured six zoo-geographical regions established by Sclater and
Wallace represent fairly well the main continental divisions:
Xorth America, South America, Africa, Australia, and the large
northern continental mass of the Old World, with India as a
tropical appendix. There is no correlation and no subordination

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70 AMPHIBIA

in this scheme. Huxley's division (1868) into Notogaka and
Arctogaea (see p. 74) is of fundamental importance. The next
improvement was the combination of the Palaearctic and Nearctic
" regions " into one, an advance originally due to Professor Newton,
carried out by Heilprin (1887) as the Holarctic region. I have,
in 1893, substituted for it the more appropriate term Periarctic,
meaning the whole mass of land which lies around the indifiTerent
Arctic zone. The want of further co-ordination and subordination
required the combination of the African and Oriental or Indian
countries into a Palaeo tropical region (1893); the Ethiopian or
African and the Indian or Oriental regions of Sclater and Wallace
thereby assuming their proper subordinate rank of subregions.

The two primary divisions Notogaea and Arctogaea are
fimdamental. The four secondary divisions, namely the A%is-
trcdian and Neotropical, Periarctic and Palaeotropicai regions,
also stand the test of application to the various classes and main
groups of Vertebrata ; but naturally, under the present con-
figuration of the world, the Palaeotropicai region is nothing but
the Southern continuation of the Eastern half of the Periarctic
mass of land. This is especially obvious so far as India is con-
cerned. There is, however, that broad belt of desert, sand, and
salt-steppes, which extends from North -West Africa to Manchuria,
and this belt is one of the most important physical features of
the Old World. It is complicated by the system of mountain-
chains which, broadly speaking, centre at the Pamirs, and radiate
westwards through the Caucasus and Alps into Spain, eastwards
through the Himalayas into China, and north-eastwards to Kamt-
schatka ; interrupted by Bering's Sea, it is continued as the back-
bone of both Americas to Patagonia.

The tertiary divisions, the subregions, have no real existence.
They depend upon the class, or even order, of animals, which we
happen to study. The faunistic distribution of the Urodela is
not that of the Anura, and both follow separate lines of dispersal,
different from those of the various orders of Reptiles, Birds, and
Mammals. This must be so. There is no doubt that the dis-
tribution of land and water was totally different in the Coal Age
from what it is now. The face of the globe at the Jurassic Age
can scarcely be compared with the aspect which the world has
assumed in the Miocene period.

This leads to another consideration, often neglected. We

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Ill GEOGRAPHICAL DISTRIBUTION 7 1

know that the various classes, orders, families, etc., of animals
have appeared successively upon the stage. A group which
arose in the Coal Age followed lines of dispersal different from
one which was not evolved until Jurassic times, and post-
cretaceous ci^eatures could not avail themselves of what assisted
their ancestors, and vice versd. The Amphibia are bound abso-
lutely to the land and to fresh water ; transportation across salt
water is not excluded, but must be accidental, and is not a case
of regular " spreading/' Speaking generally, the older a group,
the more likely is it to be widely distributed. If it appears
scattered, this may be due to extinction in intermediate countries
or to submergence of former land-connexions.

There is great danger of arguing in a circle. It is one of the
most difficult tasks to decide in cases of great resemblance of
groups of animals between their being due to direct affinity or
to heterogeneous convergence, or parallel development. It is the
morphologist who is ultimately responsible for the establishment
of faunistic regions, not the systematist, least of all he who
accepts an elaborate classification, and then mechanically, mathe-
matically, by lists of genera and species, maps out the world.
Let us take an example. The Neotropical region and Mada-
gascar, but not Africa, are supposed to be faunistically related to
each other. In both namely occur Boa and Corallus amongst
snakes, Dendrobatinae amongst Eanidae, and of the Insectivora
Solenodon in Cuba, Centetes in Madagascar. More cases can
no doubt be found which would strengthen tliis resemblance,
perhaps in support of the startling view that Madagascar
and South America have received part of their fauna from the
famous Antarctica. But the value of the Insectivores has been
disposed of by their recognition as an extremely ancient group,
or as a case of convergence, and the two genera are no longer put
into the same family as Centetidae. The Dendrobatinae {Man-
idla in Madagascar, the others in South America) are decidedly
not a natural group, but an instance of very recent convergence
(cf. p. 272). About the members of the ancient Boidae we do
not feel quite so sure.

It is therefore advisable to eliminate for zoogeographical
purposes groups about which there can be any reasonable
doubt, otherwise we may argue that certain genera must con-
stitute a very old family, because they are now restricted to widely

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72 AMPHIBIA

separated countries, or ou tlie strength of their distribution we
may conclude that the genera in question cannot be related to
each other, and do not belong to the same sub-family or family
as the case may be. Such groups are the Engystomatinae
and the genus Sj^elerpes ; amongst reptiles the Eublepharidae,
Helodermatidae, Anelytropidae, Ilysiidae, Amblycephalidae.

It is customary to represent the various regions and sub-
regions as if they had boundaries as fixed as .political frontiers.
Such limitations are quite arbitrary, and what is of more im-
portance, they differ in reality according to the class or order of
animals with which we happen to deaL Moreover, there lias
been, and is probably still going on, an exchange or overlapping
of faunas. Such debatable grounds are Central America and
the highlands of North-western South America. The famous
Wallace's line, between Borneo and Celebes, Java and Lombok,
is absolutely inapplicable to the Anura. From their point of
view the Austro-Malayan countries, Papuasia and Polynesia do
not form a sub-region of the Australian, but rather of the Palaeo-
tropical region. Concerning the Urodela, the division into Palae-
and Xe-arctic sub-regions is unjustifiable since Eastern Asia has
emphatically American aflSnities (cf also p. 96). The Sahara
and the rest of Northern Africa are intimately connected with
Arabia, Persia, Afghanistan, and Northern India, just as equatorial
Africa and Madagascar possess strong faunistic relationship with
Southern India and tlie Malay islands.

Limiting factors of distriJmiion. — Common salt is poison to
the Amphibia ; even a solution of 1 per cent prevents the develop-
ment of their larvae. Consequently seas, salt lakes, and plains
encrusted with saline deposits act as most efficient boundaries to
normal "spreading." But undoubtedly many individuals have
made long and successful voyages across the seas on floating
trees. Solutions of lime are likewise detrimental to many
species, and it is a general fact that limestone-terrain is poor in
Amphibian life, unless, of course, sufficient accumulation of humus
counteracts or prevents the calcareous impregnation of the springs
and pools in meadows. Scdamaudra maculosa is, for instance,
absent in Central Germany on the Muschelkalk, but it occurs
in abundance in neighbouring districts of red sandstone or
granite ; nor can the larvae be reared successfully in very
*' hard " water. On the other hand, Proteus lives in the sub-

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jii GEOCJRAPHICAL DISTRIBUTION 73

terranean waters of Carniola, where the whole country is nothing
hilt limestone.

Cold is another powerful limiting factor. The absolute
northern limit of Amphibian life coincides rather closely with
tlie somewhat erratic line of 0** Centigrade of annual mean
temperature, a little to the north of which line the ground
remains permanently frozen below the surface. The surface-
crust, which thaws during the summer, engenders an abundance
of insects as food-supply, but its freezing down to the icy bottom
makes hibernation impossible. There are, of course, some ex-
ceptions, for instance the occurrence of Urodela in the Schilka
river and in the district of Lake Baikal.

Banges of mountains are far less effective barriers tlian is
generally supponed. In many cases the fauna is the same on either
slope, and they act rather as equalising or dispersing factors,
especially wlien they extend from north to south. Witness the
Andes, owiiii^ to which Ecuador and Peru tear a great resem-
blance to the Central ^\jnerican fauna, and differ from the
tropical parts of Soutli America. The existence of an AiiMy-
stoma in Siam is another instance.

The more specialised a family the more intimately is it con-
nected with the physical features of the country. Typically
arboreal frogs are dependent on the presence of trees. Some have
undoubtedly spread into treeless countries and have changed
into prairie-frogs, e.g. Avris. They come out, so to speak, as
something dififerent at the other end, and it is unlikely that
these modified descendants redevelop exactly the same features as
their ancestors before the migration. Baldwin Spencer^ met
with only six species of frogs in Central Australia, JAintiody-
nastes, Chiroleptes, Helew^ioruSy and Hyla. They are in the main
identical with certain forms found in the dry inland parts of
New South Wales and (Queensland. They are to be regarded as
immigrants from the latter regions, which have been able in the
majority of cases to adapt themselves to unfavourable climatic
conditions by means of a marked development of the burrowing
habit, to which in certain cases has been added a capacity for
absorbing and holding water.

' The Horn Scientific Expedition, 1897. p. 15.5.

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74 AMPHIBIA CHAP.

Faunistic divisions of the Amphibia.

NOTOG^EA.— South World.

Characterised by the Cystignathidae* and by the predomi-
nance of Arcifera, which form nearly 90 per cent of the Anurous
population.

I. Australian region. — Absence of Apoda and Urodela.
All the Anura are arciferous, with the exception of one species of
Raria in the Cape York peninsula. The fauna of the Australian
continent and of Tasmania consists chiefly of Cystignathidae and
Hylidae {Hyla and HyleUa) and several small genera of Bufon-
idae {Psevdophryne* Notaden* and Myohatrachus*).

It is customary, and from the study of other Vertebrata quite
justifiable, to divide the Australian region into several sub-
regions, but the Amphibia lend no support to this. The only
Amphibian in the Sandwich Islands is a Biifo, closely related to
North American species. The only Amphibian in New Zealand
is Ziopehna* one of the Discoglossidae whicli are otherwise con-
fined to Europe, North-east Asia, and North-west America, and,
to judge from their low organisation, had formerly a much wider
distribution. New Caledonia possesses no Amphibia. The Fiji
Islands are inhabited by one or two species of Coriiufer, a genus
of Ranidae. The same genus is typical of the Austro-Malayan
and Papuasian islands, the fauna of which consists of Bafuf
and Cornufer, Ceratobatrachus, several genera of Engystomatinae,
Hylidae, and Pelobatidae.

II. Neotropical region. — Characterised by Apoda, Aglossa
{Pipa), abundance of Cystignathidae (Hemiphractinae,* Cystig-
nathinae, and Dendrophryniscinae*), Hylidae (Hylinae and
Amphignathodontinae*), numerous Bufonidae and Engystoma-
tinae ; DcndrohatiTiae* ; the Eaninae are represented by a few
peculiar genera, mostly restricted to the Andesian pro\'ince ;
the genus Rana occurs there in a few species only.

Absence of Discoglossidae, Pelobatidae and Dyscophinae.

Several species of Urodela, of the genus Spelerpes, extend
from Central America into the Andesian province, one occurs
in Hayti, and Plethodon platense in Argentina.

This region is by far the richest in the number of families,
* indicates Amphibia which are peculiar to the resjiectiye regions or sub-regions.

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in GEOGRAPHICAL DISTRIBUTION 75

genera and species ; the total number of the latter being, accord-
ing to Boulenger, about four-ninths of the known species. The
region comprises South America^ Central America, and the West
Indiiin islands. Central America is naturally debatable ground ;
one species of Hylodes and one Engystoraay besides about twenty
Hylidae, extend into North America proper, while possibly the
Baninae have entered the Neotropical region from the north.
Bufo is too cosmopolitan to assist our conclusions. The occurrence
of four species of Hylella in South America, one in Australia, and
one in New Guinea indicate that this is not a natural genus.

From the point of the Amphibia the whole region can be
divided into two sub-regions only : (1) The West Indian
islands with Central America and the north-western Andesian
province ; (2) the rest of South America.

AECTOGAEA.— North World.

Characterised by the absence of Cystignathidae.

I. Periarctic region. — Characterised by the Urodela, these
being almost peculiar to the region (cf. p. 96). Absence of
Apoda. Presence of Discoglossidae, Pelobatidae, Bufonidae,
Eaninae. Few Hylinae occur.

The whole region can be subdivided into three sub-regions.

1. Western Palaearctic. — Prevalence of Salamandrinae
(Salama7idra,* Chioglossa* Salamandrhia* Triton); Proteidae
{Proteus angtfinus*) ; Spelerpes fuscus* — Discoglossus, Bom-
hincUor, Alj/tes* Bufo, Hyla arborea, Felobates* Felodytes* JRana.

2. Eastern Palaearctic. — Amphiumidsie (Cryptohra'tichiLs) ;
Amblystomatinae ; Salamandrinae {Triton, Fachytriton* Tyloto-
triton*); Amblystomatinae. — Bombiruitar, Bufo, Hyla arborea,
liana, JRhacophonfs.

3. Nearctic. — ^Amphiumidae {Cryptobranchus, Amphiuma*)]
Proteidae {Typhlomolge^ Necturus*) ; Sirenidae* ; Amblystoma-
tinae : most Plethodontinae ; Desmognathinae.* — Discoglossidae,
Pelobatidae (Scaphiopvs*) ; Bufo ; Hylidae {Hyla, Acris, Choro-
philu^) ; Rana.

II. Palaeotropical region. — Characterised by the presence
of AjKKla and by the great prevalence of Firmisternal Anura,
which amount to nearly 90 per cent of the total population.
* indicates Amphibia which are jieculiar to the respective regions or sub-regions.

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76 AMPHIBIA

Absence of XJrodela (except Amblystoma persimile*), of Cystigna-
thidae, and' practically of the Hylidae, only two of which occur
in the Himalayan district But tliis great chain of mountains
should not be included within the region, while the outlying
spurs in Upper Burma (with Amblystoma) are debatable ground.
The subdivision of this widely extended region is beset witli
difficulties, chiefly on account of Madagascar and Papuasia. The
fauna of Madagascar is very remarkable. All its Amphibia are
Firmisternal, a mixture of African and Indian forms. The island
agrees with Africa, in opposition to the Oriental countries, in no
special point ; all the Baninae, except ifegalixalus, Bapjna, and
two rather common species of Rana, belong to different genera.
Madagascar differs from Africa by the absence of Apoda, of
Aglossa, and Bufonidae. On the other hand, it agrees with India
or with the Malay islands, in opposition to Africa, by the possession
of Dyscophinae, of the Banine genus Hhacophorus, and the Engy-
stomatine genus Calophrynus.

Africa and India agree with each other, and ditier from
Madagascar by the possession of Apoda, the genera Bufo and
Nectophryne, and by the close resemblance of several genera of
Baninae.

India, the Malay islands, and Papuasia with Melanesia
possess Pelobatidae {Leptobrachiuvi* Batrackopsis* Asterophrys*),
and thereby differ considerably from Africa and Madagascar.
Batrachylodes* of the Solomon Islands has unmistakable affinities
with Phrynodenna* of Karen, between Burma and Siani ; Oreo-
hatrachus* of Borneo much resembles Fhryjiobatrachus^ of West
Africa ; and Cornufer, typical of the Malay and Melanesian
islands, occurs also in West Africa. All these Baninae indicate
that the Austro-Malayan and Melanesian islands belong to the
Palaeotropical region. CeratobatrachvSy* t}^e of a sub-family, is
peculiar to Melanesia.

There are consequently several possible modes of subdivision,
all with a different result, according to the group of Amphibia,
which we may select as of leading importance, e.g, Apoda or
Pelobatidae, or Dyscophinae and Bhucophorua. The Engy-
stoniatinae and Baninae are to be eliminated, since they ocrcur in
all the countries in (question. We have either to leave the
whole region undivided — and it is a significant fact that the

* indicates Amphibia whicli are peculiar to the respective regions or sub-regions.

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GEOGRAPHICAL DISTRIBUTION

Indian countries possess not one sub-family of their own — or
we must break it up into four provinces, not sub-regions : —

1. Ethiopian, or continental African, with Agloesa and Apoda, no Pelo-

batidae, no Dyscophinae, few Bufonidae, and many Raninae.

2. Indian and Malayan, with Apoda, no Aglossa, but with Pelobatidae,

Dyscophinae, many Bufonidae and Raninae, amongst which Rhacophorun.

3. Malagasy, without either Apoda or Aglossa ; with Firmistemal Anura

only, chiefly Dyscophinae, ^d Rfiacophor-us and other Raninae.

4. Papua^ian, without Apoda, Agloflsa, Dyscophinae, and Bufonidae, but with

Pelobatidae and Ranidae.

DISTRIBUTION OF FAMILIES AND SUB-FAMILIES OF THE AMPHIBIA.

Palae-arctic.

^ 1

K! 1

5?;

West.

East.

5t '
'A

Aniphiumidae

Proteidae .

-H

Sirenidae

■'r

Apoda.

\
1

-i-

Aglossa

Discoglossidae

+ i

Pelobatidae .

-f

Bufonidae .

'' -H

Hylinae

1 -4-

Amphignathodontinae

-f

' Hemiphractinae .

! 1

' Cystignathinae .

1 +

1 i

Dcndrophryniscinae
Genyopiiryninae .

Engystoioatinae .

Dyscophinae

Dendrobatinae

-f-

Raninae

-\-

Ceratobatrachinae

+ !

1 signifies the occurrence of only one species of an elsewhere numerouis group.
X ManUlla, cf. p. 71 and p. 272.

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CHAPTER IV

STEGOCEPHALI OR LABYRINTHODONTS LISSAMPHIBIA APODA

Sub-Glass L STEGOCEPHALI or PHRAGT AMPHIBIA

With a considerable aiiwunt of denned armour, especially
on the head.

The earliest known terrestrial four-footed creatures occui- in the
Carboniferous strata of Europe and North America. They and
their immediate allies, which extend through the Permian into
the Upper Trias, are now comprised under the name of Stego-
CEPHALi, so called because the whole of the dorsal side of the
cranium is covered, or roofed over, by dermal bones (<rT€7o?, roof;
Ke^aXrjy head). That these creatures, of which naturally only
the skeletal parts are known, were not fishes, is shown by
the typically pentadactyloid limbs; but to recognise them
as Amphibia, and as distinct from Reptiles, is difficult, especially
if the incipient Reptilia, which have sprung from some mem-
bers of this Stegocephalous stock, are taken into account.
However, they possess either two occipital condyles, or none, and
their vertebrae are either pseudocentrous or notocentrous, but not
gastrocentrous. Moreover, the whole skeletal organisation is still
so ideally generalised, that it is easy to derive directly from it
the arrangement prevailing in the Apoda and Urodela.

The vertebral column always comprises a well - developed,
sometimes a very long tail. The vertebrae exhibit three t}T>es,
two of which are fundamentally distinct, while the third is a
further development of the second.

1. Lepospondylous and pseiuiocentrous. — The vertebra con-
sists of a thin shell of bone surrounding the chorda dorsalis, and
is composed of two pairs of arcualia, which meet each other.

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CHAP. IV STEGOCEPHALI 79

forming a suture, along the lateral side of the vertebra, both
partaking in the formation of a transverse process which carries
the rib.

2a. Temnospondi/lovs. — The vertebra is composed of three
pairs of units, which remain in a separate, unfused state.
Two of them are dorsal arcualia, one of which tends to form the
centrum of the vertebra, which then carries the neural arch.

2b. Stereospondylous. — The three component units fuse by
co-ossification into a solid, amphicoelous vertebra.

The ribs are one- or two-headed, rather strong, but short,
rarely reaching half-way round the body. They occur on all the
vertebrae of the trunk and on most of those of the tail. One
pair of ribs connects one vertebra, the sacral, with the pelvis, of
which the ilium and ischium are generally ossified, rarely also
a portion of the pubic region.

The shoulder-girdle is very primitive, greatly resembling that of
the Crossopterygian fishes. It consists of the following bones : —
a median, rhombic, or T-shaped interclavicle, a pair of clavicles,
of cleithra, of coracoids, and of scapulae. The limbs show the
typical pentadactyle plan, but even in these earliest Tetrapoda
the hand possesses only four fingers, with 2, 2, 3, 2 phalanges
respectively. The foot has five toes, with 2, 3, 4, 4, 3, or
2, 2, 3, 4, 3 phalanges.

Many Stegocephali were possessed of a dermal armour, covering
either the whole body or only the under parts. Hence the term
Phractamphibia (^pa/cT09, armoured). The armour consists of a
great number of small cutaneous scales, partly ailcified, or
perhaps ossified, and arranged in many more or less transverse
rows. We can only surmise that these scales were covered by
corresponding epidermal sheaths. The skull is ideally complete
in the number of separate bones which appear on its surface.
Besides the outer nares and the orbits there is always an
impaired, small, interparietal foramen. The whole temporal
region is completely roofed over. The following bones are
present : — nasals, frontals, parietals, supra- and latero-occipitals ;
lacrymals ^unless fused with the jugals ?), prefrontals, postfrontals,
postorbitals, squamosals, and epi- (or opisth-) otics ; premaxillaries,
maxillaries, jugals, quadrato-jugals, and supra-teraporals ; quad-
rates, pterygoids, palatines, vomers, and an unpaired para-
sphenoid. — The lower jaw is composed of a pair of dentaries,

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STEGOCEPHALI

articulars, angulars, and splenials. The dentaries and apparently
sometimes the splenials, the palatines, maxillae, and vomers cany
teeth. The eyes possess a ring of sclerotic bones.

Order L STEOOOEPHAU LEPOSPONDYLI.

Vertebrae pseudocentrous.

Sub-Order 1. Branchiosauri — The young had several pairs
of gill-arches, whicli, to judge from their size and from the

Fig. 12. — A, Dorsal and B, ventral views of the cranium of limnchiosaurus salaman-
droide^j x abont 4. (After Fritscli.) C, Posterior view of the cranium of Tremnto-
saunisj X about h. (Aft«r Fraas.) /ir, Branchial arches : C, condyle ; Epy epiolic ;
Fy frontal ; J, jugal ; L.O, lateral occipital ; J/, maxillary ; iV, na>al ; iVc, nostril;
Pa, parietal ; P/, palatine ; Pm, premaxillary ; /'.<>, postorbital ; J*rf, ))refrontal ;
7*5, parasphenoid ; Pt, pterygoid ; Ptf, postfrontal ; Qy quadrate ; f,>/, quadrato-jugal ;
S.n^ snpraoccipital ; ^Sy, squamosal ; »S/, supratemporal ; T, vomer.

fact tliHt they are beset with numerous nodules, denticles, or
irregular little processes like gill-rakers — seem to have been
exposed to the surface and to have carried gills. In the adult
the arches and gills seem to be absent.

One of the commonest genera is Braiichiosdurus, including
Pi'otriton. B. salamandroides of the Lower Red Sandstone of
Europe is known in every stage, from larvae of 16 mm. to the
full grown animal of 64 mm. in length. The whole body was

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IV BRANCHIOSAURI ^AISTOPODES 8 J

covered with little cutaneous scales. Pelosaurus and perhaps
Mela7ierpeton are allied genera.

The following genera are small newt-like creatures of the
Carboniferous age of Europe and North America. In Keraterpeton
of Bohemia, Ireland, and Ohio, the dermal scales were restricted to
the under parts ; and the ribs were rather long, reaching halfway
round the body. Gills have not been observed. K. crassum, a
Eui'opean species, reached more than one foot in length, two-thirds
of which fall to the tail. The ventral side is covered with a most
elaborate armour, which consists of about eighty chevron-shaped
rows of little scale-shaped nodules. The epiotic bones end in
strange processes, carrying a pair of spikes, giving the skull a
" homed " appearance, hence the generic name. Urocordylus is
an allied genu&

Sub-Order 2. Aistopodes. — Body snake-like and without
any limbs, hence the name atorov, unseen ; ribs long, and
reaching half way round the body ; from Carboniferous strata in
Ireland and Bohemia, with allied, or perhaps identical forms in
Ohio. Dolichosoma longissimum possessed more than 150 verte-
brae, and was about a yard long. The epiotics end in obtuse
projections, recalling those of Keraterpeton. These marvellous
creatures had strange appendages, extending from behind the
sides of the head, which were possibly the supports of external
gills ; since the upper end of one of the visceral arches, probably
the hyoidean arch, is attached to the labyrinth ic region, and from
this arch starts a bony rod which carries long skeletal filaments.
The body seems to have been naked.

Ophiderpeton had a compound ventral shield, while the skin
of the back contained granular scutes. Although the Aistopodes
have, not without reason, been looked upon as greatly resembling
the Coeciliae or Apoda in organisation, especially in that of the
vertebral column, the total absence of any other fossils which
might bridge over the enormous gulf between the Coal Age and
recent times, makes the attempt to derive the Apoda from these
creatures very hazardous.

Order n. 8TEOO0EPHALI TEMNOSPONDYLI.

Mostly with rather long ribs and with chiefly ventral
armour.

VOL. VIII (;

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82 STEGOCEPHALI

Chelydosaurtis from the Lower Ked Sandstone of Bohemia was
3 feet long, and possessed a beautiful, complicated, ventral armour,
consisting of about sixty chevron-shaped rows, about three times
as numerous as the vertebrae in the corresponding region.
Sphenomurus from the same strata and localities must have been
2 yards long. The trunk-vertebrae of both these genera were
composed of four pairs of arcualia. Trimerorhachis from the
Permian of Texas is very imperfectly known, but its trunk-
vertebrae, as the name implies, consist of three pairs of separate
arcualia, one of which, the interdorsal pair, tends to form a kind
of centrum.

Dissorophus muUicinctus, also from the Permian of Texas, has
been described by Cope ^ as a " Batrachian Armadillo," and con-
sidered allied to Trimerorhcwhis, Ten vertebrae are known, of
an aggregate length of 93 mm. ; the length of the creature was
perhaps one yard. The neural spines are elevated, and the apex
of each extends in an arch on each side to the ribs. These
spinous branches touch each other, forming a carapace. Above,
and corresponding to each of them, is a similar dermal and
osseous element, which extends from side to side without inter-
ruption in the median line, forming a dermal layer of transverse
bands which correspond to the skeletal carapace beneath it. This
creature remotely approaches the genus Zatod^hys, Cope, where a
dermosteous scute is co-ossified with the apex of the neural
spine. The systematic position of this genus is at least
doubtful.

Archegosaurus decheni from tlie Lower Red of Germany,
known by many well-pi eserved specimens, reached a length of 4
or 5 feet. Tlie trunk vertebrae are tri-partite, those of the tail
(luadri-partite, like those of tlie trunk of Chelydosawnis, Young
specimens show traces of gill-arches. The thoroughly terrestrial
walking limbs have four fingers and four toes ; the arrangement
of the tarsalia, most of which are ossified, lend support to the
view that the morphological axis went through femur, fibuUi,
intermedium, the centralia, the second distal tarsale, and the
second toe. The dentine and enamel of the teeth are much
folded, and this feature, which applies to most members of this
Order, to a lesser degree also to others, has caused them to be
comprised under the name of Labyrinthodonta. The upper

^ Amcr. Natural, xxix. 1895, p. 998.

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IV LABYRINTHODONTA 83

surface of the head shows very characteristically arranged
grooves, which probably contained slime - canals and possibly
sensory organs,

Actinodon and HiLchirosaurus are closely allied forms, chiefly
from the Lower Eed Sandstone of France ; GondwaTWsaurus occurs
in the Permian of India.

Order UL STEOOOEPHALI STEREOSPONDYLI.

These are the most highly developed members of the typical
Labyrinthodonta, characterised by their much-folded teeth, and
by their solid, bi-concave vertebrae. Loxomma occurs in the Upper
Carboniferous of England and in the Lower Eed of Bohemia :
Trematosaurus, Capitosaurus, and Metopias from the New Red or
Lower Trias to the Keuper of Grermany. Mastodonsaurus from
the Trias of England and Germany is the most gigantic
Amphibian known, with a skull of nearly 1 yard in length.

Lahyrinthodon from the Keuper of Warwickshire is one of
the latest members of the group. Labyrinthodont creatures have
also been described from the Trias of South Africa, e.g. Rhyti-
dosteus ; those from North America are insufficiently preserved.

Many of these and allied genera have left their footprints in
slabs of Sandstone, both Lower and New Red, in Europe, Africa,
and America. But although their spoors are common enough,
only a few can with certainty be referred to Stegocephali, e.g.
Sauriehnites salamandroides of the Lower Red of Germany.
The spoors of Chirotherium, common in the New Red of
Germany and England, for instance in Cheshire, belong to
unknown owners; both the large hind feet (which measure
nearly half a foot in length) and the much smaller fore feet, had
five digits, the first of which stood off like a thumb. Five-
fingered Stegocephali are unknown.

There is an almost complete absence of fossil Amphibia from
the Upper Trias to the Oligocene. The Stegocephali as such
seem to have died out with the Trias. The recent Amphibia, of
course, must have had ancestors in the Mesozoic age. There is
one little skeleton, from the Wealden of Belgium, which belonged
to a newt-like creature, called Hylaeohatrachus croyi. Scarce
fragments, described as Megalotriton, are known from the Oligo-
cene of France, and Triton itself seems to be indicated by

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84 APOD A OR COECILIAE chap.

remnants in the Lower Miocene of France and Germany. But
fairly complete specimens of large creatures, much resembling
CryptobranchuSy have been found in the Upper Miocene of
Oeningen, Canton Solothum, Switzerland. The first known
specimen, now at Haarlem, indicating a total length of 3 feet or
more, was described and figured in the year 1726 by Scheuchzer,
in a learned dissertation entitled " Homo diluvii testis"

Betriibtes Beingeriist von einem alten. Siinder
Erweiche Herz und Sinn der neuen Bosheitekinder.

Which may be rendered as follows : —

Oh, sad remains of bone, frame of poor Man of sin
Soften the heart and mind of sinful recent kin.

This was the motto attached to the illustration, and it remained
a warning to mankind until Cuvier declared the skeleton to be
that of some large newt. Tschudi named it Aifidrias schewchzeri,
but it is scarcely generically distinct from Cryptohranchvs, being
almost intermediate between C. cdleghanieTisis and C. japoniciis,
see p. 97.

Sub^Cla ss IL LISSAMPHIBIA.
Amj>hibia loithout dermal armour.

Order I. APODA or LIMBLESS AMPHIBIA.

The Amphibia Apoda, Coeciliae or Gymnophiona, are a small
group of worm-shaped, burrowing creatures, restricted to the
Xeotropical and Palaeotropical regions, excluding Madagascar.
Tliey have no limbs and no girdles. The tail is extremely
short ; the vertebrae are pseudo-centrous, and most of them
carry rather long ribs, none of which, however, meet to form a
sternum. The whole snake-like body is covered with a smooth
and slimy skin which forms numerous transverse folds or rings.

The most remarkable feature of the skull is its solid com-
pactness, wliich stands in direct correlation with the burrowing
liabits of these creatures. The whole dorsal surface of the
cranium is practically roofed in by bone, so that, in this respect,
it gieiitly resembles tliat of the Stegocephali ; but this resemblance
is pnnluced chietiy by a broadening of those bones which exist

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GENERAL ANATOMY

also in the other Lissamphibia, while supratemporals and supra-
occipitals are absent. There is, however, a pair of bones whicli
represent either the postorbitals or the postfrontals, perhaps
both, of the Stegocephali The qnadrato-jugal arch is enormously
developed, and by reaching the parietal, frontal, and postorbito-
frontal bones (which latter occur only in Ichthyophis and
Uraeotyphlvs) and the maxilla, extends over the whole of the
orbito- temporal fossa. The squamosal is completely fused with

Pig. 13. — ^v\\ o{ Tchtkyophia glutinos<i, x 8. (After Sarasin.) A, Lateral, B, veutral,
C, dorsal view. A, Posterior jirocess of the os articnlare ; Co, carotid foramen ; C'A,
choana or posterior nasal opening ; Fy frontal ; ./, jugal ; Lo^ lateral occipilal ;
Mx^ maxillary ; N^ nasal ; No, nostril ; O, orbit ; /', parietal ; Pw, palatine : P//<,
premaxiUary ; Pofy postfh>nta] ; Prfy prefrontal ; P/, pterygoid ; Q, quadrate ; *V,
squamosal ; SI, stapes ; 7*, tentacular groove ; To, vomer ; A", exit of vagus nerve.

the quadrato-jugal. The stapes has the typical stirrup-shape, is
even perforated by an artery, and articulates distally with the
shaft of the quadrate (as in the snakes). The maxilla is very
large and broad. Owing to its broad junction with the quadrato-
jugal arch, the prefrontal and frontal, the or>)ital fossa is reduced
to a very small hole, or the maxilla completely covers tlie eye.
Somewhere between the latter and tlie nares the maxilla is
perforated by the tentacular groove. The periotic bones are
represented by the prootics ' and epiotics ; they fuse with
the lateral occipitals and with the parasphenoid. The whole

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86 APODA CHAP.

orbito-ethmoidal region of the primordial skull is also turned,
into one mass of bone.

The angular element of the lower jaw forms a thick and
large process which projects upwards and backwards from the
mandibular joint. The former possession of a splenial bone
is indicated by the occurrence of a second series of teeth in
the mandibles of Ichthyophis and Uraeotyplihis. Other genera
have vestiges of this second row, or it may be completely
lost.

The hyoid and branchial apparatus is more primitive than
in any other recent Amphibia. In the larva the hyoid and the
first and second branchial arches are connected with each other
by a median copular piece. The third branchial arches are free
from the rest, but are fused in the middle line, the fourth are
loosely attached to the previous pair. In the adult both fuse
into one transverse, curved bar, and the second pair of branchials
lose their connexion with the basal longitudinal piece and likewise
form a transverse bar.

The vertebrae are built upon the pseudocentrous type, are
amphicoelous, and the chorda is intravertebrally destroyed by
cartilage, as in the majority of the Urodela. The number of
vertebrae is great, amounting in some species to between 200
and 300, of wliich a few belong to the tail. The first vertebra
is devoid of an odontoid process. The ribs are proximally
])ifureated as in the Urodela.

The eyes are practically useless, being either more or
less concealed under the skin, or they are covered by the
maxillar}^ bones. All Coecilians possess a peculiar tentacular
sensory apparatus, which consists of a conical flap-shaped or
globular soft tentacle, which is lodged in a special groove or
canal of tlie maxilla, between the eye and the nose, whence it is
frequently protruded while the animal is crawling about. These
tentacles in the young iHphoiiops lie, according to the Sarasins,
(^uite close to the eyes, but are later transferred nearer to the
nose. The organ consists of a peculiarly rolled up and pointed
fold which arises from the bottom of the sac or pit, where it
receives a nerve. It is protruded by becoming turgid with
l)lood, and is retracted by a strong muscle. Into the lumen of
the sac are poured secretions from the large orbital (Harderian)
gland, to keep the apparatus clean. Hence arose the mistaken

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IV GENERAL ANATOMY 87

notion of its being, a poison -organ. The whole structure is
possibly an offshoot of the naso-laerymal duct.

The skin is mpst remarkable. In the ripe embryo the
epidermis passes smoothly over the surface. Beneath follow two
layers of soft cutaneous connective tissue, bound t<^ether by
transverse or vertical lamellae, so that ring-shaped compartments
are formed, and in these are embedded slime -glands. In the
adult each compartment is modified into an anterior glandular
belt and a posterior space, from the bottom of which grow
several scales. The number of cutaneous rings agrees originally
\nth that of the vertebrae; but later, and especially in the
hinder portion of the trunk, each ring breaks up into two or
more secondary segments, and these no longer agree with those
of the skeleton. Each saile is beset with numerous smaller
scales which consist of hardened cell-secretions infiltrated with
calcareous matter. The whole scale is consequently an entirely
mesodermal product of the deeper layers of the cutis. The
usual statement that the skin forms imbricating lamellae, on
the inner side of which appear the scales, is wrong. The
'• lamellae " can be lifted up only after the general epidermal
sheath has been broken artificially in the constrictions between
the rings. No scales exist in the Indian genus Gegenophis and
in the American Sijphono'ps, Typhlonectes, and Chthonerpeton, a
secondary loss which does not indicate relationship. The scales
develop lat« in embrj'onic life, and they are reasonably looked
upon as inheritances from the Stegocepliali. The glands either
produce slime, whose function seems to be the keeping clean of
the surface of the body, or they are squirt-glands. Tlie latter
kind are also numerous and are filled with a fluid which is squeezed
out by muscular contraction, and seems to ])e poisonous, as it
causes sneezing to those who handle or dissect fresh si)ecimens.

The Coecilians live in moist ground and lead a burrowing
life. Their developmental history has only recently been studied,
and in but a few species, see IchtJuj aphis, p. 91, and Hi/pogeojyJns,
p. 92. The female is fertilised internally, copulation taking place
by means of eversion of the cloacal walls in the shape of a tube.
The spermatozoa possess an undulating membrane ; the eggs
undergo meroblastic division and the embryos have three pairs
of long external gills. Some are viviparous.

The snake-like, limbless shape of the body (Fig. 15) is, as in

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88 APODA CHAP.

snakes, correlated with an asymmetrical development of the
lungs ; the left is reduced, while the right is drawn out into a
long cylindrical sac The liver is likewise very long, and partly
constricted into a great number of lobes. Owing to the great
reduction of the ribs progression is eflTected in an almost earth-
worm-like fashion by the peristaltic motion of the skin, assisted
by its numerous ring-shaped constrictions.

The systematic position of the Coeciliae has been, and is
still, a controversial matter. The Sarasins took up Copes
suggestion, that their nearest allies are the Urodela, especially
Amphimna, and they went so far as to look upon Amphiuma as
a neotenic form of the " Coecilioidea," which they divided into
Amphiumidae and Coeciliidae; the Coecilioidea and Salaman-
droidea forming the two su})-order8 of the Urodela. They based
this startling conclusion chiefly upon remarkable resemblances
between Amphiuvia and Ichthyophis, namely, (1) the mode of
lajdng the eggs on land and coiling themselves around them ;
(2) the existence of remnants of a tentacular apparatus in
Amphiuma ; (3) Coi)e'8 statement that Amphiuvia alone among
the Urodela possesses an ethmoid like the Coeciliae. This latter
point is, however, erroneous ; it has since been shown by Davison '
that Amphiuma possesses no ethmoid bone, but that, instead of it,
descending plates of the frontals join below the premaxilla and
function as a nasal septum, with a canal for the olfactory nerves.

We look upon the Apoda with more reason as creatures
which of all the Lissamphibia have retained most Stegocephalous
characters and at the same time form a highly specialised group
equivalent to the iri-odela and the Anura. The following are
Stegocephalous inheritances peculiar to the Apoda in opposition
to the other recent Amphibia : retention of cutaneous scales
with calcareous incrustations, greatly resembling the scales of
the Carboniferous Microsauri ; occasional retention of post-
frontal and lateral nasal or lacrymal bones, and of a second row
of teeth in the mandible. To these may be added the presence of
epiotic bones, and the primitive character of the branchial aix'hes.
The loss of all these characters would turn the present Apoda
into limbless I'^rodela, but this assumption does not justify their
inclusion in this Order. The possible homology of the tentacular
apparatus has been discussed elsewhere, p. 45.

' J. Morphol. xi, 1895, p 375.

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COECILIIDAE 89

Fossil Apoda are not known ; their subterranean, life does
not favour preservation.

Only family, Ooeciliidae. About forty species are known.
These have been placed in seventeen genera, mostly on com-
paratively slight grounds, and several of these genera are probably

■ICOECILIAC.

Fio. 14.— Map showing the distribution of the Coeciliae or Amphibia Apodiu

unnatural, the distinctive characters having undoubtedly been
developed independently in various countries. We have to
remember that the recent species are the remainder of a formerly
much more numerous group ; it is also likely that more will be
discovered in the tropical forests of South America and Sumatra.
Boulenger ^ has distinguished them as follows : —

I. Cycloid scales embedded in the skin.
A. Eyes distinct, or concealed under the skin,
a. Two series of teeth in the lower jaw.

a. Quadrato-jugal (squamosal) and parietal bones in contact.
Tentacle between eye and nostril.

Ickthyophis, 2 species, India and Malay islands, p. 90.
„ below and behind nostril.

Hifpogeophisy 3 species. East Africa and Seychelles, p. 92.
„ below and in front of eye.

Dermophdsy 5 species, America and Africa, j). 93.
. „ below the nostril. Coeciliay 6 species, America.
^, Qnadrato-jugal separated from parietal.

Tentacle close to the eye. Bhinatremoy 2 species, America.
„ below and behind nostril.

Geotrypetes, 1 species, West Africa.
„ below nostril.

Uraeotyphlusy 3 species. West Africa and India.
h. One series of teeth in the lower jaw.
Tentacle in front of the eye.

CryptopsopkiSf 1 species, Seychelles.

1 P. Z, S. 1896, p. 401.

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90 APODA CHAP.

B, Eyes below the cranial bones. Quadrato-jugal in' contact with
parietal.

Tentacle near the nostril.

Gymnophis, 4 species, South America.
Herpele, 2 species, Panama and Gaboon.
II. Without scales.

A, Eyes distinct, or concealed under the skin. •
a. Two series of teeth in the lower jaw.

a. Quadrato-jugal in contact with parietal.

Tentacle behind nostril ; end of body laterally compressed.

Typhlonectes, 3 species, America, p. 93.
/^. Quadrato-jugal separated from parietal.
Tentacle between eye and nostril.

Chthonetyetoyt, 2 species, America.
6. One series of teeth.

u. Quadrato-jugal and parietal in contact ; tentacle in front of the

eye .... SijJwnops, 4 species, America.

/?. Quadrato-jugal 8ej)arated from parietal.

Bdellophis, 1 species, East Africa.

B. Eyes below the cranial bones.

a. Two series of teeth. Quadrato-jugal and parietal in contact;
tentacle behind and below nostril.

Gegenophisy 1 species, India.

b. One series of teeth. Quadrato-jugal separated from parietal.

Scolecomorphus, 1 species, East Africa.
BouU?}genda, 1 species. East Africa.

Ichthyopkis glntinosa extends from the slopes of the Hima-
layas to Ceylon, the Malay islands, and into Siam. A second
species, /. monochrous, occurs in Malabar, Malacca, Borneo, and
Java. /. glutitiosa reaches about one foot in length, with a
greatest thickness of a little more than half an inch. Tlie
general colour is dark brown or bluish black, with a yellow band
along each side of the body.

This species has been studied extensively by the Sarasins.^
It breeds in Ceylon after the spring monsoon. The ovarian egg
is oval, measuring 9 by 6 mm. The yolk is yellow ; the
blastoderm lies towards one of the poles. The strong vitelline
membrane l)ecomes surrounded in the oviduct by a dense
albuminous membrane, which forms twisted chalazae, just like
those of birds' eggs, and by these two cords the eggs are strung
together. Around all this lies another mantle of albumen.
The female digs a hole close to the surface in moist ground near

* P. and F. Sarasiu, " Zur Entwicklungsgeschichte der ceylonesischen Blind-
wiihle, IeJithyoj)hw glutinosa,'' Ergchnisse natunciss. Forschungen an/ Ceylon,
1887-1890, vol. ii.

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COECILITDAE

running water, and there lays about two dozen eggs. The egg-
strings become glued together, entangled into a bunch, and the
female coils herself round the bunch and remains in that posi-
tion, probably to protect the eggs against other burrowing
creatures, as blind snakes (Ti/phlop$ and Hhinophis) and certain
limbless lizards, with which the ground literally swarms. During
this kind of incubation the eggs assume a round shape, and grow
to twice their original size, and the mature embiyo weighs four
times as much as the newly laid egg.

The external gills are delicately fringed and red, and they
mov^e up and down in the fluid of the egg. The body of the

Fk;. 1 iK—Ichthifophis ghUinom x 1 . (After P. ami F. Sarasin. ) 1 , A nearly ripe embryo,
with gills, tail-tiu, and still with a considerable amount of yolk ; 2, female guard-
ing her eggs, coiled up in a hole underground ; 3, a bunch of newlv laid eggs ;
4. a single egg, enlarged, schematised to show the twisted albuminous strings or
chalazae within the outer membrane, which surrounds the white of the egg.

embryo is at first white, but becomes pigmented with dark grey,
A strong line of lateral sense-organs is formed, and a ring of tliem
lies around the eye and others on other parts of the head. The
short tail develops a fin. Of tlie three pairs of gills the third is
the shortest, and is generally turned dorsalwards. In embryos
of 4 cm. in length the longest gill measures as much as 2 cm.
Yolk is still present in embryos which have reached the
surprising length of 7 cm. Then the gills begin to shrink a
little, and at this time one pair of gill-clefts breaks through at
the base of the third external gill.

When the larvae are hatched the gills are lost. The young
larva takes to the water in a gill-less state, and moves about
like an eel. At the bottom of the gill-hole on each side two
arches are visible, and there are at this statue neither inner nor

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92 APODA CHAP.

outer gills. The larvae frequently come up to the surface to
breathe. The eyes are large and clearly visible, but the tentacles
are still undeveloped The epidermal sense-organs are numerous,
and appear as white spots in the grey skin ; about fifty extend
from the gill-opening to the tip of the tail.

Iclithyojphis seems to live a long time in the larval state. At
last the gill-clefts close, the tail-fin disappears, and the tentacles
come to the surface. The whole skin assumes a totally new
structure, and the fish-like larva turns into a burrowing, sub-
terranean creature so teiTestrial that it gets drowned when made
to remain in the water.

Hypogeojyhis. — According to A. Brauer^ three species of
Coecilians are found in the Seychelles: Cryptopsophis multipU-
ccUuSy which is rare, Hypogeophis rostratns and ff. altemajis.
They live in moist ground, near the coast in swamps, higher up
in humus, under rotten trees and rocks, down to the depth of
one foot. In the island of Silhouette, Brauer found them in
brooks, at least during the dry season, from May to September.
The natives call them " vers de terre." They seem to propagate
during the greater part of the year, provided there is sufficient
moisture. The female coils round the eggs, which vary from
half a dozen to thirty in number, those of H. rostratus measuring
7-8 mm., those of H, alterrians only 4-5 mm.

The embryos undergo their whole development in the egg.
Four pairs of gill-clefts break through, the first between the
hyoid and the first branchial arch, the fourth between the third
and fourth branchial arches. There appears also a spiracular
cleft between the quadrate and the hyoid arch ; this cleft is,
however, only developed dorsally, and persists for a shorter time.
The external gills appear at the same time as the clefts, upon
the first three branchial arches ; the third gill is the latest, and
remains in a vestigial condition covered up by the two others.
Tlie gills, of which the second is the longest, are not (as stated
by the Sardsins) direct prolongations of the gill-arches, but they
begin as button -Uke growths upon the arches. They begin
to disappear with the absorption of the yolk, getting actually
smaller. In embryos of 6 cm. they are 6 mm. long, while in
embryos of 6.5 cm. they are reduced to 4.5 mm. in length. The

" Beitrage zur Kenntniss der Kntwicklungsgeschichte und der Anatomie der
ymnophionen," ^ool. JcUirh, JiuiL x, 1897, ji. 389, and xii. 1899, p. 477.

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COECILIIDAE 93

first to disappear is the third gill, of course by being resorbed ;
and the clefts are closed before the creature leaves the egg. Hypo-
gtophis not leading an aquatic larval life possesses no tail-fin in
the embryonic state, the gill-holes are closed, and the epidermal
sensory organs disappear long before the time of hatching.

Vestiges of gills appear also on the hyoid and on the man-
dibular arch, but on the latter they are of very short duration.
Those of the hyoid gradually fuse with the first of the branchial
gills, and these also concentrate with their bases so that they
ultimately seem to spring from one common stem. Brauer
remarks that the distinction between internal and external gills
seems to be one of degree only ; the hyoidean and mandibular
gills namely start from the hinder margin of the arches, just
like the internal gills of Torpedo according to Ziegler, while the
other gills start from the sides of the branchial arches. He also
found a pair of little swellings behind the last -gill-cleft, and an
unpaired swelling (corresponding with a double one in Ichthyophis)
in front of the vent. Not unreasonably he sees in these swellings
the last, very transitional vestiges of the paired limbs.

Typhlonectes compressicauda of Guiana and Venezuela is one
of the largest Coecilians, reaching a length of 18 inches, with a
body -diameter of ^ inch. The general colour, as in most of these
creatures, is olive brown to black. A sort of adhesive disc sur-
rounding the vent occurs in this genus. Peters, who described
this species, found in one female six embryos of comparatively
enormous size, one of them being 157 mm. (more than 6 inches)
Jong, and 12 mm. thick, and devoid of a tail-fin. Instead of
lateral gill-openings there is a " bag " on each side 5 5 mm. long,
upon which is distributed a blood-vessel. The Sarasins have
examined the same specimen : The gills are not a bag, but con-
sist of two flat, unbroken membranes which are closely connected
with each other. In fact the outer gills of all Amphibia may be
said to begin in the shape of small bags, whence sprout
secondarily the gill- fringes ; but in Typhlonectes they form these
flaps instead of growing into the usual three gills. The embryos
have no epidermal sense-organs, but plenty of skin-glands. Prob-
ably when born they take at once to terrestrial life, the flaps are
possibly shed at birth, and there remains a little cicatrix.

Dermophis thomensis of West Africa (its other relations live
in East Africa, South and Central America) is also viviparous.

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CHAPTER V

LISSAMPHIBIA {COXTIN UE D)—VB.0J)ELA

Order n. URODELA or TAILED AHPHIBIA.

The recent tailed Amphibia, Salamanders and Newts in the wider
sense, have been grouped into four families which can be con-
veniently diagnosed by the following characters : —

Both the upper and lower jaws are furnished with teeth. Fore- and hind-
limli are always pi-esent
Maxillary bones present.

Eyes free and devoid of lids . Amfhiumidae, p. 97.

Eyes with movable lids^ Salamandridae, p. 102.

Maxillary bones absent

Eyes without lids. Perennibranchiate Proteidae, p. 132.
Both jaws are toothless. The hind-limbs, the maxillary bones and eyelids
are absent Perennibranchiate . . Sirenidab, p. 136.

These four families are closely allied to each other, especially the
Amphiumidae and the Salamandridae.

The geographical distribution of the Urodela is essentially
Periarctic, except that about one dozen species each of Ainhlystoina
and of Spelerpes extend southwards into Central America, and in
the case of the latter genus even into the Andesian parts of
South America, Pletiwdon platense inhabits Argentina.

The Urodela afford good reasons for dividing the Periarctic
region into three co-ordinate sub-regions, namely, Nearctic,
Eastern and Western Palaearctic. The difiference between the
European and the Eastern Asiatic fauna is well marked ; the two
are — at least witli our present knowledge — separated by a wide
stretch of country very poor in I'rodele forms; while, lastly.

' The existence of such a form as Typhi otriton, in the adult of which the eyes
become closed up, makes such short diagnoses of the families defective, although
there is no doubt about the Desmognathine afliuities of this genus. See p. 103.

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GEOGRAPHICAL DISTRIBUTION

there are not a few resemblances between this Eastern Asiatic
and the American fauna. The Urodela thus lend no support to the
usual division of the Periarctic into a Palaearctic and a Nearctic
sub-region. Nor is it possible to divide the Palaearctic into a
Eurasian and a Mediterranean province. We have in this case
to distinguish between an American, an Asiatic, and a European

r^SALAM. LECHRIODONTA.

Fig. 16.

\ S. MECODOMTA . .$^^ ICHTH VOOEA.

Map showing the distribution of the Urodela. " Ichthyodea ' ' — Amphiumidae
4- Proteidae + Siren idae.

fauna. The Asiatic or Eastern I^alaearctic sub-region assumes the
central position, at least from a merely geographical point of view.
Tt would be imjustifiable to assume a spreading from this centre
into Europe, and, on the other hand, into America. The centre
existed more probably in the Arctic circle, now devoid of Urodela.

So far as mere numbers of species are concerned the huge
Asiatic or Eastern Palaearctic region is the poorest, but it is
also the least explored, and China will probably yield a good
many new forms. We know at present only 15 species, nearly
all from the eastern half These 15 species represent no less
than 11 genera, 8 of which (=73 per cent) are peculiar to the
sub-region. Next comes the Western Palaearctic or European
sub-region with about 21 recent species of 5 genera, 4 of which
are peculiar. America is by far the richest, with no less than
66 species (36 eastern, about 16 western, and the rest Central
American, etc), belonging to 19 genera, 17 of which ( = 90 per
cent) are peculiar to the New World. But this richness in species
is due mainly to the abundance of the two genera Amhly stoma and
Spelerpes, just as Europe is characterised by its many Tritons.

One of the most striking features of the Asiatic sub-region is

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URODELA

its difference from the European. They have verj little in
common. Pachytriton, Tylototriton, and two species of Triton
(T. pyrrhogaster djiA T. sinensis) are the only Salamandrinae, while
all the rest are Lechriodont (see p. 102), like the American

(lEOGRAPHICAL DISTRIBUTION OF THE URODELA

Westeni PaUearctic.

Sirenidat'
Proteidae
Amphiiimidae

6 S

^6"

Salamandridae \

Eafltern Falanirctic.

1 Proteus

(1 AndriaSy Miocene)

1 Speler]>e.s

14 Triton
1 Salamaudriua
1 Chioglossa
3 Salamandra

21 species, 6 genera

1 CryptobranchuB

American.

1 Ambly stoma

1 Batrachy|)erus
1 Ranidens
1 Geoniolge

1 OnychodactvluR

2 Salaniandrella

3 Hynobius

1 Pachytriton

1 Tylototriton

2 Triton

1 Siren

1 Pseudobnnchus

1 Necturus
1 Typhlomolge

1 Amphiuma

1 Cryptobranchus

1 Thorius

1 Haptoglossa

3 Desmognathus

21 Snelerpes

2 Manculus
7 Plethodon

3 Batrachoseps

1 Typhlotriton

2 Autodas

16 Ambly stoma
1 Diuamptodon

2 Triton

15 species, 11 genera 66 species, 18 genera

! I

Urodela, excepting tlie two American Tritons, T, torosus and
2\ vlrldescens. The occurrence of an Amhly stoma, A. persimilf,
in the mountains of Siam and Burmah, is most suggestive, and
thers will in all probability be found. It must also be borne in

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AMPHIUMIDAE 97

mind that the diflferences between the genera of Amblystomatinae
are in reality very slight ; and the same applies to the sub-
families themselves. The presence or absence of teeth on the
parasphenoid, the possession of amphi- or opistho - coelous
vertebrae, do not mean much, and certainly does not forbid the
notion that all the recent Urodela are the offspring of one common
generalised stock which inhabited the northern portion of the
globe. Xothing is gained by hiding the solitary European species
of the essentially American genus Spelerpes under the name of
Geotriton. It is a Spelerpes in all characteristic points. Speaking
broadly, each of the three principal sub-families of Salamandridae
is characteristic of a sub-region; the Salamandrinae of the Western
Palaearctic, the Plethodontinae of the American, while the Ambly-
stomatinae are chiefly Asiatic, at least so far as diversity of
genera is concerned.

Fam. 1. Amphiuxiiidae. — Without gills in the perfect state.
The gill-clefts are in a vanishing stage, being either reduced to
one pair of small holes or being altogether absent. The maxillary
bones are present. Teeth occur in both jaws ; those of the
vomers form transverse rows. The vertebrae are amphicoelous.
The fore-limbs and hind-limbs are present, but small. The small
eyes are devoid of lids.

This family is now represented by two genera, with only three
species, found in the United States and in Eastern Asia.

Crtfptobranchus. — The limbs are fimctional, with four fingers
and five toes. The outer digits and the sides of the limbs are
bordered with folds of skin. The head and body are stout and
depressed ; the tail is short, laterally compressed, and provided
with a fin. The skin is very glandular and slimy, and forms a
thick, irregularly-shaped fold along the side of the body.

C. {ifenopoma) cUleghaniensis, — The gill-clefts are normally
reduced to one pair, individually to the left cleft, the right
closing up. There are, however, four branchial arches and vessels.
The general colour is brown or grey above, sometimes with
darker patches, lighter below. The " Hellbender " readies a
length of nearly 18 inches (about 46 cm.), is entirely aquatic,
and is apparently restricted to the rivers and streams of the
mountainous districts of the Eastern United States. It is very
voracious, living on worms and on fish, being much disliked by
the fishermen, as it takes the angler's bait, and destroys great

VOL. VIII II

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CHAP. V AMPHIUMIDAE 99

quantities of the valuable food-fish Coregonv^ alhus. Although
rather common and easily kept, its larvae still remain un-
known.

C. japonwus s. maximus. — The Giant Salamander of Japan
differs from its American relation in one essential point only,
namely, by the absence of gill-openings and of the modifications
of the branchial apparatus connected therewith. It has but
three branchial vessels, and the skeletal arches are reduced to
two. It lives in Japan and in China, from 600 to 4500 feet
above the level of the sea, in small streams of mountain-meadows.
It feeds upon fishes. Amphibia, worms, and insects. It is easily
fished with the hook and is eaten by the Japanese.

The first living specimen was brought to Europe in 1829 by
Th. von Siebold, its discoverer. It grew within a few years from
1 foot to 3 feet in length, and died in 1881, at least fifty- two
years old Another specimen lived in the Hamburg aquarium
for fourteen years, during which time it is said to have grown
36 cm. (more than 14 inches), having attained a length of nearly
4^ feet, or 134 era. The largest specimen known measures 159
cm = 5 feet 3 inches.

The life- history of this species is still imperfectly known.
Japanese picture-books contain drawings of the adult and of
larvae, the latter showing three pairs of fringed external gills.
Young specimens of 16 cm. length have already lost the
gills, but still retain a cleft on either side of the neck, in the
shape of a horizontal slit, and this is soon after closed up by
the skin.

The best account has recently been given by Sasaki.^ Accord-
ing to him the Giant Salamander leads a soUtary life, concealed
in dark places, under rocks in swift-flowing, tliickly shaded small
brooks of clear and cold water.

The animal may be easily captured with a fish-hook, baited
with a fish, frog, or several earth-worms, and tied to a string a
few feet in length. This is thrust by the aid of a small bamboo-
stick into the salamander's retreiit. The string is not tied to
the stick, but the point of the loaded hook is forced into one end
of it, far enough to keep it in place while this end of the rod is
pushed under the rock. When the bait has been tlius brought
near the salamander, any bite will be instantly felt through tlie

^ J. Coll. Japan, i. 18S7. p. *^69.

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I OO URODELA chap.

rod. The latter is then withdrawn as quietly as possible, the
hook and bait being left. As soon as a jerk of the string is
noticed, a pull is made, which generally ends in the capture of
the unfortunate animal. If the first pull should fail, the bait is
replaced as before, and a second opportunity is offered, which the
unwary creature accepts as readily as the first. The fisherman,
having obtained one bite, is sure of ultimate success, as the sala-
mander does not learn by experience to refuse the proffered
morsel. When captured, it emits a peculiar slimy secretion,
having an odour much like that of the leaves of the Japan
pepper {Xanthoxylon peperitum). This secretion hardens into a
gelatinous mass after a short exposure to the air.

Temminck and Schlegel state that the act of inspiration is
ordinarily performed once every 6-10 minutes. This is true
for specimens kept in tubs; but Sasaki is inclined to think
that they perform this act less frequently in their native brooks.
The eyes are so small that they are obviously of little import-
ance ; the salamanders capture their prey not by pursuing, but
by waiting for its near approach, whereupon they seize it with
their teeth by a swift lateral movement of the head. The eggs
are said to be laid in August and September, and they form a
string resembling a rosary. Each egg floats in a clear fluid,
inclosed in a bead-shaped gelatinous envelope, and this is con-
nected with the next by means of a comparatively small string.
The Qy^g measures about 6 mm. by 4 mm., and is yeUow
everywhere except at the upper pole, where it is whitish. All
attempts to make Cryj^tohranchus breed in captivity have failed
liitlierto, owing no doubt to the difficulty of obtaining the cool
temperature of its mountain streams. Sasaki's smallest specimens
measured 19 to 20 cm. These had three pairs of very short
brancliial processes, from 3 to 5 nun. in length, attached just
inside the branchial orifice. Each process was somewhat
flattened and tapering, most of them still with branchlets. In
another specimen, 20*5 cm. in length, the giUs had almost
wholly disappeared, but the branchial slits were still visible.
One of 245 cm. length showed no trace of gills, and the
branchial orifice was completely closed, but still marked by a
light streak.

Amphiuma means s. tri(hwtj/Ia. — The limbs are very much
reduced, and end in two or three little fingers or toes. Just in

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AMPHIUMIDAE

lor

front of the fore-limbs lies the pair of small gill-clefts, each
guarded by two flaps of the skin. There are four branchial arches.
The general colour of this eel-shaped creature is black, lighter
I>elo\v. The head is covered with numerous pores, arranged in
several rows, which imite in the region of the neck, so that only
two rows extend along the sides of the body. It reaches a
length of three feet, and lives in swamps or muddy waters,

FiQ. 18. — Amphiuma viea'ns, x \.

for instance in the ditches of rice-fields, burrowing occasionally in
the mud, feeding on crayfishes, molluscs, small fishes, etc. It is
confined to the south-eastern States of North America, from
Carolina to Mississippi. According to Davison,^ copulation takes
place in May. The rather hard-shelled eggs are deposited in the
following August or September, and are connected by a twisted
conl. The female lies about them in a coil. The embryos, which
are liatched in the month of November or December, have well-
developed external gills. By the following February they have

1 J, Morphol. xi. 1895, p. 375.

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102 URODELA

reached a length of from 68 to 90 ram. (about 3 inches), living
in damp localities mider rocks or rooted stumps, and have already
lost their gills. The legs are said to be relatively longer than
they are in the adult.

Fam. 2. Salamandridae (Salamanders and Newts). — Without
gills in the perfect state. Maxillaries are present. Both jaws are
furnished with teeth. The eyes are protected by movable lids,
except in Typhlotriton. Fore- and hind-lirabs present, although
sometimes very much reduced.

To this family belong by far the greater number of tailed
Amphibia. They have been, for the sake of convenience, grouped
into four sub-families, the determining characters of which are all
internal and of comparatively slight importance. Little better is
the division into Mecodo7ita, with the teeth of the palate in two
longitudinal rows diverging behind and inserted upon the inner
margins of the two palatine processes, which are much prolonged
posteriorly, and Lechriodonta, in which the series of palatal teeth
are restricted to the posterior portion of the vomers and form
either transverse or posteriorly converging rows.

I. Series of palatal teeth transverse, restricted to the posterior portion
of the vomers. Parasphenoid beset with dentigerous plates.
Vertebrae opisthocoelous : Desmognathinaty p. 102.
„ amphicoeloiis : Plethodontinae, p. 103.
II. Series of palatal teeth transverse or posteriorly converging, restricted
to the posterior portion of the vomers. Parasphenoid toothless.
Vertebrae amphicoeloiis : Amblystomatinae, p. 109.
III. Series of palatal teeth in two longitudinal series, diverging behind,
inserted on the inner margin of the long palatine processes.
Parasphenoid toothless. Vertebrae amphicoeloiis: Salaman-
drinae, j). 115.

Sub-Fam. 1. Desmognathinae. — Comprising only three genera,
with five species, in North America. Five toes.

IJesmognathus. — The tongue is attached along the median
line, free behind, oval in shape. Three species in the eastern
half of the United States. D. fuscus is one of the lungless
Urodela, for which condition see p. 46. The skin is nearly
smooth ; parotoids prominent, gular fold strongly marked,
(jeneral colour above, brown suffused with pink and grey, some-
times with a dark lateral band ; under parts mottled brown. The
vomerine teeth are frequently absent. Total length, about 4 to 5
inches. They live, carefully concealed in the daytime, under

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SALAMANDRIDAE DESMOGNATHINAE

103

stones in or on the edge of the banks of little mountain streams.
The eggs are laid in two long strings, and are wrapped round the
body of the female like a rosary, the female having resorted to a
hollow in the mud, below a stone or other suitable place. The
out^r enveloi)e of each egg tapers out into a short stalk, and the
several stalks all converge, or are glued together into one common
knot, " much like a bunch of toy balloons held in the hand of a
street vendor.'
said to be meroblastic.

The egg is

larvae

the e<i<^

The
seem to remain in
until they are
nearly adult, and they
emerge at midsummer, with
the gills already much re-
duced. The complete meta-
morphosis takes place in
the autumn of the same
year. These little newts ^'°- ^^•—^^**^^^"^^'"*/"^^"*' f^™*^®^^*^®'?^

^ ' -. -.TT-i 1 1 in a hole underground, x 1. ( After Wilder. )

can, according to Wilder/

be collected all the year round, in Massachusetts from March
to December, except during the time of deep snow. They are
nocturnal and are easily kept.

Thorias pennatulus, from Orizaba, Mexico, the only species, is
noteworthy for its extremely large nostrils, and for the tongue,
which is supported by a central pedicle, free all round, and ending
in a thick knob, which can probably be protruded. The limbs are
weak, and the digits are also much reduced. Total length, under
2 inches, or 50 mm.

Typhlotriton spelaeus, of the Rock House Cave in Missouri, is
blind, the eyes becoming concealed by the skin during metamor-
phosis, when the gills are lost.

Sub-Fam. 2. Plethodontinae. — The five genera of this almost
entirely American sub-family (only one species of which, Sjielerpes
fu.scvs, occurs in Europe) can be distinguished as follows : —

I. Tlie tongue is attached by its central pedicle only, is free all round,
ends in a soft knob and can be shot out to a considerable distance.
With 5 toes : Spelerpes, p. 104.
With 4 toes: Mancnlits, p. 106.

* Amer, Natural, xxxiii. March 1899, p. 2'i\.

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I04 URODELA

11. The tougue is attached along the middle line and cannot be ])ro-
truded out of the mouth.

Jaws with numerous small teeth.

With 5 toes : Plethodon, p. 106.
With 4 toes : Batrachoseps.
Maxillaiy and mandibular teeth few in number but very large
With 5 toes : Autodax, p. 107.

Spelerpes, — Except in a few species the limbs are well de-
veloped and possess 4 fingers and 5 toes, which are either free or
webbed. But in the Colombian S. parvipes, still more in *S'.
lineolus of Orizaba and S. uniformis of Costa Rica the limbs and
digits are reduced to mere vestiges, and are practically without
function, the body, with the extremely long tail, having assumed
a wormlike shape. The young of many, if not all, species have
a pair of short balancers below each nostril; in the adult
these organs are reduced to little swellings or lost completely.
Several species are lungless, see p. 46.

The geographical distribution of this genus, of which some
twenty species are known, is very remarkable. The majority live
in Mexico and in the United States, a few are found in Colombia
and Northern Peru {S, altamazonicus and Plethodon platense
being the only Urodeles hitherto recorded from south of the
equator), one in Hayti {S. inf^iiscatus), two (S. subpahnatits and
S, uniformis) in Costa Rica, and S. fuscus in Europe.

S, hilineatus is a little newt under 4 inches in length — 60-95
mm. — found in the Atlantic States. It is brownish-yellow above,
with a black lateral line extending from the eye to nearly the
end of the tail. The under parts are bright yellow. It lives on
land, in damp places, concealed during the daytime under stones
or old trees, whence it emerges after a rain or in the dusk of
evening.

According to H. H. Wilder,^ " the eggs are deposited in May and
June in a single layer upon the lower side of submerged stones,
each batch containing 30 to 50 eggs. The stones which are
suitable for this purpose nmst be in the form of an arch, allowing
the water to flow beneath. They are generally in the more rapidly
flowing portions of the brook, but the depth of water must be
such that the eggs are at all times entirely submerged. They are
attached to the stone by gelatinous threads, proceeding from the
outer envelope, and although they are generally contiguous, they

1 Amer. Xatural. March, 1899, ]). 235.

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V SALAMANDRIDAE — PLETHODONTINAE 1 05

are each attached separately." The eggs are holoblastic. The
larvae hatch early and continue for a long time in the larval
state, probably two or three years.

S, porphyriticiis s. salmoneus. — Yellowish-brown or purplish -
grey above with tiny darker dots and markings. The sides of
the body are salmon-coloured, with a tinge of yellow. The under
parts are whitish, turning into salmon-pink on the tail. This
beautiful newt reaches about 6 inches in length and has a very
moist, slimy skin, which, combined with the lively motions ol
the creature, make it as slippery as an eel. It is found in the
AUeghany range, from New York to Alabama.

Specimens which I am keeping prefer the wettest part of the
cage, where they lie concealed in the moss and mud, leaving their
hiding-places at night in search of insects. One of them escaped
into the greenhouse and was discovered after nine months, having
established its permanent home in a cleft between mossy stones :
when the sweepings of a butterfly-net are emptied near its hiding-
place it peeps out and with a flash of its long, forked, white-
coloured tongue it secures its prey. Occasionally it goes into a
tank, when it swims with rapid, undulating motions, the limbs
being laid back and remaining inactive ; it sometimes rises to the
surface to emit and to take* in air, but, although mostly resting
half in the water, upon a rotten stump, it often lies for hours at
the bottom without stirring. When kept in dry surroundings,
the skin soon dries and wrinkles, and the animals show every sign
of sufifocation and general discomfort. The respiration of this
lungless species by means of rapid movements of the throat is
very limited, most of the necessary oxidisation of the blood being
effected through the skin.

S, fnscus. — This, the only European species, is thoroughly
terrestrial It is found in the mountains bordering the Gulf of
Grenoa, and in Sardinia. Its total length remains under four
inches. The smooth, very delicate and easily broken skin is
brown above, light below, and speckled with lighter and darker
markings. Below each nostril is a slight swelling, the remnant
of the cirri or balancers common to the young of many species.
It lives in shady surroundings, under stones, in old trees and in
limestone-caves, glued to the walls with spread -out toes, belly
and tail, quietly waiting for insects and spiders which it catches
by flashing out the long tongue.

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io6

URODELA

CHAP.

According to J. Berg/ it keeps well in cool, moist and well-
ventilated places. It lives on flies, small beetles, and maggots :
ants are also taken at once, probably owing to their lively
movements, but a few minutes later the newts roll about in
spasms and soon die. Towards the end of March one of Berg's
specimens gave birth to four young, whiclf were 36 mm., or nearly

Fio. 20. — Spderpes fuscus,
showing the position and
Khape of the partly and
fully protruded tongue.
The figure on the right side
shows the tongue and the
skeleton of tlie hyoid ap-
paratus. Bf the threatilike,
elongated, first branchial
arch ; iT, hyoid, in reality
attached by its outer end
to the vicinity of the quad-
rate ; r, tongue. About x 2.
(After Berg and Wieders-
heim. )

1^ inclies long, and differed from the adult only by their exception-
ally large nostrils, therel)y resembling the Mexican Thorius. The
little ones shot out their tongues about 10 mm., feeding on
Aphides.

Maiicvlus, — The two species of this genus live in Carolina and
Florida. M, quadridigitatus is a very slender, graceful little
animal, about 3 inches in length, the long and thin tail being
cousidembly larger than the rest of the body. Yellowish,
minutely speckled with brown above and on the sides, greyish -
white below. Life entirely terrestrial.

Plethodon. — About seven species in North America. This
genus has given its name to that of the subfamily, which might
with more reason be called Spelerpinae.

r. glutinosvs is slaty or bluish -black, with small whitish
specks, e8i>ecially on the sides of the trunk, where they are large
and often confluent. The skin is smooth and shiny. Total length
about 5 inches, half of which belong to the tail. Holbrook con-
sidered this as one of the commonest of the North American
newts, and mostly widely distributed, from Ohio to the Gulf of
Mexico. It usually lives concealed under stones, but prefers
fallen trees, probably on account of the insects upon which it

^ ZooL Garten, 1896, p. 88.

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PLETHODONTINAE 1 0/

preya When taken in the hand it gives off a great quantity of
slime.

P. erythronotus extends into Canada and is much smaller.
Brown or grey above, mostly with a broad, reddish-brown band
over the head, back, and tail. The under parts are white, with
grey and brown specks.

Autodax s. Anaides. — The large tongue is attached along the
median line. The jaws are furnished with few, but surprisingly
large, knife-shaped teeth, about ten in the upper and fewer in the
lower jaw. The small teeth of the vomers form a chevron-shaped
series behind the choanae, those of the parasphenoid stand in one
elongated patch. The tail is round ; number of toes, five. Three
species in Western Xorth America, from California to Oregon.

A. luguhris. — The eyes are very large and prominent. The
upper jaw shows a peculiar recess on either side for the reception
of the large lower teeth. The skin is smooth, devoid of parotoid
f^lands, but has a strong gular fold. The upper parts are dark
brown or lead-coloured, with whitish dots on the sides ; under
parts white. Total length some 6 inches, about half of which
belongs to the tail. The fingers and toes are very rich in sub-
cutaneous venous sinuses.

The habits of these creatures are in many respects peculiar.
Van Denburgh ^ says of A. iecanus " that it usuaUy moves quite
slowly, moving one foot at a time, but is capable of motion
surprisingly rapid for a salamander. When moving rapidly, it
aids the action of its legs by a sinuous movement of its whole
body and tail. The latter is prehensile. Several individuals,
when held with their heads down, coiled their tails around my
finger, and, when the original hold was released, sustained them-
selves for some time by this means alone. One even raised
itself high enough to secure a foothold. This animal's tail is
also of use in another way. When caught, it will often remain
motionless, but if touched, will either run a short distance with
great speed, or quickly raising its tail and striking it forcibly
against the surface on which it rests, and accompanying this
with a quick motion of its hind-limbs, will jump from four to
six inches, rising as high as two or three."

Eitter and Miller^ have made extensive observations on the life-
history of A. libgvhris. When wishing to pass from the hand to

* P. Calif. Ae, (2) v. 1895, p. 776. * Ainer. Natural, xxxiii. 1899, p. 691.

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I08 URODELA CHAP.

the table, the creature will frequently execute a well co-ordinated
spring and alight on its feet some distance away, instead of
falling over the edge in the typical salamander -fashion. This
species is nocturnal and entirely terrestrial, and seems to be
indifferent even to proximity to water. Eotten stumps and logs
are the habitations preferred, and wherever these occur in the
region about San Francisco Bay, even though at the places
remotest from water, specimens are sure to be found.

The eggs are laid in a hollow under ground, and the female
seems to remain curled around them until they are hatched,
which takes place in two or three weeks. The specimen observed
by Ritter and Miller laid 19 eggs. Each was contained in
a gelatinous capsule 6 mm. in diameter, and was firmly anchored
to a clump of earth by a narrow peduncle about 8 mm. long.
The embryos developed very large gills, each being composed of
three broad membranous lobes, the latter being thin and delicate,
much expanded, highly vascular and widely confluent at their
bases, so that the gills of each side really form one three-lolx^d
mass. Their dorsal surfaces are applied to the inner surface of
the egg-capsule. The amount of food-yolk is considerable. The
whole larval life is passed through within the egg. Before the
young is hatched the gills wither and cease to be functional, and
the gill-slits close up. The tail is round, and shows no indica-
tion of a fin at any time during the larval period. Newly hatclied
individuals appeared much distressed when put into water, and
were quite unable to swim. They immediately sank to the
bottom and remained there until they were removed. The
integumentary sense-organs, so well developed in the aquatic
larvae of Urodeles, are entirely wanting. When hatched the
young creature is about 32 mm. long; its general colour is
blackish-grey, finely sprinkled with bluish-silver. During the
second yciir tliis garb is changed to the dusky brown of the
adult, and tlie fine silver speckling is replaced by much larger
and less numerous yellow spots.

Although one of the most terrestrial of Urodeles, this species
is lungless, but the skin remains delicately smooth and moist
throughout life. According to the observers quoted, the pharynx
plays an important part in respiration. From 120 to 180 or
even more vibrations are made by the throat in a minute, and
in some cases these movements are grouped into series of about

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V SALAMANDRIDAE AMBLYSTOMATIN AE 1 09

20 to 25 extremely rapid vibrations, with periods between each
two series.

Subfam. 3. Amblystomatinae. — Composed of seven closely
allied genera, the distinguishing characters of which are the
grouping of the palatal teeth and the number of the toes,
which varies between 4 and 5. The geographical range of the
subfamily extends over the whole of North America and
Mexico and over the whole of Northern Asia, from Kamtchatka
and Japan westwards to the Ural, and southwards into China.
The occurrence of one species, Amhly stoma persimile, in the moun-
tains of Siam, makes it highly probable that bther species and
genera exist in the hitherto unexplored intervening countries.

Boulenger gives the following synopsis : —

I. The series of palatal t«eth converge backwards, forming a V-shaped
figure.
With 5 toes : Htpiobius, 3 species in Japan.
With 4 toes : Salamandrellaj 2 species Lake Baikal, Ussuri and
Schilka rivers, and Kamtchatka, p. 109.
II. The series of palatal teeth form an uninterrupted, doubly arched
V-shaped figure.
The 4 fingers and 5 toes are furnished with black, homy claws :
Onychodactylus japonicus.
III. The series of palatal teeth form two arches, convex tbrwards, separated
by a wide interspace.
The two series are short, confined to the space between the
choanae.

With 5 toes : Ranidens sibiricvsj Eastern Siberia and N.E.

China.
With 4 toes : Batrachyperus dnends, Moupin in China.
The series are long and converge backwards, 5 toes : Dicamp-
todon euMtUtS, California.
lY. The palatal teeth are arranged in a nearly straight, transverse line,
or they form an angle which points slightly forwards ; they are not
separated by a wide median space: With 6 toes : Amblystoma.
Some 16 species in North and Central America, one in Siam, p. 1 10.

SalamaTbdrella keyserlingi. — The mode of propagation of this
newt-like species has been observed by Shitkow near Jekaterin-
burg in the Ural mountains. The eggs were laid at the end
of April and were deposited in bags, which were attached to
a plant, with one end about an inch below the surface of the
water. The bag measured 15 cm. in length and 2 cm. in width
and contained 50 to 60 eggs. The larvae were hatched in
1 4 days in a sunny aquarium ; in another with a northern

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1 lO

URODELA

aspect the hatching took 23 days. The larvae were 10 mm. loii»;,
and remarkable for the length (1 mm.) of their balancers.

Amhly stoma opacum, — The general shape is very much like
that of the European Spotted Salamander. The head is short
and broad, the snout is rounded. The eyes are very pro-
minent, with a black pupil and a dark-grey iris. The neck has
a well-marked gular fold. The tail is thick and almost round.
The hind-limbs are considerably larger than the fore-limbs. The
general colour of the shiny, moist skin is a purplish -black with
light grey, transverse, partly confluent bars, giving the creature

a pretty appearance ; the under parts
are paler, bluish-grey. Total length
between 3 and 4 inches, or 9 cm.

This beautiful species inhabits
many of the United States east of
the Rocky Mountains, from Xew
Jersey to Florida and Texas. In the
perfect state it is thorbughly terres-
trial and easily kept. My specimens
prefer the holes of rotten and moist,
moss-covered stumps, or holes beneath
stones, which they leave, at night
only, in search of earthworms and
insects.

A. talpoideum is- closely allied,
somewhat stouter and almost uniform
brownish -back. According to Hol-
brook, " it chooses light soil in which
it will bury itself in a few seconds like a mole, and there continue
its course concealed from view ; but its track can often be followed
by the elevation produced on the surface of the soil, similar to
that seen in fields infested by moles."

A, punctatum is bluish-black, with a row of roundish yellow
spots on each side of the body and tail and upon the limbs.

E. A. Andrews ^ has made observations upon the breeding of
this species. Near Baltimore the eggs are very abundant in
March and even in February, in small pools in the woods, but
the adults are then rarely seen. Even when small pools, but
4 feet wide and 9 inches deep, were thoroughly raked out

^ Anicr. Natural, xxxi. 1897, p. 635.

Fig. 21. — E^f:^sac of tSaiainaitdrcIla
schrenki. x\. (After Shitkow.)

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AMBLYSTOMATINAE

before and after the eggs appeared, no adults were found, so that
it is to be inferred that the laying takes place in the night and
that the adults leave the water every day to conceal them-
selves under stones. One female was found moving away from
a bunch of eggs early in the morning. This specimen was
kept isolated, and laid many eggs, and as these developed into
normal larvae, the existence of internal fertilisation was proved.
Previously to the laying of the eggs white spermatophores were
found in the small pools, on the dead twigs and leaves covering
the bottom.

A. jeffersonianum. — This very slender and slippery species,
reaching a length of 6 inches, is remarkable for its long fingers
and toes, and its rather compressed tail. The general colour is
brown above, dirty whitish below, generally with numerous,
small, light blue and pale brown spots on the sides of the neck,
body, limbs, and taiL There are several colour-varieties, one of
them with white specka It is a very active and surprisingly
good climber, easily escaping out of high-walled bell-glasses, hiding
in the daytime in dark and moist localities. Its range extends
from Indiana and Virginia to Quebec.

A. perdmile. — This species is remarkable on account of its
geographical distribution. It is the only non-American species,
inhabiting the higher mountains of Siam and Upper Burmah.
There is no doubt about its belonging to the genus AmhJystoma,
although it had originally been described as a Plethodon. It
closely resembles A, jeffersonuinum in most of its characters,
notably in the arrangement of the palatal teeth, general propor-
tions, slender toes, and even in the presence of whitish spots,
which are scattered over the sides of its blackish, smooth skin.

A. tigrinum. — This, the commonest species, is conspicuous for
its large, depressed head, which is as broad as it is long, its
width being enhanced by the unusually large parotoid glands.
The mouth is very wide. The large, prominent eyes are golden,
and reticulated with brown. The gular fold is strong. The
limbs are stout, the fingers and toes short. The trunk is
strongly constricted by twelve intercostal grooves. The tail,
which is as long as the rest of the body, is somewhat compressed
laterally, but bears no trace of a fin. The general colour is
more or less dark brown or bluish black, marked with numerous
yellow spots and large blotches; the under surface inclines to

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I i:

URODELA

CHAP.

grey. The length of the adult male is about half a foot ; the
females, as usual being larger, sometimes reach the leugth of
9 inches. The range is from New York to California and to
Central Mexico.

The larva of this species is the famous Axolotl. It is
provided with three pairs of delicate and much-branched external
gills, a flat, long tail with a broad ventral and dorsal fin, the latter
extending along the back almost to the neck. The limbs,
although comparatively slender, are fully developed, and the head
is much more pointed than it is in the perfect form. The larvae
usually reach 8 or 9 inches in length ; exceptional specimens

^"^^w^^'^iysyy

Fig. 22. — Axolotls or larvae of tkmhlystoma tigrinum, x ^.

have been recorded of one foot in length, and have been described
as Triton ingeiis.

These larvae were found by the Spanish conquerors to occur
in great numbers in the lakes near Mexico City, and were called
Axolotl by the natives, a word signifying " play in the water."
They were, and are still, eaten, either roasted or boiled, with
vinegar or cayenne pepper.

For many years these creatures were looked upon as a
species of the Perennibranchiata, under the generic name of
Siredon {JS. axolotl^ s. jylsciforiyiis, s. mexicaiius, etc.), althougli
Cuvier suspected that they were but the larvae of an otherwise
unknown terrestrial Urodele. The mystery was not cleared up
until the year 1865, when some Axolotls which had been kept
for a year in the Jardiu des Plantes at Paris, suddenly began

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AMBLYSTOMATINAE I I 3

to pair, and laid eggs which within six months developed
into full -sized Axolotls. This certainly looked as if these
creatures were not larvae, but a true Ferennibranchiate species.
But to the general surprise several of these joung Axolotls
gradually lost their gills, the clefts closed up, the fins of the
back and tail disappeared, the head became broader, the crea-
tures left the water permanently, and in fact turned into the
already well-known terrestrial Amhlystoma tigrinum. The other
brothers and sisters of the same brood remained aquatic Axolotls,
which thereby revealed themselves after all as the larval and not
as the perfect stage of this remarkable species.

At the suggestion of Kolliker and Weismann, FrL Marie
von Chauvin ^ undertook, at the University of Freiburg, long and
carefully conducted experiments, showing (1) that little Axolotls
can comparatively easily be caused to develop further into the
perfect Atnhlystoma if they are induced to breathe air more
frequently than usual ; shallow vessels, perhaps also insufficiently
aerated water, will produce the desired result ; (2) that the
commencing metamorphosis can again be checked, the shrinking
gills then imdergoing fresh development ; (3) that they can
be forced to remain Axolotls; (4) that the cutting off of the
gills has no influence upon their possible metamorphosis, the
gills being easily and quickly renewed. The same lady found
also that AmUystoma, the perfect form, lives in the water
during the pairing time and behaves in the same way as the
Axolotls.

The latest observations have been made by MetzdorfiF.* Axo-
lotls, at least those which are kept in captivity in Europe, are
ready for propagation several times in the year, either in the
spring, from* April to June, or in December. The male deposits
spermatophores, which in the following night are taken up by
the female into the cloaca. On the following day, preferably in
the i^ftemoon, she grasps a suitable leaf, for instance that of
Vallisneria, with the hind-limbs, and presses it against the vent.
The eggs are expelled by strong wriggling movements of the
body, and are formed into three or four packets of six to ten
eggs each, so that about thirty eggs are laid at one sitting.

* ZeiUchr. wiss, Zool. xxvii. 1877, p. 522 ; xli. 1891, p. 365 ; Zool. Am.
1882, p. 513.

- Zoolog. Garten, 1896, p. 114.

VOL. VIII I

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114 URODELA

Then she takes a rest before proceeding again ; the whole process,
in which the male takes no further interest, lasting about two
days. The most suitable temperature is one of 18-20° C, or
about 68** F. The water must be well aerated. Sterile eggs
turn white on the second day. The little larvae are hatched in
about a fortnight. Eggs which are kept in a higher tempera-
ture, from 22-24° C, develop more quickly, but the resulting
young are smaller; they show already on the fifth day head,
tail, and the beginning of the gilla According to Bedriaga,
they live at first upon Infusoria and Daphnia ; when they are
20-25 mm. long they eat Tuhifex rivvlorum; later on they take
scraped meat and are liable, when hungry, to nibble off each
other's giUs, but these are easily reproduced. Wlien 20-25 cm.
long, at the age of about six months, they are able to breed.
The chief point of interest is the fact that this species of AvMy-
stoma frequently remains throughout life in the larval state, except
that it develops generative organs. The natural causes of this
retention are not completely known. According to Shufeldt,
who observed them under natural conditions near Fort Wingate
in New Mexico, plenty of food, the drying up of the swamps, and
the increasing temperature of the diminishing water, hurries on
the metamorphosis, while deeper water retards it. Weismann *
suggested that the specimens in the Mexican lakes which
remained Axolotls were prevented from becoming perfect
Amblystomas on account of these lakes, after the disappearance
of the surrounding forests, having receded from their former
boundaries, which are now covered with a saline, uninhabitable
crust. This may be an explanation, although Axolotls do not
live in brackish water. But Weismann went farther, and with
his well-known dialectic powers has succeeded in spreading the
belief not only that the Axolotl is a case of reversion to an
ancestral stage, but that the present Amhlystoina, instead of
being the progressive, perfect form, is likewise a casfe of reversion.
A reversion from a reversion ! The whole line of evolution
would then be as follows: Amhly stoma; its young, owing to
adverse circumstances, revert to the stage of the Perennibranchiate
ancestors of all Urodela ; if some of these Axolotls lose their gills
and fins, they revert tbereby into the original Amhlystoma.

^ Zcitschr. unss. Zool. xxv. 1875, p. 297. See also Hahn, Jiev, Quest. SeL
(2), i. 1892, p. 178.

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V SALAMANDRIDAE SALAMANDRINAE I I 5

Surely a roundabout way of explaining a curipua but after all
rather simple process of Neoteny ; cf. p. 63.

Observations on the metamorphosis of Siredon lichenoides
into AmbU/8to7na mavortium have been made by Marsh, who also
gives figures of the larval and adult forms.^

Snb-Fam. 4. Salamandrinae. — The six genera of this sub-
family fall into two natural groups : I, True Salamanders, with
the palatal teeth arranged in a pair of 8-shaped figures, and
without a fronto-squamosal arch. II, Tritons, with the palatal
teeth in the shape of a A, i-^- the right and left series meet at
an angle ; the fronto-squamosal arch is present, either bony, or at
least ligamentous. Triton cHstatiis is, however, exceptional, in
that the two palatal series often do not meet and that the arch
is absent. The number of fingers is universally four, that of the
toes is five except in Salamandrina, which has only four.

The geographical distribution of the sub-family, entirely Peri-
arctic, may be said to be the reverse of that of the Amblystomatinae.
Of the twenty-five species namely, only two are American, four'
are Eastern Asiatic, and of the remaining nineteen, two are Algerian,
while the rest live in Europe or in Asia Minor. It is in fact an
essentially Palaearctic group.

The six genera can be distinguished as follows : —

I. The palatal teeth are arranged in two S-shaped curves. True Sala-
manders

Tongue short and thick. Salamandray p. 115.
Tongue long and projectile Chtoglosmy p. 121.
II. The palatal teeth are arranged in a A shape. True Tritons.
With only four toes. Salamandrina^ p. 122.
With five toes.

Pterygoids separated from the maxillary and quadrate

bones: Triton, p. 122.
Pterygoids touching the maxillae and quadrates. Himalo-
Chinese : Tylototriton, p. 132.
•s Pterygoids united broadly with the maxillae. Chinese :
Pachytriton, p. 132.

Sala7na7idra,—Witho\xt fronto - squamosal arch. Five toes.
Tail round. Three species in Europe and Western Asia.

S. macvlom, — The Spotted or Fire Salamander. General habit
stout. Usual length about 5 to 6 inches ; the females are mostly
larger than the males; specimens of more than 8 inches in

' Amer, Joum, Sci. (2), xlvi. Nov. 1868, p. 364.

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I 1 6 URODELA CHAP,

length are giants. Head as broad as it is long, snout rounded.
Limbs and digits stout and short. The skin is smooth,
shiny and full of pores, with a strong gular fold. The parotoid
glands are large and covered with large pores. A series of distinct
swellings, or cutaneous glands, each with a distinct opening,
extends along either side of the back, and a shorter series along
the flanks. The general colour of the Spotted or Fire-salamander
is black, with irregular, large yellow patches on the back and
limbs. These markings vary extremely, so much so that scarcely
two * specimens, collected at random, are alika In some the
yellow patches form two more or less regular bands, in others
they are partly confluent ; again the yellow may be preponderant
on the back or much restricted. Occasionally the chrome-
yellow is replaced by orange. The under surface is as a rule
bluish grey-black. This combination of shiny yellow and black
is a good instance of warning coloiu*s. The creature is
poisonous,* cf. p. 38. When left in peace, or handled gently,
it is perfectly harmless, but when treated with violence, or
submitted to severe pain, a milky white fluid exudes from
the glands and is, under violent contractions of the muscular
skin and body, sometimes squirted out in fine jets to the distance
of a foot. Burning pain and subsequent inflammation result if
this poison gets into the eye. The same applies to the mucous
lining of the mouth and throat. A few drops of this poison
introduced into the blood or into the stomach of a small animal
are sufficient to cause its death. Cold-blooded animals are as
susceptible as warm-blooded creatures.

I once put two American bull-frogs into the same outdoor en-
closure with a large number of salamanders. Next morning the
huge frogs were foimd dead, each having swallowed a salamander,
which they were not acquainted with and had taken without
suspicion.

The Fire-salamander has a wide range, namely the whole of
Central, Southern, and Western Europe with the exception of
the British Isles. It extends southwards into Corsica and
Algeria, eastwards through Asia Minor into Syria. Where
it does occur it is ratlier common, provided the terrain is
mountainous or hilly and covered with vegetation. There it lives
under moss or rotten leaves, in the roots of old trees, in the
cracks and clefts of the ground, of rocks or of ruins of buildings ;

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SALAMANDRIN AE I I J

in default of anything better under heaps of stoneSi or in the
holes dug by mice or moles. One chief necessity for its happiness
is moisture.

The salamander does not occm* everywhere, but is rather
local. On certain kinds of limestone it is rare or absent ; granitic
terrain and red sandstone seem to suit it best, for instance the
Hartz Mountains, Thuringia, and Heidelberg are favourite
localities. But even there we may spend days and weeks and
never come across a single specimen. We may turn stones, rake
up the moss and leaves, pry into cracks, and we unearth perhaps
a few sorry-looking, listless, dull and dry, half-emaciated creaturea
The same place after a thunderstorm will be literally swarming
with sleek, lively salamanders, in search of earthworms and all
kinds of insects, especially at dusk or during the night. They
disappear in the autumn, in October, to hibernate in the ground,
out of the reach of frost, and they reappear again in April.
I^ter on they congregate at little springs, always at running
water, to reach which they have often to make long migrations.
This is the only time when these thoroughly terrestrial creatures
approach water, in which they easily get drowned.

Although this species is so common its mode of r^roduction
has been satisfactorily discovered only quite recently. There are
some puzzling facts which it took a long time to observe correctly
and to interpret. The larvae are born in April, May, or June,
while there are no eggs in the oviducts, but in July these are
full of fertilised eggs before copulation takes place. This seems
contradictory. The explanation is as follows. In July there is an
amplexus of the sexes, short, and often on land — a sort of pre-
liminaiy exciting performanca Both sexes then descend into
the water, but generally remain on land with the fore part of
the body. The male deposits a spermatophore and the female
takes part of this into its cloaca. In the case of a virgin female
the eggs are fertilised in the oviduct and ripen until the autumn,
but the larvae nearly ready for birth remain within the uterus until
the following May, i.e. about ten months. The mother then crawls
half into the water, mostly at night, and gives birth to from a
few to fifty young, fifteen being perhaps the average. The young
are surrounded by the egg-membrane, which either bursts before
or shortly after expulsion. This species is consequently viviparous
in the proper sense. If she produces a few young only, say from

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I 1 8 URODELA CHAP.

two to five, these are much larger and stronger than those of a
large litter. Occasionally a few addled or only partly developed
eggs are also expelled.

In the case of old females which have produced offspring
before, the whole process is more complicated. The sperma^ takes
up in July, remains in the receptaculum of the cloaca until the
May or June following, i,e. until the previous larvae have passed
out of the uterus and are born. Then the spermatozoa ascend
to the upper ends of the oviducts, where tliey meet and fertilise
the new eggs. After these have descended into and filled the
uterus, and are already developing into embryos, copulation takes
place again in July, preparatory for next year's eggs.

The new-bom salamanders have three pairs of long external
gills, a long tail furnished with a broad dorsal and ventral
fin, and four limbs, althougli these are small. The total
length is about 25 mm. or 1 inch. The general colour is
blackish with a pretty metallic golden and greenish lustre.
The little creatures are very active, and at once eat living or dead
animal matter. In captivity they are liable to nibble each
other's gills and tails. During the first six or eight weeks they
assume a row of dark spots on the sides ; these spots enlarge, and
the whole skin becomes darker. Yellow spots appear next, first
above the eyes and on the thighs, later upon the back ; the
ground-colour at the same time becomes black, until at the
beginning of the fourth month they look like the parents.

The metamorphosis is very gradual. The tail-fin diminishes
first, but the gills grow until shortly before the little creatures
leave the water. Darkness, cold, and insufficient food retard the
metamorphosis, sometimes until October. It is easy to rear them
artificially provided they are well fed, kept in a light place, and
in clean, well aerated water. If prevented from leaving the
latter, for instance when kept in a glass vessel with vertical walls,
or if hindered by a piece of gauze from rising to the surface
and taking in air, they can be kept as larvae well into the
winter.

Very young, perfect little salamanders, of from 1 to 2
inches in length, are excessively rare; even specimens of 3
inches are far from common. They probably spend the first two
or three years of their life in careful seclusion.

A few adults can be easily kept for many years in shady

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SALAMANDRINAE I 1 9

places provided with moss, rotten stumps and stones, to afford
them suitable moist and cool hiding-places, and they readily take
earthworms, larvae of beetles, snails, woodlice, etc. But any
attCTipt to keep them in large numbers ends in failure. They con-
gr^ate together in clumps, all making for the same cavity or recess,
as if that were the only one in existence (very likely they are
right in so far as that place is probably the best), and they get
rapidly enlarging sores, chiefly on the elbows and knees. These are
soon infested with fungoid growths, and this disease spreads
like an epidemic and soon carries them off. .

S, atra, — The Alpine Salamander differs from the Spotted
Salamander by its uniform black colour and smaller size, which
averages between 7 and 5 inches. It is restricted to the Alps
of Europe, from Savoy to Carinthia, at from 2000 to as much
as 9000 feet elevation, living with predilection near waterfalls,
the spray of which keeps the neighbourhood moist, or in mossy
waUs, in the shade of forests near brooks, or under flat stones on
northern slopea The most interesting feature of this species
is that it produces only two young at a time. These are
nourished at the expense of the partially developed eggs in the
uterus, and they undergo their whole metamorphosis before
they are born. By far the best and most complete accoimt
of this mode of propagation has been given by G. Schwalbe.^
The length of the ripe embryos is about 45 mm. ; they lie
mostly bent up, with their heads and tails turned towards the
head of the mother. The gills are beautiful, delicate red organs,
the first pair being generally directed forwards and ventralwards,
the second upwards, the third backwards; they are longest
when the creature is about 32 mm. long, while there is still
much yolk present. At this stage the gills are so long as to
envelop nearly the whole embryo. There is rarely a second
embryo in the same uterus, and an extra foetus is generally
smaller, frequently a monstrosity not fit to live ; it is probable
that it is not used as food, but that it is expelled at parturition.
The embryo passes through three stages, (1) still enclosed within
its follicle and living on its own yolk, (2) free within the
vitelline mass which is the product of the other eggs, (3) there
is no more vitelline mass, but the embryo is possessed of
gills 10-12 mm. in length, and is still growing. During the

1 Zeitachr, Biol, xxxiv. 1896, pp. 340-896.

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1 20 URODELA

second stage the yolk is directly swallowed by the mouth.
The walls of the maternal uterus are rather red. The ex-
change of nutritive fluid takes place through the long external
gills, which thereby function in the same way as the chorionic
villi of the Mammalian egg. Each gill contains a ventral artery
and a dorsal vein, each of which looks like thB midrib of a pinnate
leaf; there is also a fine nerve and a weak bundle of striped
muscular fibres. Each gill- filament receives a capillary artery'
which extends to the epithelium of the tip, where it turns
into a capillary vein. . The epithelium of these filaments, which
are full of blood, is ciliated, the resulting current being directed
from the base towards the tip. In older larvae this ciliation
becomes restricted to the tips. The body of the gills is furnished
with flat epithelium, these non-ciliated portions alone are closely
appressed to the uterine wall, and it is here that the exchange
of gas takes place between mother and larva. The nutrition
takes place through the gills, as they are bathed by the yolk-
mass.

Schwalbe also explains the whole question of the reduction
of the number of embryos. He says rightly that in S. m^icuJosa,
which gives birth to many young, there are in the oviduct many
eggs which have only partly developed into embiyos, and these,
perhaps from want of room and nourishment, degenerate into the
irregularly shaped whitish-yellow bodies which are occasionally
found packed in between the developing embryos. Consequently
all those eggs had been fertilised near the ovaries. S, atra exhibits
a further stage in so far as most of the eggs, fertilised above in the
oviduct, degenerate, and only two or three become fully developed.
These few embryos live on the degenerating eggs, which together
produce the vitelline material spoken of above. The two full-
grown and metamorphosed embryos, each measuring about 50
mm. in length, are equivalent to the numerous new-bom larvae
of ^''. mavitlosa, esi)ecially if the smaller size of the adult Alpine
Salamander is taken into consideration.

Mile, von C'hauvin ^ has experimented with the unlx)rn larvae
of this Salamander. She cut out 23 larvae and put them into
water. One of them, already 43 mm. long, took earthworms
on the next day, and the beautifiil long, red gills became pale
and shrunk, and on the third day were cast off close to the

* Zt'tfschr. w/t*. ^tW. xxix. 1877. ppt 824 f., pi. xxii.

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SALAMANDRINAE I 2 I

body. New gills sprouted out on the same day, first in the
shape of three tiny knobs on either side. After three weeks
they had become round globes, which gradually sprouted out
into several branches, far shorter and more clumsy than the
original gills. During the whole time the larva was lying
quietly at the bottom, in the darkest comer, but showed a good
appetite. The fin of the tail disappeared and was supplanted
by a stronger one. In the sixth week the skin was shed in
flakes, and this process took fifteen days. This larva lived in the
water for fourteen weeks and grew to 6 cm. in length ! When
the new gills gradually shrank, the compressed and finny tail
assumed a round shape, the skin became darker and shinier,
and after the larva had again shed its skin, there appeared the
dark rugose skin of the typical S, atra. The gills were reduced
to useless appendages — not cast oflF — and the creature crawled
out of the water. A fortnight later the gill-clefts were closed.
A 'second larva behaved . similarly, first casting off the featherj'
gills, substituting a new and stronger set, which, however,
fourteen days after, excision from the uterus, shrank again, and
on the nineteenth day the gill-clefts were closed. The lady also
observed that nearly ripe larvae, when cut out, rushed about
in the water and ate, just like the new-born larvae of the Spotted
Salamander.

A third species, S, catu^asica, is found in the Caucasus. It
rather resembles the Spotted Salamander in coloration, but has
a larger tail and lacks the lateral warts. The male is remark-
able for the possession of a soft permanent knob or hook at the
top of the root of the tail. This pommel possibly prevents the
slipping off during the amorous amplexus, provided the sexes then
entwine like certain Tritons.

Chioglossa lusitanica. — The. only species of this genus is
restricted to the north-western third of the Iberian peninsula.
This graceful, slenderly-proportioned and beautiful Salamander
is apparently very rare and local, having hitherto been found
at a few places, namely, near Coimbra, Oporto and Corufia. It
lives under moss, and runs and climbs with an agility surprising in
a Urodele. The tongue is long, ending in a fork, and is supported
by a median pedicle so that the tip can be quickly protruded
to the distance of more than an inch. The whole length of
the animal is about 5 to 6 inches, two-thirds of which belong to

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122 URODELA

the long tail, which is compressed at the end. The skin is smooth
and shiny, with a gular fold and large parotoids. The general
colour is a rich dark brown, with a pair of broad reddish-golden
bands along the back and tail^ the bands being separated by an
almost black vertebral line.

The few specimens which I have been lucky enough to
observe made little holes or passages in the moist moss of their
cage, peeping out with their heads in wait for little insects,
which they caught with flash-like quickness. They seem to be
crepuscular.

Scdamiuidrina perspicillata. — This genus, represented by one
species, a native of Liguria and Northern Italy, possibly extend-
ing into Dalmatia, is the only Salamander which has but four
toes. The akin is not shiny and smooth, but is finely granular
and dry, forms no gular fold, and is devoid of parotoid
glands. The tail is more than half the length of the animal,
which measures from 3 to 4 inches. The genei-al colour is
black -brown with a broad V-shaped orange - yellow mark
extending from eye to eye over the occiput. A faint irregidar
yellowish line extends along the middle of the back and tail.
The throat is black, with a diffused white patch in the middle ;
the belly is white, with black dots ; the anal region, the inner
sides of the legs and the under side of the tail are carmine-red.

This slender and pretty Salamander is diurnal, and feigns death
when discovered. Only the female goes into the water, in March,
to glue the eggs on to submerged rocks or water-plants. The
young finish their metfimorphosis by the month of June, and
reach full size during the winter, the climate of their home
being sufficiently genial to make hibernation scarcely necessary.

TrifMti s. Mo/(/e. — The tail is strongly compressed and
frequently has a permanent fin. The fronto- squamosal arch is
variable, it l)eing either bony as in the South European, Eastern
and American species, or reduced to a ligament, or lastly absent
as in 7\ crisfatus. The males of all the English Newts, of T.
vittntus and of 2\ vuinnoratus, develop a high cutaneous crest
on the back and tail during the breeding season, and this crest
acts not only as a swimming organ and ornament, but also as
a sensory organ.

The whole genus comprises some eighteen species, twelve of
which are European, although some of these extend into Western

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SALAMANDRINAE I 2 3

Asia ; T. pyrrhogaster and T. sinensis are found in N.E. China,
the former also in Japan ; T, poireti and T. hagenmuelleri live in
Algeria, and only two, T. torosus and T, viHdescens, are North
American. Some of the species have a limited range; thus
T. mo^Uanus is confined to Corsica, T, rttsconii to Sardinia, T.
hoseai to the north-west of the Iberian peninsula and T, asper to
the Pyrenees.

Newts all prefer moisture without heat. During the pairing
season they take to the water, mostly to stagnant pools, which
sometimes implies long migrations. During this period, which
is in some cases rather prolonged, they become thoroughly
aquatic and undergo some important changes. The tail-fins are
much enlarged ; in the males of some species a high cutaneous
fold gi'ows out on the back, devoid of muscles, but rich in sense-
organs. The whole skin, instead of being dry, possesses numerous
mucous glands and, what is of more importance, specialised
sensory apparatuses which are arranged chiefly along the lateral
lines of the body and part of the tail

After the breeding season Newts become terrestrial, hiding in
cracks, trees, or in the sandy soil. Some species aestivate during
the hot and dry season. They hibernate either in the ground,
or occasionally in ponds. T, wlgaris is difficult to keep in
the water beyond the pairing season, while this is easily done
with T, alpestris and T» cristatus ; 2\ waltli can live in the
water for years. The food consists of all kinds of insects, centi-
pedes, worms, snails, etc, which are searched for chiefly at night.
It is astonishing to see a little Triton getting hold of and gradu-
ally swallowing a wriggling earthworm almost as thick and as
long as itself. When two newts seize the same worm, as these
voracious and jealous creatures often do, each gets hold of one
end, and swallowing as much as it can, twists and rolls round
in a direction opposite to that of its rival, until the worm breaks,
or until the jaws of the two newts meet and the stronger of
the two draws it out of the weaker one and swallows the
whole worm. They do not drink, but soak themselves in the
water.

The skin is shed periodically, and rather often by the rapidly
growing young; by the adult, during the life in the water,
rarely during the sojourn on dry land. The skin breaks round
the mouth ; assisted by the fingers and by contortions of the

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1 24 URODELA

body, it is then slipped backwards over the trunk and tail,
whereupon the newt seizes the skin with the mouth, draws the
shirt off entirely, and — swallows it. Such freshly shed skins are
very delicate and pretty .objects when suspended in water or
some preserving fluid. The shed skin, consisting only of the
outermost layer of the epidermis, is entire, but turned inside out,
with fingers and toes complete, the only holes being those for
the mouth, eyes, and vent.

None of the Tritons are viviparous. The eggs, which are
glued singly or in small numbers on to stones or water-
plants, are hatched in about a fortnight, sooner or later accord-
ing to the species and the prevailing temperature. The larvae
are always provided with three pairs of branched external gills ;
the fore-limbs appear much earlier than the hind-limbs. Most,
perhaps all, larvae develop two pairs of thread-like protuberances
on the sides of the upper jaw, by means of which they attach
or anchor themselves on to water-plants shortly after they are
hatched. Thus moored they remain motionless in a slanting
position, now and then wriggling their tails and shifting their
place, or sinking to the bottom. The metamorphosis is finished
during the first summer, and the little newts, often partially
transparent, leave the water to hide under stones. Not unfre-
quently the metamorphosis is retarded and not finished by the
autumn. The larvae of T, cristatiis, especially when reared in
ponds with abrupt or overhanging banks, so that they cannot
leave the water, retain considerable remnants of the gills, still
more frequently the clefts, although breathing chiefly by the
lungs. Such individuals reach a length of 3 inches, and are
.larvae so far as the finny tail and the gills are concerned. They
hibernate in this condition, and in exceptional cases reach sexual
maturity ; — at least the females, which develop ripe eggs ; the
males are not known to produce spermatozoa.

Much has been written on the amorous games of newts,
but it is only recently that the mode of fecundation has been
actually observed. Gasco^ placed the newts in glass vessels
suspended from the ceiling of his laboratory. The antics of the
enamoured male around the female, rubbing the latter with its
head, or lashing it gently with the tail, and playing around ij
in its often beautiful nuptial dress, are meant to excite tho

* Ann, Mu8, Oenova, xvi. ]880, p. 83.

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SALAMANDRINAE

125

female. The male then at intervals emits spermatophores,
which sink to the bottom, and the female takes them up into its
cloaca. For further information see p. 54.

Triton eristcUus. — The Crested Newt has a slightly tubercular
skin with distinct pores on the head, on the parotoid region and
on a line along the side of the trunk. There is a strong gular
fold. The general colour above is dark or black-brown with an
olive tinge, interspersed with darker spots; the sides of the
body bear irregular white spots. The under parts are yellow,
almost always vrith large black spots. The iris is golden yellow. —
The nuptial dress of the male is very striking. A high, serrated

Fig. 2S.— Triton cristatwi. 1, Female; 2, male in nuptial dress, x §.

crest occurs on the head and body ; the upper surface of the head
is marbled with black and white ; the under parts are orange-
yellow with black spots, and the sides of the tail are adorned
with a bluish-white band. — The female, always devoid of a crest,
generally exhibit^ a yellow line along the middle of the back.
— The average length of fully adidt specimens is about 5-6
inches or 13-15 cm.; the females are as usual larger than the
males; 144 and 162 mm. for an English male and female
respectively are exceptional records.

Propagation takes place in April. The newly hatched larvae
are yellowish-green, with two black dorsal bands, and with a
whitish edge to the tail-fin. By the middle of July they are
about 5 cm. long, and the white-margined tail now ends in a

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1 26 URODELA

thread 1 cm. in length. The general colour above is light olive-
brown, dotted with black ; the flanks and belly have a golden
shimmer.

The Crested Newt has a wide distribution, extending from
England and Scotland through Central Europe into Trans-
caucasia ; the northern limits are Scotland and Southern Sweden.
Although found in Greece and Lombardy, it does not occur in
the Iberian peninsula nor in the South of France, where it is
represented by the next following speciea

Triton marmoi^atus, — The Marbled Newt is of the same size
as the Crested Newt. Its ground colour is grass -green above,
brown below, with numerous large and small irregularly shaped
marbling patches, spots and dots of black. The crest of the
neck and trunk is entire, not serrated, adorned with dark
vertical bands, and separated from the high dorsal fin of the tail
by a deep indenture or gap. The female has an orange line,
slightly sunk in, instead of the crest. This newt is confined to
France and the Iberian peninsula. In the North of Portugal
and in Galicia it is frequently seen in little streams and ponds
during the months of March and April. The rest of the year it
spends on land. In France occur hybrids of this species and
T, cristatics. They have been described as T, blasii,

T. alpestris. — The Alpine Newt is easily distinguished by the
rich orange colour of its under parts, which are unspotted, except-
ing a few dark specks across the throat, below the gular fold.
Specimens with many ventro-lateral black spots are exceedingly
rare. All the upper parts are dark, but vary individually. The
prettiest specimens are dark purplish grey, with black marblings ;
others incline more towards brown ground -tones, the blackish
markings then appearing more prominent. The sides are often
stippled with tiny whitish dots. The iris is golden yeUow. —
The nuptial male has a low, not serrated crest, which extends
uninterruptedly from the nape into the dorsal fin of the tail.
The crest is pale yellow, with black vertical bands and spots.
The ground-colour of the upper parts inclines to blue, especially
on the sides. The lower fin of the tail assumes an irregular
band of bluish- white confluent patches.

This newt is rather small, females rarely exceeding 100 mm.
or 4 inches in length. Its home is chiefly the hilly and
mountainous parts of Central Europe, from Holland to Lombardy,

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SALAMANDRINAE 1 27

Austria-Hungary, and Greece. Although it ascends the Alps to
between 6000 and 7000 feet, it is also found in the Nether-
lands, but not in the North German plain.

T, wlgaris (s. taeniatus, a functatus). — The Common or
Spotted Newt usually reaches 3 inches (7-8 cm.) in length.
Boulenger's record-specimen measured 104 mm. It is charac-
terised by the yellow, partly orange under surface, which is
always spotted with black. The upper parts are olive-green or
brown, inclining to white on the flanks ; the black spots of the
back, sides, and especially of the tail, are arranged in more or
less distinct lines, giving a somewhat banded appearance to
some femalea — The breeding dress of the male shows a non-
serrated, but "festooned" high and very wavy crest, which
extends from the neck without interruption into the likewise
wavy tail -fin. The tail is adorned with a lateral, glittering
blue stripe, interrupted by vertical dark spots. The larvae are
marked by a series of yellow dots, which extend over the lateral
line and the tail, which latter temporarily possesses a terminal
filament like that of the larvae of T. cristatus.

The distribution of the Spotted Newt is the same as that of
T. cristatus, namely Europe, with the exception of the Iberian
Peninsula and Western Asia.

T, palmatus s. helveticus. — This is the smallest of all the
European newts, rarely reaching more than 3 inches in length.
It is distinguished by several specific characters. The tail ends in
a thread which is in some males 10 mm. in length, but is only just
indicated in the female. The breeding male develops a cutaneous
fold along each side of the back, and a low, enjbire, vertebral
crest; the toes are fully webbed. The under parts are pale
yellow, inclining to orange towards the middle of the belly, and
with a few blackish dots. The lower caudal crest has its edge
blue in the male, orange in the female. The general colour
of the smooth skin is olive-brown above, with numerous dark
spots, which are arranged in more longitudinal streaks on the
head.

The Webbed Newt is a native of Western middle Europe,
ranging from Great Britain and Northern Spain to Switzerland
and Western Germany.

Closely allied to the last species are 1\ boscai of Spain
and Portugal, T. itcUicus, T. montadoni of Moldavia, and the

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128 URODELA chap.

beautiful T. vittatus of Asia Minor. From China and Japan
are known T, pyrrhogaster and T. sinensis.

The North American species are T. torosus and T. viridescens.
The former, of Western North America, is one of the largest
newts, reaching a length of more than six inchea The head is
much depressed and broad, and has very prominent parotoid
and other glands. The limbs are strong, especially in the
male. The skin of the upper parts is very granular, uniform
dark brown, without a crest. The tail, which is larger than
the head and body, is strongly compressed, with a low dorsal
and ventral fin. The under parts and the lower edge of
the tail are uniform yellow or orange red. The iris is green.
A specimen in my keeping spends most of its time in the cracks
of rotten stumps or on the top of moss in the darkest shade.
It lives on earthworms but despises insects. Like most of the
other newts it becomes lively at dusk.

T. inridescens is common throughout the Northern and
Eastern parts of the United States. Large females are about

1 1 cm. long, the males 1 cm. less. The
general colour above is brown, with a
tinge of green ; on each side of the
trunk, with a row of bright vermilion
spots; the under parts are orange,
studded with small black dots. Half-
grown specimens are brownish red,
with the same lateral red spots as the
adult. According to Jordan,^ this
voracious species lives chiefly on the
larvae of insects, on small molluscs
Fio. 2i.^Triton riridescens. 1, such as Cyclas and Planorhis, on earth-
^ iut; mL'^e t;*:^'; worms and on smaU Crustacea. It is
X 6 ; 2, a spermatophore just eminently aquatic in the adult stage.

discharged showing its gelatin- mi i -j i» a -i ^ x

ous base with a projecting spike The cggs are laid from April to June,
which bears a tuft of spermato- the period lasting for One individual

zoa, x2. (After Jordan.) /. . • ?

tour to SIX weeks, or even longer.
One female laid 108 eggs in all from 20tli April to 30th
May. After having selected a suitable plant, for instance an
Anacharis or a bunch of Fontinalis leaflets, she bestrides the
plant and gathers in the surrounding shoots with her hind-

^ Joum. Morphol. viii. 1893, p. 269.

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SALAMANDRINAE 1 29

limbs, pressing the leaves closely around the cloaca. She next
turns on her side, or occasionally on her back ; with fore-
limbs outstretched and rigid, with hind-limbs and leaves com-
pletely hiding the cloaca, she remains perfectly motionless for
six to eight minutes. Then she slowly leaves the " nest," which
now holds an egg well protected by a tangle of shoots glued
together by the gelatinous secretion poured out of the cloaca.
Jordan concludes, from the fact that he never found spermatozoa
in the oviducts, that the eggs are fertilised just before they are
expelled, when passing the receptaculum seminia

The metamorphosed young pass their life on land under
stones and logs as the so-called red variety, which is merely a
stage in the life-history of the species. It seems to take them
several years to reach maturity, and to become again typically
aquatic. Young, red individuals which I have myself kept,
have behaved for more than a year like the young of other
newts, spending their time under moss and bark without going
into the water.

The change from the red-spotted stage has been exhaustively
studied by Grage.^ He remarks that this species is very common
near Ithaca, in an upland forest and along the head-waters of
the Susquehannah. The transformation takes place either in
the autumn or in the spring, either while the newt is still on
land, or after entering the water.

Of two which were kept in a jar with moist wood, one was
especially brilliant, but within two weeks it assumed, in the
middle of September, the characteristic coloration of the viri-
descent form. The two specimens were in the jar until the
following July, when they were placed where they could enter
the water. This they did with great readiness, and they re-
mained submerged for a considerable time at first. The time
under water increased in length, until within two or three days
the pharyngeal respiration under water was fully established.
On the other hand, viridescent specimens never reassume the
red garb when kept out of the water.

Eed specimens entering the water in the spring, changed
into the greenish form within a few weeks, and established
the pharyngeal respiration, losing the ciliated oral epithelium.
Branchiate larvae and the adult aquatic forms hjive non-ciliated

' Amer. Xatural, 1891, p. 1084.
VOL. VITI K

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1 30 URODELA CHAP.

epithelium, and the cilia are re-established when a green speci-
men is forced again to live on land. Ciliation always exists in
the red stage, and in the green stage before the newt has taken
to the water. The cilia sweep towards the stomach.

The three following South European species belong to the
Euproctus group, so called on account of the mostly conical,
backward directed, and vividly coloured vent.

T, asper s. pyreruieus. — The Pyrenean newt has hitherto been
found only in the Pyrenees, for instance in Lac Bleu and Lac
d'Oncet, which latter lies about 7000 feet above the level of the
sea. According to Bedriaga,^ it prefers lakes which are supplied
during the whole summer with water from glaciers. It is
very sluggish, only moving to breathe and when in search of
food, which consists of worms and insects. The general colour
is greenish brown, dark above ; the under side of the head and
body are bright orange red in the female, yellow in the male ;
dark spots separate this bright colour from the flanks. The tail
has a narrow ventral stripe of briglit red and yellow. The
cloaca of the female is bright red, that of the male dull grey.
The total length amounts to about 4 inches or 10 cm.

The pairing time is the end of June, or later in cold seasons.
The male gets hold of the female by forming a noose with its
tail round her ; it lies underneath, the cloacae being pressed
together so that the spermatozoa can be taken in directly. The
larvae have large yellow-green spots on the back and sides,
and a bright red ventral tail -fin ; when metamorphosed the
greenish spots become more confluent on the back, producing a
broad spinal band. Larvae which live in deep water are dark,
while those in sunny places are light-coloured and spotted with
yellow.

T, montambs in Corsica and T, rusconii in Sardinia are allied
forms, but the males are distinguished by a spur-like process or
dilatation at the end of the fibula.

2\ xvaltliy the Iberian Newt, is olive-brown above, yellowish
with blackish markings below. Tlie tail has a yellow or orange
ventral line. There is no crest. A remarkable peculiarity of
this species (which it shares only with Tylototriton aTidersoni
of the Loo-Choo Islands) is its ribs, which are very long,
sharply pointed, and frequently perforate the skin. Before

» P.Z.S, 1895, p. 150.

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SALAMANDRINAE

131

jierforation the point of the rib lies in a lymphatic space. This
surprising feature has by raany authorities been considered as
abnormal or pathological Certainly young, and even many
adult, individuals are found in which the skin is not perforated,
but when these are handled the wriggling motions of this strong
newt force the points of the ribs through the skin, and they
remain sticking out to the extent of several millimetres. The

SEliujrltX^

Fig. 25. — Triton walUL Spanish Newt aclult and larvae x \

wounds heal up, the skin forming a neatly linished-off hole
through which the spike projects, not as a formidable, but as a
sufficiently awkward, protective weapon.

Large females reach a length of 10 inches. The larv^ae
metamorphose, as a rule, when they are between 2 and 3 inches
long, but those which have been bred in tanks often reach
double this length. These newts are frequent inhabitants of
the rain-water cisterns common in the South of Portucral and
Spain, into which they tumble without ever being able to get
out again. This species spends most of its time in the water.

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132 URODELA chap.

preferring ponds, among the vegetation of which they can 1«
watched lying motionless, with their limbs hanging down and with
the head close to the surface ; but they are lively during the night
When their ponds dry up they leave them, crawling into the
most unexpected places, to aestivate under rocks, or even in the
walls of old buildings, where they are found by accident only.
The range extends from Central Spain and Portugal into
Morocco.

Tylototriton verrucosus lives in the Eastern Himalayas and in
the mountains of Yunnan. The skin is tubercular, with large
parotoids ; above uniform black-brown, pale below ; the tail has
a ventral yellow or orange line. Total length about 6 inches.
T, andersoni of the Loo-Choo Islands is remarkable for the
pointed ribs which perforate the skin.

Fiichytriton hrevipes, discovered in Kiansi, Southern China,
has a smooth skin, olive-brown above, with many black dots;
the under parts are yellowish, dotted with black. Total length
about 7 inches.

Fam. 3. Proteidae. — The three pairs of fringed external
gills persist throughout life. Both fore- and hind -limbs are
present. The eyes are devoid of lids. The maxillaries are
absent. Teeth are present on the premaxillaries, on the Vi>mers,
and on the mandible. The vertebrae are amphicoelous.

This family consists of only three genera, with one s^iecies
in each.

Necturtis miiculatus s. Meiwhranchus lateralis. — The eyes are
functional, being covered by the thin transparent skin. The
limbs, although short, are well developed, and have four fingers
and four toes. The whole animal, which reaches the lengtli of
one foot, is quite smooth and slimy, brown with irregular dark,
blackish spots and patches, which frequently form a dark lateral
band extending from the mouth to the tail. The latter, which
measures about one-third of the whole length, is strongly com-
pressed, curries a thick dorsal and ventral fin, and is rounded otf
at the end. The skin of the throat forms a strongly-marked
transverse fold. The thick stalks of the gills are brown, while
the numerous and delicate fringes are dark red in life ; Ijeneath
and behind them are two gill-clefts. K ^naculatus is found in
the eastern half of the United States, chiefly the eastern part
of the basin of the ^lississippi and the Canadian lakes.

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PROTEIDAE 133

These creatures are rather dull ; they remain mostly at the
bottom of the water, more or less concealed in the weeds or
between rocks during the daytime. Mine, which are kept in a
roomy, light-coloured tank, lie motionless, with their gills spread
out transversely. Every now and then the gills contract
suddenly and become i>ale, whereupon they are filled again with
blood. Very i-arely they rise to the siu:face, but tiny air-
bubbles are let out more frequently, especially when the animals
are disturbed. Then the gills collapse, are laid flat against the
neck, and the creature darts about with quick, eel-like motions.
At night thSy leave their hiding-places, swim about or creep
along the ground with slow, undulating movements, the limbs
l)eing scarcely used, in search of food, which in their wild state
consists of rather large Crustacea, small fishes, worms, insects and
frogs. They are most voracious, and absolutely indifferent to
cold. The spawning takes place in the months of April and
May.

Proteus arufuinMS. — The fore- and hind-limbs are fully de-
veloped and possess only three fingers and two toes. The eyes
are completely hidden beneath the opaque skin. This peculiar
creature is restricted to the subterranean waters of Carniola,
Carinthia, and Dalmatia. The vast caves of Adelsberg not
far from Trieste are especially celebrated for the occurrence of
the " 01m," the German name of this animal. The river
Poik, a moderate mountain -stream, but a large, tierce torrent
during the rainy season, disappears into the limestone-hills, and
rushes through enormous stalactite-grottoes, most of which have
been only partially explored, until several miles farther on it
reappears on the surface. There, deep down below the surface,
in absolute darkness, in an almost constant temperature of about
50" F. is the home of Proteus.

Their total length is scarcely one foot. The whole body is
white, occasionally suffused with a slight fleshy, rosy tinge,
wliile the three pairs of gill -bunches are carmine -red. They
are easily kept in captivity, and live for many years, provided
three conditions are strictly adhered to, viz. fresh and clean
water, an equable low temperature of about 50° F. = 10° C. and
darkness. The question of food is not so very important, since
specimens are known to have existed for years, although they
refu.sed to take any nourishment. How far darkness is an

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134

URODELA

absolute necessity is not known. Anyhow, the white skin is
almost as susceptible to light as is a photographic plate. If
light is not absolutely excluded the white skin becomes in time
cloudy, with grey patches, and if kept exposed to stronger light,
the whole animal turns ultimately jet-black. Mr. Bles has
succeeded in producing several totally black specimens, having
kept them for several months in a white basin under oixiinary
conditions of light. Xo experiments have yet been made to
find out if the black pigment deposited is lost again in darkness.
Those which are kept in a tank in ah absolutely dark cellar of
the Cambridge Museum, with permanent water-supply, are doing
very well When approached with a candle they become rest-

Fir.. 2^. — Proteus anguinvs. x I. Front view of the month in the left upper corner.

less or remain partly hidden in all sorts of seemingly most un-
comfortable attitudes, squeezed in between the sharp-edged tiles
and drain-pipes with which tlieir lodgings are furnished. But
the introduction of a wriggling worm, a little crustaoean or
other live bait draws them from their liiding-places, and, guided
by the motions of the prey in the water, possibly also by the
sense of smell, they snap it uj) and devour it.

If the water is not sufficiently well aerated, they rise to the
surface, emit a bubble of air, and take a new supply into their
lungs. As a rule they remain motionless under water, but the
gills contract spnsmodically and become paler, whereupon they
till again with blood and darken ; the contrast Ix^tween the pure
white body and tlie carmine-red feathery gills is very beautiful

Until recently the mode of propagation was quite unknown.
Several Proteus, kept by E. Zeller, laid, in the middle of April,

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PROTEIDAE 135

a number of eggs which were then fastened singly on to the
under side of projecting stones in the water. The pale yellow
yolk measured 4 mm. in diameter and was surrounded by a
cover of 1 mm. in thickness, besides an outer gelatinous mantle,
so that the whole egg measured about 11 mm. The larvae
were hatched after 90 days ; they were 22 mm. long, and
already much like the adult, except that the fin was not
restricted to the tail, but extended over the last quarter of the
trunk, and that their eyes were still visible. The fore-limbs
were already typical in shape, but the hind-limbs were still toe-
less little stumps.^

Typhlomolge rathhuni. — It is of the greatest interest that
a subterranean Perennibranchiate newt, in all respects closely
allied to Protetcs, has recently been discovered in Texas. There
can be no doubt that similar conditions of life have produced
these closely resembling forms from Necturvs -like ancestors,
one in Europe, the other in North America, absolutely
independently of each other. The limbs of Typhlomolge are
long and very slender, the four fingers and five toes are thin,
free and pointed. The head is large, the mouth square. The
eyes are completely hidden and the whole animal is colourless
and white. The tail is furnished with a dorsal and a ventral
fin. The very deep gular fold is nothing but tlie pair of
united but large opercular flaps. The three pairs of gills are
remarkable for their blade -like stalks, while the gill -lamellae
proper are short and restricted to the tapering ends. Total
length about 75 mm., of which the head measures 15, the tail
32 mm.

This peculiar creature inhabits subterranean caves in Texas,
to judge from the fact that all the specimens hitherto known
have come up with the water of an artesian well 188 feet
deep, near San Marcos. According to Blackford,- " the legs are
used for locomotion and the animals creep along the bottom of
the aquarium with a peculiar movement, swinging the legs in
irregular circles at each step. They climb easily over the rocks
piled in the aquarium> and hide in the crevices between them.
All efforts to induce them to eat have been futile, as lias also
been the case with blind cave-fish in captivity, and they are

^ See also M. von Chauvin, ZcUschr, xciss. ZooL xxxviii. 1883, p. 671.
2 Nature, Ix. 1899, p. 389.

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136

URODELA

CHAP.

either capable of long fasts or live on infusoria in the water."
It seems more reasonable to suppose that these newts live upon
Crustacea, four kinds of which, all new to science, also came up
with the water.

Fam. 4. Sirenidae. — The three pairs of fringed external
gills persist throughout life. The body is eel-like. Hind-limbs
are altogether absent, while the fore-limbs are short and have
three or four fingers. The maxillary bones are absent. With
the exception of small teeth on the vomer the mouth is tooth-
less, but the jaws are furnished with homy sheatha The eyes
are devoid of lids, but shine through the skin.

The Sirenidae are the most degraded members of the Urodela
and are represented by two closely-allied genera, each with one

Fig. 27. Siren lacertirw. x ).

species, in the south-eastern parts of the United States. Their
most interesting feature, which bears upon the question of
neoteny, is their retrograde metamorphosis as described by
Cope.^ The gills atrophy in tlie young and are subsequently
redeveloped. Cope therefrom concludes rightly that the ultimate
or persistent gills of Siren are signs of maturity and not a larval
character. In young specimens of Siren of 5 to 6 inches in
length the gills are functionless ; in one of 3 inches they were
found to be entirely vestigial and " subepidermal," ix, covered by
a common dermal investment. Unfortunately really young
larvae are still unknown. Old Sirens can live without gills, as
has been shown by aquarium-specimens. In the adult Pseudo-
hranchus all the gills are normally covered up by an investment
of the skin so as to be quite without function and movability.
Siren larertina, the " uiud-eel/* is distinguished by the
* Amer. Natural, xix. 1885, p. 1226.

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SIRENIDAE 137

possession of three pairs of gill-clefts and by its four lingers.
It reaches a length of 70 cm., or about 2^ feet, of which about
one-third is taken up by the tail, which is strongly compressed
and finned. The skin is smooth, mostly blackish, lighter below,
sometimes with whitish specks all over the body. This creature
is frequently found in ditches and ponds, where it burrows in
the mud. When swimming the limbs are folded back. They
are said sometimes to leave the water and to crawl about on the
moist ground.

Pseudobranchus striatus has only one pair of gill-clefts and
only three fingers. The slightly granular skin is dusky brown
above, with a broad yellow band on either side and with a paler,
narrower stripe below. Total length about 7 inches.

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CHAPTER VI

LISSAMPHIBIA (^CONTIXUED) ANURA

Order m. ANURA or TAILLESS AMPHIBIA

The recent tailless Amphibia, or Frogs and Toads in the widest
sense, contain such a great number of species (about 900), with
such a diversity of characters, that it is necessary, if only for the
sake of mere convenience, to group them into a considerable
number of families and sub-families. The characters avaikble
for this purpose are few.

1. The possession of a tongue characterises the Phaneroglossa, the abeenc*'

of a tongue the Aglossa.

2. The character of the shoulder-girdle. — Overlapping of the two

halves of the shoulder -girdle on the ventral side characterises the
Arcipera, while in the Firmisternia the two ventral halves meet in
the middle line and form a firm, median bar. See, for details, \\ 24.

3. The shape of the transverse processes or diapophyses of the sacral

vertebra which carries the iliac or hip-bonea These processes are either
dilated or cylindrical.

4. The presence or aljseuce of teeth in the upper and lower jaws. This is

indicated by a formula in which 0 means absence of teeth ; max. means
presence of teeth in the upper jaw ; mand. means presence of teeth in the
lower jaw.

5. The terminal joints or phalanges of the fingers and toes are some-

times claw-shaped. See p. 26.

6. The shape of the centra of the vertebrae. — Opisthocoelom, if the

posterior end is cui)-sha])ed or concave, procoelous if the anterior end Is
concave and the posterior is convex. See p. 19.

By means of these charactei^s we can arrange the Anura in
the following key :- —

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CLASSIFICATION 1 39

^ja&itS: W^<^^^> "• ^"-

I. Aglossa. Sacral <

Vertebrae op"
II. Phanero^Iossa.
A. Arcifera.

a. Sacral diapophyses dilated.

a. Terminal phalanges not claw-shaped.

Opisthocoelous, with ribs, '^^ }dISCOGLOSSIDAE, p. 162.

Procoelous, withotlt ribs, ^JBUFONIDAE, p. 166.
Procoelous, or opisthocoelous, "j

witKout rib«. "5^ JPELOBATIDAE, p. 160.

[ max. AmphigTuUhodan-

fi. Terminal phalanges claw-shaped— J maud- ' ^***^» P- ^®®-
HYLfDAEj^max,^^^^ p ,gj^

max. Hemiphractinae^

h. Sacral diapophyses cylindrical —

CYSTIGNATHIDAE

B. Finnistemia.

a. SiAcral diapophyses dilated —

ENGYSTOMATIDAEl

mand. p. 210.
max. Cytlignathinae,
~0 p. 211.

0 Dendrophryniscinae,
0 p. 227.

^^^^^ Dyacophinae, p. 235.

0 OenyophryniTUUy
SiEd. p. 236.

- Engystomatviiaef p. 225.

/ max. CeratobcUraMjiar,
nmnd". P- 287.

b. Sacral diapophyses cylindrical— j "^<*^- Raninae, p. 238.

RANIDAE I 0 ^

' - Betidrobatinaet p. 272.

Concerning the evolution of the classification of the Anura,
it is interesting to follow the changes of the value attached
to the various anatomical characters by systematists. At first
the presence or absence of teeth and of adhesive discs on the
fingers and toes were considered to be of prime importance for
the division of the Phaneroglossa.

Dmndril et Bibron, 1 84 1. "Erp^tologie gen^rale."

L Phrtnaglosses = Aglossa of Wagler: Pipa and Xenopus,
II- PHAN^ROOLOdSEs. I. With teeth, a. Without discs : Raniformes.

6. With di'^cs : Hylaeformes.

2. Toothless .... Bufoniformes.

Stannins, 1856 (see p. 8), separated the Engystomatidae as " Systomata,'

and used the presence or absence of the "manubrium stemi" (omo-

stemum) as a character of distinction between his Bufoninae and Raninae.

1 1

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I40 ANURA

Giinther, 1858, "Catalogue of the Batrachia Salientia." No progress was
made by his scheme, which relied upon the tongue and digit*.
Aglossa with Myohatrachus,
Opisthoglossa. a. Oxydactyla. 6. Platydactyla.
Proteroglortsa : Rhinophrynidae.
Cope, 1864. "Oil the limits and relations of the Raniformes." ^ He
introduces the shoulder-girdle and the sacral diapophyses, and drop*
the discs as too adaptive and misleading. He distinguishes between
Raniformes and Arciferi.
Cope, 1865. "Sketch of the primary groups of the Batrachia Salientia." ^
Aglossa. •

Bufoniformia (Bufonidae).

Arcifera (DiscpgLossidae, Scaphiopodidae, and Hylidae).
Raniformia.
In 1867 Cope separates the genus Hemisus as Gastrechmia on account of it3

jieculiar pectoral arch.^
In 1875, "Check-list of North American Batrachia and Reptilia," Cope
elaborates his system :

Class Batrachia. Order Anura.

1. Raniformia.

2. Firmistemia. [Dendrobatinae and Engystomatidae.]

3. Gastrechmia : Heni'Uus,

4. Bufoniformia. [Bufonidae.]

5. Aglossa. Pipa.

6. Odoutaglossa. Xenopus.

7. Arcifera, [Cystignathidae, Hylidae, Pelobatidae and Disco-

glossidae.]
Cope consequently considered the characters of the pectoral arch as equi-
valent to those of the dentition.
Houlen^'er, 1882, "Catalogue of the Batrachia Gradientia & Ecaudata,''
recognises that the pectoral arch is of greater systematic value than the
dentition. The latter is used, together with the shape of the sacral
•liapophyses, for the seiwration into families.

f 1. Ranidae.

2. Dendroljatidae.

3. Engystomatidae.

4. Dyscophidae.

5. Cystignathidae.

6. Dendrophryniscidae.

7. Bufonidae.

8. Hylidae.

9. Pelobatidae.

10. Discogloesidae.

11. Hemiphractidae.

12. Amphignathodontidae.
II. Agloss. . . . (^3- Dactylethpidae.

I. Phaneroglossa. A. Firmistemia,

Jj. Arcifera.

114. Pipidae.

Pruc. Ac. Phihui. 1864, p. 181.
The Katnral History Peview, No. xvii. 1865, ]►. 97.
Journ. Ac. Nat. Hist. Philod. vi. p. 189.

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VI CLASSIFICATION I4I

This emendation of the Arclfera and Firmisternia was accepted
by Cope in his synopsis of the families of Vertebrata {Amer.
Xatural. xxiii., 1890), except that he atill retained his suborder
Grastrechmia.

Since the publication of Boulenger's great work a number of
forms have been discovered which, from the characters of their
dentition, have necessitated the establishment of certain new
families, namely, Ceratobatrachidae and Genyophiynidae ; and
Boulenger was the first to recognise that the taxonomic value
of the mere presence or absence of teeth in the jaws liad been
overestimated. I therefore propose using it as a character
^stinctive of the sub-families only, thereby reducing the number
of families, relying first (leaving the Aglossa aside) upon the
firmisternal or arciferous condition of the pectoral arch, secondly
upon the dilated or cylindrical shape of the sacral diapophyses,
thirdly upon the dentition. Blindly consistent application of
these principles would reduce the Phaneroglossa to four families
only, namely Eanidae, Engystomatidae, Cystignathidae and a
fourth family comprising all the Arcifera with dilated sacral
diapophyses. This would obviously be wrong. We have there-
fore to resort to other additional characters or rather peculiarities.
The opistfaocoelous character of the vertebrae and the possession
of distinct ribs, together with the disc-shaped tongue, sepa-
rate the Discoglossidae and justify their retention as a family.
The Hylidae are marked off by the claw -shaped terminal
phalanges, but the remaining forms, comprising the Bufonidae
and Pelobatidae, cannot be separated except by their dentition,
and I plead guilty of inconsistency in i-etaining them as separate
families.

After all, our classification may not represent tlie natural
system, and it may be nothing but a convenient key.

When we have eliminated the characters of the vertebrae, the
dentition, the claw-shaped phalanges and the adhesive discs, it
may well be asked what characters remain. The firmisternal is a
further, higher modification of the older, more primitive arciferous
condition. The difference between the dilated and eylindiical
shape of the sacral diapophyses is in not a few cases very slight,
and there are various, most suggestive exceptions. The presence
or absence, size and shape of the omosternum and metaaternuni
are of very limited taxonomic value, not always applicable to all

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142 ANURA

the members of the same family. The fact is, that the Anura
are a very recent and a most adaptive, plastic group. The earliest
known fossils are scarcely older than the Middle Eocene.

Almost every one of the greater families has produced terres-
trial, arboreal, aquatic, and burrowing forms. Their habits have
modified, and are still shaping their various organs, first of course
those by which the animals come first and most directly
into contact with their surroundings {e.g, adhesive discs, denti-
tion, general shape of the body, length of limbs, wartiness of
the skin, tympanic disc). These are the so-called adaptive chanic-
ters, sometimes decried as merely physiological ; as if habits,
use, and requirements did not likewise influence and ultimately
model every other organ {e.g, tympanic cavity, Eusttu^hian tubes,
vertebrae, ribs, coccyx, pectoral arch, etc.). There are true Toads,
Bufonidae, which are as smooth, wartless, slender-bodied and long-
legged as the most typical of " Frogs"; true Eanidae, like Rhaco-
phorus, which by their green colour, large adhesive discs and
arboreal habits may well put any of the Hylidae to shame.
Ceratohyla has developed the claw-shaped terminal phalanges
which are otherwise typical of, and peculiar to, the Hylidae, but
this genus reveals itself by various details as a close relation of
the other Hemiphractinae ; and tliese fall in with the Cysti-
gnathidae on the strength of their cylindrical, not dilated, sacral
diapophyses.

In sketching the phylogenetic tree of the families of the
Anura we have to proceed with great caution.

There is not much doubt about the Aglossa. They have
retained some of the most primitive character, but liave by now
been so much modified and specialised that they are to be looked
upon as an early side-branch.

Among the Phaneroglossa the Discoglossidae are with certainty
the oldest, but are now scarce in genera and species, and much
specialised. The Pelobatidae connect them with the Bufonidae.
The Cystignathidae form a rather ill-defined assembly which
points downwards to the Pelobatidae, upwards to the Hylidae.
There is no divergence of opinion about the Eanidae being the
highest of all the Anura, and amongst them the Raninae the
most typical, the Dendrobatinae the most specialised. If we
assume that moderately dilated sacral diapophyses represent a
more primitive stage tlian cylindrical processes, we shall natu-

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AGLOSSA

143

rally look to the Engystomatidae as the connecting link between
the Eiiuidae and the Arcifera, through Bufonoid creatures still
with teeth in both jaws. If, on the other hand, we take the
dilatation to be a further development from more or less
cylindrical processes, then the Kanidae can be considered as
having sprung from Cystignathoid creatures, which have con-
solidated their pectoral arch into the firmistemal condition ;
and in this case the Firmisterniu would not be a natural
group, the Engystomatidae pointing, to the Bufonoid stock.
This would, to a great extent, mean a reversion to Cope/s
idea.

Sab-Order 1. Aglossa. — The two diagnostic peculiarities
of the few members of this group are : first, the absence of
a tongue; secondly, the union of the Eustachian tubes into
one median pharyngeal opening in the posterior portion of
the palate.

The pharyngeal opening and the tubes themselves are wide,
the tympanic cavities are present, but the tjnnpanic discs are
not distinct from the

XENorus.

rest of the skin. The
fronto-parietal bones are
fused into one mass, a
rare feature in the
Anura. The nasals are
large. Fipa and Jfyme-
Twchirus have no teeth,
Xen^pus has teeth on
the upper jaw. The
vertebrae are opistho-
coelous and typically
epichordal in their de-
velopment ; the second, third, and fourth carry long ribs, which in
old specimens fuse with the supporting diapophyses. The sacral
diapophyses are enormously dilated, and the sacrum is fused with
the OS coccygeum. The serial number of the sacral vertebrae
exhibits a most interesting gradation. In Xenojpus the ilium
is carried by the diapophyses of the 9 th, in Pipa the 9 th and
8 th, in HyiTunochirus the 7 th and 6 th. In these cases the
two diapophyses of each side are fused together into a single
broad blade, and their original duplicity is indicated only by the

!!l«inPA.

Fig. 28. — Map showing distribution of Aglossa.
Uynvenoehirus to he added in Equatorial Africa.

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144 ANURA CHAP.

holes for the spinal nerves. Hymenochirus has consequently
only 5 presacral vertebrae, the vertebral column being shortened
to the greatest extent known amongst Vertebrata. For further
information see p. 22. The ilia are much broadened vertically,
and are firmly attached to the sacrum. The shoulder -girdle
is sometimes described as of the arciferous type, but this is quite
unjustifiable. The epicoracoid cartilages do not overlap each
other, but meet, and partly fuse in the middle line. The
three genera exhibit some differences. In Pipa and Hymeno-
chirus the bony portions of the coracoids are much expanded
dorsally, and there is a considerable amount of epicoracoid
cartilage, that of the precoracoid bars extending backwards as
a broad-based and blunt omosternum. Xeno2)U8 is devoid of
an omosternum, and the configuration of the whole apparatus
is more slender. The metasternum of Xeriopvs and Hymetw^
chirus broadens out laterally. Hymenochirus greatly resembles
Breviceps, a genus of Engystomatinae, in the relative position
and size of the various parts of the shoulder -girdle and
sternum.

The tibio-fibula of Hymenochirus has a wing-like expansion of
thin bone on each side, forming a deep groove on the outer aspect.
The astragalus and calcaneum are united by a similar bony
expansion with wing-like projections

The lungs are remarkable for the prominent development of
trabecular projections and niches, so that their free lumen is
much restricted; they have thereby reached a much higher
stage than in any other Amphibia or even many Autosauri. The
persistence of an arteria sacralis s. caudalis, a vessel absolutely
absent in the adult JRana, is a primitive feature, and the same
applies to the presence of a true first spinal or suboccipital
nerve.

The skin of the back and belly is supplied by two great
branches from the arteria anonyma, one arising proximally, the
other distally from the subclavian ; herewith is correlated the
almost complete absence of the arteria cutanea magna, which as a
branch of the ductus pulrao-cutaneus plays such a prominent
role in the other Anura. Only in Fipa, but not in Xenopus, is the
great cutaneous vein represented by a very small branch. Both
these genera possess a much more complicated " diaphragm " than
tlie other Anura, chiefly owing to a special muscle which arises

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AGLOSSA 145

from the anterior end of the ilia and spreads out fan-like to
the oesophagus and to the bases of the lungs.^ This diaphrag-
matic arrangement is correlated with the great development of
the lungs, and is not a primitive but an advanced feature. It is
reasonable to suppose that this has caused the reduction of the
usual arteria pulmo-cutanea, and that the other two cutaneous
arteries have been developed secondarily. The Aglossa are
generally considered as the lowest Anura, and only Cope looked
upon Fipa and Xenopus as two convergent terminal branches.
Beddard came to the conclusion that both are closely related to
each other, chiefly on account of their peculiar diaphragmatic
arrangement. The whole question has entered upon a new stage
since the recent discovery of Hymeiiochirus, which is in many
ways intermediate between the two other genera. Moreover, the
mid-Tertiary Palaeobatrachvs of Europe is undoubtedly related to
them, and we conclude now that all these four genera belong to
one group with a distribution formerly much wider than Africa
and part of South America. But this does not necessarily mean
that the Aglossa are in all respects the most primitive group
of living Anura. On the contrary, they possess few decidedly
primitive chai-acters, namely, the long typical ribs, the presence
of the first spinal nerve, the unimportant persistence of the
arteria sacralis, and lastly, the possession in the tadpoles of a
right and left opercular " spiracle." The absence of the tongue
cannot possibly be an archaic feature, considering its universal
presence in all the other Amphibia, including the Apoda, and
the suggestive circumstance that this organ is least developed in
the entirely aquatic members of the Urodela. In fact, thoroughly
aquatic creatures, which seize and swallow their prey under water,
require no elaborate tongue ; and since we know that the Anura
must owe their typical formation to terrestrial life, it follows that
those which have again taken to the water and are tongueless,
have lost this organ. As I have shown elsewhere,* the epichordal
development of the vertebrae is likewise a secondary feature, far
from primitive; and the tendency of the shortening of the
vertebral column, which has reached its extreme in Hymeno-
chirus, points to the same conclusion. The apparatus of the
shoulder-girdle and sternum is in the last transitional stage from
the former arciferous to the typically consolidated firmisternal

> Beddard, P.Z.S, 1895, p. 841. » p^^;, j^rans. B. 136, 1896, p. 1.

VOL. VIII L

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146 ANURA

type. In fact there is little left which is primitive, but much
that is very specialised and highly developed in the Aglossa,
mostly in adaptation to their absolutely aquatic life, to which they
must however have taken very early. They are in a position
somewhat analogous to the Eatitae among Birds, which are like-
wise an old group, although many of their most striking features
have been acquired secondarily.

Xenopus s. Dactylethra, The upper jaw is furnished with
teeth. The ilia are attached to the ninth vertebra. The pupil is
round. The terminal phalanges are pointed. The fingers are
free, the toes broadly webbed, and the first three are covered with
sharply pointed, horny, black -brown nails, a feature which is
alluded to by the alternative generic names. A cutaneous
tentacle projects from below the eye and naturally invites com-
parison with the tentacle of the Apoda and of Urodela. The
skin is smooth, rich in mucous glands, besides certain tube-
like apparatuses, possibly sensory, which are scattered over the
body, especially on the head, and form a conspicuous series of
white dots along the dorso-lateral line, from the eye to the vent.
The general colour of the upper parts is olive brown, mottled
darker, while the under parts are whitish. The female has
three cutaneous flaps closing the vent. The male develops black
nuptial brushes along the inner side of the fingera There are
several species, all African (Ethiopian).

X. laevis, ranging from the Cape to Abyssinia, is distinguished
by the absence of a metatarsal spur. The tentacle is very short.
Size about 3 inches. X. mudleri of Zanzibar and Benguella,
is smaller. The tentacle is conspicuous, as long as the diameter
of the eye. The inner metatarsal tubercle carries a sharp claw.
X cal curat us of tropical West Africa is only 2 inches long, and
has strong metatarsal claws, short tentacles and very minute eyes.

The habits and oviposition of the " Clawed Toad " have been
described by I^^slie.^ The Boers call it " Plathander," i.e. flat
hand. Entirely aquatic, it rests floating in the water, with the
nostrils exposed, and leaves the water only if it has to change the
locality on account of drought or scarcity of food. The pairing
takes place, at least at Port Elizabeth, in the early spring, t.^. in
the month of August. The only sound which is emitted is heard
during this time, a very slight and dull tick-tick, audible at only

1 P.Z.S, 1890, p, 69.

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148 ANURA

a few feet distance. The male grasps the female by the loins ;
the eggs are extruded singly, measuring only 1.5 mm. in diameter,
but swell to double that size. They are attached singly to stones
or wateisplants.

Latterly these creatures have frequently been brought over to
England. They stand confinement very well, even in a little
aquarium with sufficient water-weeds to keep the water fresh ; and
they do not require special heat. They greedily snap up worms,
strips of liver, or meat, and poke the food in with their hands.
A few kept by Boulenger in a glass jar have lived for the last
eleven years in the ordinary temperature of a room in London.
Curiously enough they are often in amorous embrace, regardless
of the season, but they have never shown any signs of spawning.

Some of those in the Zoological Gardens in London laid ^gs
on Saturday the 27 th of May, and on the morning of the follow-
ing Monday the larvae were already hatched. They have been
described by Beddard.^ The larvae are provided with an unpaired
circular, ventral sucker. The tentacles begin to sprout out on
the sixth day after hatching, at first not in connexion with the
cranial cartilage, but soon a cartilaginous rod runs into the
tentacle from the ethmoid "just above the joint with the under
jaw." Boulenger has most reasonably compared these organs
with the "balancers" of Triton and Avihlystomn (cf. p. 46 for
the possible homologies of the balancers). The tentacles soon
reach a great length and give the tadpole a curious appearance.
In tadpoles of X, calcarcUus, 65 mm. long, the tentacles
are 30 mm. long, and are inserted just at the angle of the
mouth. By the time that these tadpoles show their fore-limbs,
the feelers are reduced to 4 mm. in length, and their relative
position has been shifted to a little above the angle of the gape,
and whilst the latter gradually extends further and further back,
the feelers come to lie, or rather remain, below and a little in front
of the eyes.

The tadpoles have no traces of horny teeth. External gills
project as low conical or lamellar processes from the first three
branchial arches, but so-called internal gills are not developed.

Amongst a number of Clawed Toads imported in the spring
one female became swollen with eggs, but as they did not show
signs of wanting to breed, a pair was put into the tropical tank
1 P.z.s. 1894, p. 101.

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AGLOSSA 1 49

in the Cambridge Botanic Gardens, a transfer which had the
desired effect. Eggs were laid, and more during the following
nights ; they hatched out within thirty hours. The whole brood
was lost, before any of them were older than a few days, since they
were attacked, beyond the possibility of a cure, by a Sajyrolegnia
or some similar pest.

Si/menochirtcSy represented by one species, If. boettgeri, has
been discovered in the Ituri, German East Africa, and in the
French Congo, and has no doubt a much wider distribution.
It is scarcely 1^ inch long, and is easily recognised by the
toothless mouth, the half-webbed fingers (hence the generic
name), the incompletely webbed toes, the third of which is
longer than the fourth, and the absence of sensory muciferous
canals in the skin. The three inner toes are, as in Xenopus,
furnished with small black claws. The skin is rough, beset with
small granular tubercles. The general colour above and below
is olive-brown. The vent is, as in Xenopus, produced into a
spout or semi-canal, but is devoid of dorsal flaps of skin.

PipcL — This Neotropical member of the Aglossa is quite tooth-
less, but the jaws of the adult have horny substitutes. The only
species is P, americana, the famous Surinam Toad, chiefly
known from the Guianas, but undoubtedly extending much
further, having recently been reported from the neighbourhood
of Para.

The general shape of this creature is veiy peculiar. The head
is much depressed and triangular ; the eyes are ver}^ small ; the
skin forms several short, irregularly-shaped flaps and tentacles
on the upper lips and in front of the eye, and at the angle of
the mouth. The tympanum is invisible. The pupil is round.
The fingers are very slender and free, ending in star-shaped tips ;
the toes are broadly webbed. The whole skin is covered with
small tubercles and is dark brown above, while the under parts of
the very flat and depressed body are whitish, sometimes with a
dark brown stripe along the middle line. In the female the skin
of the back forms growths for the reception of the eggs, and in
these the young undergo their whole metamorphosis.

The most characteristic feature of the skin,^ which has
exactly the same structure in both sexes, is the papillae, which

* Groenberg und Klinckowstroein, *'Ziir Anatomie der Pipa americana,'* Zool.
Jahrb. Anal, vii. 1894, p. 609.

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CHAP. VI AGLOSSA 1 5 1

are spread over the whole surface, except on the webs of the toes,
on the cornea and on the star-shaped points of the fingers. Each
papilla carries a little horny spike, and a poison-gland frequently
opens near its basa Larger poison-glands exist on the dorsal
and ventral side in four rows, and smaller glands open upon the
sides of the body, but there are no parotoid complexes. Slime-
glands occur all over the surface. The epidermis consists of the
usual layers, namely the Malpighian, the stratum corneum, and
the part which is shed periodically. The latter is completely
horny, appearing to be structureless like a cuticle, but it is in
reality composed of polygonal cells with flattened nuclei ; each
little spike is one modified horny cell. The whole outer-
most layer contains black-brown pigment. The upper portion
of the cutis is devoid of pigment, then follows a layer of
clusters of ramified dark pigment -cells, and lastly the rest of
the cutis.

Each of the four fingers ends in a four-armed star, the tips of
which again carry four or five sensory papillae. The cartilage of
the terminal phalanges is correspondingly star-shaped.

According to Klinckowstroem these toads, which are entirely
aquatic, are easily collected at the end of the long dry period,
when they are all confined to the half-dried-up pools. But
they do not spawn there. This happens after the rains have
inundated the forest, and then it is very difficult to get the
females with eggs on their backs. Each of the eggs, when
once they have been glued on to the back, sinks into an in-
vagination of the skin. The initial stages are probably the
same as those caused by the eggs on the belly of Rhacophorvs
reticulatus (see p. 248). Later, each egg is quite concealed in a
cavity with a lid. These cavities are simply pouches of the skin,
and are not formed by enlarged glands as has been suggested
by some anatomista Each cavity consists of the epidermal
pouch and the lid. How the latter is produced is not known.
According to the €^thors quoted above, the lid looks like a shiny
or sticky layer which has hardened into horn-like consistency.
It lies exactly like a lid upon the rim of the pouch itself,
and is certainly not in structural or organic continuity with
the epidermis. Most probably it is produced by the remnant
of the egg-shell itself, which, after the larva is hatched, is
cast up to and remains on the top of the cup.

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152

ANURA

Bartlett^ has described the spawning of specimens in the
Zoological Gardens in London.

" About the 28th of April 1896 the males became very Uvely,
and were constantly heard uttering their most remarkable
metallic, ticking call-notes. On examination we .then observed
two of the males clasping tightly round the lower part of the
bodies of the females, the hind parts of the males extending
beyond those of the females. On the following morning the keeper
arrived in time to witness the mode in which the eggs were
deposited. The oviduct of the female protruded from her body
more than an inch in length, and the bladder-like protrusion
being retroverted, passed under the belly of the male on to
her own back. The male appeared to press tightly upon this
protruded bag and to squeeze it from side to side, apparently
pressing the eggs forward one by one on to the back of the female.
By this movement the eggs were spread with nearly uniform
smoothness over the whole surface of the back of the female
to which they became firmly adherent. On the operation
being completed, the males left their places on the females, and
the enlarged and projected oviduct gradually disappeared from
one of the females. In the other specimen, the oviduct appears
not to have discharged the whole of the eggs."

Boulenger, who examined this second specimen, which died,
confirmed this egg-bound condition. He remarks further : " The
ovipositor formed by the cloaca (not by the prolapsed uterus),
was still protruding and much inflamed. It may be deduced
from the observation made by the keeper, that fecundation must
take place before the extrusion of the eggs, and it is probable
that the ovipositor serves in the first instance to collect the
spermatozoa which would penetrate into the oviducts, the
eggs being laid in the impregnated condition, as in tailed
Batrachians."

Sub-Order 2. Phaneroglossa — ^Fam. 1. DiscogloBsidae.—
The tongue has the shape of a round disc, adherent by nearly
the whole of its base, and it cannot be protruded. The vertebrae
are opisthocoelous, and in the aquatic genera are of the most
exaggerated epichordal type ; the diapophyses of the second to
the fourth vertebrae carry short, free ribs, and those of the sacral
vertebra are dilated. The metasternum behind is forked. The

1 P.Z,S, 1896, p. 595.

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PHANEROGLOSSA DISCOGLOSSIDAE I 5 3

upper jaw and the vomers are provided with teeth. The males
have no vocal sac. The tadpoles are distinguished by having
the opercular spiracle placed in the middle of the thoracic region
(see general anatomical part, p. 44).

The few members of this family have a peculiar distribution.
Liopelma is confined to New Zealand, where it is the solitary
representative of the Amphibia. Ascaphus is found in North
America. The other genera, Discoglossus, Bombinator, and Alytes,
are typical of the Palaearctic sub-region, and are, with the excep-
tion of Bomhinator, confined to the Western Provinces (cf. Map,
Fig. 32, on p. 161).

I>iscoglo88U8. — The tympanum is indistinct, being more or
less concealed by the skin. The pupil is round or triangular.
The omosternum is small. The vertebrae are of the epichordal
type.

D. pi4;tus, the only species, has a smooth and shiny skin,
provided with numerous small mucous glands. The palms of
the hands are provided with three tubercles, of which the inner-
most is the largest, and is carried by the vestige of the thumb.
The coloration of this species is very variable. The ground-
colour of the upper parts is a rich olive brown with darker, light -
edged patches, which are either separate or confluent in various
ways, forming broad, longitudinal bands, or a few larger asym-
metrical patches, separated in some individuals by a broad and
conspicuous light brown or yellowish vertebral strii)e. An
irregular reddish band frequently extends from the eyes back-
wards along the sides. The under parts are mostly yellowish
white. This variability is purely individual, the most differently
marked and variously coloured specimens being found in the same
locality and even amongst the members of one and the same brood.
The male develops various nuptial excrescences, consisting of
minute, dark, horny spines, notably on the inner palmar pad, on
the inner side of the first and second finger, on the chin and
throat, and smaller and more scattered spicules on the belly and
legs.

This pretty and extremely active little creature, which
measures between 2 and 3 inches in length, is confined to
the south-western corner of the Palaearctic sub-region, being
found in Algiers and Morocco, Sicily, Sardinia, Corsica, and the
southern and western parts of the Iberiair Peninsula. Curiously

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154 ANURA

enough it is absent in the Balearic Isles. Bather aquatic in its
habits, frequenting pools and streams, it is also often found
on land.

The male has a feeble voice, which sounds like"ha-a, ha-a-a," or
" wa-wa-wa," uttered in rapid succession. The pairing season lasts
a long time, in Algeria from January to October, but a much shorter
time in the north of Portugal, where it extends over the spring
and summer months. Boulenger has made extensive observations
on many specimens kept in captivity. The embrace, whicli
never lasts long, is lumbar. The eggs are small, 1 to 15 mm.
in diameter, dark brown above and greyish below, each sur-
rounded by a gelatinous capsule of 3-7 mm. in diameter. The
eggs are laid singly, and a set amounts to from 300 to 1000, the
whole mass sinking to the bottom of the pool. Each female lays
several times during the season. The eggs are developed very
rapidly, the larvae escaping sometimes after thirty -six hours,
but usually from the. second to the fourth day. The external
gills are lost on the seventh day, when the tadpoles are 11 mm.
long ; the hind-limbs apj^ear on the tenth, and after four weeks
the tadpoles reach their greatest length, namely from 25-o0 mm.
The fore-limbs appear on the thirtieth day, and a few days later
the most precocious specimens leave the water and hop about.
Others, however, of the same brood took from two to three months
in metamorphosing.

This species lives on insects and worms, and can swallow its
prey under water.

Bomhinator. — The tympanum is absent and the Eustachian
tubes are very minute. The pupil is triangular. The omosternum
is absent. The vertebrae are absolutely epichordal. The fingers are
free, the toes are webbed. The upper parts are uniformly dark, and
are covered with small porous warts. The general shape of the
head and body is depressed or flattened downwards. The habits
are eminently aquatic. This genus consists of three s]3ecies, two
of which are European, the third Chinese.

B. igneus. — The under parts are conspicuously coloured bluish
blac^k with large irregular red or orange-red patches ; the upper
parts are more or less dark grey or olive black. The iris is
golden, speckled with brown. The male has a pair of internal
vocal sacs by which the throat can be inflated ; nuptial ex-
crescences are developed on the inner side of the fore-arm and the

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niSCOGLOSSIDAE

155

first two fingers. Total length from 1^ to 2 inches, the males
l>eing generally smaller than the females. This " Fire-bellied
toad," the " Unke '* of the Germans, is essentially a native of
lakes, ponds, and other standing waters of the plains.

It ranges through the whole of North Germany, Bohemia,
and Hungary into Eussia, eastwards as far as the Volga. The
latter river, the Danube, and the Weser form, roughly 8[)eaking,
its boundaries : northwards it extends into Denmark and the
southern extremity of Sweden.

Fi(j. 31. — liombincUor u/netu. x 1. Fire-bellie<l Toad.
•• warning " attitnde.

Two of them in

B. pachyp^us. — The under parts are yellow instead of red. The
male is devoid of vocal sacs, but has nuptial excre.scences on
the under siurface of most of the toes, in addition to those
on the fore-arm and fingers. The " Yellow-bellied Toad " is the
representative of the red-bellied species in Southern and Western
Europe, preferring, although not exclusively, the hilly and
mountainous districts. It ranges from France and ]»elt;iuni
through South-Western Germany, continental Italy, and the whole
of Austria and Turkey in Europe. Where Ijoth species meet, for
instiince in the hilly districts between the Weser and the lUiine,
in Thuringia and in Austria, tlie predilection of the yellow-l»ellied

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IS6 ANURA

species for the hills, and that of the other for the plains, is well
marked. ,

AVhile B, igmus prefers standing waters with plenty of
vegetation, B. pachypus is often found in the smallest occasional
puddles produced by recent rain, for instance in the ruts of
roads. Botli species have otherwise much in common. They are
essentially aquatic They hang in the water, with their legs
extended, nose and eyes just above the surface, and bask or lie in
wait for passing insects, the fire-bellied kind preferring to con-
ceal itself in the vegetation of the margins of ponds. During
the pairing season, in Grermany in the month of May, they are
very lively and perform peculiar concerts, one male beginning with
a slowly repeated note like " hoonk, hoonk," or " ooh, ooh," in which
all the other males soon join, so that, when there are many, an
almost continuous music is produced. This sound is not at all
loud, a little moiu'nful and very deceptive. It appears to be a
long way off, certainly at the other end of the pond, until by
careful watching you see the little creature almost at your very
feet. But on the slightest disturbance the performance ceases,
they dive below and hide at the bottom. The yellow-bellied
kind, when surprised in a shallow puddle, skims over the mud,
di8turl)s it, and allows it to settle upon its flat body, so that
nothing but the little glittering eyes will betray its concealment.
AVhen these toads are surprised on land, or roughly touched, they
assume a most peculiar attitude, as shown in Fig. 31. The
head is partly thrown back, the limbs are turned upwards
with tlieir under surfaces outwards, and the whole body is
curved up so that as much as possible of the bright yellow
or red markings of the under parts is exposed to view. The
creature remains in this strained position until all danger seems
passed. In reality this is an exhibition of warning colours, to
show the enemy what a dangerous animal lie would have to deal
with. Tlie secretion of the skin is very poisonous, and the
fii-e-toiids are thercby well protected. I know of no creature
which will eat or even harm them. I have kept numbers in a
large vivarium, together with various snakes, water-tortoises, and
crocodiles, but for yciirs the little fire-bellies remained unmolested,
although they shared a pond in which no other frog or newt could
live without being eaten. Himgry water-tortoises stalk them
under water, touch the intended prey with the nose in order to

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DISCOGLOSSIDAE I S /

get the right scent, and then they withdraw from the Bomhinator,
which has remained motionless, well knowing that quick move-
ments, or a show of escape, would most likely induce the tortoise
to a hasty snap, with consequences to be regretted by both.

After they have been handled frequently, they do not readily
perform, but simply lie still, or hop away. Miss Durham ex-
perienced considerable difficulty in inducing her tame specimens
to assiune and to keep up the correct warning attitude. The
statement that they " turn over on the back " is a fable, graphic-
ally fixed in various illustrated works.

It has been said that these two species are diurnal and
thoroughly aquatic. They are certainly active in the daytime,
sing in full sunshine, and spend most of their time in the water,
but they display much more 'liveliness towards the evening and
during the night, especially when there is a moon. My fire-toads
live by no means always in the water, but conceal themselves
in the daytime under stones, while they are regularly all astir
at night in search of worms and all kinds of small insects.

The spawning takes place several times during the spring
and summer. The amplexus is lumbar, and the eggs are
extruded singly. They sink to the bottom, or are attached to
water-plants. The oviposition takes a long time, perhaps the
whole night, and several dozen eggs, not hundreds as in the
allied genera, make a set. The egg, with its swollen gelatinous
capsule, is large for so small a creature, namely 7-8 mm. in
diameter. The embryos escape after a week, and the tadpoles
reach two inches in total length. Those of B. igrieus have a
triangular mouth, but in B, pachypus this is elliptical, as in
Alytes and Discoglossus. Metamorphosis is completed in the
same autumn; the little toad is then about 15 mm. long, and
differs from the adult by the absence of the conspicuous colora-
tion of the under parts. In reasonable conformity herewith it
does not take up the warning attitude. The colour appears
gradually during the second year, but full growth is generally
not reached until the third year. They do not hibernate in the
water, but hide on land out of the reath of frost.

Alytes. — The tympanum is distinct, the pupil vertical, the
omostemum is absent. The only two species live in South-
Western Europe. The male attaches the eggs to its hind limbs,
and nurses them until they are hatched.

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IS8 ANURA CHAP.

A. obstetricans, the " Midwife-toad," has the general appearance
of a smooth toad. The upper parts are rather smooth, some-
times almost shiny, in spite of the numerous more or less
prominent warts, of which those of the lateral lines, and those
above the ear, are generally most marked. The colour of the
upper parts varies a great deal according to the prevalence of
greenish and reddish spots upon the grey or brown ground-
colour. The red is sometimes, especially in the breeding males,
rather conspicuous on the parotoid region and on the upper sides
of the body. The under parts are whitish grey. The iris is
pale golden, with black veins. The male has no vocal sac, and
is as a rule smaller than the female, the latter reaching a length
of two inches.

This species occurs in the whole of the Iberian Peninsula and
in France, extending into Switzerland and beyond the Bhine
valley inio Thuringia. Altitude above the sea does not seem to
have any influence upon its range, which reaches fix)m sea-level to
the tops of subalpine mountains. I have found great quantities
of its tadpoles in Portugal on the Serra d'Estrella, nearly 6000
feet high, and they are recorded from 6500 feet in the Pyrenees.
They seem to be ubiquitous in Spain and Portugal, not that they
are often found or seen, but they are heard everywhere ; besides,
tadpoles are sure to be in the clear cold lakes on the tops of the
mountain-ranges, in the dirty puddles caused by the village
fountains, and in the sun-heated swampy ditches on the road-
side witli scarcely enough water to hold the wriggling mass.
Wherever there is water within easy reach, on the lonely
mountains, in fertile valleys, in the gardens of the busy towns,
you hear during the whole night, from March to August, the
double cjiU-note of the male, sounding like a little bell ; but to
see the performer is quite a different matter. He sits in front
of his hole, dug out by himself or appropriated from a mouse, in
a crack of the bottom of a wall, imder stones, or in a similar
place into which he withdraws for the day.

The pairing and the peculiar mode of taking care of the
^'ggs l»y the male, which ♦habit has given it the specific name
ohatefrivinis, the midwife, have been most carefully observed by
A. de risle du Drc^neuf, near Nantes. A condensed account h»»
been given by Boulenger. Several males collect around a
female on land, not in the water, and the successful one graspi^

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DISCOGLOSSIDAE I 59

her round the waist. For nearly half an hour the male lubricates
the elocu^al region of the female by more than one thousand
strokes of his toes, whereupon the female extends the hind-limbs,
forming with the bent hind-limbs of the male a receptacle for
the eggs, which are then expelled with a sudden noise. The
eggs are yellow and large, up to 5 mm. in diameter, and are
fastened together in two rosary-like strings, several dozen making
one set. During the expulsion of the eggs the male shifts its
body forwards, clasps his fore-limbs round the female's head, and
fecundates the eggs. After a rest he pushes first one hind-limb
and then the other through the convoluted mass of eggs, which
then have the appearance of being wound round the hind-
limbs in a figure of 8. Then the sexes separate and the male
withdraws with its precious load into its hole, which it, however,
leaves during the following nights, in search of food, taking
this opportunity to moisten the eggs in the dew, occasionally
even immersing them in the water. After at least three weeks,
when the larvae are nearly ready, he betakes himself to the
nearest water, and the larvae burst the thereby softened
gelatinous cover of the eggs. Not infrequently the same male
ventures upon a second pairing, and adds another load to the
one which already hampers its movements. The eggs being
large, owing to the great amount of yellow food-yolk, the embryos
are enabled to be hatched in a more advanced stage than in most
other Anura. The larva develops only one pair of external gills
within the egg. These appear first in the shape of oval bags
upon the third branchial arch, which sprout out secondary
branches, soon in their turn to be resorbed and replaced by the
so-called internal gills before hatching.

Fischer-Sigwart ^ gives the following account of the growth
of this species. The male took to the water, with its load of
twenty to thirty eggs, on the 6th of June. The larvae escaped
out at once, 16-17 mm. long, the body measuring 5 mm. On
the 14th they had reached 32 mm. in length, whereupon they
grew very slowly, although they were well fed, in a temperature
of about 50* F. This same brood did not metamorphose until
May of the next year. The growth took place as follows : —
The hind-limbs appeared on the 8th of September, when the
tadpoles were 50 mm. long; by the middle of the next May they

* ZooL Garten, 1885, p. 299.

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l6o ANURA CHAP.

had reached their greatest length, 76 mm., the hind-limbs being
18 mm. long, whilst the fore-legs were just indicated. On the
21st of May the hind-limbs were 27 mm. long, and the whole
creature was practically metamorphosed, except for the tail. The
latter was resorbed on the 13th of July, and the little toads,
25 mm. In length, were actually smaller, certainly far less bulky
and heavy, than the tadpoles, which had required one year and
a quarter for their metamorphosis.

The early broods probably finish their development by the
autumn of the same year, but those which are born later, in
July and August, certainly hibernate in the water. I have
found very small tadpoles, scarcely 1 5 mm, long, on the Cantabrian
mountains as late as the end of September, and rather large
ones in the spring at the time of first pairing ; the fact that
this takes place during the whole summer explains the occurrence
of tadpoles in all stages of development almost the whole year
round.

A. cisternasi has only two palmar tubercles, the middle or
third one of A. obstetricans being absent ; the outer finger is short
and thick. Instead of a very long and wide fronto-parietal
fontanelle, the fronto-parietal bones diverge only in front so that
there are two fontanelles, a small one in the parietal and a
latge triangular one in the frontal region. The limbs are
relatively shorter and stouter in conformity with the habits
of this species, which prefers to burrow in sandy localities.
Otherwise it leads the same kind of life as A. obstetricans, and
the male carries the eggs. It has hitherto been found in Central
Spain and in the middle provinces of Portugal.

Liopelina ia intermediate between Alytes and Bombinator,
agreeing with the latter, in conformity with its essentially
aquatic life, in the absence of a tympanum, while the Eustachian
tubes are entirely suppressed. The tongue is disc-shaped, but
is slightly free behind. The pupil is triangular. The male is
devoid of a vocal sac. L. hochstetteri is the sole representative
of the Amphibia in New Zealand, where it is apparently rare.
The upper parts are covered with smooth tubercles, and are dark
brown with blackish spots ; the under parts are whitish. Total
length only 1^ inch.

Fam. 2. Pelobatidae. — The upper jaw and, as a rule, the
vomers are provided with teeth. The tongue is oval, slightly

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PELOBATIDAE

l6l

nicked, and free behind, so that it can be thrown out, except
in Asterophrj/s turpicola of New Guinea, which has a large but
entirely adherent tongue. The vertebrae are procoelous, except
in Asterophrys and the Malay genus MegalophrySy where they
are opisthocoelous. The sacral diapophyses are strongly dilated.
The omosternum is small and cartilaginous. The metastemum
has a bony style, and ends in a cartilaginous, rounded or heart-

"-■^■^

^ CtSTISWATHIOAE. IIDIH QISCOCLOSSIOAE. ^'f^ PELOBATIOAI,
Fig. 32. — Map showing distribution of Cystiguathidae, Discoglossidae, and Pelobatidae.

shaped disc, but in Scaphiopus it forms an entirely cartilaginous
plate. The tympanic disc is mostly hidden or indistinct, and is
quite absent in Pelohates. The Eustachian tubes are very small in
Felohates, and exceedingly minute in Scaphiopus stagncdis of New
Mexico. The pupil is vertical. This family contains seven
genera with about twenty species, with a rather scattered
distribution.

A. Toes extensively webbed, sacrum and coccyx confluent

a. Metastemum a cartilaginous plate. America Scaphiopus^ p. 164.

b. Metastemum with a bony style. Eiu*ope

B. Toes nearly free. Metastemum with a bony style.

a. Vertebrae procoelous.

a. Sacral vertebra articulating by one con-
dyle with the coccyx.
{Europe
New Guinea
j3. Sacral vertebra with two
India and Malaya

b. Vertebrae opisthocoelous.

{Ceylon and Malayan Islands .
New Guineii .
VOL. VIII

PelobateSj p. 162.

PelodyteSy p. 165.
Batrachopsis.
condyles.

Leptobrachium, p. 166.

Mi'f/alophrySy p. 60 (Fig. 11).
Asterophrys.

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I 62 ANURA

Pdohates (" Spade-foot "). — The tympanum is absent ; the toes
are webbed. The inner tarsal tubercle is large, and is transformed
into a shovel which is covered with a hard, sharp-edged, honiy
sheath. The skin of the upper surface of the head is partly
co-ossified with the underlying cranial bones, giving them h
pitted appearance. The general shape is toad-like.

P. fuscus, — The smooth skin is brown above, with darker
marblings, while the under parts are whitish, but the coloration
varies greatly, from pale to dark brown or olive-grey with more
or less prominent irregular dark, sometimes confluent, patches.
Some specimens are adorned with numerous red spots. The
tarsal spur is yellow or light brown. The iris is metallic red
or golden. The male has a long oval gland on the upper
surface of the upper arm, and although possessed of a voice, has
no vocal sacs. The total length of full-grown females is nearly
3 inches, that of males half an inch less.

The " Spade - footed Toad," which occurs throughout the
whole of Central Europe, extends from Belgium and the
middle of France to North-Western Persia, and from the southern
end of Sweden to Northern Italy. It prefers sandy localities,
in order to dig its deep hole, in which it sits concealed
during the daytime. Owing to the looseness of the sand,
the hole is filled up so that no trace of its inhabitant is
left. The digging is done by means of the spades, and in
suitable localities the animal soon vanishes, sinking backwards
out of sight. Except in the breeding season, or at night, it is
therefore found only accidentally. The sand-loving habits do
not, however, prevent it from enjoying moist localities. Several
which I have kept for years dig themselves into the wettest
moss in preference to the drier parts of the^r habitation. Being
thoroughly nocturnal, they hunt after nightfall, the food consist-
ing of all sorts of insects and of woims. When captured they
utter a startling shrill cry, and their skin becomes covered with
a dermal secretion which smells like garlic, a peculiarity which
has given them in Germany the name of " Knoblauch skrote,"
" garlic-toad." Although they become very tame, so that they
no longer smell when handled, they can be made ill-tempered
by being pinched or otherwise teased, whereupon they take up
a defiant attitude, and with open mouth continue to cry for
several minutes. Some such scenes occur now and then, without

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PELOBATIDAE 1 63

my interference, with the specimens which share their abode with
several species of Amhlystoma and Spelerpes; there are heard
now and then sudden loud yells, like the squeak of a cat or the
yapping of a little dog.

In the spring the Spade-footed Toads take to the water for
about a week, and the male's call-note is an ever-repeated cluck-
ing sound, which can also be produced under water, with the
mouth shut, the air being shifted backwards and forwards through
the larynx. The male grasps his mate below the waist ; the eggs
are combined into one thick string, which is about 18 inches
long, and is wound round and between the leaves and stalks of
water-plants. The eggs measure 2-2*5 mm., and are very
numerous, a large string containing several thousands. The
larvae are hatched on the fifth or sixth day in a very unripe con-
dition. They are only 4 mm. long, quite black, and still devoid
of gills and tail. They attach themselves to the empty gelatinous
^g-membranes, which they possibly live upon. On the following
day the tail begins to grow ; two days later fringed external gills
sprout out and serve for about ten days, when they in turn give
way to new, inner gills. The little tadpoles then leave their moor-
ings and become independent. The hind-limbs appear in the
ninth week, the fore-limbs in the twelfth. At the age of three
months they begin to leave the water. The most remarkable
featmre is the enormous size of the full-grown tadpole, the body of
which is as large as a pigeon's egg; the usual total length,
including the tail, amounts to about 4 inches or 100 mm., but
occasionally regular monsters are found. This was the case some
thirty years ago, when the Berlin Museum received a number of
tadpoles, the largest of which measured nearly 7 inches. They
were found in the month of December near Berlin, in a deep clay-
pit with high, steep walk, so that the tadpoles were prevented
from leaving the water. Similarly hemmed-in broods probably
hibernate in the water under the ice, and such instances have
been recorded. Normally they metamorphose into the much
smaller toad within the same year.

P. ctUiripes. — This is the Spade-foot of the whole of Spain and
Portugal and of the southern and western parts of France. It
is similar in habits to P. ftisctis, from which it differs but
slightly. The tarsal spur is black, and there is a parieto-
squamosal bridge which completely roofs over the temporal fossa

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164

ANURA

and closes the orbit behind. — Boulenger has discovered the rare,
individual occurrence of minute teeth on the parasphenoid and
on the pterygoids of this species. These teeth are unquestion-
ably the last reminiscences of a condition almost entirely super-
seded in the recent Anura.

P. syHacus from Asia Minor and Syria agrees with P, cvJtripes
in the cranial configuration, but has the yellow or brown spur
of P, fuscus.

Fig. 33. — Pelobates culirijpeSf Spade-foot Toad, x 1, and under surface of left foot.

Scaphiopiis. — The Spade- foot of North America and Mexico
differs slightly from those of Europe, chiefly by the presence of a
more or less hidden tympanum and of a subgular vocal sac, and by
the sternum, which forms an entirely cartilaginous plate without
a special style. The close relationship of these two genera is
further indicated by the occurrence of peculiar large glandular
complexes in some of the species, pectoral in S. solitarius, tibial
in S. muUiplicatiis of Mexico. At the same time this genus
approaches Pelodytes. — About eight species are known, two of
which inhabit the United States, the others Mexico.

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PELOBATIDAE 1 6$

S. solitarius is the commonest species of the Southern States.
It is brown above, with darker patches ; its total length is about
2 inches. According to Holbrook it excavates small holes half a
foot deep, in which it resides, seizing upon such unwary insects
as may enter its dwelling. It never leaves the hole except in
the evening or after long -continued rains. It appears early in
March, and soon pairs; as an instance of hardiness Holbrook
mentions that he has met it whilst there was still snow on the
ground. When teased they assume^ a humble attitude, bending
the head downwards with their eyes shut, as illustrated by
Boulenger.*

Pelodytes is, like the rest of the genera, devoid of the tarsal
digging spur. The tympanic disc is rather indistinct ; the
male has a subgular sac. The general appearance of the slender
body with long hind-limbs and toes is frog-like. Two species
only are known, one in South -Western Europe, the other in the
Caucasus.

P. jmnctatus. — The " Mud-diver " has the upper parts covered
with ismall warts, and is about 1^ inch in length. Its
coloration is variable, and changes much. One day it may
appear greenish brown, the next day pale grey ; in the daytime
perhaps with many bright green spots, and in the evening spot-
less and unicoloured. The under parts are mostly white, some-
times with a fleshy tinge. The male has a voice like " kerr-kerr "
or " creck-creck," uttered during the breeding season, which lasts
from the end of February until May, according to the temperature
and the more Southern or Northern locality. Occasionally they
breed a second time in the summer or autumn. The male
develops nuptial excrescences, chiefly three rough patches on the
inner side of the fore-limbs or on the inner side of the first two
fingers, while the belly and thighs are covered with small
granules. In the mode of copulation, the laying of the small
and numerous eggs, the hatching of the larvae in a tail- and gill-
less condition, this genus closely resembles Felobates ; but the
tadpoles never reach a colossal size, the usual length being 2
inches, and even this is comparatively large for so small a species.
It inhabits the greater part of France, most of Portugal, and the
southern half of Spain, avoiding, however, the central plateaux
and the mountain-ranges. Its habits are essentially nocturnal,

1 P.Z.S. 1899, p. 790.

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1 66 ANURA

living in the immediate vicinity of the water, into which it hops
with a long jump in order to hide in the mud. Easily kept,
it breeds regularly in captivity, according to circumstances at
almost any time of the year.

P. caucasicus has been discovered in the Caucasus at an
altitude of 7000 feet. The remaining genera of this family
contain only a few species each, and are restricted to South-
western Asia, the Malay and Papuan Islands. The commonest
is Zeptohrachium, which ranges from the Himalayas to Borneo
and Java. Pupil vertical. Vomerine teeth sometimes absent.
Tongue roundish, very slightly nicked behind. Tympanum in-
distinct. Omosternum small, cartilaginous. Male with internal
vocal sacs. Tarsus with a roundish tubercle. Some of the
species, e.g. Z. carinense from the Karen Hills, attain to a large
size, namely, 6 inches ; they seem to live on rats and mice, and
one specimen contained a young squirrel.

Pam. 3. Bufonidae (Toads). — The formula : — no teeth in the
upper and lower jaws, vertebrae precocious and without ribs, sacral
diapophyses dilated, — is sufficiently diagnostic of this cosmopolitan
family. The generally entertained notion that toads have a
rather thick-set, short-limbed, warty appearance, does not apply to
all the members of the family. The majority are quite terrestrial,
many are burrowing, the Javanese Nectes is aquatic, the Afro-Indian
Nectophryne is arboreal, while the Mexican Myohatrachus and the
Australian Rhinophi^ynus eat termites and are correspondingly
modified ; lastly, Bufo jerboa is a slender, long-legged creature.

Teeth are almost entirely absent, except in Notaden, which
has teeth on the vomers. The omosternum is mostly absent,
except in Engystomops and in some species of BufOy while in
Notaden it is merely vestigial. The metasternum shows more
variety. The tympanum is usually distinct, but varies even
within the same genus, being hidden beneath the skin or being
entirely absent. The terminal phalanges are modified according
to the habits of the species, but they are never claw-shaped.

The Bufonidae are connected in various directions. The Neo-
tropical Engystomops greatly resembles the likewise Neotropical
Cystignathoid Paludicolay and the Australian Pseudophryne closely
approaches the Australian Cystignathoid Crinia, It is therefore
all the more remarkable that a similar approach, in another
direction, namely, towards the Firmisternal family of the Engysto-

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BUFONIDAE

167

matidaejs indicated by the Mexican Rhmophi^s and the Australian
Myohatrachus, However, since there are no true Engystomatidae
in Australia, although several genera occur in Papuasia, these
cases may be instances of convergence without necessarily im-
plying relationship. An unmistakable line of connexion leads,
according to Boulenger, to the Pelobatidae, the link being the
Himalayan Cophaphryne, with very strongly dilated sacral
diapophyses, with a single condylar articulation of the coccyx
with the sacral vertebra (as in some Indo- Malayan Pelo-

^= auro. W// FORMS with fimger discs . a\\\ ^ orws besides bufo.

Fio. 34. — Map showing distribution of Bufonidae. The vertical lines indicate the
occurrence of Bufonidae, but not of Bvfo.

batidae), while this articulation is bicondylar in all the other
Bufonidae.

The whole family is divided into eight genera with more than
a hundred species, of which only about fifteen do not belong to the
genus Bufo. The distribution of the family is well-nigh cosmo-
politan, with the remarkable exception of Madagascar, Papuasia,
and the small islands of the Pacific ; Bvfo has been wrongly said
to inhabit the Sandwich Islands. The greatest number of
species, chiefly Bufo, occur in the Neotropical region, the greatest
numl)er of genera in Central America, where Bufo is rare, and in
Australia, where it is absent.

A. Pupils contracted to a horizontal slit. Typically arciferous

a. Australian. Tympanum invisible. Fingei-s and toes not dilated.

1. With vomerine teeth. Both the onio- and meta-stenium are

rudimentary. East Australia : . . Kotaden bennetti.

•1. Without vomerine teeth. Omosternmn aV)sent. Metasternum
cartilaginou- : ..... /V»Wo^^n/?<f', p. 168.

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1 68 ANURA CHAP.

b. Not Australian.

1. Omostemum narrow and cartilaginous. Metastemuni with a

bony style ending in a cartilaginous disc. Fingers and toes
slightly swollen. Neotropical : . Engystamops, p. 168.

2. Omosternum absent Metastemum cartilaginous.

a. Fingers and toes webbed ; terminal phalanges T-shaped
and with adhesive broadened tips. Africa and India :

Nectophryne, p. 169.

p. Fingers free, toes webbed ; terminal phalanges simple,
not dilated. Tympanum distinct. Java : Nectes, p. 169.

3. Metastemum cartilaginous, sometimes ossified along the middle.

Fingers free ; toes more or less webbed ; tips simple or dilated

into very small discs : . . . . Bufo, p. 169.

B. Pupil a vertical slit The epicoracoid cartilages are narrow and scarcely

overlap. Omostemum absent except in Cophophryne. Vomerine teeth

absent Sacral diapophyses strongly dilated. The terminal phalanges

are simple and the tips are pointed.

a. Aastralian. Tympanum distinct The metastemum is calcified
along the middle : ..... My ohatrachicSy i). 184.
h. Mexican. Tympanum absent. Metastemum rudimentar}' :

Rhinophrynusy p. 185.

c. Himalayan. Tympanum absent Metastemum with a slender bony

style :..... Cophophryne Hkkimensi.s.

Engystomops is interesting because it closely resembles the
Cystignathoid genus Paludicola, and thereby seems to connect
these two families. It differs from Pcdudicola chiefly by the
absence of teeth, by the moderately dilated sacral diapophyses
and by the slightly swollen tips of the fingers and toes, the end-
phalanges of which are, in one species, E, petersi, T- or anchor-
shaped. The tympanic disc is either distinct or hidden. The
males have a large subgular vocal sac. The generic name refers
to the small head with a prominent snout. Three species are
known from Central America and Ecuador.

Pseudophryne appears to be another link with the Cysti-
gnathidae by its resemblance to the Australian genus Crinia, from
which it diflfers by the absence of teeth and by the absence
of an omosternum. The sacral diapophyses are but moderately
dilated. The males have a flat oval gland on the hinder side of
the thighs, and they are provided with a subgular vocal sac
The 3 or 4 species of this genus which live in Australia, both
East and West, are not unlike Bomhinator in their general shape,
short limbs and coloration. The skin of P. australis and P.
hihroni is covered with small smooth warts and is blackish
brown, while the under parts are blackish with large yellow

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VI BUFONIDAE 1 69

patehea Total length little more than one inch. Concerning
the breeding habits, see p. 223.

Nectojphryne. — -The sacral diapophyses are strongly dilated.
N, afra, without a tympanum, but with fully-webbed digits and
several broad, cushion-like or lamellar pads on the fingers and
toes, inhabits the Cameroons, N. tuberculosa of Malabar, and
iY guentheri and N, hosei of Borneo, have a visible tympanum
and the fingers are webbed at the base only. These slender
and long-legged species are most probably arboreal, as indicated
by the broadened, but truncated, tips of their fingers and toes.
iV; hoseri is about 4 inches long, iVi misera is a little creature of
only |- inch in length. Nectes, hitherto known by one species,
iV. subasper of Java, is a swimmer and exceeds 6 inches in
length. The tympanum is very distinct ; the small nostrils look
upwards. The toes are long and webbed to the tips ; the hind-
limbs are very long. The sacral diapophyses are strongly dilated.
The skin of the upper parts is very rugose, covered with round
warts, and dark brown ; the under parts are granular and uni-
formly light brown.

Bufo. — The great number of species, more than 100, renders
a strict definition of this genus difficult. The tongue is pear-
shaped, thicker in front, entire, not cut out, but free behind, so
that it can be projected. The fingers are free, the toes more
or less webbed although never completely so. The terminal
phalanges are obtuse and sometimes carry tiny discs. The
omostemum is absent or merely vestigial. The metasternum
is a rather large cartilaginous plate with a v^aist, which is
sometimes incompletely calcified. The sacral diapophyses are
moderately dilated. The tympanum is distinct or hidden. The
skin of the upper parts is always rich in specific poison -glands,
a concentration of which forms in many species very conspicuous,
thickened ' parotoid glands. The surface of the skin may be
smooth, moist and slimy, or rough and warty, sometimes covered
with tiny, sharp, horny spikes and quite dry.

The genus is cosmopolitan, with the exception of the whole
Australian region and Madagascar, from which we may perhaps
conclude that its original centre was not in Notogaea, in spite
of the diversity of species in the Neotropical region, whicli now
contains about half of all the species known. Next to Central
America the Indian region is richest in species of Bvfo.

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I70 ANURA CHAP.

B, vulgaris. — The Common Toad of the Palaeaictic region.
The skin of the upper parts is much wrinkled and beset with
numerous round warts or poison-glands, the openings of which
can be seen with the naked eye, especially on the large parotoid
complexes. The outermost layer of the epiderm, in feet all that
portion which is periodically shed, is elevated into numerous
little coraified spines. The extent of their development varies
much ; southern specimens, especially those from Portugal, being
perhaps the roughest. Others appear quite smooth to the touch,
and this is the case with many English specimens. The skin
of the under parts is more granular and devoid of specific glands.
The general colour of the uppfer parts is olive grey to dark
brown, more or less mottled ; the under parts are whitish, often
with a brown, yellow or reddish tinge.

The coloration of this species varies considerably and is more-
over very changeable. These changes depend chiefly upon the
surroundings and the locality, in which certain styles of coloration •
seem to be the fashion, not necessarily to the absolute exclusion
of otliers. Some specimens are of a rich brown colour, with or
without dark brown spots and patches, and these are sometimes
confluent, forming irregular, longitudinal bands. The ground-
colour of other individuals is olive grey, with or without darker
patches, and these paler tones prevail in toads which live
on light-coloured soil, for instance on chalk. I recently found
one between two dark-coloured slates, and this creature was so
black that it gave the impression of having soiled itself with
coal-dust. One and the same specimen will appear paler or
darker according to its mood and the leading tones of its
immediate surroundings, but it cannot change its dominant
ground-colour. A third colour-variety occurs more frequently
in the mountainous districts of Southern Em-ope. I have ob-
taineil the most handsome specimens in the Serra Gerez, in North
Portugal. Their ground-colour is pale brownish-yellow, with
many large and small, rich brown patches, or if the latter colour
predominates, these patches and spots are separated from each
other by creamy seams, with the occasional effect of dark brown,
yellow-ringed eyes. Eastern Asiatic specimens often have a
fine yellow vertebral line and the under parts are inclined to be
marked with dark spots.

The iris is red or coppery, mottled with black. The male

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BUFONIDAE 171

has no vocal sacs, and, besides being smaller than the female,
is distinguished by slight nuptial excrescences in the shape
of little homy brushes on the inside of the inner palmar
tubercle and the three inner fingers. The full size of this toad
varies extremely. Taking the standard of everyday experience
in England and Central Europe, one would call any female
beyond 3^ inches in length, and any male of more than 2^ inches,
unusually large. But occasionally they grow to a much larger
size, especially in the mountains of Southern Europe, provided
there is a rich vegetation of meadows and deciduous trees so
as to insure a variety of plentiftd food. Although Fatio ^
mentions a toad 153 mm. = 6 inches long, and Boulenger suc-
ceeded in getting a toad from Paris which measured 132 mm.,
i.e. almost 5^ inches, one of my specimens from the Serra Gerez
seems to hold the record with a total length from snout to vent of
135 mm. or more than 5^ inches. Jersey is also famous for its large
toads, possibly on account of the many large greenhouses. These
large specimens do not constitute a special race. The monsters
among them are without exception females, often but not always
st^erile, as I have often found large masses of eggs in them. Food is
the chief cause. At least I have observed that the more voracious
of some Spanish and Portuguese specimens, which were already
3 J- inches long, and therefore entitled to respect, continued to
grow rather rapidly, adding about half an inch within a year.
Again, if the growth of a promising toad is arrested for
a season — not necessarily by starvation, but by uncongenial
surroundings, sameness, and unvaried nature of food — they
consolidate so to say, or settle down, and no amount of future
good feeding will turn them into exceptionally big specimens.
There are no data to tell how old such monsters really are. At
least ten years are required by the Southerners to reach four
inches. The usual length of life attained by a toad is likewise
unknown. Boulenger kept one in a box provided with a sod,
a pan of water and plenty of varied food, but twelve years of
close captivity did not make any appreciable difference in its
appearance. A number of large Spanish and Portuguese speci-
mens in my greenhouse were at first very shy, and tried every
possible means of escape or sullen hiding, but gradually they
condescended to take food when lifted on to the slate-covered

* Faune Vertebr. Suisse^ iii. 1872, p. 587.

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172 ANURA CHAP.

stage upon which their food was spread. After a few weeks
they had learned this so thoroughly that, towards the usual
hour of feeding, they climbed most laboriously on to the slates,
lying in wait between the flower - pots, and coming forward
when we entered the house. The rest of the day and night
they spent on the ground, under stones or plants, each in its
individual lair. The biggest of all, and several others, became
so tame that they took food whilst sitting on the hand, and
then they looked up for more. The food must be alive and

Fig. 35. — Bh/o vulgaris. Portuguese si>ecimeu. x |.

show movement. Mealworms, snails, beetles and other small
creatures are first carefully inspected with bent-down head, and
are sometimes followed for a few inches ; then comes an audible
snap, a flash of tlie rosy tongue and the prey has disappeared.
T^rge earthworms are nipped up with the jaws and laboriously
poked in with the hands, the fingers being so placed as to clean
the worm of adherent soil and other impurities. Very largo
wonns are shaken, twisted, pressed against the ground ami
gulped down with convulsive movements, but not unfrequently
the tip-end remains for some minutes sticking out of the tightly
shut mouth. Several are taken at one sitting, until the toad
is gorged. One of the biggest took full-grown mice, which

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BUFONIDAE I73

were not " fascinated by the fiery eyes " but were stalked into
a corner and then pounced upon immediately when they moved.
The shells of snails can for half a day be felt through the body ;
they then dissolve or are disgorged. The dung, which is passed in
large, long masses, is often full of fine earthy matter, the contents
of the earthworm's intestines, and sometimes it contains the
chitinous remains of certain beetles which are supposed to be
excessively rare. I know of no instance of slugs being eaten.

The regular hunting -time begins w^ith the evening and
is continued throughout bright nights, the toads crawling
and hopping about. They are expert , climbers of rocks, and
succeed in reaching apparently inaccessible places by shoving
themselves up between vertical walls, and taking advantage of
any roughnesses for foothold. Every few weeks they shed their
skins. Without any preliminary symptoms or loss of appetite
or liveliness, the body makes a few twisting motions, the back
is now and then curved, and the skin splits down the middle
line. Owing to the more forcible Contortions of the body it
slides down to the right and left of the back, whereupon the
toad gets hold of the peeling-ofiF skin with fingers and toes,
scraping the head and sides, and conveys the thin, transparent,
slightly tinged skin into the mouth, slips out of it backwards
and swallows it. The new surface is then quite wet and shiny,
but it soon dries and hardens.

Many toads, for instance the Common Toad and the Pantherine
Toad, assume a peculiar attitude when surprised. Instead of
blowing themselves up by filling their lungs with air, they
raise themselves upon their four limbs as high as possible, but
turning the back towards the enemy in a slanting position, either
to the right or to the left side, apparently in order to present as
much surface as possible, in other words to look their biggest.

Some of my specimens hibernated regularly for a few months,
burying themselves completely in loose, dry soil, under leaves, or,
— a favourite place, — in a heap of cocoa-nut fibre. Others, and
this applies also to English specimens transferred from the
garden into the greenhouse, are lively all the year round, but
even they withdraw for an occasional sleep of a few weeks at ■
any time of the year.

The whole family of large toads came to a sad end after four
years, when they were put into new temporary quarters, a slate-

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[74 ANURA

bottomed terrarium. Being kept during my absence m wrmging
wet moss, which became fouled by their own excretions, they
contracted a mysterious disease from which they never recovered
They are rather averse to wet surroundings, and except dunng
the short pairing season they live in cool, shady places, pre-
ferably with just a little dampness. Occasionally they take a
soaking bath. One specimen, living in the garden, repair«l
during the hot and dry summer nights to a standpipe m the
garden, enjoying the occasional drips of water.

Considering the amount of snails and other noxious creaturea
destroyed by them during their regular nocturnal hunts, toada
are eminently useful creatures. Nevertheless, they suffer much
through the stupid superstition of people who ought to know
better. It is difficult to find a gentle, absolutely harmless and
useful creature that is more maligned than the European toad.
It brings ill-luck to the house, the " slimy toad " spits venom, sucks
the cows' udders and after that destroys their power of giving
milk ; it poisons the milk in the ceUar, and a certain builder's
horse, which was grazing in the grounds of the Cambridge
Museums, and died there from a large concrement obstructing
its bowels, was solemnly declared to have swallowed one of my
toads. Silly superstitions, owing to faulty, or rather entire want
of, observation ! The toad is not slimy, but dry; it is often
found in buildings, where it keeps down the woodlic« ; it cannoi
suck, nor does it drink at all; it does not spit venom, but
becomes covered with milky white and very strong poison when
in acute agony, for instance when trodden upon ; and unless the
big skin-glands be forcibly squeezed, there will be no squirting.
Therefore, leave it alone, or put down food on its evening beat,
and it will soon come to know and to recognise its friends.

The Common Toad can exist without food for a long time,
provided the locality is cool and damp, but it wastes away almost
to skin and bones. In order to disprove the persistently crop-
ping up fable and sensational newspaper-accounts of toads having
lieen discovered immured in buildings, where they were supposed
to have lived for many years, Frank Buckland put a dozen
specimens into separate holes bored in a block of porous lime-
stone, covered them up tightly with a glass plate and buried
the block a yard deep in the soil. A second dozen were treated
similarly, but were put into a block of dense sandstone. After a

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BUFONIDAE 1 75

year and two weeks all the toads enclosed in the latter block
were of course found dead and decomposed, but most of those in
the porous block were still alive, with their eyes open, and did
not succumb to starvation until eighteen months of confinement.
These poor creatures could of course not move about, and were
practically undergoing enforced continuous hibernation. Other-
wise they would soon have wasted away and have died within
six months. Those which tumble into deep and dry wells remain
rather small, but generally manage to keep alive for years on
the spiders, woodlice, earwigs and other insects which likewise
tumble in.

Toads hibernate far from the water in dry holes or clefts, retir-
ing in the middle of October in Central Europe, and they do not
reappear before March. Soon after, and this depends naturally
upon the season, they congregate in ponds or pools, and the
males, which far outnumber the females, for whom they fight,
make a peculiar little noise, something like the whining bleat of
a lamb, uttering this sound day and night. The male having,
after much wrestling with competitors, secured a female, which
is often several times bigger than himself, clasps her tightly, by
pressing his fists into the armpits, and the pair swim or crawl
about in this position sometimes for a week before the spawning
takes place. The number of eggs laid at one sitting is enormous,
varying from 2000 to 7000. They are very small, only 1-5-20
mm. in diameter, and are expelled in two double rows or strings,
one coming out of each oviduct. These strings consist of a soft
gelatinous mass, in which the double rows of entirely black eggs are
imbedded, and they measure in the swollen condition about 6 ram.
or ^ inch in diameter, and from 10 to 15 feet in length. The
strings are wound round and between water-plants by the parents,
which move about during the laying and fertilising process.
According to the coldness or warmth of the season the larvae are
hatched in about a fortnight, and for the next few days they hang
on to the dissolving gelatinous mass of the egg-strings. They then
leave the slime and fasten themselves by means of their suckers
to the under side of grasses and water-plants or sticks, with
their tails hanging downwards, still in a rudimentary condition,
but henceforth progressing rapidly.

Fischer-Sigwart ^ found the time of development as follows :

^ Zool. Garten f 1885, p. 299.

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176 ANURA

— ^The eggs were laid on the 6th of March ; the larvae left the
jelly on the 16th, being 4 mm. long. On the 2nd of April they
measm^ed 13 mm.; on the 25th, 20 mm. On the 7th of May
the hind-limbs appeared. On the 18th of May the tadpoles had
reached their greatest length, namely 24 mm., and this is a rather
small size for so large a species. The fore-limbs broke through
on the 28th, and the metamorphosis was completed eighty-five
days after the eggs were laid, the creatures leaving the water
on the 30th of May. Tlie tadpoles showed a preference for
rotten pieces of Agaricus, which were floating in the water. The
little baby-toads are surprisingly small, scarcely 15 mm. long,
and live in the grass, under stones, in cracks of the ground, and
hop about in much better style than their heavier and more
clumsy-looking parents. Where many broods have been hatched
they can be met with in myriads, the ground literally swarming
with them, and as they are naturally stirred up by a sudden
warm rain, perhaps after a drought, people will occasionally state
it as an observed and well-ascertained fact that " it has rained
toads."

What becomes of all these hopeful little creatures ? Although
it takes them fully five years to reach maturity, one would expect
that the whole country would be swarming with toads ; but since
this is not the case, there being not more toads now than there
were before, it follows that their enormous fecundity is only just
sufficient to keep the race going. Adult toads seem to have
scarcely any enemies except the Grass Snake, which takes them in
default of anything better. But how about the reduction where
there are no snakes ? We know nothing about epidemics which
might carry them off, but elderly toads are liable to a horrible
disease produced by various kinds of flies, notably by Lucilia
hufo7iivora and Calliphora silvcUica, the maggots of which some-
how or other eat their way from the nostrils into the brain and
into the eyes. Those which reach the brain at first produce
effects similar to those of Coenwnis cerebralis, the hydatid or
bladder-worm of sheep. The toad inclines its head towards one
side, and cannot crawl straight, but walks in a circle. By eating
away the brain they gradually destroy the host's life. But
if none enter the brain, and a few only find their way into the
eye, they only impair or destroy its sight. Such toads show
signs of pain, poking at or stroking the affected eye, which becomes

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BUFONIDAE 1 77

inflamed, and ultimately remains enlarged, with the iris partially
or entirely destroyed by the maggot, which does not develop
further, but dies in the eye-chamber, this being really an unsuit-
able place for it. The eyesight is of course affected, and is mostly,
but not in all cases, lost Such half -blind individuals — the
disease affecting sometimes one eye only — recover their health,
and except for a little awkwardness, behave like normal specimens.
This applies to Bufo vulgaris as well as to B. calamita. Australian
Anura are cursed with a fly of their own, called Bairachomyia}

B. vulgaris inhabits almost the whole of the Palaearctic region;
— the whole of Europe, with the exception of Ireland, the Balearic
Islands, Sardinia and Corsica. Northwards it extends to Trondhjem,
and thence along a line drawn across Eussia and Siberia to the
Amoor. Its southern Umit in Asia is indicated by a line drawn
from the Caucasus through the Himalayas into China. In Asia
Minor and in Persia it is absent. South of the Mediterranean
it occurs only in Morocco and Algeria.

B, melanostictus is the common toad of the whole Indian
region and of the Malay Archipelago. The epidermis of the fingers
and toes is thicker and more cornified than usual, and is stained
black brown, hence its specific name. The male has a subgular
vocal sacw In other respects the Indian species much resembles
the more spinous or rough-skinned and brown varieties of the
European species. According to S. S. Flower this toad is very
conmion in the Straits Settlements, hiding by day under stones
or logs, or in holes, coming out shortly before sunset, and remain-
ing abroad till dawn ; it may be met with on the roads and in the
grass, hopping or crawling about in search of ants, bees, and
similar food. It utters a rather feeble, plaintive cry when handled
for the first time. It can change its colour from light yellowish
to dark brown. The spawn, which resembles that of B. vulgaris,
may be seen in March and April in ponds, in long strings twined
about the water-weeds. The tadpoles are very like those of the
common English toad in form, size, colour, and structure of
mouth. The largest adult found in Penang measured 115 mm.
(about 4 inches) from snout to vent.

* For further information, cf. Portschinsky, "Biologie des mouches copro-
phages et ndcrophages, 2me partie. Etude sur la Ltteilia bufonivora, parasite
des batraciens anoures.*' — Horae Soe. ctU. Jiosa, xxxii. pp. 225-279 (in Russian).
German summary in Zool, Centralbl, y. 1898, pp. 855-859.

VOL. vni N

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178 ANURA CHAP.

B. lentiginosiis a ainericanvs is the common toad of North
America, from Mexico to the Great Bear Lake. It is worth
noting Lhat this species resembles in its coloration the Eastern
races of B. vulgaris, in so far as they generally have a light
vertebral line, and frequently dark spots on the under surface.
Tlie upper parts are brown and olive, with darker spots, two of
which form a chevron behind the eyes. But the tympanum is
large, and the male has a subgular vocal sac ; the inner metatarsal
tubercle is very large, and is used as a kind of digging spur.
During the pairing time they take to the pools in great numbers,
uttering their music, which consists of a prolonged trill, continued
by different individuals, both day and night. Holbrook knew
an individual which was kept for a long time, and became per*
fectly tame. During the summer months it retired to a comer
of the room into a habitation which it had prepared for itself in
a small quantity of earth placed there for its convenience.
Towards the evening it wandered about in search of food. Some
water having been squeezed from a sponge upon its head one hot
day in July, it returned the next day to the spot, and seemed
well pleased with the repetition, nor did it fail during the extreme
heat of the summer to repair to it frequently in search of its
shower-bath.

Several varieties of this widely distributed species, whose
average length is 2|^ inches, have been described. The prettiest
was called B. quercinus by Holbrook — ^according to whom it is
mostly found in sandy places covered with a small species of
oak — which springs up abundantly where pine-forests have been
destroyed. It is called the " oak-frog," as it spends most of its
time in concealment under fallen oak-leaves, or partially buried
in the sand.

B. maHnus s. agua is the giant among toads, and is one of the
commonest species of the Neotropical region, ranging from the
Antilles and Mexico to Argentina. It frequently reaches a
length of 6 inches, with a width of 4 inches when squatting
down in its favourite attitude. The upper parts are rough, owing
to the prominent warty glands, of which the parotoid complex is
enormous. The general colour above is dark brown, with sooty
dark patches ; below whitish, often with blackish patches. This
creature appears at dusk, often in large numbers, especially during
the rainy season, hopping about, not crawling, with surprising

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VI BUFONIDAE 1 79

activity. The voice of the male, strengthened by a subgular sac,
is said to be a kind of loud snoring bark. The pairing time
begins, according to Hensel,^ yrith the winter rainy season, especi-
ally June, and lasts several months, until October, but it is^
interrupted by the cold, which in the hills of South - Eastern
Brazil covers the ponds with ice. Tiien the tremulous bass
voice of the males is heard no longer ; they have all withdrawn
beneath stones and trees in the neighbourhood of the water. The
eggs are laid in strings. The larvae are at first quite black and
very small, and the young baby-toads are only 1 cm. in length.
They differ considerably from the adult until they are more than
1 inch long ; the upper parts are yellowish brown, with darker
ocellated patches, each with a light seam, most conspicuous along
the sides of the head and back. The under parts are grey, finely
stippled with yellow.

Budgett'^ remarks that B, marinus feeds on all kinds of insects.
'•' One half-grown specimen sitting by a man's foot picked off fifty-
two mosquitoes in the space of one minute, picking them up with
the tongue as they settled. The call of this very common toad
consists of three bell -like notes; the middle one being the
highest. The enormous parotoid glands are discharged like
squirts when the creature is roughly handled. When wet weather
comes on it hops out from its hiding-place to sit in a puddle,
with its head out."

In many species of Bufo the crown of the head forms more or
less prominent ridges, especially strong in the region between the
eyes ; for instance, in B, melanostictus and B, lentiginosus. The
skin overlying these ridges is liable to be involved in the cranial
ossification, and this reaches its greatest extent in the two Cuban
species B. empusus and B. peltocephalus. It is a curious coincidence,
to say the least, that such dermal ossifications should be best
developed in Neotropical species, in those very countries which
amongst the Cystignathidae have produced the abnormal genera
Triprio7i, CalyptocephcdvjSy and Pternohyla. The most peculiar
and odd -looking species is Bufo ceratophrys, a native of
Ecuador, which has the upper eyelid produced into a horn-like
appendage, the two sharply-pointed cones standing out trans-
versely, reminding us of several species of the Cystignathoid
genus Ceratophrys ; there is also a series of four small pointed
^ Arch, NiUurg. xliv. 1868, p. 141. * Quart. J. Micr. Sci. xlii. 1899, p. 3.

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l80 ANURA CHAP.

appendages on each side of the body. Protective concealment
is possibly the reason of these queer outgrowths. »

B, viridis s. variabilis, the Green or Variable Toad, reaches a
length of about 3 inches, and is the prettiest toad of £uropa
The skin is distinctly smooth, the numerous porous, large and
small warts being flattened. Parotoid glands are well developed,
and a similar pair of glands sometimes occurs on the inner side
of the calf, especially in Central Asiatic and in Algerian specimens.
The coloration is very variable and changeable. The ground-
colour of the upper parts is creamy, with large and small, partly
confluent and irregularly shaped spots and patches of green, here
and there interspersed with vermilion-red specks, especially along
the sides of the back. The under parts are whitish, sometimes
spotted with bleujk. The iris is brass-coloured, greenish-yellow,
with fine dark dots. The male does not differ from the female
in size, but has an internal subgular vocal sac, a conspicuous
callosity on the inner side of the first finger, and nuptial brushes
on the first three fingers and on the inner palmar tubercle.

The changing of colour afifects mainly the intensity of the
green; the same individual which now looks almost uniformly
dull, almost grey, with dusky olive patches, will, if put into
grass and sprinkled with water, within a few minutes appear in
a tastefully combined garb of grass-green on a creamy ground.
Some Southern and Eastern specimens have a creamy stripe along
the vertebral line, thereby closely resembling B, calamita, from
which, however, they can always be distinguished by the little
pads below the joints of the toes ; these pads being single in B.
viridisy and double in B, caiamita and in B. vulgaris.

The Green Toad spends most of the day in holes, although it is
not averse to daylight, and it roams about chiefly in the evening.
It can jump well, much better and oftener than the Brown Toad
The food consists strictly of insects of all kinds, and most
individuals prefer slow starvation to eating an earthworm.
Although continuing to live four or five years in captivity, they
do not readily become tame ; they are indeed no longer wild, and
when handled they no longer emit their peculiar insipid smell, but
on being approached they still crouch deeply into the grass, or
withdraw into their holes, just as they did when recently caught.
The voice is heard during the pairing season, and sounds like the
slow creaking of a door, or a combination of a spinning top and

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BUFONIDAE I 8 1

rattle. In Germany, during the months of April and May, they
take to the ponds, or, improvident like the common frog, to a
roadside ditch. The male sits upon the female and grasps her
below the arms, his hands on her breast, and in this position
they remain for days. The eggs are laid in two strings, twisted
around water-plants, and are very, numerous. H^ron-Eoyer
has calcidated them at 10,000 or more in one set. The
embryos are hatched, like those of the Common Toad, before
the appearance of the external gills and of the tail. In this
imperfect condition they remain in the jelly of the egg -strings
for a few days, while their external gills sprout out like un-
branched little stumps, only to disappear again. In about eight
weeks the tadpoles, which reach a length little more than 1^
inch or 40 mm., have metamorphosed and leave the water as
baby-toads scarcely half an inch in length.

This species has a very wide range, namely, the whole of
Middle Europe excepting the British Isles, France and the
Iberian Peninsula ; the region between the Elbe and Bhine being
its western limit ; southwards it extends over all the Mediter-
ranean idands and the north coast of Africa, eastwards through
the whole of Bussia, Western and Central Asia, not entering India,
but spreading along the Himalayas into China. Stoliczka men-
tions its having been found in the Himalayas at an altitude of
15,000 feet, the highest record of any Amphibian, at least in
such latitudes.

B. calcnnita, — The Natterjack is practically the representative
of the Green Toad in Western Europe, but both species occur
together in Denmark, Southern Sweden, and nearly the whole
of Germany- Its southern limit is Gibraltar. In the British
Isles it occurs in South- Western Ireland, in Co. Kerry, and in
England and Wales, l^eing however local, and preferring sandy
localities, where it is found in considerable numbera This
predilection is shown by its frequency on the sandy dunes of
most of the islands off the German and Dutch coast, where it
may be seen running about in glaring sunshine.

Besides in the coloration, it differs from B, viridis in the follow-
ing points. The little subarticular pads of the toe-joints are
I)aired, not single, and the hind-limbs are decidedly shorter, so
much so that this species cannot hop. But it runs well, like a
mouse, generally in jerks, stopping every few seconds, and owing

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1 82 ANURA CHAP.

to this habit it is called the "running toad" by the field-
labourers of Cambridgeshire. The skin is smooth, but less so
than in B, viridis, owing to the slightly more prominent warts ;
the parotoids are small; a similar pair of glands lies on the
upper surface of the fore-arm and another on the calf. The
tympanum is rather indistinct. The ground-colour of the upper
parts is light brownish yellow, with a green tinge and scattered
green spots ; most specimens have a narrow yellow stripe along
the vertebral line and over the head. The under parts are
white, more or less speckled with black. The iris is greenish
yellow and speckled. The male, which is of the same size as
the female, — very large specimens reaching 3 inches in length, —
has a large subgular vocal sac, and develops nuptial brushes on
the first three fingers, but the first lacks the thickened pad of
B, viridis.

The yellow vertebral line is sometimes absent in specimens
from the south of France and the Iberian Peninsula ; and since
these southerners are as a rule more handsomely marked, the
green being more pronounced and arranged in larger patches,
interspersed with red spots, they much resemble B, mridis.
Boulenger, who has paid especial attention to this vertebral
streak, which is a not uncommon design in various species of
different families, has made the interesting observation that the
streak has never been found in Danish and German specimens
of B. viridiSy where B, calamita occurs also, while it is not
uncommon in B, viridis of Italy, South-Eastern Europe, Asia,
and North Africa, where B. calamita is not found. Lastly, he
remarks that in Eastern Asia, where neither B, viridis nor
B. calamita with such a line occurs, the same character is
assumed by some specimens of B. vulgaris. The only conclusion
we can draw from these facts is, that for some unknown reason
the streak is a desirable, but not necessary, possession, but that
it is not kept by two species in the same country, B, viridis
dropping it entirely where the typically streaked species,
B. calamita, also occurs. The breeding season does not begin in
England and Middle Europe until the end of April, in cold springs
not before May, but it lasts for several months. The males, con-
gregating in pools in great numbers, make a loud noise, each
individual uttering a rattling note which lasts a few seconds,
the repetition distending its bluish throat into the shape of a

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BUFONIDAE 1 83

globe as large as its head. As the note is taken up by all the
other males, a continuous chorus is established, which on warm
and still nights can be heard nearly a mile o£f. Single croaks are
uttered at any time of the day. The embrace, the male digging
its fists into the armpits of the female, often takes place on
land, near the edge of the water, to which they resort in
the night for spawning. The egg-string3 are slung around water-
plants, unless the water is a mere puddle, and are much shorter
than those of B, viridis, measuring only 5 to 6 feet, and contain-
ing altogether 3000 to 4000 eggs. The larvae, when hatched,
are very small, imperfect, and blackish ; the external gills last a
very short time. The young tadpoles live on mud, subsisting on
diatoms and low Algae ; they are the smallest tadpoles of all
the European kinds, scarcely reaching more than one inch in
length, and they metamorphose quickly, the baby-toads leaving the
water and running about in less than six weeks, when they are
only 10 mm., scarcely three-eighths of an inch, in length. By the
end of their second summer they are still only three-quarters of an
inch long, and they do not reach maturity until the fourth or
fifth year, with a size of 1^ to 2 inches ; still smaller young
males become mature several years before they are full grown.

Natterjacks stand captivity well and become very tame.
When discovered, they first do their best to run away, instead of
hiding or squatting down, and when caught they become covered
with a slightly foamy lather, the exudation of their glands, which
has a peculiar smell, reminding some people of gunpowder, others
of india-rubber. They are not very particular as to food, all
sorts of insects and earthworms being taken. Natterjacks are
great climbers and diggers. Many of mine have established
themselves in the peat with which the walls of the greenhouse
are covered, where they have dug out, or enlarged, holes in
which they pass the daytime, just peeping out with their bright
eyes ; others sit high up, always in dry places, and bask. In
the evening they descend, hunting about on the ground, and
occasionally they go into the water, whereupon they become
quite flaccid and soft. When taken up and held between two
fingers, being slightly pressed under the armpits, both sexes
utter little jerky notes, as — by the way — most toads and frogs
do under similar conditions.

In Cambridgeshire they frequent certain clay-pits sun-ounded

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1 84 ANURA

by high and steep walls of sand, the breeding places of large
colonies of sand-martins. During the months of May and June
they are found in the shallow water, running about on the mud,
sometimes swimming, in which they are not very proficient, and
rarely diving. But they spend most of the time on land. Early
in October they climb up and enter the holes of the sand-martins,
or they dig large, deep burrows for hibernation, and the old
males are the first to disappear.

B, mauritanica s. pantherina,- — The " Pantherine Toad " is one
of the few African species, and is one of the prettiest of all toads.
The skin is almost smooth, although provided with porous glands.
The parotoids are large, but flat ; large glandular complexes on
the legs or arms are absent. The tympanum is very distinct
The upper parts are adorned with a delicate pattern of dark-
edged, rich brown or olive patches upon a light, buff-coloured
ground ; the under parts are uniform white ; the male has a
subgular vocal sac. The total length is 3 to 4 inches. This
beautiful species is one of the gentlest, and it becomes tame enough
to lap up food whilst sitting on one's hand. It lives entirely
upon insects, prefers shade and dusky light, and utters a sound
like " kooh-rr." It is a native of North-Western Africa, Algiers,
and Morocco. In the rest of Africa, from Egypt to the C'd\ye,
Seneganibia to Abyssinia, it is represented by B, regvlaris. This
species has often little spiny tubercles upon the warts, and
occasionally a light vertebral line ; the colour of the upper parts
either closely resembles that of the previous species, or it is
uniform liglit brown, while the under parts are whitish, or
variegated with brownish patches. West African specimens are
the smallest, only 2 inches long; those of the Cape are the
largest, reaching 5 to 6 inches.

The next two genera approach the Engystomatinae, and
therel)y lead from the arciferous towards the firmisternal type.
The epiconicoid cartilages are narrow^ and they scarcely overlap,
so that by a further step in this direction they could easily fuse
into the firmisternal condition. Another bond between these
two genera and the Engystomatinae is their habits, they being
aiit- eaters of an extremely stout appearance, with exclusively
short limbs and very small heads.

Myohatrachus gouldi, living in Australia, has a smooth skin,
brown above, lighter beneath, and is about 2 inches lone.

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HYLIDAE

185

Rhinophrynus dorsalis of Mexico is remarkable for its tongue,
which is elongated, subtriangular and free in front, so that it
can be protruded directly — not by i-eversion as in other toads
— and can be used for licking up the termites which seem to be
its principal food. The body of this ugly creature is almost egg-
shaped, and the head is merged into this mass, only the narrow
truncated snout protruding. The limbs are very short and stout.
The toes are more than half webbed, and there is a large oval,
shovel-like metatarsal tubercle, covered with horn and used for
digging. The general colour is brown, with a yellow stripe

the

along the spine and with irregular spots and patches on
Hanks and limbs. Total length 2 to 2^ inches.

Fam. 4. Hylidae (Tree-frogs). — The upper jaw — in Ampin

rp^^ SB

EHYLA

Fio, 36. — Map showing distribution of Hylidae. The vertically shaded countries are
inhabited by Hyla and by other genera of Hylidae ; the horizontally shaded countries
only by Ilyia.

gnaihodon the lower jaw also — and the vomers carry teeth ;
Triprion and Diagleim alone have teeth on the parasphenoid also,
and the latter genus is further distinguished by possessing
palatine teeth. The vertebrae are procoelous and have no ribs ;
the sacral diapophyses are dilated. The omo- and meta-stemum
are cartilaginous, the latter forming a plate with scarcely any
basal or style-shaped constriction. The terminal phalanges are
invariably claw-shaped and swollen at the base, and carry a
flattened, roundish, adhesive cushion. The tympanic disc is
variable in appearance, being either free, or more or less hidden
by the skin. The tongue is also variable in its shape and in the
extent to which it can be protruded.

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1 86 ANURA CHAP.

Most, if not all, Hylidae are climbers, and manjr lead an
arboreal life, but it does not follow that all the " Tree-frogs " are
green.

Their distribution is very remarkable. To say that this
family is cosmopolitan with the exception of the African region,
is literally true, but very misleading. There are in all about
150 species, and of these 100 are Notogaean; one-half of the
whole number, or 75, being Neotropical; 23 are Central
American, 7 Antillean, and about 18 are found in North
America. One species, Hyla arborea, extends over nearly the
whole Palaearctic sub-region, and two closely allied forms occur in
Northern India and Southern China. Consequently, with this
exception of three closely allied species, the Hylidae are either
American or Australian. We conclude that their original home
was Notogaea, and that they have spread northwards through
Central and into North America. The enormous moist and
steamy forests of South America naturally suggest themselves as
a paradise for tree-frogs, and it is in this country, especially in
the Andesian and the adjoining Central American sub-regions,
that the greatest diversity of generic and specific forms has been
produced. It is all the more remarkable that similar forest-
regions, like those of Borneo and other Malay islands, are
absolutely devoid of Hylidae (while there are about a dozen
species in Papuasia), whose place has however been taken for all
practical purposes by correspondingly modified Banidae, notably
the genus Rhdcophorus. Lastly, the fact that tropical evergreen
forests of Africa and Madagascar possess no Hylidae, but are
inhabited by several kinds of tree-climbing Bhacophoriis, points
with certainty to the conclusion that the origin of this large and
flourishing family of Hylidae was not in Arctogaea.

The versatility and the wide distribution of the Hylidae
has naturally produced cases of convergent analogy, and the
various species of one " genus " may be in reality a heterogeneous
assembly. Such an instance is probably the genus Eylella, of
which four species live in the Andesian and Central American
provinces, while the two others occur in New Guinea and
Australia.

The two North American genera Chorophilus and Acris, and
the Brazilian Thoropa, connect the Hylidae with the Cysti-
gnathidae, in so far as their finger-discs are very small, or even

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VI HYLIDAE 187

absent, and their sacral diapophyses are only slightly dilated.
On the other hand, it has to be emphasised that the possession
of adhesive discs on the fingers and toes does not necessarily
constitute a member of the Hylidaa That requires the further
combination of an arciferous sternum, with dilated sacral dia-
pophyses and teeth in the upper jaw. Finger -discs are easily
developed, and still more easily lost. Those of the typical
Hylidae are constructed as follows. The terminal phalanx is
elongated, claw-shaped, swollen at its base. Between it and the
penultimate phalanx lies an interphalangeal cartilaginous disc
which projects ventrally below the end-phalanx, thus assisting
the formation of the ventral pad, and the turning upwards of the
whole disc-like phalanx like the claw of a cat. This peculiar
motion can be well observed in Tree-frogs which are at rest
upon a horizontal leaf, or, better still, upon a rough stone, when
the creatures take good care to adjust their discs into a safe
and easy position. The pad or disc itself is furnished with
unstriped, smooth muscular fibres, the contraction of which pro-
duces one or more longitudinal furrows on the under side. When
the disc is in action or adhering, being flattened to a smooth
surface, the end-phalanx sinks into the cushion; when not in
action, the cushion swells and the phalanx appears as a slight
dorsal ridge. The disc is rich in lymph-spaces, and its surface
contains mucilaginous glands.

Various suggestions have been made to explain the function
of these discs. Suction, adhesion, and glueing -on have been
resorted to. Suction, through production of a vacuum, is quite
imaginary and does .not exist. The question has been thoroughly
studied by Schuberg.^ Adhesion is due to the molecular attrac-
tion of two closely appressed bodies. The less air remains
between them the stronger it is. Consequently it can be in-
creased by the interference of a thin layer of fluid, which as
everyday observation shows, possesses both adhesion and cohesion.
The more sticky the fluid, the more effective it is, as shown
experimentally by Schuberg, who moistened the under surface
of a glass plate, and pressed it against a little disc of glass from
which was suspended a weight A disc of 1 6 square millimetres,
approximately equal to the aggregate surface of the 18 discs of
a European tree-frog of 4 grammes in weight, carried with water-
1 ArbeUen InsUL WUrzlmrg, x. 1895, p. 57.

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I 88 ANUKA

adhesion no less than 14 grammes, with glycerine-sohition 20
grammes, — more than sufficient to suspend the frog. The sticky
secretion of its glands greatly enhances the adhesive power.
Tree-frogs, when hopping on to a vertical plane of clean glass,
slide down a little, probably until the secretion stififens, or dnes
into greater consistency. After a few days I find the glass-walls
of their recently cleaned cage quite dirty, covered everywhere
with their finger-marks. On the other hand, wet leaves or moist
glass- walls affi)rd no hold. The adhesion of these froga is
assisted in most cases by their soft and moist bellies, just as a
dead frog will stick to a pane of glass.

All Hylidae have a voice, often very loud, and enhanced by
vocal sacs, which are either internal, swelling out the throat, or
external, paired or unpaired.

The various Hylidae resort to all kinds of modes of rearing
their broods. Most of them lay many eggs, up to one thousand,
in the water, not coherent in strings but in clumps ; others lay
only a few, attach them to various parts of the body, or, as in
the genus Nototrema, the female receives them in a dorsal pouch.
These raodiiacations will be described in connexion with the
different species.

Sub-Pam. 1. Amphignathodontinae.— i?o^/t dipper and hucer
jaw with teeth.

Amphignathodon, of which only one species is known, A.
guentheri of Ecuador, agrees with Nototrema in all important
characters except that it possesses teeth in the lower jaw
in addition to those in the upper. There are further differ-
ences, but tliey are of degi-ee only. The . sacral diapophyse3
are more strongly dilated and the omosternum is absent. The
tympanum is distinct. The pupil is horizontal; the roundish
tongue is slightly free behind. The terminal phalanges are claw-
shaped and cany large discs. The female has a dorsal pouch
opening backwards. The skin of the head is involved in the
ossification of the cranial bones. The skin of the back is smooth,
slightly tubercular, non-granular below. The middle of the
upper eyelid carries a small, pointed, cutaneous appendage, and
even this little character occurs also in some species of XototreMth
e.g, in N, longipes and in N. cornutum. The heel carries a
triangular little flap. The upper parts are olive in spirit-
specimens, probably green in life ; the borders of the doi-sal pouch

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HYLIDAE 1 89

are black. The sides of the body are adorned with a black,
white-edged streak, the limbs are whitish, with black cross-bars.
The total length of the female type-specimen is 3 inches.

Snb-Fam. 2. Hylinae. — Lower jaw toothless.

The Hylinae are divided by Boulenger into 13 genera, which
can be recognised by the following key, without reference to
their natural afi&nities : —

A. The contracted pupil forms a horizontal slit

a. Tipe of the fingers and toes with laige disca

"a. With vomerine teeth.

Female without a donal pouch . Hyla, p. 189.

Female with a dorsal pouch . . Nototremay p. 202.
p. Without vomerine teeth . Hyldla^ p. 203.

b. Tips with very small discs. Tongue free l^ehind.

Tympanimi distinct North America

and Peru ..... ChorophihiSj p. 208.
Tympanum indistinct North America Acris grylliLa, p. 207.

c. Tips simply swollen, not dilated into discs.

Brazil ....... Thoropa miliaris,

p. 209.

B. The contracted pupil forms a vertical slit Tropical America.

a. Tips with large discs.

a. Tongue extensively free behind.

Inner finger and toe opposable . Phyllomedum, p. 203
Inner finger and toe not opposable . Agalychnis, p. 206.
p. Tongue scarcely free behind. Ecuador NycHmantis rugicq)s^

p. 206.
With parasphenoid teeth and peculiar

helmet-shaped head. Yucatan . Triprion^ p. 207.
Ecuador . Dtaglenay p. 207.
Without parasphenoid teeth. Head
peculiarly helmet-shaped. Pupil

rhomboid. Brazil . . Corythomantis green-

ingi, p. 207.
6. Tips without discs. Without parasphenoid

teeth, but head peculiar in shape. Mexico Ptemohyla fodiensy

p. 207.

Hyla. — The pupil is horizontal. The tympanum is distinct or
hidden. The tongue is entire or slightly nicked in its hinder
margin, which is more or less free behind. The fingers and toes
are provided with typical adhesive discs.

This is the largest genus of all Amphibia, containing about
150 species, and its distribution coincides with that of the whole
femily. Many of the species are very closely allied to each

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I90

ANURA

CHAP.

other, differing only in small points, for instance in the extent
of the webs to the lingers and toes, the configuration of the
vomerine teeth, the size and appearance of the tympanic disc,
and the relative length of the hind-limbs. In some of the West
Indian, and in one Brazilian species, H, nigromacvlata, the upper
surface of the head is rough, owing to the cutis being involved
in the cranial ossification. Bony or perhaps only calcareous
deposits in other parts of the skin are rare, but are notably
developed in ff, diisynotus of Brazil, in which they extend from
the head to the sacrum, rendering the skin immovable.

Many are capable of changing colour to a great extent, and
it is a popular error to suppose that all tree-frogs are green,

s^.^;i^^'^V%.':">''^^''i»^

.mBD

Fig. 37.— //y« arborea, var. meridioncdis. South European Tree-frog, x 1.

although this colour is perhaps the most common in the arboreal
kinds.

H. arhorea. — The tongue is rather round, slightly nicked
behind, and can be protruded but little. The tympanum is dis-
tinct, but small. The upper parts are grass-green, quite smooth
and shiny owing to the skin being covered with a film of
moisture ; the under parts are yellowish-white and granular, flesh-
coloured or rosy on the thighs. Total length of large females
- inches. This, the Tree-frog of Europe, has an enormous
range, namely, from Morocco, France, and the south of Sweden,
South ^^.^'""^^ ^^ ^^^^1^ a^d Asia Minor to Japan and

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HYLIDAE 191

Several varieties have been described : the typical or European
form is ornamented with a narrow black stripe, which, beginning
at the nose, extends backwards along the side of the body to the
groin, where it generally forms a hook turned upwards. This
black colour forms the ventral boundary of the green, and is
itself narrowly seamed with white on its upper border.

In the south of France, the Iberian Peninsula, Morocco, and
the Canary Islands the black lateral stripe is often absent ;
this is is the var. meridionalis. In Spain and Portugal both
forms are found in the same localitiea

In the Asiatic, chiefly in the eastern specimens, the lateral
stripes tend to break up into irregular spots, vanishing altogether
towards the groins ; this var. aavignyi s. japonica occurs also on
most of the Mediterranean islands.

IT. arhorea can change colour to a great extent, mostly in
adaptation to its immediate surroundings, but ill health and
moulting may also influence it. The change is slow. The usual
colour is green, brightest on bright, sunny hot days, dull when
the sky is overcast, or when it is windy and showery. Day and
night have no influence upon the colour-changes. The hue of
the green agrees mostly with that of the foliage on which the
frog happens to take its rest, for instance a field of Indian corn,
birch-trees, or oak-trees. I once received a consignment from
Saxony. When the box with moss was unpacked, they were of
the dullest greenish-grey ; they were put into a wired-oflf corner
of the yard and were given the freshly cut branches of a lime-tree
to sit upon. On the following morning I at first looked for
most of the frogs in vain. The leaves had withered and all those
frogs which sat upon the dark brown branches had put on a light
brown garb, mottled with darker patches.

Another specimen, one of several which were at liberty in
a greenhouse, took to resting on the frame of the window-pane,
in a corner where putty, glass, and discoloured white paint met ;
in the morning it was always of a mottled leaden colour, but
during the nocturnal hunting it was green. In the winter, the
window-corner being of course cold, the frog remained stationary
for several months, but kept the leaden grey colour, until one
day in the early spring it was mottled with green, and soon after
it joined its green mates.

Liebe observed a half grown tree-frog which he kept in Gera

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192 ANURA CHAP.

during the winter in a glass with water-cress. While the temper-
ature was near freezing the frog sat in the water, very lethargic,
breathing perhaps once every quarter of an hour. Its colour
was light green. When the water-cress was cut and removed,
the frog darkened and became at laat quite a discoloured grey.
When the waternjress was put back, the creature reassumed the
light yellowish-green colour, remaining in its lethargic condition
until it became lively in the spring sunshine.

The European tree-frog spends most of its time in the summer,
after the pairing is over, in trees, often in the very crowns ; but
the neighbourhood of even a small patch of Indian com has still
greater attractions. There are all sorts of green insects to be
caught, there are fair chances of coming across the common
Cabbage White, a butterfly which the tree-frog loves, and last
not least the large luscious leaves afford a firm foothold, and the
axillae between stalk and broad-based leaves are just the places
for the frog to slip into, where nobody can find it. During the
day they mostly sit still, on the keen look-out for passing insects,
which, when they settle within reach, are jumped at ; otherwise
they have first to be stalked. The jump is quite fearless,
regardless of the height above ground; there is the leaf upon
which the prey sits, and even if this leaf be missed, there are
others, and one of them is sure to be struck by some of the discs
of either fingers or toes. If the fall is broken by the toes, and
the new leaf or branch is very elastic and bends down, then there
are some frantic antics to be gone through until the frog has
settled itself again. Then the large blue-bottle, or the butterfly,
is devoured at leisure, wings and all being poked in with the
assistance of the little hands. But the real hunting-time is the
night.

During a shower the frog shifts its position to the under side
of the leaf, or into a less slippery position, and during continuous
wet it descends into the grass, or it takes to the water. Its
greatest enemy is the Grass Snake, which prefers it to anything
else, not minding the poisonous secretion of the skin, which is
sharp enough to produce sneezing or even temporaiy blindness
when incautiously brought into the human eye.

The male h«ts an internal vocal sac, which, when inflated,
bulges out the whole throat into a globe, much larger than the
head. The voice is a sharp and rapidly-repeated note, something

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HYLIDAE 193

like " epp-epp-epp," or " creek, creek, creek," with more or less
of an a sound. It is uttered at any time of the day, more fre-
quently at dusk, and of course chiefly during the pairing season.
This tree-frog suffers from the reputation of being a good weather-
prophet, and it is for this reason often kept in confinement, the
orthodox abode being a muslin-covered glass jar, with a hole to
put flies through, water and plants at the bottom, and a little
ladder to sit upon. The prophesying is of the usual popular
unreliable nature, although the little creature, provided it is
a male, often sounds its voice on the approach of a shower, or
when there is a thunderstorm in the air. During continuous fine
weather it sits on the top of the ladder, or is glued on near the
rim of the glass, while on wet and dull days it is less active, and
may keep nearer the ground or in the water. There is a German
rhyme which well expresses the prophet's reliability by its am-
biguity : —

Wenn die Laubfroeche knarreD,

Magst du auf Kegen barren.

When the tree-frogs croak, you may wait for rain. Sometimes
it does come true.

Tree-frogs are not very intelligent, although they have a keen
sense of locality ; but they are nice pets, being easily kept, and
have a pretty appearance. There is a record of one which lived
for twenty-two years in confinement.

The pairing begins soon after the frogs reappear from their
hibernation in the ground ; in Germany in the month of May.
The congregating males make a great noise and take to the water
before the females, which join them when ready to spawn.
The male grasps his mate near the shoulders, and the pair swim
about together, sometimes for days, imtil the eggs are expelled.
These are laid in small clumps of 800 to 1000, which soon
swell up and remain at the bottom of the pond. The larvae
are hatched in ten days; two days later the adhesive sucker
below the throat appears, and after another two days a pair of
thread-like external gills are developed. The tadpoles, which
reach a length of 2 inches, owing to the long tail, which is
nearly three times as long as the body, metamorphose in about
twelve weeks, and the baby tree-frogs, scarcely half an inch in
length, hide in the grass for the next two years, until they are
about half grown, not reaching maturity until the fourth year.
VOL. viii 0

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194 ANURA <^HAP'

Since many pairs congregate in the same pool, and each
produces up to one thousand eggs, most of which are hatched,
the neighbouring meadows sometimes literally swarm with tiny
tree-frogs. Nevertheless the adults are comparatively rare and
are very local.

H. carolinensis a lateralis of the South-Eastern States of
North America greatly resembles K arborea in general appear-
ance, size, and habits. But the head is more pointed, and the
vivid green of the upper parts is separated from the yellowish
white under surface by a conspicuous, pure white line, giving the
little creature a very smart and neat appearance. According to
Holbrook, it ascends trees, but most commonly lies upon broad-
leaved water-plants, like JVymphaea, and in fields of Indian corn.
Motionless during the daytime, they emerge in the morning and
evening from their hiding-places, and become very brisk and
noisy, often repeating their single note, which is not unlike that
of a small bell. When one begins, hundreds take it up from all
parts of the corn-field.

Among other tree-frogs of the South-Eastern States may be
mentioned H. squirella, 1^ inch in length, which is very change-
able in colour, generally olive above with darker spots and bars on
the limbs, and with a white upper lip. It lives in trees, sheltering
in the bark. H, femoralis of the same size, without the whit«
lip, lives high up in the trees of the dense forests of (Jeorgia and
Carolina.

H. versicolor is one of the most delicately coloured species of
Eastern North America, extending northwards into Canada. It
is about 2 inches long. Its colour passes within a short time
from dark brown or olive grey to pale delicate grey, almost
white, occasionally retaining a few large darker patches on the
back, and delicate cross-bars on the limbs. A small portion of
the sides and the posterior part of the belly are bright yellow.
The %kin is granular, owing to the presence of small warts which
produce an acrid secretion. It is said to be found in trees, or
about old stone fences overgrown with lichens, the colour of
which it resembles to perfection. It becomes very noisy towards
the evening, in cloudy weather or before rain, the voice consisting
of a liquid note, terminating abruptly, like " l-l-l-l-luk." My
own captives fully bear out this statement of Holbrook's. Settled
motionless during the day upon a piece of bark in a shady

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VI HYLIDAE 195

corner, but occasionally uttering the quaint and rather faint
note, they become very lively in the evening, catching insects by
long jumps, or investigating the hollows of decaying mossy
stumps. Their general colour is then spotless, almost silvery
grey. In the day-time they are sometimes suffused with delicate
green.

The propagation has been studied by Miss M. H. Hinckley.^
They pair in shallow pools, in Massachusetts, in May. On the
10 th of that month eggs were attached singly, and in groups,
on the grasses resting upon the surface of the water ; first drab-
coloured, they became lighter in a few hours. Some larvae
escaped from the gelatinous envelopes on the following day, the
others on the third day ; they clung to the grasses by means
of their prominent suckers. The head and body were cream-
coloured, with olive dots, and averaged ^ inch in length. Gills
appeared on the fourth day, to disappear again during the four
following days, first those of the right, then those of the left,
side ; the suckers became less conspicuous, and the general colour
turned into deep olive-green, with fine golden dots on the upper
and lower surfaces. The eyes were of a brilliant flame-colour.
On the eleventh day the suckers or " holders " had disappeared,
and the hind-limbs were indicated by small white buds. By
June 5 th, i.e. the twenty-seventh day, the toes developed the
terminal discs ; tlie mottling of gold had given way to a uniform
olive or pea-green. Movements of the future arms beneath the
skin appeared on the 28 th of June, at the age of seven weeks.
The arms, mostly the right one first, were thrust out on the 2nd
of July ; the fins of the tail were absorbed rapidly, and towards
the end of the seventh week the nearly transformed creatures
began to leave the water. The young frogs changed colour
rapidly, in adaptation to their surroundings, but the four
specimens which survived were never all found to be of the
same colour during the next three months. They first Rved
upon Aphides, later upon flies, and they were alert nocturnally.
About the beginning of October they left the fronds of their
fernery and nestled away in the damp earth, which they left
only when the temperature rose above 60° F.

ff. vasta of Hayti is the giant of the tree-frogs, reaching a
length of 5 inches. In order to support its great weight the

1 Pfoc. Bo8l. Soe. Aa^. Hist. xxi. 1883, p. 104.

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ig6 . ANURA

adhesive discs of the fingers and toes are of a surprising size,
about as large aa a threepenny piece. The skin is covered with
small warts, and forms a peculiar fold on the hinder surface of
the fore-arm and on the tarsus, and small flaps near the vent
The colour is grey above, blackish on the head, with a brown
band between the eyes ; the under parts are flesh-coloured, the
throat with black spots.

H. maxima, of the forests of British Guiana, is scarcely less
gigantic, and is distinguished by a projecting rudiment of the
pollex, while the adhesive discs are smaller than the tympanum.
The skin forms folds on the arms and tarsus, like those of K
vasta, in addition to a triangular flap at the heeL The general
colour is reddish-brown above, sometimes with a dark vertebral
Une, the under parts are whitish and covered with large
granules ; the throat of the male, which has an inner vocal sac,
is brown.

H.fdber of Brazil is closely allied to the last species, but the
skin of its upper parts is quite smooth. There is a small tarsal
fold, and one extending from the upper eyelid to the shoulder.
It is light brown above, with darker marks which form a con-
spicuous vertebral line, transverse bars on the hind-limbs, and
a few irregular, scattered, vermicular or linear marks on the
head and body. The adult, when put into a strong light, will
rapidly turn pale ; at night the longitudinal stripe on the back
and the bars on the hind-limbs become very distinct ; the under
parts are white, and exhibit a beautiful orange tinge. This is
the famous " Ferreiro " or " smith." As will be seen from tbe
following graphic account by Dr. Goeldi^ of Para, this species
doubly deserves its name o^faber, not only in virtue of its voice,
but also because of the marvellous nest-building habits recently
discovered.

" The Ferreiro is common in the Province Eio de Janeiro,
more frequently still in the mountain regions of the Serra dos
Orgjlos than in the hot lowland. Its voice is one of the most
characteristic sounds to be heard in tropical South America.
Fancy the noise of a mallet, slowly and regularly beaten upon a
copper plate, and you will have a pretty good idea of the concert,
given generally by several individuals at the same time and with
slight variations in tone and intensity. When you approach the
^ P.Z.S. 1895, p. 89 (with a sketch of a pond, with nests, in Dr. Goeldi's garden).

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VI HYLIDAE 197

spot where the Tree-frog sits, the sound ceases. But keep quiet,
and it will be resumed after a few moments. You will discover
the frog on a grass-stem, on a leaf of a low branch, or in the
mud. Seize it quickly, for it is a most wonderful jumper, and it
will utter a loud and shrill, most startling cry, somewhat similar
to that of a wounded cat."

The "Smith" makes very regular pools, in the shallow
water of ponds, or nurseries for the tadpoles surrounded by a cir-
cular wall of mud. Dr. Goeldi has watched the building process
during a moonlit night : " We soon saw a mass of mud rising to
the surface carried by a Tree-frog, of which no more than the
two hands emerged. Diving again, after a moment's time, the
frog brought up a second mass of mud, near the first. This was
repeated many times, the result being the gradual erection of a
circular wall. From time to time the builder's head and front
part of body appeared suddenly with a load of mud on some
opposite point. But what astonished us in the highest degree
was the manner in which it used its hands for smoothing the
inside of the mud wall, as would a mason with his trowel.
When the height of the wall reached about 4 inches, the frog
was obliged to get out of the water. The parapet of the wall
receives the same careful smoothing, but the outside is neglected.
The levelling of the bottom is obtained by the action of the lower
surface (belly and throat principally) together with that of the
hands."

The male takes no active share in the construction of the
nest, but will suddenly climb up the wall of his home, and then
upon the back of his busy mate. The building operation may
take one or two nights, and is performed in the most absolute
silence; the croakers around are all males clamouring for a
mate.

The eggs are laid during one of the following nights, and are
hatched some four or five days later, the parents keeping hidden
in the neighbourhood of the nurseiy. Heavy rains may destroy
the walls, and thus prematurely release the tadpoles.

It is only owing to such keen observers and lovers of
nature's fascinating ways that the breeding habits of some
Brazilian Hylidae have become known.

H. nebvlosa, a luteola also living in Brazil, is yellow above,
with brown dots ; the sides of the belly and thighs have trans-

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198 ANURA CHAP.

verse bluish bars, the under parts are whitish. Its size is under
2 inches. Goeldi has often found it in the sheaths of decaying
bapana-leaves. It glues the lumps of eggs on to the edges and
to the inside of the withered leaves, where even during the hot
hours of the day sufficient coolness and moisture are preserved.
These lumps are enveloped in a frothy substance, in which the
nearly metamorphosed tadpoles can be watched wriggling. If
these are put into water, all will die in a few hours.

H. poly taenia deposits its eggs in free lumpy masses on water-
plants. It is a small creature, little more than 1 inch in length,
light olive above, with numerous brown parallel longitudinal
bands on the body and limbs. A dark, white - edged band
extends from the nose along the side of the body. The heel has
a short flap of skin. The male has an internal vocal sac.

H, goeldii is a most interesting form, leading to the allied
genus Nototrema, Boulenger ^ has described a female which was

captured by Goeldi on the
5th of January, near Para.
It is about 1^ inch long.
The whole surface of the back
is occupied by a layer of
twenty -six pale yellow ^^'g^
Fio. ss.—/fyia goeUiii. x 1, Female with which are 4 mm. in diameter.

eggs in the incipient dorsal brood-pouch. r^^^ ^j^.^^ ^^ ^^^ ^^^j, j^

expanded into a feebly reverted fold, which borders and supports
the mass of eggs on the sides, thus suggesting an incipient stage
of a dorsal brood-pouch. Owing to the great amount of yolk,
the young are probably able to remain upon the mother until
they are nearly metamorphosed.

H, coerulea s. cyanea is one of the largest Australian green tree-
frogs, ranging from the South to the very North of Australia.
The discs are as large as the fully-exposed tympanum. There
is no projecting rudiment of the pollex, but a slight cutaneous
fold borders the inner side of the tarsus. The skin is smooth
and shiny, always a little moist, and studded with numerous
rather large pores on the nape and shoulders ; this somewhat
thickened region forms a prominent fold which begins behind
the eyes. The belly and the under parts of the thighs are
granular as in most Hylidae. The male has an internal vocal

» P.Z.S. 1895, p. 209.

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HYLIDAE

199

sac ; and during the breeding season, which seems to occur during
our autumn and winter, develops brown rugosities on the inner
side of the first finger. The tongue is round, slightly notched
behind and free enough to be protruded a little.

The alternative specific names are most unfortunately chosen,
as they apply only to spirit-specimens. During life this tree-
frog exhibits a considerable amount of colour -changes. The
normal colour is bright green above, white below. A conspicuous

Fig. 39. — Uyln coeruUa. Australiau Tree-frog (from photogrRphs). Length of the
large specimen 4-2 inches. Tlie upper right specimen with vocal sac inflated.

feature of this species is the frequent occurrence of white specks
or spots, which are probably due to the deposition of guanine,
a peculiar white colouring matter. The spots appear in any part
of the green skin, and are quite irregular in their distribution.
Sometimes they remain for weeks in the same place, or they dis-
appear after a few days and others appear. They are in no way
connected with the shedding of the skin, nor do they indicate ill-
health. R, coerulea lives well in confinement, and becomes tame
enough to take food from one's fingers, even when sitting upon the
hand. Some of mine took to living during the daytime in a small
box, preferring a crowded condition in companionship with Natter-

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200 ANURA CHAP.

jacks. Others squeeze themselves into the most uncomfortable
cracks, while others again prefer the broad leaves of Philodeiidron.
A favourite plac« for two or three at a time is the funnel-
shaped spaces formed by ^rom^/ia- plants. Those specimens
which are hidden in the box or in the hollows of rotten
stumps are, almost without exception, dull, very dark brownish
olive, while those on the Bromelias assume exactly the sombre
dull green of its leaves. Lastly, those which sit in the light,
exposed places, no matter if upon a leaf, on a white stone, or
upon a board, are emerald -green, especially beautiful on hot,
sunny days ; — and they are not always averse to the full glare of
the sun. When squatting upon a flat surface, such as a broad
leaf, they tuck the fore-paws under the head like a cat, and with
half open eyelids, the pupil contracted to a tiny slit, so that the
golden iris is exposed, they remain motionless during the day.
They take food when offered, but at night they roam about,
either hopping on the ground, or making enormous leaps from
leaf to leaf, sometimes deliberately stalking some choice insect,
and patiently climbing up a stem, hand over hand. At night
their whole aspect is changed. The colour is saturated green,
the eyes are transformed into round, projecting shiny black
beads, and the head is erect. The ludicrously dreamy, complacent
look has given way to wide-awake alertness. They take all
kinds of living food. When they find an earthworm, they first
look at it, bending the head sharply down, lift themselves upon
the fore-limbs and then pounce upon it, nipping the prey with
the jaws, and then poking it down deliberately with the hands.
Cockroaches are simply lapped up, and disappear in the twinkle
of an eye. Mealworms, wood-lice, butterflies and moths, flies and
spiders are taken. The stomach of a specimen in the Dresden
Museum, from the Aru Islands, contained some four or five young
freshwater Crustaceans of the genus Sesarma. They fortunately
do not molest smaller frogs of their own kind and of other
species. Like many Amphibia they like a change of diet, and
ultimately refuse their food if it is unvaried. To my surprise my
largest specimen, which measures a little more than 4 inches,
takes snails. Helix virgata, half-a-dozen at a time, and on the
following day, not during the night, vomits the sucked-out shells
in a lump, like the pellets of birds of prey. During this rather
painful-looking procedure the whole tongue and about half an

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HYLIDAE 20 1

inch of the everted gullet are protruded out of the mouth, and
are then slowly withdrawn. After having roamed alx)ut all
night, they return to their respective resting-places, where each
individual is sure to be found in exactly the same spot, day after
day. They do not mind being looked at, but if taken up and
put back they avoid that place for perhaps a week, taking
shelter somewhere else.

Both sexes have a voice, but that of the female is only a
grunting noise, while the male inflates its gular sac and sends
forth a sharp cracking sound, which can turn into a regular
bellowing like the gruff barking of an angry dog. They bellow
at any time of the year, freqiiently on the approach of a shower
or during a thunderstorm. Certain noises will also induce them
to bark. The rattling produced by the syringing of the
greenhouse, sawing of wood, hammering, the raking of the gravel,
or even the scraping of boots on the gravel -path is liable to
start one of the males, and the others are sure to chime in.

According to Fletcher, H. coervlea and H. aurea lay their
eggs in round white frothy patches, which float in the water,
chiefly during the months of August and September ; but when
the spring months are very dry, the pairing is delayed until
the following January. Several other Australian species of
Hyltty e.g. H. ewiiigi, spawn at any time of the year if the
conditions are favourable. They attach their eggs to sub-
merged blades of grass or to twigs.

H. aurea is one of the commonest and most beautiful species,
occurring throughout Australia and Tasmania, excepting of
course in the large deserts. It has the appearance and restlessness
of a water-frog, is not unlike RaTia esculenta, and grows to about
three inches in length. The tympanum is very distinct, but
rather small The fingers are without a poUex-rudiment, the
tarsus has a fold along its inner edge. The adhesive discs are
decidedly small. The male has two internal vocal sacs, which
bulge out sideways. The skin is smooth and shiny. The
under parts are white ; the upper parts are, speaking generally,
a mixture of blue and oUve, with blue or brown spots, but spirit-
specimens give no idea of the beauty which this changeable
species can assume. Sometimes the same individual is saturated
blue and green, with several longitudinal stripes of burnished
copper along the back ; a few minutes later the stripes glitter

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202 ANURA cilvp.

like gold, and in other moods the whole upper surface is mottled
blue, green, and brown. My specimens often went into the
water and did not climb. The food is said to consist chiefly of
other small frogs in preference to insecta

Nototrema differs from Hyla in so far as the female has a
pouch on the back for the reception of the egga This bag is
formed by an infolding of the skin ; it opens backwards in front of
the vent, it has a sphincter and is permanent, although it distends
to larger dimensions when in use. An initial stage of such a pouch
is possessed by Hyla goeldii (Fig. 38). The pupil is horizontal,
the tongue can be protruded but little ; the tympanum is free,
and the adhesive discs of the fingers and toes are well developed.
These " marsupial frogs," of which about half-a-dozen species are
known, live chiefly in the tropical forest-region of South America,
notably from Peru to Venezuela.

K, marsv.piatum is green with darker blue-green spots, or
with longitudinal patches which are each surrounded by a
whitish or yellow seam of little dots. The limbs have cross-bars.
Total length about 2^ to 3 inches. The eggs of this species
are comparatively small and numerous. The very small tadpoles
have no external gills, and escape from the pouch to finish their
metamorphosis in the water.

X. tfstudineum, about 3 inches in length, is of a uniform lead-
colour, but is lighter beneath. The skin of the back is studdeii
with stellate calcareous deposits, a peculiarity which is alludeti
to in the specific name.

.V. oviferum is brown above, with darker patches on the sides
of the body and with cross-bars on the limbs. The last two
species and K Jissiiics of Brazil, near Pernambuco, carry their
young in tlie pouch until the metamorphosis is completed. This
long nursing-period necessitates a great amount of food-yolk in
the eirtrs^ and this enlargement in turn implies a considerable
reduction in their number. The female's load consists of about
fifteen eggs only, but these are of a great size, namely one-eighth
of the length of the mother's body.

X, pyfjmaeuvi, in Venezuela, is a tiny creature. The female,
just one inch in length, carries only from four to seven eggs. It
looks then " as if it carried a sac filled with a few gigantic baUs."
This species is further worthy of note on account of the opening
of the brood-pouch, which is a longitudinal slit, whence a kind

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HYLIDAE 203

of thin and slightly elevated ridge or fold of the skin extends on
to the neck. The suggestion, that this seam is burst open, in
order to set the full-grown young free, instead of their passing
through the existing opening, is scarcely credible.

These Neotropical tree-frogs seem to be rare, and females with
embryos are of course still more uncommon, so that the best
account of their structure is still that given by Weinland ^ of N,
ovi/erum. How the eggs get into the pouch has not yet been
observeil, but it is most likely with the help of the male, im-
mediately after fertilisation. The pouch forms two blind sacs
which extend forwards over the sides of the back. The eggs are
large, 1 cm. in diameter, and the enclosed embryos, or rather
tadpoles, had a length of 15 mm., with a large amount of yolk still
contained in the spirally wound intestine. The first two gill-arches
carried each a double thread, which expanded into a funnel-
shaped membrane, not unlike the flower of a Convolvuhis, and
furnished with a capillary network ; the stalk contained muscular
fibres. These most peculiar structures are of . course the much
modified external gills. Those of K testudineum and N. cornutum
are likewise bell-shaped.

Hyhlla differs from Hyla chiefly by the absence of vomerine
teeth, and consists of about half-a-dozen small species, about one
inch in length. The fact that two species live in Queensland
and New Guinea, while the others are natives of tropical America,
suggests that this genus is not a natural but an artificial
assembly, an instance of convergent evolution.

Phyllomedusay composed of about one dozen species of tree-frogs,
is characterised by the vertically contracted pupil, large adhesive
discs, and the opposable nature of the inner finger and of the
hallux, the last joints of which are like thumbs. The sacral
diapophyses are strongly dilated. The range of the genus
extends from tropical Central America to Buenos Aires. Most
of the species are about 2 inches in length, blue -green to
violet above, with white purple-edged patches on the sides of the
body ; the under parts uniform white, or with purple or brown
patclies. The male has a subgular vocal sac. Some have more
or less distinct parotoid glands. Ph. dacnicolor of Mexico is
uniform green above, whitish below, and attains a size of more
than 3 inches. In Ph. bicolor of Brazil, the skin of the upper

^ Arcfi. Anat. und Phys. 1854, p. 449. Also Boulenger, P.Z.S. 1898, p. 107.

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204 ANURA

parts is studded with calcareous deposits, and the parotoids are
large. It is blue-green above, purplish white below, the sides of
the body and limbs with white purple-edged spots.

Ph. hypochondricdis has been found breeding freely in the
Paraguayan Chaco by Budgett,^ from whose account the
following notes have been extracted. This brilliantly coloured
frog is green above, which colour may become brown -grey or
bluish at will ; below, white and granular. The flanks are
scarlet, with black transverse bars, and the plantar surfaces are
deep purplish black. Total length about 1| inch.

The " WoUunnkukk," as it is called by the Indians, from the
call of both male and female at pairing time, is extremely slow
in its movements, and is active only at night. At this time, if it
is seen by the aid of a lantern as it slowly climbs over the low
bushes and grass, it is very conspicuous. In the daytime, how-
ever, nothing is seen but the upper surface of the body as it lies
on the green leaf of a plant. It has a remarkable power of
changing its colour to harmonise with its surroundings, and can
efieet a change from the brightest green to light chocolate in a
few minutes. The skin is also directly sensitive to light ; for if
the frog is exposed to the sun while in a tuft of grass in such a
way that shadows of blades of grass fall across it, on removal it
will be found that dark shadows of the grasses remain on the
skin, while the general colour has been raised to a lighter
shade. Its food consists largely of young locusts. The species
on each side are divided into five distinct clusters. The creature
has a large saccular diverticulum, which is very heavily pig-
mented.

In the breeding season — December to February — this beauti-
ful frog collects in considerable numbers in the neighbourhood of
pools. During the night-time they call incessantly to one
another, and produce a sound as of a dozen men breaking stones,
well imitated by the native name.

The eggs are enclosed in batches in leaves near the margin of
the water. Budgett has been able to watch the whole process of
oviposition and fertilisation. He found, at ll P.M., a female
carrying a male upon her back, wandering about in search of a
suitable leaf. At last the female, climbing up the stem of a
plant near the water's edge, reached out and caught hold of the

* Quart. J. MicT. Set. xlii. 1899, p. 313.

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HYLIDAE 205

tip of an overhanging leaf, and climbed into it. Both male and
female held the edges of the leaf together, near the tip, with their
hind-legs, while the female poured her eggs into the funnel
thus formed, the male fertilising them as they passed. The
jelly in which the eggs were laid was of sufBcient firmness to
hold the edges of the leaf together. Then moving up a little
further, more eggs were laid in the same manner, the edges
of the leaf being fastened together by the hind-legs, and so on up
the leaf until it was full. As a rule, two briar-leaves were filled
in this way, each containing about 100 egga The time occupied
in filling one leaf was three-quarters of an hour.

Development proceeds rapidly. Within six days the embryo
increases from the 2 mm. of the egg-diameter to 9 or 10 mm.
When it leaves the leaf it is a transparent glass-like tadpole, whose
only conspicuous parts are the eyes. These are very large and
of a bright metallic green colour, so that when swimming in the
water all that is seen is a pair of jewel-like eyea The newly-
hatched tadpole has also a bright metallic spot between the
nostrils somewhat in front of the pineal spot. This is the point
which touches the surface of the water when the tadpole is in its
favourite position. Whether it is a protective coloration, or
some mechanical arrangement for holding the surface, Budgett
could not make out.

The egg contains a great amount of yolk ; the rest of the
jelly-like contents of the egg becomes fluid, so that towards
the end of embryonic life the larva comes to lie quite freely
within a membranous capsule. The external gills app^r
on the third day, and reach their greatest size on the fifth,
when these bright red filamentous organs extend beyond
the vent. By the time the tadpoles are ready to be hatched
these gills have quite disappeared, there is a median spiracle,
and the lungs are shining through the transparent body-
wall. Five weeks later, i.e. six weeks after the eggs were laid,
the tadpole is 8 cm. long, glossy green above, rosy and silvery
below, and the hind-Umbs protrude. The young frog at the
close of its metamorphosis is two-thirds the length of the adult,
and at this time acquires the red flanks barred with black.

The first account of the breeding of Phyllomedusa was
given by v. Ihering^ concerning Ph, iheringi of Southern Brazil.

1 Ann. Mag. Nat. Hist. (5) xvii. 1886, p. 461.

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206

ANURA

" Phyllomedum does not lay its eggs in the water, although
the larva develops in that element, but in the open air in masses

50 millim. long by 15-20 broad,
between leaves hanging over the
water. Willows are frequently used
for that purpose. The e^^ - mass
contains rather large white ova,
wrapped up between two or three
leaves in such a way as to be com-
pletely enveloped save an inferior
opening. My attempts at rearing
the eggs failed owing to the leaves
drying up ; but I am assured that
the tailed larvae may be seen wrig-
gling in the gelatinous mass. As
at a later period the latter is found
Fio. 40. -A branch with eggs of empty, we must infer that the larvae

Phyllomedusa iheringi, x 1, en- drop intO the Water bclow. The
veloped in the leaves. (After v. i» j i i

ihering.) ^ggs ^^c found Only on plants

hanging over stagnant water."

" The adult animal is a stupid creature, and will let itself be
taken without attempting to escape. Their moderately loud
voice resembles somewhat the sound produced by running the
finger nail over the teeth of a comb. Only during the breeding
season, in the month of January in Rio Grande do Sul, do these
frogs make their appearance ; at other times not one is to be seen,
probably because they establish themselves high up in the trees."

Agalychnis, with two species in Central America, is practi-
cally like Hyla ; but the pupil is vertical, and the tongue is
extensively free behind.

Nyctimantis differs from either by its round tongue, which is
not nicked behind, and is almost completely adherent, much
resembling that of the Discoglossidae. The sacral diapophyses
are but slightly dilated. The only species, N. rugiceps, lives in
Ecuador, and grows to nearly three inches in length. The head
is large and rough owing to the skin being involved in the
cranial ossification. It is further peculiar in its coloration, the
under parts being chestnut -brown instead of whitish. The
upper parts are olive-grey or brown.

The following four genera, each represented by one or two species

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VI HYLIDAE 207

only, much resemble each other in the curious shape of the head,
which forms a flat projecting snout, used probably for digging in
rotten wood in search of insects. There is a peculiar degradation
in the extent of dentition of the palatal region. Diagleria and
TripHon are the only Anura which possess a longitudinal row of
parasphenoid teeth. Diaglena petasata of Mexico and D. jordani
of Ecuador have, moreover, a transverse row of teeth on the
palatine bones in addition to those on the vomer.

Triprion petasattcs of Yucatan has parasphenoid and vomerine
teeth. The head is a bony casque, with strong superciliary
ridges, the skin being extensively ossified. The mouth forms
a flat snout, owing to the long projection of the upper over
the lower jaw. The skin of the back is smooth brown with
darker spots ; the under parts are uniform whitish. The male
has a subgular vocal sac. Like Diaglena and Corythomantis they
possess adhesive discs on the fingers and toes, and climb trees.
The total length of this curious creature is 2 inches.

Corythomantis greeningi of Brazil has a similar head. The
vomers alone carry teeth, besides of course the maxillae. The
pupil is rhomboid. The tongue, as in the
two previous genera, is roundish, scarcely
free. General colour above olive, with
darker freckles ; the sides are studded with
whitish tubercles; the under parts are
whitish. The male is devoid of vocal sacs. ^ ,, « , r ^ ...

, Fig. 41.— Head of Corytko-

Total length 3 inches. mantis greeningL x 1.

Pternohyla fodiens of Mexico ap- ^bmT''^^''^^'^ ^''"'
proaches the previous three genera by the

curious shape of the head and prominent upper jaw, although
these features are not so exaggerated. The dentition agrees
with that of Corytliomantis and other normal tree-frogs. The
lingers and toes are not provided with discs, in conformity with
the burrowing, not climbing, habits of this creature. The
next following three genera connect the Hylidae with the
Cystignathidae. The sacral vertebrae are but slightly dilated.

Acris. — The adhesive discs are very small, the tympanum is
indistinct. A. gryllus, the only species, inhabits the greater part
of Eastern and Central North America, extending northwards
into Canada. It attains a length of 1^ inch. The colora-
tion is very changeable, in adaptation to the surroundings. As

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208 ANURA

a rule it is brown, with a more or less reddish or grey ground-
tone, ornamented with dark brown or blackish irregular, longi-
tudinal patches,one of which is bordered with light green,and there
is often a light vertebral streak. The legs are cross-barred, the under
parts are whitish brown and yellowish. The male has a subgular
vocal sac, and its most remarkable feature is the voice, which
closely resembles the noise of a cricket or of certain grasshoppers.
Holbrook describes it as a merry little frog, constantly chirping
like a cricket, even in confinement. It frequents the borders of
pools, and is often found on the leaves of aquatic plants, rarely
on the branches of such low shrubs as overhang or dip into the
water. When disturbed it takes long jumps, and hides at the
bottom of the pond. Insects are secured by leaps. It can
easily be domesticated, and takes food readily from the hand.
Sprinkling them with water never fails to make them more
lively and noisy. Appearing in April in great numbers, they
are said to vanish early in the autumn for hibernation. The
tadpoles are metamorphosed by the end of August.

Chorophilus. — The fingers and toes are provided with very
small adhesive discs. The sacral diapophyses are very slightly
dilated. About seven species occur in North America, chiefly in
the Southern States, one, Ch, cuzcamis, in Peru. Ch. ocularis is
the smallest of the frog-kind known, and lives in South Carolina,
frequenting damp places, the vicinity of stagnant pools, water-
plants or low shrubs, for instance the " myrtle," Myrica cerifera,
I once had two of these tiny creatures less than three-quarters of
an inch in length. They were very active, and took surprisingly
long leaps, jumping distances of 2 feet, but could not be kept
through the winter, although they took minute insects readily
enough. The head is narrow, long and pointed ; the upper parts
are of a rich chestnut-brown with a bronzy gloss. The upper
jaw is white ; a black band extends along the sides of the head
and body. The under parts are yellowish white.

Ch, ornatus is another inhabitant of the South-Eastern
States ; its name refers to the dark brown patches on the back
and sides, bordered with golden yellow, upon a reddish-brown
ground-tone, while the under parts are silvery white with fine
grey spots. This frog, a little more than one inch in length,
lives on land in dry places, preferably in corn-fields, has no voice,
and, except during the pairing season, carefully avoids the water.

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C YSTIGN ATHIDAE 2 09

Th&ropa. — The fingers and toes are free, the tips simply
swollen and not dilated into discs. Closely allied to Chorophilus,
Th, miliaris, of Brazil, the only species, has very long toes. The
head is broad and flat. The upper, nearly smooth surface of the
body is flesh-coloured, with brown marblings'; the limbs are
cross-barred ; the under parts whitish, granular on the belly. The
male is devoid of vocal sacs. The total length may be 2 inches.
Hensel has published the following notes of this species, under the
name of Hi/lodes dbbreviattis. The tadpoles are quite flat, their
bellies forming a kind of sucking disc, so that these creatures,
even before the appearance of the hind-limbs, can quickly wriggle
up vertical walls of stones, provided these are covered with a
little water. In correlation with this habit, the root of the tail
is not compressed laterally, but is as broad as it is high, and the
usual vertical fin is restricted to its distal third. On the prox-
imal portion of the tail the ventral fin is flattened and broadened
out so as to form almost the continuation of the peculiar disc-
like belly. The anal opening is not a projecting tube, but is a
flattened transverse slit.

FanL 6. Cystignathidae. — This is one of the largest families,
and also one of the least satisfactory. Its numerous members,
more than 150, exhibit such a versatility in adaptation to
circumstances (there are aquatic, terrestrial, arboreal, and burrow-
ing species), with a corresponding development or loss of
anatomical characters which we should like to rely upon as
taxonomic marks, that the numerous genera not only run into etich
other, but also get entangled with those of other families. In
fact the whole family is ill defined. It can be characterised as
follows : — The shoulder-girdle is arciferous ; the sacral diapophyses
are cylindrical or but slightly dilated; the metasternimi has
either a bony style or it forms a cartilaginous plate; the
terminal phalanges, although they sometimes carry adhesive
discs, are never claw-shaped.

The last statement is, of course, intended to separate the
Cystignathidae from the Hylidae, of which, however, the three
genera Thoropa, Chorophilus, and Acris stand on debatable
ground (cf. p. 186, Hylidae), while, on the other hand, most of
the Australian genera, notably Chiroleptes, have unmistakably
dilated sacral diapophyses. The difference from the Pelobatidae
can in this case be one of degree only.

VOL. vni p

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210 ANURA CHAP.

The Cystignathidae may be said to i^present the Eanidae in
Notogaea. Some of them can be distinguished from the true,
typical frogs solely by the arciferous type of the shoulder-
girdle and sternum. There is in both families the same adaptive
versatility, the same amplitude in the formation of the finger-
tips, the occasional slight dilatation of the sacral diapophyees,
the same range in the configuration of the omo- and meta-stemum.
In fact, young Eanidae, before the firmisternal character is assumed,
are indistinguishable from Cystignathidae, and the latter would
turn into Eanidae if they could be induced to consolidate their
sternal apparatus.

The geographical distribution of the Cystignathidae is
suggestive of their being an old family, most of whose meml^ers
have reached a high stage of morphological development. The
overwhelming majority inhabit the Neotropical region, a few
forms extending into tropical Central America and into the
Antilles ; the rest, some twenty species only, are confined to the
Continent of Australia and to Tasmania.

The family name is rather a misnomer. It is taken from the
genus Cy8tignath%L8, which is, or rather was, characterised by the
peculiarly broadened lower jaw, hollowed out by the vocal sacs ;
but this generic name had to give way to that of Leptoddctyhn, in
obedience to the often senseless rule of priority. The family
is composed of three sub-families.

Sub-Fam. 1. Hemiphractinae. — Teeth are carried by both
jaws, tlie vomers and the palatine bones ; or by the palatines ami
parasphenoids in Amjphodus, The vertebrae are opisthocoelous,
devoid of ribs, and the sacral diapophyses are not dilated. The
shoulder-girdle and sternum are strictly arciferous. The omo-
sternum is very much reduced ; the metasternum forms a
cartilaginous plate. The tongue is slightly free behind. The
tympanum is distinct. Three genera, with eight species, all
inhabitants of South America.

Hemiphrdctus. — The head is large ; the upper surface of all
the cranial bones appears pitted, owing to most of the covering
skin being involved in the ossification. The temporal fossa is
bridged over or roofed in by the fronto - parietal and the
squamosals, so that the orbit is completely encircled by bone,
as in Pelohates cv.ltripes. The terminal phalanges are simple
and are not dilated into discs. The teeth of the lower jaw are

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VI CYSTIGNATHIDAE HEMIPHRACTINAE 2 I I

very small and numerous. The tongue is round and very small.
U. scutatus, the only species, living in Ecuador and Colombia, is
a frog-like creature, with a large helmet-shaped head. Total
length 2^ inches.

Ceratohyla has the same kind of helmet-shaped head, and the
orbit is likewise enclosed by bone, but the terminal phalanges
are claw-shaped and carry regular adhesive discs. This genus,
the five species of which live in Ecuador, bears undoubted
resemblances to the Hylidae. In C. proboscidea the upper eyelid
is produced into a little upright fold, as in Amphignathodon
and some species of Nototrema and Ceratophrys among Cysti-
gnathidae. The snout is produced into a long, compressed, bifid
appendage, and the heel carries a triangular flap. In C. luhcdua
the partly ossified helmet sends out a pair of diverging processes,
formed by the squamosals, extending backwards and sideways
from the concave and ridged interorbital spaces. The tip of the
snout and the tips of the divergent horns form an equilateral
triangle, and the whole head bears a striking resemblance to
some of the fossil Eeptiles from the Elgin Sandstone, e.g.
Triceratops. Total length 3 inches.

Amphodus wucheri. — The only species of this genus has been
found near Bahia. It has teeth on the palatine bones and five
series of small teeth on the parasphenoid, but none on the vomers.
The teeth of the mandible number about eleven on each side
and decrease in size towards the symphysis. The tympanum is
distinct; the heart-shaped tongue is free behind. The cranial
bones are only slightly pitted. The skin is smooth above,
chocolate-brown, spotted with yellow, and with a yellow band
on the sides of the body beginning with the upper eyelid and
ending in a broad patch above the vent. The under parts are
yellowish white.

Sub-Fam. 2. Cystignathinae. — The upper jaw alone is provided
with teeth. Vertebrae procoelous. The twenty-seven genera of
this sub-family have been arranged in the following key, merely
for convenient determination.

I. American genera.

A. The metastemum forms a cartilaginous plate without a narrow

handle. The pupil contracts into a horizontal slit

a. The terminal phalanges are bifurcated, Y -shaped, and provided

with large discs ; the tympanum is distinct ; the omostemum

is absent . . . Centrolene geckoideum, Ecuador.

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212 ANURA CHAP.

h. The terminal phalanges are T-shaped and carry disc& The
omostemum is cartilaginous.
a. Discs divided by a dorsal groove.

With vomerine teeth . Elosia, 3 species in Brazil

Without „ SyrrhopuSy^ 9 speciei^, South America.

p. Discs undivided.

With vomerine teeth . Hy lodes, p. 214.

Without „ .... Hylopsis,

c. Terminal phalanges simple, pointed, or with very small diaoi.

First finger opposed to the others . . Pseudis, p. 213.

d. Terminal phalanges simple, without discs.

a. Tympanum hidden. A large, flat gland on each side of
the body . Gyclorhamphm fuligtnosus, Brazil

p. Tympanum distinct. Head rough, entirely bony.

Calyptocephahts, p. 215.
y. Tympanum hidden or absent Tongue roundish, not
nicked, free behind. Toes webbed.

Telmatobius, 6 species in Western South America.
& Tongue heart-shai)ed, free. Toes webbed,

GeratophrySj p. 215.
€. Tongue round, free behind. Toes webbed. With two
tooth-like projections in the lower jaw.

LepidohatrackuSy p. 218.
f Tongue entire, or slightly nicked, free behind. Toes free.
BorhorocoeteSy 1 1 species in Western South America.
?/. Tongue entirely adherent Tympanum distinct.

Zaehaenus parvulusy Brazil
B. Metastemum with a bony style.

a. Pupil horizontal

a. Terminal phalanges T-shaped, with discs.

Tympanum distinct

Plectromantis, 2 si)ecies in Western South America.
/3. Terminal phalanges simple ; tips not dilated into regular

discs.

1. Tympanum distinct

Sacral diapophyses slightly dilated.

Edalorhina, 3 species in Ecuador and Peru.
Sacral diapophyses not dilated.

LeptodactyluSy p. 218.

2. Tympanum indistinct or hidden, Palvdicola, p. 220.

b. Pupil vertical Terminal phalanges simple and not dilated.

Chili.

a. Tongue slightly nicked . Livmomedusa macroglossa^

p. Tongue entire, but free behind. Digits very long.

Hylorhina silvatica.
II. Australian genera. The terminal phalangss are simple and not dilated.
The omostemum and metastemum are cartilaginous, the latter forming
a plate, semi-ossified only in Heleioporus.

= PhyUobates (part) Bibron ; of. Boulenger, P,Z,S. 1888, p. 207.

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VI CYSTIGNATHIDAE CYSTIGNATHINAE 2 I 3

A, Pupil contracted into a horizontal slit

a. Omostemum rudimentar}'. Vomerine teeth present.

a. Tympanum distinct . . Phanerotis fletchei-i,

p. Tympanum hidden .... Cryptotis hrevis.

h. Omostemum present. Vomerine teeth vestigial

Grinia^ 4 specie«.

c. First finger opposed to the others . Chtroleptes, p. 221.

B, Pupil contracted into a vertical slit

a. Omostemum rudimentary. Vomerine teeth absent.

Hyperolia martnorata.
h, Omostomum fully developed. Vomerine teeth present
a. Tympanum distinct. Toes webbed.

Mixophyes fasciolatm.
p. Tympanum hidden. Toes webbed. Heleioporus, p. 222.
y. Tympanum indistinct. Toes free or slightly webbed.

Limnodynastes, p. 222.

Pseudis, widely distributed over South Americia, consists of
four species which have the appearance of long-legged frogs.
The fingers, of which the first is opposed to the others, are free ;
the long toes are fully webbed. The tympanum is exposed.

F, paradoxa is absolutely aquatic, floating in pools, and is
extremely shy. In life it is most beautifully coloured with
bronze, bright green, and black markings above ; underneath it is
shiny yellow, with brown spots on the body and stripes on the
thighs. Within a few minutes after death all the brilliant
colours of the smooth skin of the back turn into dull uniform
brown, with indistinct dafker spots. Total length of the adult
from 2 to 2^ inches. The specific name refers to the peculiar
shape and monstrous size of the larva or tadpole.

One of the larvae described and figured by Parker measures
10^ inches in length, the head and body taking up 3^ inches.
The spiracle lies on the left side and the hind legs are ^ inch
long, just breaking through the skin. The vent is median.
The huge tail is very thick and muscular, and is furnished with a
high, irregularly shaped dorsal and ventral fin, the whole organ
measuring 4 inches dorso-ventrally. Another larva, or rather
tadpole, in the national collection is older, and although still very
large, namely, 7 inches long, has fully developed hind-limbs 3
inches long ; the fore-limbs are less than half that size, the left
protrudes through the spiracle, while the right has broken through
the skin. The dorsal and ventral fins of the tail have much
shrunk ; the whole organ, 5 inches long, is gradually tapering to
a point like the tail of ordinary tadpoles. By the time that the

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214

ANURA

CHAP.

tadpole is nearly ready to leave the water, its whole bulk is re-
duced to less than one-fifth that of the largest tadpole. It measures
from snout to vent only 1^ inch (in the 7-inch tadpole this
distance is fully 2 inches), and the tail, devoid of fins, is
reduced to 2 inches in length. Instead of the solitary left
spiracle there are now two, one on the ventral side and a little
in front of the base of each arm, the border of each hole being
continued by a peculiar semilunar fold.

Hy lodes. — The numerous species, nearly fifty, of this tropical
American genus exhibit several anatomical differences. The

tympanum is sometimes indistinct
or hidden, in which -case the
Eustachian tubes are generally
very narrow. The fingers are free,
and carry discs, like the toes,
which are sometimes slightly
webbed. The males have a sub-
gular vocal sac, producing a loud,
or whistling, voice. The general
appearance is that of land- and
tree-frogs ; the size is small, mostly
between 1 and 2 inches.

H. martinicensis is about 1^
inch in length. The ground-

FiQ. i2.^Hi,iodes viuHinicefisis. 1, ^olour is pale yellow-grey, with a

an egg with embryo about seven large brown patch on the nape,

ofd-^ftiieyon^Frog^^^ which colour is continued over the

all xf ; 4, adult maiexi. (After back in the shape of more or less

^^" coherent or dissolved patches. A

dark brown stripe runs along the middle of the sides. The

limbs are barred with brown, the under parts are whitish. This

species, known by the vernacular name of " coqui" inhabits many of

the West Indian islands, e.g. Barbadoes, Martinique, Porto Rico, and

Hayti. It has become famous, as it was the first instance known

of a frog which undergoes its whole metamorphosis within the

egg. The pairing takes place on land, in the months of May

and June, when the female lays about twenty eggs, which are

enveloped in a foamy mass and glued on to a broad leaf, or

hidden in the axillae of Iridaceous plants. The mother seems to

remain in the neighbourhood watching the eggs, which are

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CYSTIGN ATH I N AE 2 I 5

large, measuring 4-5 mm. in diameter. Dr. Gundlach, a resident
in Porto Eico, was one day, in the month of May, attracted by
sounds like those of a young bird, and found three males and
one female of this species sitting between two large leaves of an
orange-tree. He put them all into a glass vessel and soon saw
a pair in embrace. The female laid about twenty-five pale
straw-coloured eggs. The embryo develops neither gills nor
gill-openings, but a large well-vascularised tail, by means of
which, being immersed in the watery fluid contained within the
egg, it seems to breathe. After twenty-one days the tadpole,
having used up all the available yolk and fluid, and most of its
own tail, bursts the egg-shell and hops away as a little frog of
5 mm. in length, but still with a stumpy white tail, which is
quite absorbed within the same day.

This species has several times made its appearance in the
tropical houses of Kew Gardens. It seems to have bred and
vanished again.^

Citlyj^iocephalas is remarkable for the dermal ossification of
the cranium, which has assumed the greatest possible extent.
It affords a curious parallelism to Triprioii and other Hylidae,
which are likewise . Central American forms. Only two species
are known ; C. gayi of Chili, and C. testudinicejys of Panama.
They are large, thoroughly aquatic creatures, 5 to 6 inches in
length, with huge heads. The tadpoles grow to an enormous
size. One specimen of C. gayi in the National Collection is
more than 6 inches in length, the tail taking up more than half
of the total : the spiracle lies on the left side, the vent on the
right, and the hind-limbs are still half . en veloped in a kind of
fold of the skin.

Cerato2)hry8 is a genus of some ten toad-like species, living in
South America, from Guiana to Argentina. The generic name
alludes to the peculiar modification of the eyelid, which in most
species is developed into a triangular, upright, but flexible
appendage. The head, in conformity with the huge mouth, is very
large. The tympanum is rather indistinct, sometimes quite
hidden. Several of the species have a large dorsal shield, which
is produced by a thick ossification of the cutis, but is not fused
with any of the vertebral processes. The male has a vocal sac.
C. dftrsata s. haiei of equatorial Brazil is a monster toad, reaching
> See Gunther, Nature, lii. ISP.*), p. 643.

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2l6 ANURA CHAP,

a leiijiTtli tA' (j niches. The upptr eyelid ia tranaformed into a
tiian<;uhir liui'D, wlieuce a uutaneoiis ridge extendi all alonj^ tiie
side of the hack, meeting that of tlie otlier wide ahove the vent.
There is iiu ost^eoiis sliield on tlie back. The tym|mnuia is
hidden, (Ironnd-eulourH, omnge or green, witli ^haq^ly niarke<l
dark brown or blackish patehes,

C\ cornntff, In Northt^n Brazil, lacks the dorsal sliield, but hfl«
horned eyelids ami a vi.sihle tyiupanuni. Its ctjloratioo rendei^
it one of the must heautifid toad-like creatures known. The
ground-eolonra are green, lilack and lirown, with an oraoge-yellow
strijie over the head and hack. All tlieso eoloure are most pleasingly
Identled and arranged in marbled patches or s^tripes radiiitinjz

fnim various centres, as, for itisi;ince, from the eyes towards the
circumference of the month, the slit of which they pass, the
same line of the pattL'rn ]»eing continued npon the lower jaw.
The whole Kurfacc makes tlie impreHsion uf a gay but ex-
4uisit<^ly hataii>nious carpet. The under parts are yellow, in-
cliiiiii^ tn white townnls tlie iiiiddle.

C. onutfa has a diirsal shielil Tlte tympaniuji is ju.st visible,
and lli<- eyelids form i>nly h>w but ,sh si rp -edited ]krojeetions. This
is lik< wisi^ a Iti'Euiliful toad, living tbielly in Urnguay, Northem
Argentiiiii, ;Hid r:iragiiJiy, where it is nrnversiilly known as the
" rsrui rzo." one of the Spanish words signifying a toad. Its siie
rarely surpasses 4^ indies. The ground -cok»urs are greenish
and vcIIkw, with laig*' diirk grren patches on the biick, decreasing

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CYSTIGNATHINAE

217

in size on the flanks. Each of these insular patches is sur-
rounded by a narrow line of white and yellow dots, interspersed
here and there with lines of rusty brown or red. The object of
this elahomte carfjet-like pattern is conceal tiieiit. These toads,^ —
and this applies to all the speciei^, — hnry theniaclveH half in
the ground, preferably in the grass, where they are well-ni^li
invisibla If there is not enough green vegetation, tiiey throw,
with their feet, little lumps of earth upon their backn, the skin

FfO. ii. — VertUuphrj^g t'rnnta. Fnmi .^'^lff'rt\]

f*f which l»ecomes at the same time iimrt; c liiikini riiid nssiniiLS
duller tones. There the creature liert, ]ieri'n'tly roHrrMlrt!,
l»etrayed only by the metallic glittering eyes, waiiiii^r fnr ^nmv
unfortunate creature to pass into the tnqi repn-si'iiti ^l by
the enormous mouth, whicli opens and Kliuts witli li^Hitning-
rapi*lity amt with an audible snap. Tlu^ seem tu live
chieflj- on frogs, and sometimes they turn euiiiiibals. T\vt>
K[jecdmen8 were l^roiight over to me frnia raa-rms Aires hy a
friend, in a well-ch>8ed basket with moi.st soil at the bottom, but
only one was visible on arrival The other wns in si tit- tlie

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2 1 8 ANURA CHAP.

larger one, and could still be felt through the soft body. This
same cannibal took large-sized frogs greedily, one or two for a
meal, swallowing them whole and then sinking back into its
lair, which it scarcely ever left, except for an occasional soaking
bath in its water-pan, especially before shedding its skin. It
lived for many months in the same enclosure with a pantherine
toad, Bufo mauritanicay of equal bulk, until one morning I found
the Moroccan half swallowed and almost lifeless in the mouth of
the American, whence it was rescued with difficulty. It came
round after a few hours, but never fully recovered, lingering on for
weeks ; the skin was changed to a lead-colour so far as it had
been swallowed and partly dissolved by the gastric juices, and
soon began to develop festering ulcers.

These " homed toads " make a squeaking noise when teazed,
not at all loud or strong in proportion to their size. Ill-tempered
individuals jump at their aggressor and can inflict rather painful
nips. They hibernate during the dry season in the ground.

Lepidohatrachtis, — Large teeth in the upper jkw, and two
large tooth-like projections in the lower jaw near the symphysis.
Vomer toothless. Sacral diapophyses not dilated. Tongue
round, and free behind. Tympanum distinct. Great develop-
ment of the membrane-bones on the head, and a weaker ossifica-
tion in the skin of the back, recalling that in Ceratophrys.
The eyes are closely set together, and the nostrils take up the
most elevated portion of the head.. Pupil horizontal. The
two species of this genus were discovered by Budgett^ in the
Paraguayan Chaco. Z. as2)er lives continually in muddy pools,
floating with just the eyes and nostrils above the surface. If
disturbed it slowly sinks to the bottom, leaving no ripple. It
feeds largely on Bufo granulosus. Total length from about 3
inclies. The skin of the upper parts is tubercular, tough, and of
a dull leaden colour ; the tips of the toes are horny. L, laevis
is smooth and slimy, " with the organs of the lateral line showing
clearly upon it," a feature elsewhere known to exist in Xejwpus
and Ze2?tohra4:hium only.

Leptodactylus = Cystignathus. — Some twenty species inhabit

tropical America, from Central Mexico to Buenos Aires. The

fingers and toes are not webbed and end mostly in points ; only

a few species, e.g. i. hylaeodactylus, having small adhesive discs.

* Quart. Micr. Sd. xlii. 1899, p. 329.

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CYSTIGNATHINAE 2 I 9

The legs are long and the general appearance is very much like
that of an ordinary frog.

One of the commonest and prettiest Brazilian species is L.
ocellatuSy which is characterised by a number of longitudinal
glandular folds on the back and flanks. The colour of the
upper parts is olive -brown, that of the prominent folds is
yellowish white, interspersed with black spots. The under parts
are yellowish white, with blackish marblings on the throat.
The males have a sharp black spur on the inner carpal edge
and one on the rudiment of the thumb. Total length about
4 inches.

According to HenseP the spawning takes places in Eio
Grande do Sul after hibernation. The voice of the male is then
very loud, resembling the sound made by a carpenter chopping a
beam. They repair to ponds and produce a cup-shaped puddle,
about 1 foot in width, by raising a wall of mud, which
separates the inner water from that of the pond. The tadpoles
remain in this nursery until the spring-rains demolish it and set
the young ones free. Drought causes the drying up of these
water-pans and subsequent destruction of the brood.

L, mystacinus is another Brazilian species, about 2 inches in
length. Its specific name refers to the dark brown stripe which
runs from the tip of the mouth through -the eye to the tympanum.
This species is thoroughly terrestrial, and never enters the water.
It digs a cavity, the size of an ordinary tea-cup, under stones or
rotten trunks, always in the neighbourhood of ponds and just
so high above the water. that the latter can rise up to the
nest in the rainy reason. The straw-coloured eggs are laid in
this cavity, and are enveloped in a foamy, sticky mass, like the
well-beaten white of an egg. The young tadpoles seem to live
on this froth until the rains set them free. When, however, the
rains delay and a drought kills the broods of other less circxm)-
spect species, these tadpoles, still provided with gills and long
tails, remain in their moist nest or withdraw further beneath the
rotten stumps, huddled together in large numbers until the next
rainy season.

Similar nursing habits have been recorded of L. albilahris,
which inhabits Mexico, Cuba, and several other West Indian
islands. The same applies to Z. typhonius. (rundlach found eggs

» Arch. NcUurg, .xxxiii. 1867, p. 124.

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220 ANURA

of this " Sapo " in Puerto Eico on the 4th of November ; on the
25 th the young showed the first signs of hind-limbs, on the 3rd
of December of fore-limbs, and on the 7 th of the same month
they began to climb out of the water.

Paludicola is a semi-aquatic genus with some eighteen species,
ranging from Mexico to Patagonia and across the Andes into
Chili. Some of them have a peculiar gland on the Imnbar region,
or large, flat warts on the back, sometimes arranged in longi-
tudinal folds. The toes are slightly webbed, or free, according to
the more or less pronounced aquatic habits.

P. fuscomacidatay an inhabitant of Southern Brazil, Paraguay,
and Uruguay, is a short-limbed frog, with spreading slender toes
and a small head. There are shovel -
shaped, black, horny tubercles on the
metatarsus. The general colour is olive
above, with darker markings and con-
fluent white -edged spots; the limbs are
cross -barred; the lumbar glands are
black, with a white margin in front. The
male has a vocal sac. Budgett ^ gives
the following account of its habits : —

The peculiar cry, which is so con-
stantly heard in the neighbourhood of
\^-^SC" shallow pools in the Paraguayan Chacb,

Fig. ^i^- PcUudicoia j\u.co. ^^^ resembles that of a kitten, is pro-
macuiata, x 1, with vocal duccd by the alternate inflation of throat
sacs partly filled. ^^^ abdomen. WTicn fully inflated, the

frog appears to be the size of a golf -ball, but, if startled,
instantaneously shrinks to one-fifth of that size, so that it seems
to have vanished. It has also the power of ventriloquising.
The food consists largely of water-beetles. In the spawning time
it was found at night floating on the surface of pools in the
distended condition, and crying to the females in a most
mournful way. On coming to the surface it fills its lungs with
a few gasps, greatly distending the walls of the abdomen, and
then drives the air into the vocal sacs, causing them to become
distended as the body collapses, and giving rise to a kitten -
like cry.

Tlie eggs are chiefly laid in Januarj^ and are found embedded

^ Quart. J. Mier. Sci. xlii. 1899, p. 309.

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CYSTIGNATHIN AE 2 2 I

in a frothy mass floating upon the surface of the water. The
eggs measure only 1 mm. and are without pigment, and with ex-
tremely little yolk. The larvae become free-swimming within from
eighteen to twenty-four hours after the first segmentation. When
ready for hatching they wriggle their way through the froth to
the water below, and hang into it from the floating froth.

P. hiligonigera s. notata, in Brazil, lacks the lumbar gland, the
place of which is marked by a black spot. The upper parts are
olive, with darker marblings and a dark lateral stripe. The male
has a black throat and two external vocal sacs. Hensel found
the eggs, in Eio Grande do Sul, in September, forming a frothy
mass of the size of a fist, floating between grass upon the water
near the margin.

The following three genera may serve as Australian examples,
especially since we are indebted to Baldwin Spencer for interesting
observations made on their habits in Central Australia.^

Chiroleptes, of which six species are known, is easily recog-
nised by the first finger, which is opposed to the others. The
sacral diapophyses are slightly dilated. The general shape is that
of a thick-headed, rather stout land-frog or of a tree-frog. The
tympanum is distinct, and the toes are only half webbed, or even
less, except in Ch. platycephaluSy in which the toes are entirely
webbed and the tympanum is indistinct. This species is about
2 inches long, uniformly olive-green above, with a few tubercles
on the otherwise smooth skin. Other species rather resemble the
European Natterjack in coloration.

Spencer's account is as follows : — " In Central Australia Ch.
platycephalus seems to prefer the hard clay pans rather than
sandy creeks, as the sand-beds of the latter are too loose for the
formation of the burrow. We came across the animal first when
encamped by the side of a very shallow clay pan, the floor of
which was deeply cracked with the sun's heat. Around the edge
were withered shrubs of Clieno2Jodium nitrariaceumj and it was at
the base of these that the black fellows looked for the burrow.
In the hard-baked clay were imprints made by the frog as it
burrowed, and about a foot underground we came across the
animal, puffed out into a spherical shape, and just filling up a
cavity, the walls of which were moist but not wet. The ground

' Report on the Work of the Horn Scientific Expedition to Central Australia y
pt. ii. "Zoology," 1896, p. 164.

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22 2 ANURA

was 80 hard that it had to be chipped away. When one side of
the burrow was opened, the frog remained perfectly still ; its
lower eyelid was drawn up over the eye and was very opaque,
giving rise to the belief amongst the blacks that the animal is
blind. In the sunlight, after a short time, it opened its eyes.

" On squeezing the body, water was forced out of the cloaca ;
this was accumulated principally in the urinary bladder. * On
cutting the body open it was seen that there was a certain
amount of water in the subcutaneous spaces, but that the greater
portion, which caused the great swelling-out of the body, was
contained in the body-cavity itself; and it was also observed that
the lungs were considerably distended and lengthened, their
apices lying right in the pelvic region. They contained air and
not water, but their outer faces were bathed with the water in
the body -cavity." The larvae and tadpoles probably develop
with extreme rapidity, soon to aestivate as very small frogs.

HeleiopoTus has a calcified metasternal plate and slightly
dilated sacral vertebrae. The two species have a toewi-like ap-
pearance, owing to their stout bodies, short limbs and conspicuous
parotoid glands. H. (dbopunctatus is mottled whitish red
and brown above; it extends from Western into Central
Australia, ff. pictus is olive, with darker marblings, and is distin-
guished by a light vertebral line. It is likewise found in Central
Australia, and it extends into Victoria and New South Walea
Spencer found, it in swarms after heavy rains, the specimens
being much swollen and distended with caterpillars and beetles.
They looked as if they were simply gorging themselves with
food preparatory to returning again to their long aestivating
condition.

Limnodyruistes is one of the commonest genera in Australia.
The six species have the habits and appearance of stout frogs or
smooth toads. L, dorsalis seems to range through the whole of
Australia, from east to west, and looks like the European
Felobates. The skin is smooth, but with an elongated white
gland extending from beneath the eye to the shoulder, and
another glandular complex on the thigh. The upper parts are
mottled olive-brown, often with a light vertebral line. The
under parts are whitish, with brown spots. The male has a
vocal sac. One of the specimens in the National Collection con-
tained a half-grown Heleioporus alhopunctatus in its stomach.

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CYSTIGN ATHINAE 223

Concerning the pairing and the other habits of the Anura of
New South Wales we have some valuable notes by J. J. Fletcher.^
He observes that Australian frogs spawn whenever they are ready,
and when the very irregular conditions of moisture will allow
it, but that they are not all ready at the same time, i.e. they
have no fixed period of the year. Limnodynnstes, Hyla mirea,
and ff. coerulea deposit their spawn in the water, in more or less
irregular floating patches, which look white and frothy. The
period extends from July to May, and is at its height in August
and September ; but if there is a spring-drought vigorous spawn-
ing may be looked for about the middle of January, when heavy
showers are likely to occur. Crinia and several Hyla, e.g.
H. ewingi, spawn at any time of the year. The eggs form small
submerged bunches, enclosed in a transparent jelly, attached to
the blades of grass or twigs of dead branches in the water.

Fseudophryn^, a genus closely resembling Crinia, but on
account of the absence of teeth in the lower jaw relegated to
the Bufonidae, spawns during the Australian summer and autumn.
The numerous ova of F. australis and P. bibroni are laid separately,
not in the water, but under stones, or in the debris of reed- and
grass-tussocks, on the edge of a pool.

The larvae of Fsevdophryne and others have often to depend
upon the next following rain, sometimes waiting for months to
be released from the eggs, wherein they have so far developed.
But the tadpoles, once hatched, probably do not bury themselves ;
they either metamorphose or die.

The males of Mixophyes and Hyla grasp the females in the
axillary region ; those of Limnodynastes, Hyperoliay Crinia, and
Fseudophryne throw their arms round the inguinal or lumbar
region.

For some three months during the winter, commencing
about May, the frogs, like lizards and snakes, resort to shelter
under logs and stones, beneath which they are then to be met
with in a more or less sleepy condition. During the hot and
very dry periods many bury themselves in the drying-up mud,
which becomes very hard, and does not release them until the
next rains. They croak during showery times of the year.
There is no evidence that any Australian species live in the high
EvxalyptuS'tre^^

^ Proe. Linn, Soc. N,S. W. (2), iv. 1898, p. 357.

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2 24 ANURA CHAP.

Hylopsis platycephalus, of South America, is of importance as
forming a link with the Dendrophryniscinae, owing to the very
small size of the teeth in the upper jaw. There are no vomerine
teeth. The fingers and toes are webbed, and furnished with
discs. The very small omosternum and the metasternum are
cartilaginous. The pupil is horizontal. Total length, about
or under 1^ inch.

Sub-Fam. 3. Dendrophr]mi8cinae. — The two Neotropical
genera of this sub-family are characterised by the entire absence
of teeth. The toothless condition of the upper jaw is really the
sole character which separates them from the Cystignathinae,
taken as a whole. The suppression of the tympanum and of the
Eustachian tubes in Batracho'phrynus, and the fully webbed toes
of B. inacrostomus indicate complete adaptation to aquatic life.
The absence of the omosternum in Dendrophryniscus, the absence
of vomerine teeth, the dilated phalangeal tips, the entire and qiiite
adherent tongue, are all features which likewise occur in some of
the Cystignathinae, and therefore cannot be urged against their
affinity. Lastly, the recently discovered South American genus
Hylopsis is, as pointed out by Werner,^ an intermediate link,
owing to the extremely small, scarcely visible teeth in the upper
jaw.

Dendroph7yni8cu8 brevipollicatus has been found in the neigh-
bourhood of Eio Janeiro. The head is depressed and triangular.
The tongue is entire, but free behind. The tympanum is sup-
pressed. The omosternum is absent ; the metasternum forms a
long bony style. The sacral diapophyses are cylindrical. The
terminal phalanges are simple, but carry dilated tips. The first
finger is rudimentar}'. The skin is nearly smooth, reddish brown
above, whitish below ; the limbs are cross-barred.

Batrachophrynus inhabits the mountains of Peru. The head
is much depressed and small, with the eyes directed upwards,
as is usual in essentially aquatic species. The tongue is large,
circular, and entirely adherent. The tympanum and the Eus-
tachian tubes are suppressed. The omosternum is cartilaginous,
and the metasternum forms a cartilaginous plate. The sacral
diapophyses are cylindrical. The terminal phalanges are simple,
and carry no discs. The four fingers are short; the toes are
webbed. The male has no vocal sac. B, brachydactylus has a
1 ZooL Anz. xvii. 1894, p. 156.

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VI ENGYSTOMATIDAE — ENGYSTOMATIN AE 225.

smooth skin, olive-brown above with darker spots. B, viacro-
stomus, 2 inches in length, is distinguished by its larger size, and
by its completely webbed toes.

Fam. 6. Engystomatidae (Narrow-mouthed Toads). — Firmi-
stemia with dilated sacral diapophyses,

Sub-Fam. 1. Engystomatinae. — Without teeth in the upper
jaw. — Although there are only about 60 species known, these have
been grouped into more than two dozen genera, many of which
are represented by one or two species only. The range of this
sub-family is peculiar, namely, Neotropical and Palaeotropical.
Scaphiophryne and Rhomiophryne are peculiar to Madagascar ;
Calophrynus occurs in the same island and in the Indian region ;
Xenobatrachus, Sphenophryne, Liophryne, Ma7Uophryne, Callulops
and Xenorhina live in New Guinea. Breviceps, Cacosternum and
Hemisus are confined to Africa, while of Fhrynomantis two species
live in Africa, and the third in the Malay island of Amboina.
Such freaks of distribution indicate either that many of these
genera are not established upon very valid characters, or that
their respective species are instances of convergent evolution, and
do not form natural genetic groups.

Many of the members of this sub-family live upon ants and
termites, and it is a well-known fact, not restricted to the Anura,
that this kind of fare has a peculiar, modifying influence upon
the structure of the mouth, teeth, tongue, limbs, and various
other organs. In the present case the tongue is not much
affected ; it is, with few exceptions, more or less oval, not nicked,
but free behind ; in the Indian Glyphoglossus and in Rhomhophryne
of Madagascar only is it modified into a rather long and grooved,
almost double, apparatus.

A very common feature is the small size of the mouth and
the formation of a snout, which projects beyond the upper rim of
the mouth and beyond the nostrils. Such a prominent and
pointed snout is well developed in Rhinoderma, Phryniscus, Calo-
pkrynus, Stereocy clops, Hypopachus and Engy stoma. The mouth
is very narrow in Cajcopus, Glyphoglossus, Rreviceps, Rhomho-
phryne, and Hemisus, all creatures which seem to be confirmed
eaters of ants and termites. However, it must not be supposed
that the mouth of all the genera is narrow, although this
character, rather marked in Engy stoma, is now embodied in the
name of the family. A peculiar development of the palatal regi6n

VOL. VIII Q

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226 ANURA CHAP.

is possibly correlated with this food. The palate is mostly
toothless, but its skin is frequently raised into a transverse fold,
between or behind the vomers, and into a second fold in front
of the oesophagus; these folds are sometimes rather hard and
serrated or denticulated. The palatine bones carry true teeth
in Rhomhophryne, and sometimes in CalltUa; in XeriohcUrachus
the teeth are reduced to two large pairs. The tympanum is
usually hidden.

The shape of the body is generally very stout. The limbs are
short, notably so in GlyphoglossiLS, Breviceps, JRhombophi'yne,
ffemisios, Stereocyclops and Cacopus. Others, for instance most
species of Microhyla, Fhryniscm, Ccdhda, and Sphenophryne, are
of a very slender build ; and their limbs, instead of being short
and well adapted to digging, are long and may even be provided
with typical adhesive discs, supported by T-shaped phalanges,
especially in the two genera last named, and in Scaphiophryrie and
Phrynomantis. However, none of the forms provided with discs
are known to be arboreal.

Exceptional diversity is shown in the shoulder -girdle and
sternum. The omosternum occurs only in Bhinoderma and
ffemisiAs. The metasternum is a cartilaginous plate, very large
in Cacoptcs, distinctly small in Breviceps, and almost absent in
Hemisus. The precoracoids and clavicles show all stages from
a well -developed condition (Breviceps, Bhombophryne, Hemisus,
Rhinodermay Phrynisctts and Brachycephaltis) to complete absence.
The circumstance that these bars are very weak in Melano-
batrachiis, Calophrynits, Scaphiophryne and Ifypopachtts, i.e.
in Palaeo- and Neo-tropical genera, indicates a widespread
tendency towards complete suppression, a feature independently
aimed at both in America (Engystoma) and in the Old
World.

Until we know something about the habits of the members of
this much diversified sub-family, it is idle to connect the various
modifications with each other, and thus, by correlation, to find out
their meaning. Those forms which possess well-developed discs
on their fingers and toes are said not to be arboreal. What is the
true meaning of !he prominent snout which is not restricted to the
digging forms ? Most of the good diggers have well-developed
precoracoid bars, and the coracoids are distinctly strengthened,
but in Glyphoglossus and in Cacopus the precoracoids are entirely

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ENGYSTOMATINAE 227

absent, and this loss is compensated for by exceptionally strong
coracoids.

On the whole, those genera are to be considered as the most
primitive which have undergone the fewest losses. Those with a
complete shoulder-girdle, with an omo- and meta-stemum and
with simple phalanges, are necessarily the older forms. One step
farther back in another direction, the possession of teeth on the
palate, and on the upper jaw, leads to those genera which have
been separated off as Dyscophinae, while teeth in the lower jaw
constitute the Genyophryninae. Lastly, the firmistemal type has
necessarily been evolved- from the arciferous condition, and there the
two Bufonid genera Myohatrachvs and Rhinophrynus, the former
Australian, the latter Mexican, with their narrow and scarcely
overlapping epicoracoid cartilages, seem to form a connecting link,
although their ant-eating habits, vdth concomitant modifications
in structure, may be nothing but cases of convergent evolution.

Key to the genera : —

I. American. A, with omoetemum .... Rhinoderma, p. 228.

B. without omofitemuni.
a. Pupil horizontal

Precoracoids present

Sacrals strongly dilated. Oreophrynella,
„ moderately „ . Phryniscus, p. 230.
„ feebly „ . Brachycephalns, p. 231.

h. Pupil vertical

OL Precoracoids feeble, Hypopachiis,

p. ,, abeent Engystoma, p. 231.

c Pupil roimd. Precoracoids

present . . Stereoq/clops, p. 231.

II. Palaeotropical. a. Pupil horizontal.

a. Precoi-acoids present

With palatal teeth. Madagascar.

Rhomhojyhryne.
Palate i\ith dermal papillae. Africa.

Breviceps, p. 232.
With palatal dermal folds. Madaga.scar.

Scaphiophryne.

With serrated palatal folds. Madagascar and

India. Calophrynm.

Palate smooth. New Guinea, Sphenophryne

and Liophryne.
fi. Precoracoids absent.

Malacca. . Phrynella, p. 233.

New Guinea . Mantophryne.

Africa .... Cacostemum.

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228 ANURA CHAP.

b. Pupil vertical

a. Precoracoids present. India. MelanohatrachHt.
^rica. Hemisu^, p. 232.
p, Precoracoids absent

Tongue ovaL India. . Cacopt^

Tongue ellipticaL India. Microhyla.

Tongue divided by a longitudinal furrow.

India. Glyphoglossui, p. 233.

Fingers and toes with discs. Africa and

Amboina. Phrynomantit,

New Guinea. CaUul(fi.

c Pupil round. Precoracoids absent. Tongue roTmi

India. Oallula, p. 234.
Tongue long, oval, with a deep groove. New Guinea.

XenorhiiM.

Note.—Xenohatrachus ophiodon. New Guinea. Palatine bones, each with two
large curved teeth. Otherwise imperfectly known.

Rhinodenna. — Omosternum and precoracoids present. Palate
without teeth. Tympanum indistinct Terminal phalanges
simple, and not dilated. Tongue heart-shaped, and free behind.
Pupil horizontal. Habitat, Chili.

Eh, darmni, the only species, was discovered by Darwin,
during the voyage of the Beagle, Its total length is only
3 cm., or little more than one inch. The shape is grotesque, as
the skin is prolonged, beyond the very small triangular mouth,
into a false nose, i.e. a nose-shaped projection, while the nostrils
remain at their original place. The skin is smooth above,
granular on the under parts, and forms a triangular flap or spur-
shaped appendage on the heel. A glandular fold extends along
the sides of the body. The general colour is brown above, black
below, with large white patches, the latter colour being sometimes
predominant on the throat and chest. The male has a pair of
internal vocal sacs, and the use of these as nurseries for the
young has made this species famous.

Espada ^ has given an elaborate account of this species, which
lives on the ground in shady woods. Its voice sounds like a little
bell, and before taking its short jumps, it erects itself vertically
upon the hind-limbs. The gular sac of the male opens by two
slits, one on each side of the tongue. Generally this sac does not
extend beyond the middle of the chest, but during the breeding
time the eggs are put into it, whereupon it becomes greatly dis-
tended, so much so indeed that it reaches back as far as the groins ;

* An. Soc. Espan. i. 1S22. See also Howes, P.Z.S. 1888, p. 231.

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ENGYSTOMATINAE 229

dorsalwards around the flanks, almost to the vertebral dia-
pophyses ; ventrally and forwards it reaches the chin. The walls
of the sac are of the same structure as the buccal lining, of which
they are in fact continuations. They adhere, at intervals, to the
cutis and to the pectoral and abdominal muscles.

The effect of the distension of the sac upon neighbouring
organs is twofold. First, the viscera are pressed back within the
abdomen ; this disturbance is temporary and does not apply to all
specimens; the feeding in no way impeded. Secondly, a per-
manent change is produced in the direction of the precoracoid
bars, in such a way that each bar is curved tailwards and rests
with its ventral half upon the coracoid; owing to this forcible
bending the clavicles do not meet each other. There is, of course,
not so much space gained by this slight rearrangement of the
shoulder -girdle as Espada implies, but we have here, perhaps,
an illustration of direct correlation between two originally
independent organs, namely, shoulder-girdle and vocal sacs. Ee-
peated distension of the throat-bag during every breeding season,
while the whole organisation of the male is in a highly excitable
condition, has pressed the clavicular bars back, or rather has
staved them in, and this at first pathological and abnormal con-
dition has at last become a fixed feature. It is to be regretted
that we know next to nothing about the habits, especially the
mode of breeding, of the other genera which likewise have reflected
or very feeble precoracoids and clavicles. Their weakness or even
complete absence must have a reason, or rather must have had
a cause.

The pairing and oviposition, and the manner in which the
eggs are conveyed into the gular sac, have not yet been observed.
Espada examined five males with young, the number of which
varied from five to fifteen. In one male with eleven embryos the
most developed tadpoles measured 13-5 mm. from the snout to
the end of the tail, and they were lying within the chest of the
father, the less advanced in the farther recesses of the bag. Three
of the tadpoles had already completely-formed fore- and hind-limbs,
while the arms were still hidden. The least developed were still
globular, a proof that the eggs are conveyed into the bag.
Another male with fifteen embryos looked as if it had gorged
itself with the almost fully -formed tadpoles, which measured
14 mm. They were quite irregularly distributed, and nowhere

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2 30 ANURA CHAP,

attached to the walls of the bag. None of them had horny jaw-
armaments, and not even the smallest specimens showed any traces
of gilk, resembling in this latter character those in the female
brood-pouch of Nototrema. The intestine of the tadpoles is short
and thick, coiled up spirally and filled with yolk, certainly not
with vegetable or other foreign matter. Consequently the entire
development from the egg to the complete stump-tailed little
creature is undergone within the pouch ; and this, after the young
have escaped, probably shrinks back to its original size and acts
as a gular vocal sac.

Phryniscus, — About ten species of this tropical American
genus are known ; they extend from Costa Bica to Buenos Aires.
They differ not inconsiderably in various details. The tongue is
elliptical, entire, and free behind. The palate is smooth. The
tympanic disc is absent. Fingers and toes more or less webbed,
sometimes with swollen tips, without, however, forming adhesive
discs. In a few species the first toe is quite indistinct. The male
has a subgular vocal sac. The mouth is small, and there is a
short snout. The general appearanc^> varies much. Ph. nigricans
of Uruguay, etc., is stout and has very short hind-limbs ; the skin
of the upper parts is black, spotted with white, and covered with
warts. Most of the other species are slender, with larger hind-
limbs and a perfectly smooth skin, the coloration of which ranges
from dull uniform brown, or black with crimson markings, to
bright green with purple spots. The under parts are, as a rule,
conspicuously coloured, a rare feature in Anura, the favourite
colours being orange, yellow, or even crimson, with or without
black patches.

Phryniscus nigricans has been observed in Paraguay by Budgett,^
who gives the following account. This is a brilliantly coloured
frog of toad-like appearance, and about 33 mm. in length. The
ground-colour is black, w^ith yellow spots or patches on the upper
parts, the under parts are black, with scarlet blotches, the palms
of the hands and soles of the feet are scarlet. At the breeding
season both sexes utter a call-note which consists of two clear
musical " rings," followed by a long descending " trill," like that
of our British Greenfinch. This frog, which at ordinary times is
the slowest and boldest of frogs, is now active and excessively
shy. Swimming rapidly between the blades of grass, it climbs a

1 Qiiari. J. After. Sci. xlii. 1899, p. 307.

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VI ENGYSTOMATIN AE 2 3 1

tuft, and dilating its throat, repeats its call ; but if in the least
disturbed, it is suddenly gone. The eggs are laid in quite tempo-
rary pools in grassy ground, and form separate globules of jelly,
which float on the surface of the water, and are heavily pigmented.
The development is excessively rapid. The segmentation be-
ginning at 10 A.M., they were hatched and wriggling about by
7 A,M. the following day. They are probably washed down into
deeper pools by the retreating waters, and for this purpose the
manner in which the eggs are laid, namely, in separate globules
of jelly, seems especially suited.

Brachycephalns tphippium in Brazil, the only species, is
remarkable for the development of a broad dorsal shield of bone,
which is fused with the processes of the second to seventh vertebrae,
an ossification which strongly resembles that of several species
of the likewise Brazilian CercUophrys, a genus of the Cystignathinae.

Stereocyclops is remarkable for the peculiar formation and
protection of the eyeball& The anterior portion of the sclerotic
is ossified into a ring, which surrounds the transparent cornea.
Another peculiarity lies in the metastemum, which is so much
broadened out that its cartilage is in wide contact with the
posterior edge of the coracoids. The epidermis is everywhere
" thickened by a chitin-like deposit." The only species, S. incras-
satus, found near Eio Janeiro, is an altogether aberrant creature.
Its general appearance recalls that of Fipa. The gape is large,
with a slightly projecting muzzle ; the limbs are so short that
the upper arms and the thighs scarcely stand out from the
broadened and flattened body, which is leathery brown, with a
narrow white median line extending dorsally from the nose to
the vent.

Ungystoma, with about five species in the Southern States,
Central and South America, is the type-genus of the whole family,
chiefly on account of priority of name. It is fairly characteristic
in so far as the mouth forms a narrow, somewhat projecting snout ;
the precoracoids, the clavicles, and the omosternum are absent,
the palate is devoid of teeth, the lining of the mouth forms a
dermal ridge across the palate and another in front of the oeso-
phagus, the tympanum is hidden, the sacral diapophyses are
moderately dilated, and the tongue is elliptical and free behind.
The pupil is vertical The fingers and toes are free, ending in
slightly dilated or blunt tips ; the terminal phalanges are simple

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2 32 ANURA CHAP.

and the hind-limbs are short. The male has a subgular vocal
sac

The most northern species is E. carolinefise, living in the
Southern United States, concealed under the bark of fallen trees
or in old fences. The skin is smooth, but forms a fold across the
head, behind the eyes. The general colour is brown, with light,
whitish dots on the under parts. Total length 1 inch.

Breviceps is a South African genus with three species. The
coracoids are very strong and directed backwards, but so broadened
tliat they form a long and strong symphysis, touching in front
that of the precoracoids, which stand transversely and are well
developed. The metasternum is cartilaginous and decidedly smalL
The sacral vertebra has much dilated diapophyses and is co-ossified
with the coccyx. The general appearance is extremely stout and
short, the head being almost drawn into the nearly globular body,
and ending in a short snout with a small mouth-opening. The
tongue is long and oval, not nicked, but slightly free behind. B,
viossambicus is about 2 inches long, and looks like an overstuffed
round bag, out of which the short arms and legs project from the
elbows and knee-joints only. The tarsus is provided witli a strong
horny, spade-like tubercle, which enables the creature to dig into
the ground, and into the nests of termites, which seem to be its
chief food. Peters found this species in enormous numbers, during
the tropical rains, coming out of the ground, whither they with-
draw again completely for the dry season. The skin is smooth,
reddish brown above, with darker patches ; the under parts are dull
white, with a large black patch on the throat.

Hemisus is another African genus, with two species, H.guttatum
in Natal, and H. sudanense in East and West Africa. This genus
is so exceptional in its shoulder-girdle, that Cope separated it
from all the other Anura as a special sub-order Gastrechmia.
The precoracoids are extremely strong, and form a broad symphysis
from which springs the long cartilaginous omosternum; the
coracoids are slender, very long, and converge backwards to a
narrow symphysis, and there is no metasternum. The two
symphyses are connected by a narrow cartilaginous median bai',
probably produced by the much modified epicoracoid cartilages.
However, except for the reverse development shown l)y the omo-
and meta-sternum, it is easy to connect this apparentlv quite
anomalous shoulder -girdle of Hemims with that of Brevxctp$.

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ENGYSTOM ATINAE 233

(cf. Fig. 5, 5 and 6, p. 25). The sacral diapophyses are slightly
dilated ; the fingers and toes are free and end in points. The
tongue is triangular, broader in front. The lining of the mouth
forms a transverse ridge across the palate, and another in front of
the oesophagus. The male has a subgular sac. The general shape
is stout, the head small and ending in a pointed snout. Colour
brown above, with whitish spots. Total length about 2 inches.

Glyphoglossus has a peculiar tongue. It is elongated, notched
behind and in front, divided into two lateral halves by a deep
groove ; moreover, the tongue is not only extensively free behind,
but also slightly so in front. The skin of the palate forms a
transverse serrated ridge. The precoracoids and the omosternum
are absent; the metasternum is a well-developed cartilaginous
plate. The sacral diapophyses are moderately dilated; the
terminal phalanges are simple. G, molossus, the only species, is
olive-brown above, marbled on the sides; the under parts are
uniformly whitish. This creature, about 2 inches in length, looks
like a roundish bag, with a ridiculous, short face. The type-
specimen, still the only one known, was taken by I)r Theobald
under the following circumstances : — " I had halted one day
within the tidal portion of the Irawaddy delta, to enable my
boatmen to prepare their dinner. One of my servants, having
cooked his rice, poured out the hot water as usual on the ground,
and some of it went down a hole that happened to be near the
spot. No sooner, however, had the hot water disappeared than
out scrambled in great haste a fine Glyphoglossus, only, alas ! to
be transferred to a collecting jar."

Phrynella. — The tongue is heart-shaped, free behind. The
palate is smooth and toothless. The fingers and toes end in
small discs, supported by T-shaped phalanges ; the fingers are
free, the toes extensively webbed. Precoracoids absent ; meta-
sternum cartilaginous. Pupil horizontal. Malay Peninsula.

Ph. pollicaris is dark olive brown above ; an oblique yellow
line runs from the eye to the angle of the mouth ; a pale yellow
mark, across the forehead, through the eyes, and down the sides
of the body. A dark-centred yellow patch on the anal region.
The limbs are banded yellow and brown. The imder parts are
brown, with paler specks, dark on the throat. Iris red brown.
The whole coloration changes considerably.

"They inhabit the hills of Perak from 3000 feet upwards,

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2 34 ANURA CHAP.

and live in holes iu trees, which are so situated as to contain
more or less rain-water. They have a loud flute-like, musical
note, which they utter at irregular intervals, principally during
the night. The form and size of the hole in which they are
seem to have a great deal to do with the loudness of the note, as
specimens when extracted from their holes have far more feeble
vocal powers than they had when in them. These frogs blow
themselves out with air, and look more like bladders than any-
thing else. When inflated they float on the surface of the water,
and will remain motionless for a long time, with legs and arms
stretched out." ^

Callula. — The tongue is round, entire, and free behind. The
palatine bones form an acute, sometimes toothed ridge across the
palate ; two dermal serrated ridges in front of the oesophagus.
Fingers free, sometimes with dilated tips, supported by T-shaped
phalanges. Precoracoids and omosternum absent ; metastemum
cartilaginous. Pupil round. About seven species iu the Indian
region.

C\ pulchra. — The following account has been extracted from
ilr. S. S. Flower's observations : ^ —

This pretty creature inhabits most of the warm portions of
the continental Indian region, from India and Ceylon to South
China and Malacca. The back is a rich dark brown, divided from
the yellow of the head by a narrow black line which extends
from eye to eye and forwards to each nostril. A conspicuous
yellow band runs from the eyes to the hind-limbs. The sides of
the body and the limbs are mottled yellow and brown. The
under parts are dirty buif ; the throat of the male is black. The
intensity of colouring varies individually and from time to time,
the contrast between the brown and yellow being occasionally
very brilliant. Total length up to 3 inches, the male being the
smaller sex.

" I have been told by both English and natives that this frog
was unknown in Singapore until some nine or ten years ago,
when it was introduced by a half-caste (why, it is not known),
and that it rapidly spread about the island. It is now well
known as the * Bullfrog ' by the English in Singapore, and
detested for the noise it makes at night. The voice of these
rotund animals can be heard every night after heavy rain ; it is
' S. S. Flower, P.Z.S. 1896, p. 910. « Ibid. p. 909.

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ENGYSTOMATINAE DYSCOPHIN AE 235

a deep guttural croak, ' wau-auhhhh/ very strident and prolonged.
The males croak while floating on the surface of the water, the
single vocal sac under the mouth inflated like a globe, and . the
arms and legs extended. They can hop well on land and are good
swimmers. The skin is excessively slimy ; the secretion comes off
profusely, and dries on the hand into a sort of white gum, with
a faint aromatic smell. This gum dissolves in hot water and
coagulates in cold. The general appearance of these frogs is very
stout, their girth being about twice the length from snout to
vent. The tongue, which is oblong in spirit specimens, in life is
very elastic, assuming, when extended, a vermiform shape and
reaching about 4 cm. in length. They appear after sunset,
crawling on old wood and feeding on white ants."

Sub-Fam. 2, Dyscophinae. — With teeth in the upper jaw.

This small group of nine genera, with scarcely more than one
dozen species, all with one exception living in Madagascar, has
been separated by Boulenger from the Engj'stomatinae merely on
account of the presence of teeth on the upper jaw and on the
vomerine margin of the palatine bonea He himself remarks that
Ccdluella may be considered a toothed Hypopachus, and Pletho-
dontohyla a toothed Callida. These are obvious cases of con-
vergent analogy. Except for the teeth, the Indian Ccdluella
would be merged into the American Hypopojchus, and this would
present an instance of the most puzzling geographical distribution.
In the case of the other two genera, one Indian and Malayan, the
other Malagasy, no such suspicion would arise, since there are
many other instances of such a coincidence of distribution. There
is the same divergence or unsettled condition in the modification
of various parts in the Dyscophinae as in the Engystomatinae.
The precoracoid bars are weak and curved backwards, and closely
pressed against the strong coracoids, in Dyscophus, Calluella and
Plaiypelis, while these elements are reduced to unossified bars,
and the clavicular portions completely lost, in Plethodontohyla
and in Fhrynocara. The omosternum is absent and the meta-
sternum is small in all except Dyscophus, in which both these
parts are exceptionally well developed and large, although re-
maining unossified. The palate of Dyscophus and Calludla is
provided with curious, serrated dermal folds like those which are
so common in the Engystomatinae ; and well-developed discs on
the fingers and toes, supported by T-shaped phalanges, are

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236 ANURA CHAP.

possessed by Platypelis, Cophyla and others. The sacral dia-
pophyses are dilated. The pupil is either horizontal or vertical
Those which are provided with discs to the fingers and toes
are climbers, and mostly slender and long-legged, sometimes of
very small size, for instance Cophyla, the body of which is scarcely
one inch in length.

The genera can be determined by means of the following
key : — ^

-4. Pupil vertical. Palatine teeth in long transverse Beries. *

a. Precoracoids ossified. Tips of fingers and toes not dilated.

Sternum very large. Madagascar . Dyscophu^

Sternum small Burmah . Calluella.

b. Precoracoids not ossified. Tips dilated . Plethod(mtohj\a.
B. Pupil horizontal

a. Palatine teeth in long transverse series.

a. Precoracoids ossified. Tips dilated.

Fingers and toes free. Precoracoids en-
tirely ossified .... Mantipus.
Fhigers and toes webbed at the base.

Precoracoids semi-ossified . . Platyhyla.

p. Precoracoids not ossified. Tips not dilated . Phrynocara,

b. Palatine teeth in one or two small groups.

Precoracoids ossified. Tips dilated.

Two small groups of palatine t-eeth . PlatypdU

One single group in the middle of the

palate ...... Cophyla.

No teeth on the jmlate . Anodontohyla.

Dyscophus antongili, — Madagascar. Greneral appearance stout,
with short legs and a wide mouth. Total length about 3 inches.
The skin is mostly smooth, and forms a broad glandular fold
which extends from the eye to the groin. The upper parts are
beautiful magenta red, with a purplish streak beneath the lateral
folds; the under parts are yellowish white, with minute grey
specks. Red or pink colours, and the lateral folds, occur also in
most of the other members of this family, for instance in the
Indian genus CallueUa.

Sub-Fam. 3. Genyophrymiiae. — With very sinall teeth on thf
anterior portion of the lower jaw.

Genyophryne thomsoni. — Pupil horizontal. Tongue obloug

and entire. With teeth on the palatine bones, and a serrated

transverse dermal ridge in front of the oesophagus. Sternum

cartilaginous. Precoracoids absent. Sacral diapophyses moderately

' Boulenger, Ann. Nat. Hist. (G), iv. 1889, p. 247.

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VI GENYOPHRYNINAE — RANIDAE CERATOBATRACHINAE 237

dilated. Tympanum hidden. Head large and much depressed.
Heel with a triangular dermal flap. The smooth skin is pink
brown above, with blackish marks ; a light line extends on each
side from the eye along the back. Under parts black. About
32 mm. in length. Sudest Island, between New Guinea and
the Louisiade Archipelago.

Fam. 7. Bsjiidae. — Frogs, in the true sense, are all well
diagnosed as Mrmisternia, with cylindrical sacral diapophyses.
According to the presence or absence of teeth in the jaws they
can be subdivided as follows : —

Sub-Fam. 1. Ceratobatrachinae, with teeth in the upper and
in the lower jaws. The sole representative is the genus Cerato-
batrachus.

Sub-Fam. 2. Baninae, with teeth in the upper, but none in
the lower jaw. These are the Eanidae of Boulenger in the
Catalogue of Batrachia Salientia.

Sub-Fam. 3. Dendrobatinae, without teeth in the upper and
lower jaws.

Sub-Fam. 1. Ceratobatracluiiae. — Teeth present in both
jaws. Those of the lower jaw, between 20 and 30 in number in
Ceratobatrachus, the only genus, are nearly all inserted upon the
articular bone ; only 2 or 3 are carried by the dentary element,
which, although large, enters into the formation of the upper
border of the jaw at the anterior end only. In the small extent
of the share of the dentary in the formation of the edge of the
lower jaw, and in its anterior " toothlike " process, Bana adspersa
of Africa bears unmistakable resemblance to this genus. The
tongue is deeply notched, and free behind. Pupil horizontal.
Vomers furnished with teeth. Tympanum distinct and large.
Precoracoids present. Omosternum and presternum with a bony
style. Sacral diapophyses cylindricaL Fingers and toes free,
with swollen tips. Outer metatarsals united. Male with two
internal vocal sacs.

C. guentheri, Solomon Islands, the only species, has an
enormous mouth and a triangular head not much smaller than
the rest of the body. The skull is furnished with prominent
ridges and a small curved spine at the angle of the jaws. The
hind-limbs are rather short. The skin of the upper parts shows
linear ridges, variously arranged ; that of the belly is granular.
A triangular dermal flap on the tip of the muzzle, one on the

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238 ANURA CHAP.

upper edge of the eyelids, others on the heel and above the
vent. The cjolour and markings are very variable, the ground-
colour is yellowish to pink, brown, grey or olive, with darker
and lighter markings. Total length of the males 3 inches, of
females 3^ inches. — Guppy, the discoverer of this peculiar
creature, remarks that " horned Frogs are very numerous in
these islands, and so closely do they imitate their surround-
ings in colour and pattern, that on one occasion I captured
one by accidentally placing my hand on it when clasping a
tree."

Sub-Fam. 2. Raninae. — The vertebrae are procoelous and
devoid of ribs. The precoracoids are always present and ossified
from the clavicles, and are parallel with the much stronger and
ossified coracoids. The omosternum usually possesses a bony
style, but in the Indian genera Nannohatrachxts and Nannophrys
and in Phyllodromtcs of Ecuador it remains cartilaginous, and
in Colosthetus of Colombia it is absent. The metasternuiu
also possesses a bony style, but it remains cartilaginous in the
Indian genera OxyglossnSy Nannophrys, Nannohatrachus and PhyllO"
dromus, in the last two genera rather reduced and slender,
while in the Ecuadorian and Colombian genera Jfylixalus,
Prostherapis and Colosthetvs, it is reduced to a membranous piece.
In quite a number of genera the normal number of phalanges
is increased by one owing to the intercalation of an extra
phalanx between the terminal and the otherwise penultimate
phalanx.^ This is the case in all the species of Cassuia,
Hylambates, Pappia, Megalixcdus, Rhdcophorvs, Chiromantis,
Ixalus and Nyctixalus, but it is doubtful if all these genera
are thereby more nearly related to each other than to the rest of
the Raninae. The structure of the tips of the fingers and
toes exhibits more variety. The terminal phalanges are mostly
simple, with slight swellings at the ends, or they are Y- or T-
shaped in conformity with more or less developed adhesive discs ;
in the African genus Hylamhates only they are claw-shaped, as
in the Hylidae.

Gampsosteonyx batesi, recently described by Boulenger from
the Gaboon, shows a unique modification of the terminal
phalanges of the second to the fifth toes. They are transformed
into sharp and curved claws, like those of a cat, but instead

1 See Boulenger, F.Z.S. 1888, p. 204.

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RANIDAE — ANINAE

239

of horny sheaths, it is the bone itself which is thus sharpened
and perforates the skin, an anomaly reminding us of the ribs
of Triton waltli. Total length of the type -specimens, about
3 inches.

Adhesive discs are common, and are best developed in Ehaco-
phoruSy IxaluSy Rajpjda, and Megalixalus. In the Neotropical
genera, excepting Colosthettcs, the discs are very peculiar, being
provided on the upper side with leathery scales which are
separated by a fissure. The fourth and fifth metatarsals either
diverge and are connected by a distinct web, or they lie close

^sRANIDAC IlllllU DENDROBATES. M^ FORMS WITH FINGER DISCS.

Fig. 46. — Map shovring distribution of the Ranidae.

together with only a groove between them, or lastly they appear
externally united.

The tympanic disc is very variable, large, small or quite
hidden. Vomerine teeth are present or absent. The pupil con-
tracts into a horizontal slit except in some Palaeotropical genera.
The tongue is universally free behind, mostly deeply notched,
and can be well protruded ; only in the Indian Oxyglossus and
in the Neotropical genera, excepting Hylixalus, its posterior
margin is entire. — There are terrestrial, arboreal, and aquatic
members in this large sub-family. The geographical distribution
of the Baninae, which comprise about twenty genera with at least
some 270 species, is almost entirely Arctogaean. None, with
the exception of three species in the Papuan subregion, occur
in the Australian region ; and only four genera, with one or
two species each, inhabit the tropical Andesian district, the

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240 ANURA

remainder of South America being without any Baninae. All
the species of the whole Periarctic region belong to the genus
Rana except in Eastern Asia, where the closely allied genus
Ehacophorus occurs also. The entire sub-family of Baninae is, in its
fulness and diversity of development, essentially Palaeotropical.

Many of the genera, even in the present more liberal sense gls
interpreted by Boulenger, are based upon imimportant characters,
and in reality run into each other. This is for instance the case
with Bana and Hhacophorus.

The following tabular arrangement is merely a key for
determination and does not necessarily express relationships.
The presence or absence of vomerine teeth is a character easily
ascertained, but it separates closely allied genera, for instance,
Rhacophorus from Ixalus and Micrixahis from Rana.

The genera with extra, interpolated phalanges are marked *.

Key for the determination op the genera of Raninae.

1. Pupil vertical.

A. With vomerine teeth.

rt. Omostemum very slender and cartilagin -
oils. Small discs. India and Ceylon,
3 species ..... Nannohatrachus.
h. Omostemum with a bony style.

a. Outer metatarsals webbed. Small

discs. South India, 2 species . NyctihatrachvA,
/3. OwtjOT metatarsals close together. Africa.
Fingers and toes with inter-
polated phalanges.

Without terminal discs.

2 species . . . Cassina*

With discs supported by
claw-shaped phalanges,
10 species . . . Hylambates.*

Fingers and toes without inter-
polated phalanges ; with-
out discSb
Toes webbed . THchobatrachus robusttis^

p 271.
Toes free, with sharp

claws . Gampsosteonyx hatesi^

p. 272.

B. Without vomerine teeth. Discs well de-

veloped. Outer metatarsals united.

Tropical Africa and Madagascar, 7 species Megalixalus*

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VI RANINAE 241

II. Pupil horizontal.

A, With vomerine teeth.

a. Outer metatarsals webbed together.

Fingers free, toes webbed . Rana^ p. 249.

Fingers and toes more or less webbed.

Always with discs . . Rhacophorus* p. 245.

Two fingers opposed to the others.

Africa Chiromantis* p. 244.

b. Outer metatarsals united, or separated

by a groove only.
Omo- and meta-stemum with a bony

style ComufeVf p. 243.

Omo- and meta-stemum slender and
cartilaginous,

Ceylon, 2 species . Xannophrys.

Mozambique . Phrynopm boulengerL

B, Without vomerine teeth.

(I. Palaeotropical.

a. Tongue narrow and entire. No
discs. Outer metatarsals
webbed. India, 3 species . Oxyglossu9.

p. Tongue oval, feebly nicked. Large

discs. Solomon Islands . Batrachylodes vertebralis,

Karin Hills . . Phrynoderma atperum.

y. Tongue deeply not<;hed. Outer
metatarsals united by a web.
Discs none or very small.

Africa, 3 species . . Phrynohatrachus,

Borneo . . . Oreohatrachus baluensis.

With regular discs.

Number of phalanges normal.

India, 6 8i)ecies *. Micrixalus.
With an extra, interpolated phalanx.

India, 18 species . Ixalus*
Two fingers opposed to the others.

Karin Hills . Chirixalus dorme*

& Tongue heart-shaped. Outer metatarsals united.
Fingers and toes free, tips blunt

Africa, 8 species . . Arthroleptis, p. 242.

Fingers and toes more or less

webbed, with regular discs.

Africa and Madagascar,

23 species . . . Rappia*

VOL. VIII

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242 ANURA CHAP.

b. Neotropical.

Metasternum small, cartilaginous or menibranouri.
With discs.

1. With a pair of dermal scales on the discs.

Omostemum i^ath a bony style.
Tongue heart-shaped.

Ecuador, 2 species . Hylixalm.
Toes free. 6 species . Phyllohates, p. 242.
Tongue entire. Ecuador
and Colombia, 3
species . . . Prostherapis.

Omostemum cartilaginous.

Ecuador . . . Phyllodromus piUchelliut.

2. Discs without scales. Omo-

stemum absent

Colombia . . . Colosthetus latinasus.

Phyllobates} — This is one of the few Neotropical genera, and
like nearly all of these has peculiar adhesive discs on the fingers
and toes, each disc bearing on its upper surface two dermal
scales. The tympanum is distinct. Vomerine teeth are absent.
The general appearance of the five species is that of tree-frogs.
One species, Ph, bicolor, yellowish above, dark brown beneath, lives
in Cuba. The others inhabit Central America and Venezuela.
They seem to have peculiar nursing habits. Ph, trimtatis of
Venezuela and Trinidad carries its tadpoles on its back, on to
which the young fix themselves by means of their suckers.
Nothing is known about their breeding habits, for instance whether
the young are hatched on the back, or, as seems more likely,
if the parents (the specimen described by Boulenger * is a male)
only give their offspring a temporary lift in order to convey
them from a drying-up pool to a healthier place. It is remark-
able that several species of Dendrobatinae, which inhabit the
Biime countries, have precisely the same habits.*

Artkroleptis. — Slender and long-limbed little frogs, about
one inch in length. The fingers and toes are free, very slender,
and end in slightly dilated tips, the supporting phalanges being
simple. The tympanum is variable. The skin is smooth or
finely granulated. The colours are inconspicuous, brown or grey
tones usually prevailing. About ten species are known, mostly

* BouleiJger hais shown {P.Z.S. 1888) that Bibron's 8})€cie8 of PhyllobnUs,
liitherto grouped amongst the Cystignathidae, are Ranoids, closely allied to
Ilylixalus and Prostherapis. The other species now form the Cystignathoid
genus Syrrhopus, Cope (of. p. 212,i. - P.Z.S. 1895, p. 209. » Cf. p. 273.

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VI RANINAE 243

from Continental Africa, a few from Madagascar and the islands
in the Indian Ocean.

A, seychellensis. — Brauer ^ has discovered the mode of nursing
of this frog. He found a specimen of A. seychellensis which carried
nine tadpoles on its back, in the month of August, in the Sey-
chelles, about 1500 feet above sea-level, upon an old tree-fern. The
little ones were ah'eady provided with long tails, the hind-limbs
were partly free, the fore-limbs still covered by the skin, and
they held on by their bellies ; not, like the young of Phyllobates,
by their " suckers." Another specimen carried young which were
still further developed. He also found
an old frog, near which was lying a little
heap of eggs, not enveloped in a common
mass of jelly. The old frog escaped, but
the eggs were taken care of in a vessel
with moist sand at the bottom. By the ^,

following morning the eggs were hatched ^^^J^^ ^^,^^/t
and the tadpoles were clinging by their ^^^ ,-;,-Arthroieptis seychei-

bellies on to the walls of the glass. Unsis, carrying Tadpoles.

Brauer concludes that the young, when ^^- (^^^er Brauer.)
hatched, creep on to the parents' back, he or she waiting near the
heap of eggs until the latter are ready. Curiously enough, he did
not find out the sex of the nurse, nor are we told if the young are
taken to the nearest water to finish their metamorphosis, or if they
remain upon the parent's back until they hop off as baby-frogs.
The yolk is very large. When the four limbs are already
developed, the gill-cavity possesses no gills and no outer opening ;
and since the lungs are only just beginning to sprout, the tad-
pole must needs breathe by means of its skin. The jaws have
no horny coveringa The adults live on the ground between
moist leaves, and eat chiefly termites.

Cornufer, with about twelve species, is an essentially Austro-
Malayan and Polynesian genus, but one species, C, johnstoniy has
been found in the Cameroons. The fingers and toes are free,
and their T-shaped phalanges support adhesive discs. The
tympanum is distinct. The general shape is frog-like, usually
with slender and very long hind-limbs and toes, the discs of the
latter being much smaller than those of the fingers. The coloration
is dull, mostly brown, more or le.ss marbled, whitish below. The

' Zool. Jahrh. Syst. xii. 1898, p. 89.

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244 ANURA CHAP.

upper eyelid of some species, e.g, of C. unicolor of New Guinea,
has a small tubercle, hence the generic name. The skin of the
back is glandular and granular, forming slight folds on the back
and on the sides of the head in some species. The male has
one or two internal vocal sacs.

C, corrugatus is one of the most widely distributed species,
inhabiting the Philippines, New Guinea, and Duke of York
Island. The granular skin forms longitudinal folds on the back,
one of which reaches from the eye to the shoulder. Brownish
above with darker markings, below yellowish, with or without
brown spots on the throat. — Three species inhabit the Fiji
Islands.

Of C, solomonis of the Solomon Islands little is known about
the propagation, although the large size of the egg, whicli
measures 5 mm. in diameter, suggests that the young undergo
most or the whole of their metamorphosis within the egg.

Chiromantis is distinguished by the peculiar arrangement of
the fingers, the first and second being opposed to the others;
their terminal phalanges are obtuse and support small knobs or
discs. The general shape is that of a frog with long and slender
hind-limbs. The tympanum is distinct.

Ch. xerampelina, the type -species, was discovered by Peters
at Mozambique; it is a middle-sized frog, about 2 inches in
length, brown above with reddish spots on the sides ; the male is
devoid of vocal sacs.

Ch, petersi, a native of East Africa, differs from the preceding by
the possession of an internal vocal sac. Ch. rufescens = guineensis
shows very little of the typical grasping arrangement of the
fingers ; the two inner ones are separated from the two outer
fingers by a wide gap, but they all lie in the same plane, are
much webbed and possess large discs, so that by the latter two
characters a link is formed with Ithacophorus, to which the
present genus is closely allied. Total length about 2^ inches.

Buchholz ^ has observed the peculiar breeding habits of this
rather large, brown, and slender tree-frog in the Cameroons. In
the month of June he found on the leaves of a low tree, standing
in the water, a white foamy mass, like the froth of a broken
egg, containing a number of newly hatched larvae and quite
transparent eggs. Within three or four days this mass l>ecame

* Monatsher. Bcrl. Ac. 1875, p. 204 ; 1876, i). 714.

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VI RANINAE 245

fluid, and the larvae, provided with external gills and a long tail,
swam about in the slime. In the natural course of events the
larvae are probably washed down into the water by the rain.
He found that the female deposits the eggs in the foamy
mass at night, during the months of June and July, on
various kinds of trees, either between the roots or iu a cavity
formed by gluing together several leaves, sometimes 10 feet and
more above the water, or near the margin. On one occasion
the mother was seen sitting upon the foamy mass, clasping the
same with its four limbs.

Bkacophorus. — This large genus, containing more than forty
species, has a curious distribution. At least one dozen species
are found in Madagascar, eight or nine in Ceylon, the rest in
Southern India, the Himalayas, the Malay Islands and Philip-
pines, extending northwards through China and Southern Japan.
Therefore this genus, with the three species of the African
ChiromarUis, extends over the whole of the Palaeotropical
region. The generic name has reference to the possession by
many species of little dermal flaps, especially at the inner side of
the heel, and it has nothing to do with the parachute-like use of
the hands and feet of certain species, to be mentioned presently.

The terminal phalanges are generally bifurcated, rarely
obtuse, and support well-developed adhesive discs. The fingers
and toes are webbed to a variable extent. The two outer meta-
tarsals are likewise connected by a web. The tympanum is
distinct. The general appearance is tliat of tree-frogs, and
many of them are green. The males have one or two in-
ternal vocal sacs. Not all the species have dermal appendages.
Hh, maximus, for instance, the largest of all, living in the
Himalayan forests, has none. A heel-flap occurs in some half-
dozen Indian species ; and Rh, madcLgascariensis has these flaps
on the heels and on the elbows. Some have queer little lappets
above the vent, or on the edges of the arms and legs; in others the
bend of the arm is fringed. The small si^e of these appendages,
in comparison with the webs and discs, makes them practically
useless so far as increase of surface is concerned, and they have
most likely some other, although unknown meaning, especially
the flaps over the vent. Lastly, in the majority of species the
fingers are not more than half-webbed, or even less, and in a
few only, the webs reach down to the discs.

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246

ANURA

Several species of this genus are remarkable for two reasons.
rii*8t, the great enlargement of the fully-webbed hands and feet,
which are then used as parachutes ; secondly, the mode of pro-
pagation.

Greatly exaggerated notions are, however, entertained about
the parachutes, ever since Wallace's description^ of the iirst
" flying frog." The creature was brought to him in Borneo by

Fia. 48. — Rhacojj/u/rns pardaliSf x about 1. {From Vi&U&ce^ Mala j/ A rcJiipelago.)

a Chinese workman. " He assured me that he had seen it come
down, in a slanting direction, from a high tree, as if it flew. . . .
The body was about four inches long, while the webs of each
hind-foot, when fully expanded, covered a surface of four sc^uare
inches, and the webs of all the feet together about twelve square
inches."

The species in question is Hh. pardalis, an inhabitant of
Borneo and of the Philippine Islands. Specimens from Wallace's
Collection are in the National Collection and the largest speci-

* Malay Arcldpelago, 2Tid cd. i. 1869, p. 38.

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RANINAE 247

men shows the following measurements. Total length 6 "5 em,
or 2^ inches, not 4 inches.

Area covered by one fully-expanded hand 3*4 square cm.
n M « foot 6-0 ,, „

9*4 square cm.

i.e. for the four limbs 18'8 square cm. = about 3 square inches,
and not 78 square cm. or 12 square inches. By some un-
fortunate oversight Wallace must have mixed up the total
expanded area with that of the four hands and feet ! In Brehm's
Thierleben the 78 square cm. have increased to 81 cm., and the
artist h£is in the somewhat larger species Rh. reinwardti improved
upon this, and has produced a truly startling picture by a further
exaggeration based upon the figure given by Wallace.

Rh. reinwardti lives in the forests of the mountains of Java
and Sumatra. It reaches 3 inches in length, and is grass-green
above, yellow below. Younger specimens are further adorned
with large blue patches on the webs of the hands and feet and
behind the armpits. Besides the flap on the heel and the
curious cutaneous fringe on tlie forearm, suggestive of an in-
cipient flying-membrane, the skin forms a projecting fringe on
the inner side of the fifth toe and a transverse flap above the
vent.

Of Rh. leucomystax, Annandale, who accompanied the Skeat
Expedition to Malacca, gives the following account : — '' This frog,
which is called by the Malays of Lower Siam either ' Berkata
Pisang ' (banana-frog) or * Berkata Ehumah ' (house-frog), lays
its eggs either on leaves of branches overhanging the water,
or on the mud surrounding buffalo -wallows. The ova are en-
closed in a round mass of yellow froth, which afterwards becomes
steel -grey, about as large as a cricket -ball. Should they be
placed judiciously in a position sheltered from the sun, the
tadpoles may either hatch, and reach a considerable degree of
development, before the mass is washed into the water, or the
froth may be melted almost as soon as it is formed and the eggs
be carried into a pool by a shower of rain. Very often, how-
ever, the whole mass is dried up by the heat of the sun before
the rain comes. During the breeding season, which seems to
; occur as often as the land is flooded under the trees, for I have
I never seen the eggs of this frog on the bank of a river, the

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248 ANURA

males croak loudly, producing a sound which can hardly be
distinguished from the chattering of the large black and yellow
squirrel, Sciurvs bieolor"

These arboreal frogs have a peculiar mode of nursing the
young and taking care of the egga - Bh. macvlatus of Ceylon,
Malacca, etc., and Bh, schlegeli of Japan, lay their eggs in a foamy
mass^ the size of a fist, on the margins of ponds, and the whole
process has recently been described by Ikeda.^ He observed
the Japanese Rh, schlegeli depositing the eggs in soft, muddy
ground covered with grass, and in wet, muddy banks of paddy-
fields, ponds, and similar localities near Tokyo. Sometimes they
are deposited between the leaves of trees, near the ground-
The breeding season extends from the middle of April to the
middle of May. Towards the evening the female, bearing the much
smaller male on her back, retires underground for the deposition
of the eggs. The spots chosen are 10-15 cm. above the surface
of the water; the female digs a spherical hole 6-9 cm. wide.
Sitting thus concealed underground, the frogs assume a dark
colour and the spawning takes place during the night, where-
upon the parents leave the nest. The eggs are enveloped in a
white mass of jelly full of air-bubbles, the whole frothy lump
looking like the well-beaten white of a hen's egg, with the
yellowish eggs scattered through it, and measuring some 6 cm.
in diameter. The air-bubbles are 2-3 mm. large. The froth
is originally very elastic and sticky, but it gradually sinks down,
becomes liquid and ultimately runs out of the hole. It is pro-
duced in the following peculiar manner. During and after the
deposition of the eggs the female puts her feet upon the sticky
jelly, part of which adheres and is then pulled out as a thin,
transparent membrane stretching between both feet. The latter
are then thrust backwards, the membrane is folded downwards
and becomes a vesicle of 5 to 10 mm. in width. By repeated
working of the limbs the successively formed bubbles ai-e trodden
and kneaded into froth, which ultimately surrounds and at the
same time separates the eggs.

The female of Jth. reticulatvs of Ceylon attaches the eggs,
about twenty in number, to the under surface of her belly, on
the skin of which they leave little cellular impressions. What
becomes of the tadpoles is not known.

^ Annotat. Zool. Jaji. i. 1897, p. 113.

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RANINAE 249

jBA. leiicomystax is found in the Malay Archipelago, Farther
India, and the Philippine Islands.

S. S. Flower ^ found the tadpoles about Singapore, from January
to April, in small ponds and in rain-water butts. The spiracle
lies on the left side, directed backwards and upwards, nearer the
anus than the end of the snout. The anus opens on the right
side. Exceptionally large tadpoles measured 46 mm. in total
length, the recently transformed young only 14-18 mm.

" A cheerful little frog of most graceful build. It comes out
^ from its hiding-places shortly before sunset, and remains abroad
all night. The males are easily found as they sit on shrubs or
trees, or on the edges of the rain-water butts under the verandahs
of the houses, and from time to time utter a single, rather
musical, short croak. In March and April they can be found
both by day and night in embrace, in the ponds. This species
changes both its colour and markings very rapidly and fre-
quently, but dark bands across the legs can always be more or
less distinguished ; the lower parts are some shade or other of
buff, but the principal variations of the upper part are as
follows: pale bronze, either uniform or with four longitudinal
dark -brown or black lines; imiform, almost orange, bright
bronze ; chocolate, with darker mottling ; pale brownish green
or olive, with irregular dark spots; yellowish green, mottled
with darker or brown." The females are considerably larger
than the males; the largest male caught was 48 mm. from
snout to vent, and the largest female 68 mm.

Bana. — The following combination of characters should be a
sufficient diagnosis : pupil horizontal ; tongue deeply notched
and free behind ; vomers with teeth ; fingers free, toes webbed,
fourth and fifth metatarsals diverging and webbed together.

In conformity with the great number of species and the wide
distribution of this genus some of the organs vary considerably,
indeed so much so that many of these modifications have been
deemed sufficient to be of generic importance. Fortunately the
species are so numerous that these characters mostly form an
uninterrupted series from one extreme to the other.

The terminal phalanges are mostly simple and pointed ;
sometimes transversely dilated or T-shaped, according to the
presence of more or less developed discs. Such discs are, for

1 r.Z.S, 1896, p. 906.

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2 50 ANURA CHAP.

instance, present in the Malay species It, erythraea and R. chul-
conota and in the Indian R, corrugata. The tympanum occurs in
every stage from a conspicuous, free disc to being quite hidden
by the skin. .The vomerine teeth either form a pair of tiny,
mostly transverse rows, between the choanae, or they are
arranged in two oblique series which extend beyond the hinder
edges of the choanae.

The vocal sacs vary greatly. Many species, e.g. R. agilis,
have none at all. Most species have a pair of internal sacs, and
in comparatively few, about a dozen, these sacs have become ex-
ternal, a feature which indicates no relationship of the species
thus distinguished, for instance the European R. esculenta, the
Japanese R. rugosa, the Indian R, hexadactyla, R. cyanophlyctis
and R. chloronotay the Bornean R. glandidosa, the African R.
oxyrhynchus and R, raascarenieiisis, the Mexican R, montezumae.
In R, esculenta, and perhaps in a few others, even the female has
some traces of these otherwise male organs, indicated by slit-
like folds of the outer skin l)elow the angles of the lower
jaw.

Nuptial excrescences on the inner metacarpal tubercle and on
the inner fingers of the male are common; they reach their
greatest development in the Himalayan R. liebigi, the male of
which is " remarkable for the extreme thickness of its arms, the
inner sides of which are studded with small conical black spines,
each supported on a rounded base produced by a swelling of the
skin. A large patch of similar spines exists on each side of the
breast."'

Specific glandular complexes in the skin are mostly restricted
to a pair of lateral or dorso-lateral folds ; they are often absent,
but a few species, e.g. R, glaridulosa of Borneo, R, temporalis of
Ceylon, R, elegans and R, albolahris of West Africa, have a pair
of large flat glands at the base or inner side of the arms.

All the species of Rana, except those in the Solomon Islands,
spawn in the water, where the development of the tadpoles takes its
course. Those of some Indian species, notably R, alticola and
R, afghana of the Himalayas, and R, ctirtipes of Malabar, are
very peculiar, being provided on either side of the shoulders with
a large oval parotoid-like gland, well defined and crowded with
pores; R alticola possesses in addition an unpaired, sharply

' Boulenger, CaL BcUracJt. Salic tUia, p. 22.

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RANINAE 251

marked glandular complex on the top of the root of the tail, or
rather upon the future coccyx. These complexes gradually
disappear with age.

The genus Rana, with about 140 species and sub-species,
is distributed over the whole of Arctogaea so far as this is
available for Amphibian life, while there are only a few stragglers
in Notogaea, namely, a few species in Ecuador and in the
Peruvian or Upper Amazon district. None exist in the rest of
the Neotropical region, including the Antilles, and practically
none in Australia; but R. arfaki and R, papiia inhabit New
Guinea and the northern corner of Australia, R, kreffti the
Solomon Islands. A few species are restricted to Madagascar,
and a few others live there and on the continent of Afiica.

So far as number of species is concerned, the home of the
genus Rana is the Palaeotropical region ; about one dozen (some of
them with a very wide range) live in the Palaearctic sub-region,
scarcely more in the Nearctic sub-region, and a few in Central
America.

R. tem})oraria (the common European Brown Frog or Grass-
frog). — The tympanum is distinct, two-thirds the diameter of the
eye in size. The first finger is slightly longer than the second,
which is shorter and weaker than the others, whilst the fourth is
the longest. All the fingers are quite free. When the hind-
limbs are laid forwards along the body, the ankle-joint reaches to a
point between the eye and the tip of the snout. The five toes,
which are about half webbed, increase in length from the first to
the fourth, while the fifth is about equal to the third. The sole
of the foot has a small, blunt, inner metatarsal tubercle ; the outer
one is scarcely visible. The skin is smooth, always moist, owing
to the minute mucous glands ; but a series of larger glands forms
a pair of folds along the upper sides of the back ; l)eginning
behind the eyes they converge slightly beyond the shoulders,
diverge a little in the sacral region, and converge again towards
the vent. Another, much feebler, A-shaped ridge lies between
the shoulders.

The male has two internal vocal sacs, which, when in use,
bulge out the skin of the thix^at beneath the angles of the mouth
like a x)air of globes. It is further distinguished from the female
by the stronger muscles of the arms and by a pair of swollen
pads on the inner side of the first finger. During the pairing

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55 2 ANURA

season these pads are enlarged into cushions covered with black
horny rugosities.

The iris is golden, with dark specks. The coloration is,
generally speaking, brown above, with black-brown irregular
spots, especially on the sides of the body, and with cross-bands
on the legs. The under parts of the male are white or pale
yellow, with a bluish tinge on the throat, while the female is
more yellow instead of white, inclining to orange. In both sexes
the under parts are mostly spotted with darker colours. A large
dark-brown patch, extending from behind the eye over the
tympanum towards the shoulder, is always present and has given
this frog its specific name. Otherwise the coloration varies con-
siderably ; more or less according to the locality and nature of the
sxirroundings, and to individual variation and temporary change
of colour.

Some specimens are almost spotless above and of a rich
brown, or almost yellow colour, the spots being restricted to the
sides below the lateral folds. Others have very few spots, but
tliese are then arranged in two interrupted streaks on the
back. The under parts, especially the flanks, may be lemon
yellow instead of whitish, and the darker markings may be
almost absent. Boulenger has figured a beautiful specimen,
almost orange red, with red spots and vermiculations on the yellow
under surface. I have foimd similar red specimens of unusually
striking appearance between Berlin and Spandau in a forest-
glade, through which run little streams with banks of red fen*u-
ginous soil Specimens which live in woods with rich black soil
are often very dark, all the brown and reddish tints being absent.
The variations are, however, really endless, and it is difficult to find
two individuals exactly alike,even amongst a great number collected
in the same locality. Moreover, they change colour. Warmth
makes them paler, cold causes the chromatophores to expand and
the whole frog appears darker. During the breeding season the
males assume a delicate bluish hue, especially on the throat, but
this film quickly fades away when they are taken out of the
water. It is caused by the swelling of the cutaneous lymph-
spaces which extend their ramifications into the epidermal layer,
and it is not a question of pigmentation or of chromatophores,
but a case of interference -colours, bhie being frequently the
result of the light passing through a cloudy, colourless, but not

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VI RANINAE 253

quite transparent and thin stratum, in this case the turgid
epidermis.

The habits of the Grass-frog are essentially terrestrial. It
spends most of its time on land, preferably in damp places, but
local fashion permits of a great deal of freedom, as these frogs
are sometimes found not only in very wet, naturally irrigated
places, but also in the water itself. However, the Grass-frog
when pursued rarely takes to the water for safety. It trusts
to flight, first by a few long and fast jumps, and then to conceal-
ment by squatting down between grass, under leaves; it
rarely creeps into a hole, even if there be one near. The
jumps soon become shorter and shorter after a few dozen repeti-
tions. It swims well, but cannot climb. The food, which
consists chiefly of insects, snails, and worms, must be mo\'ing to
excite interest ; then the frog, whose favourite position is half
squatting, half supported by the arms, erects itself, and, facing
the insect, turns round upon its haunches, adjusts its position
anew by a shifting of the legs, and betrays its mental agitation
by a few rapid movements of the throat. All this time the prey
is watched intently until it moves ; then there follows a jump, a
flap of the tongue and the insect is seen no more. As a rule
these frogs do not crawl, they jump or hop, even whilst stalking,
and this takes place at any time of the day ; in fact they are
very diurnal, although they become more active towards the
evening. When caught they are at first very wild and, like
all true frogs, very impetuous, committing acts of astonishing
stupidity without any apparent sensie or appreciation of distance
or height. The captive will not only jump off the table, whilst
a toad stops at the edge and looks carefully down, but without
hesitation he jumps out of the window, regardless of the height
above the ground. This is di^e to sheer fright ; he loses his
head. When at large in his native surroundings, nothing will
induce him, although hotly pursued, to commit suicide by
jumping down a precipice. But all this wildness calUas down
wonderfully soon. The captive no longer dashes his head
against the glass, he does not struggle or twist when taken up ;
on the contrary, he makes himself at home, watches your coming
with intense expectation, and without hesitation accepts the
proflfered mealworm, maggot, butterfly or earthworm ; in short,
he shows what a jolly and intelligent fellow he really is.

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2 54 ANURA CHAP.

The Grass-frog has many more obvious enemies than perhaps
any other Amphibian, and it is not even slightly protected by
any appreciable poisonous secretion. Nevertheless it is extremely
common. A whole host of birds eat it— for instance^ buzzards,
harriers, and above all storks. Foxes, polecats, and stoats are not
averse to it, and the Grass-snake derives its main sustenance from
it. In fact the enemies of the little frog are legion, one of the
worst being Man. In France, Italy, and other parts of the
Continent, the skinned fleshy hind-limbs are turned into a by
no means disagreeable ragoAt, or into dainty morsels when
fried in butter and encrusted with bread -crumba This frog,
together with its cousin the Water-frog, also suffers from the
distinction of being one of the chief martyrs to science. Per-
haps the story is true that Galvani was led to his investigations
into animal magnetism and electricity by observing that the
legs of a number of skinned frogs, strung up by his wife upon
the bronze railings of the balcony, jumped whenever the scissors,
which cut off the feet, touched the other metal Frogs have
suffered ever since. Easily procured and of a convenient size,
they are used in every biological laboratory, and the young
student is supposed to be initiated into the mysteries of Verte-
brate structure by the careful dissection and study of this, the
worst of all the so-called types. Next to Man there is no
animal which has been studied so minutely, and has had so
many primers and text-books written on it, as this frog. In
spite of all this it is very little understood, thanks to its rather
aberrant and far from generalised structure.

However, the frog, by reason of its fertility, holds its own. Early
in the year, sometimes while there is still ice and snow, the frogs
leave their hibernating places (mostly holes in the ground, under
moss, or in the mud), and they begin to pair in standing or slowly
flowing, mostly shallow, waters.

They are not always very careful in the selection of the
spawning locality, many of them lay their eggs in a ditch, or
even in the shallowest puddle, which is sure to dry up, and thus
to cause the destruction of the whole brood. This carelessness is
all the more surprising when there are large pools or lakes in the
immediate vicinity, perhaps only one hundred yards to the other
side of the road. The Natterjack is, by the way, equally care-
less, while other toads and the tree-frogs are very circumspect.

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RANINAE

255

Both sexes can croak, and this sound is frequently produced
under water ; but there are no regular concerts, although many
collect in the same pond or spring, which is perhaps the only
suitable place for miles around. The male puts its arms aroimd
the chest of the female, behind her arms, and the embrace is so
firm that nothing will induce him to loosen his hold. The process
becomes an involuntary reflex-action, a cramp which may last for
days, or even for weeks, if sudden cold weather sets in, until
the female is ready to expel the eggs, an act which is quick and
soon over. The usual time of spawning in Middle Europe is the
month of March, earlier in warm, later in cold seasons; in
southern countries, February or even January, but in Norway
not until May. Although the males of thia species are not
more numerous than the females, and therefore should be able
to mate without much trouble, their ardour is so great that they
occasionally get hold not only of the wrong kind of frogs, but of
toads or even fishes, and, if taken off by force, they fasten on to any-
thing else, a log or on to your
own fingers. The eggs measure
2-3 mm. in diameter, are black
with a whitish spot on the lower
pole, number from 1000 to 2000,
and sink at first to the bottom.
Their gelatinous cover soon swells
to a large globe more than 10
cm. in width, and the whole
mass, as large as a man's head,
floats on the surface, often stained
with mud and other impurities.
During the cold weather which
often prevails in the spring, the
dark brown larvae are slow in
their development ; and provided
with rather large branched ex-
ternal gills and a well-developed
tail, they wriggle about in the
dissolving slime for three or four
weeks. Fischer Sigwart^ has timed
and measured them as follows. — The eggs were laid on the 10th of

1 Zool. Qart. 1885; p. 299.

Fio. 49. — Rana temjx^raria. Eight suc-
cessive stages in the developmeut
from the egg to the almost complete
Frog. X 1.

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2 5^ AXURA CHAP.

March. On the loth the larvae were 4 mm. long and began to
leave the e;L!gs. On the 19th ther measured, body 4, tail 9, total
13 mm.; on the 5th of April 10, 16, and 26 mm. respectively.
On the 13th of May they were 40 mm, long and the hind-Umbs
axipeared: the fore-legs burst through on the 25 th, when the
tadjioles had reached their greatest length, namely 45 mm., the
\xMly mea.suring 15 mm. On the 31st of May they left the
water, still provided with a rather long tail of 20 mm., the total
length being reduced to 35 mm. The larvae of this set developed
unusually fast, perhaps owing to artificial conditions. The whole
development is, however, mostly finished in three months, so that
the little stump-tailed baby-frogs swarm about well before mid-
summer, and have time enough to grow to the size of 20 mm. or
'I inch before they begin to hibernate in October.

In higher localities and in northern countries the tadpoles
are wjmetimes obliged to winter in the unfinished condition.

In spite of the unusually hot summer of 1899 I found
plenty of tadpoles on the 10th of September in the tarns of the
hills of Xorth Wales, 1500 feet above the level of the sea ; while
thousands of little frogs, with and without stumpy tails, were
hoj^ping about in the surrounding bogs. The water of these
tarns is always veiy cooL Cold and rainy weather set in by the
middle of the month, and on the 26th the tadpoles, all rather
small, measuring only 35 mm., with the four limbs developed, but
still with a broad fin on the taU, had all settled down under
stones at the bottom of the now very cold water, prepared for
hibernation. A few were taken home and kept in a glass vessel
with wat<»r, cool, but less so than that of their native tarns.
Within two days they lost the fins on their tails ; before the
end of a week they left the water, and crawled on to the moss,
and the tails were reduced to little stumpa By the 10th of
October the metamorphosis was complete, the little frogs measured
only 13 mm. in length and showed no desire to hibernate in the
genial atmosphere of the greenhouse.

This species has a very wide distribution. It ranges from the
west of Ireland to the islands of Saglialin and Yezzo, being found
everywhere in the enormous stretch of intervening countries,
practically the whole of Central and Northern Europe and the
middle belt of Asia. Its most northern extent is the whole of
Sweden and Norway. I have found it to the east of the Dovre-

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RANINAE 257

fjeld, at an elevation of 4000 feet, well-nigh the snow-line. In
conformity herewith it ascends the Italian Alps up to 10,000 feet.
The southern limit in Europe is the Cantabrian range and the
hilly province of Galicia. In the rest of the peninsula, in Italy
and Lombardy, Greece and Turkey, and on the Mediterranean
islands it is absent.

JB. arvcdis is often confounded with B. temporaria, as it differs
from the latter only by the following characters. The snout is
rather more pointed and is narrower ; the inner metatarsal tubercle
is large, compressed, and hard ; the dorso-lateral glandular folds
are more prominent and the belly is white and immaculate ; lastly,
it scarcely reaches 3 inches in length, a size which is not rarely
surpassed by the other species. There are also some differences
in habits. B. arvcdis prefers moist, boggy, open localities, and
does not ascend beyond 2000 feet in Central Europe. • It pairs
as a rule later in the spring and the eggs are smaller, only 1^-2
mm. in diameter ; they do not swell up so much, and the whole
mass does not float but remains at the bottom of the shallow
water. The coloration much resembles that of B, temporaria,
and is likewise subject to much variation, except that the pale
vertebral stripe is perhaps more conmion. This species is distri-
buted over the whole of Central Europe, Kussia, and Western
Siberia, south of the 60th degree of latitude, living side by side
with B. temporaria. Between the rivers Elbe and Rhine it
becomes decidedly rare, and the latter river is practically its
western boundary, while the Bavarian Alps and the Danube
form its southern limits.

B. agilis is still more frequently confounded with both the
two former species. It differs from either by the absence of
the two internal vocal sacs of the male, and by the decidedly
longer hind-limbs, the tibio-tarsal joint reaching often a little
beyond the tip of the snout. The inner metatarsal tubercle is
as prominent as in i?. arvalis. Total length up to 3 inches.
The prevailing colour of the upper parts is rather yellow or pink-
brown with few and small blackish spots; a A -shaped dark
mark on the neck is often present, and the large dark patch on
the temporal regions is always conspicuous. The under parts are
white, inclining to lemon yellow on the flanks and thighs. The
iris is golden yellow in its upper half, dark brown in the lower
half.

VOL. VIII S

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258 ANURA CHAP.

This species has a much smaller range than the first two : —
from France through Middle and Southern Germany, Switzerland,
and Lombardy to Hungary and Greece. The specific name refers
to the quick and long leaps of this pretty, or rather delicately-
coloured frog, which prefers woods and wooded glens to large
open places. Their voice differs much from the croak of the
common Brown Frog, and agrees with that of R, arvalis, which
is transcribed by Boulenger, who has kept them aUve, as a
rapidly uttered "co-co-co," or "cor-cor-cor." According to the same
authority, the pairing takes place as in R, temporaria, but is of
much shorter duration, the females usually resorting to the water
only at night and when quite ready to spawn. Specimens in
embrace are therefore seldom found in the daytime. The eggs
resemble those of E. temporaria in size, but they do not swell up
so much and they do not float.

These three species of European brown frogs, difficult enough
to distinguish, have of late been increased by three more, thanks
to the sagacity of Boulenger. These latter inhabit South Europe,
and the males all lack the internal vocal sacs.

R iberica has a very small range, namely the north-western
portion of the Iberian peninsula, from the Tagus northwards into
•Galicia, but south of the main extension of the Cantabrian
chain. The rest of the Peninsula south of these mountains has
no brown frogs, the only species of Rana being R esculenta.
R. iberica is rather local, being restricted to those hilly and
mountainous districts which are well watered. A favourite haunt
is the numerous streams in the wooded parts of the Serra Gerez,
the red, disintegrated granite of which suits this little, extremely
active, and reddish frog to perfection. The prevailing ground-
colour varies according to the district, from pale to dark reddish
or orange brown, with red specks and larger, dark brown spots,
which in some specimens begin with the A-shaped mark between
the shoulders. Dark spots on the flanks are very variable ; the
hind-limbs show the usual darker cross-bars, and the temporal
region has the conspicuous dark patch. The ground-colour of
the under parts is whitish, suffused with a pink tinge, and the
throat is much speckled with brown ; the toes are pink. The
size of this pretty frog amounts to 2 inches. The breeding time
is the month of March. When caught and squeezed they emit a
slight " co-co-co."

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RANINAE 2 59

Ji. graeca inhabits Italy and the Balkan peninsula from Bosina
to Morea, together with R. agilis, from which it is very difficult
to distinguish except that it is a little smaller, remaining below
2^ inches, and is generally more uniformly pale grey brown to
yellowish and pinkish brown above, with scarcely any, or only a
few, small dark specks on the back and limbs. The temporal
patch is likewise paler than in the other species. The flanks are
spotless, their colour gradually passing into the light buff of the
under parts, which are more or less marbled with grey. The iris
is golden, speckled with dark brown.

B. latastei of Lombardy and Northern Italy down to Florence
is the last of these closely allied frogs. Its affinities lie with
M. iherica and K agilis. The latter and R. latastei^ although
living side by side in the same locality, for instance near Turin,
are said not to interbreed. The voice is a rapidly uttered " keck-
keck-keck ;" the length remains below 2^ inches. The ground
colour is greyish or reddish brown with a dark brown A-shaped
mark between the shoulders, and a few red, orange, or blackish spots
on the back. The flanks are without definite dark spots. The
under parts are whitish, with a strong pink tinge, especially along
the middle of the throat and on the chest, the paler portions
being mottled with pale grey brown.

Perhaps the least unsatisfactory way of distinguishing between
R. agilis, R. graeca, and R. latastei (R. iherica need not be con-
founded with them on account of its distribution) is the size of
the tympanum, and its distance from the eye. The tympanum is
smallest in R. graeca, its diameter being about half that of the
eye and from ;| to the whole of its width distant from the eye.
In R, latastei the tympanum is a little larger, and about ^ to |^
its own width distant from the eye. R, agilis has the largest
tympanum, measuring about f of the diameter of the eye, and
the distance between the two organs amounts to only ^ of the
size of the tympanum.

Brown land-frogs of the R. temporaria group are found in most
countries of nearly the whole Periarctic and Oriental regions, and
by the time their races and varieties have been studied as
minutely as those of Europe are now being scrutinised, the
number of species will indeed be great.

R. silvatica is the chief representative in North America. It
closely resembles i?. agilis, but is smaller, only 2 inches in length,

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26o ANURA

and possesses a pair of internal vocal sacs. Its specific name
refers to its predilection for forests of oak, among the dried leaves
of which it conceals itself so successfully that it is discovered with
difficulty. B. japonica of Eastern Asia is almost indistinguishable
from this American species and from the European Ji, agilis,

B, opisthodon of the Solomon Islands has the vomerine teeth
in two oblique series entirely behind the level of the choanae.
The general shape of this large frog is stout, the type specimen
of the male measuring 78, that of the female 125 mm. = 5
inches. The upper surface of the female is covered with small,
fiat warts, that of the male is much smoother. The upper
parts are dark brown, while the under surface is brownish white.
The male has two internal vocal sacs.

This species is interesting as affording another instance of
shortened development, the whole metamorphosis being gone
through within the egg. Mr. Guppy, its discoverer, has supplied
the following notes : " During a descent from one of the peaks of
Faro Island I stopped at a stream some 400 feet above the sea,
where my native boys collected from the moist crevices of the
rocks close to the water a number of transparent gelatinous
balls, rather smaller than a marble. Each of these balls con-
tained a young frog, about 4 inches in length, apparently fully
developed, with very long hind-legs and short fore-legs, no tail,
and bearing on the sides of the body small tufts of what seemed
to be branchiae. On my rupturing the ball or egg in which
the little animal was doubled up the tiny frog took a mar-
vellous leap into its existence, and disappeared before I could
catch it. On reaching the ship an hour after, I found that some
of the eggs which I had put in a tin had been ruptured on the
way by the jolting, and the liberated frogs were leaping about
with great activity. On placing some of them in an open-
mouthed bottle, 8 inches long, I had to put the cover on, as they
kept leaping out."

Boulenger^ lias figured and further described the eggs and
young. The egg measures 6-10 mm. in diameter, and is a trans-
parent capsule in which the young frog is coiled up in the same way
as figured by Peters in Hy lodes martinicensis ; but none of the
specimens, which are in an advanced stage of development,
show anything of a tail. There are no gills, but on each side

* Trans. Zool. Soc, xii. 1884, p. 51.

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RANINAE 26 1

of the abdomen are several regular transverse folds, the func-
tion of which is probably that of breathing organs, like the
tail of Hylodes. The tip of the snout is furnished with a
small conical protuberance projecting slightly through the
delicate envelope of the egg, and evidently used to perforate
that envelope.

R. guppyi, likewise an inhabitant of the Solomon Islands, is
a giant among frogs. It was discovered by Mr. Guppy on the
Shortland Islands. The type-specimen measures 165 mm. s= 6^
inches in length ! The skin of the upper parts is covered with
minute warts, and forms a strong fold above the distinct, but
small, tympanum. General colour dark olive brown above,
dirty white below.

R. tigrina is a common species of Eastern Asia, including
the Malay Islands. On account of the strength of its voice,
and its size, which is said to reach 7 inches, it is called the
" Indian Bullfrog." Mainly aquatic, it has a strong cutaneous
fringe along the outer side of the fifth toe. The skin of the
back is thrown into longitudinal folds, and a strong fold
marks the upper border of the tympanum. The general
colour above is olive brown, with dark spots, often with a light
vertebral line ; the under parts are white. The male has a pair
of large external vocal sa<is.

R. gracilis has the same distribution, but it remains much
smaller, and the toes are only half, instead of fully, webbed.

R. catesbiana is now the settled name of the " Bullfrog "
of North America, the much more appropriate name of mngiens
having been sacrificed to the fetish of priority. The tympanum is
extraordinarily large, at least equal to the size of the eye, largest
in the male. The 'first finger does not extend beyond the second ;
the toes are connected by a broad web down to the ends, and
there is a small inner, but no outer, metatarsal tubercle. The
upper parts are olive brown, clouded with dark brown or blackish
spots ; the xmder parts are yellowish white, often marbled with
brown, especially on the throat. The iris is reddish, with an
outer yellow ring. The male possesses two internal vocal sacs.
Total length of adult specimens about 5 inches, but there are
giants on record 7 inches in length, while the stretched
hind -limbs measure another 9 or 10 inches. Its home ex-
tends over the whole of the United States, East of the

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262 ANURA CHAP.

Rocky Mountains, southwards into Mexico, northwards into
Canada.

According to Holbrook the Bullfrogs are solitary in their
habits, only collecting together in the breeding season, when
hundreds may be seen in the same small pond ; and then the
croak uttered by the males is so loud as to resemble the distant
roaring of a bull, and can be heard on still evenings at a distance
of half a mile. The voice is a hoarse bass " brwoom," playfully
translated into " more rum." " They cannot be said to abound,
but are found commonly enough sitting half immersed in
water, or on the banks of ponds, waiting for theu' prey. If
alarmed they hop suddenly into the water, but do not conceal
themselves at once, frequently skinuning along the surface for
several yards before they dive below." They are the most
aquatic of all the North American frogs, and Holbrook has
known specimens to live in wells for years, where they could
not rest a moment on solid ground above the water.

The Bullfrog is voracious, and takes almost anything that
lives or gets into his own pond — MoUusca, Crustacea, fishes and,
above all, frogs. There is no doubt that they drag down and
swallow a good many ducklings and the young of other
water-fowl, but certainly not the half-grown birds which have
a way of disappearing from the farms wherever there are
negroes and other farm-hands about. In turn the bullfrog has
sufficient enemies to keep its numbers down, in fishes, birds,
otters, and snakes, and, in the South, alligators. Although easily
kept and growing comparatively tame, they are dull, having
to be kept in solitary confinement on account of their
greediness, 'which knows no limits. Two of our specimens each
swallowed a full-grown Salamandra maculosa, and died within
the same night, probably not understanding the meaning of
the conspicuous black and yellow warning colours of the
European.

R. clamata s. fontiiialis, likewise an inhabitant of Eastern
North America, may be called a smaller edition of the Bullfrog, its
usual full-grown size being about 3^ inches. The tympanum is
conspicuously large, but the toes are webbed to a lesser extent,
and the skin forms a glandular fold which extends from
the shoulder in a curve to the flank. This species is partial
to the neighbourhood of running streams; it is said to be

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RANINAE 263

exceedingly timid, and to utter a short cry when disturbed and
making its enormous leaps.

Another North American relation is B. hcdecina s. palvstris,
frequenting the neighbourhood of ponds and rivers, very lively
and capable of jumping 8 to 10 feet. The tympanum is

Fio. 50.—Ra7ia clamata^ x |.

smaller than the eye, but there is the same glandular lateral
fold as in R. clamata. The vocal sacs are internal and decidedly
small.

B. esculenta. — The common Water-frog of nearly the whole
Palaearctic region is closely allied to the American Water-frogs
described above, and, like most of them, has the vomerine teeth
in two small oblique rows between the choanae and extending a
little beyond their posterior border. But the males have a paii-
of external vocal sac«. The tympanum is distinct, about two-
thirds the size of the eye. The first finger is slightly longer
than the second. The toes are entirely webl^ed. Besides the
usual subarticular phalangeal tubercles, the sole of the foot is
provided with two metatarsal tubercles, the outer of which is very
small, while the inner is much larger, althougli varying in size
from a soft oval to a long, curved, shovel-shaped structure. The
skin is smooth, except for a pair of prominent glandular folds
which extend from behind the eye along the dorso-lateral line.
The coloration varies considerably. Tlie upper parts are mostly
greenish brown, with black brown spots on the back, and larger
patches on the limbs. Most specimens have three lighter stripes
along the back, the middle one mostly green, the two lateral
bronzy brown and coinciding with the glandular folds. The
tympanum is brown, and there is occasionally a dark temporal
patch. The posterior aspect of the thighs is invariably

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264 ANURA

spotted with black and white or yellow, in opposition to the
K temporaria group, where these parts are never spotted.

The total length of this species varies much. Specimens 2^
inches in length are certainly mature, those of 4 inches are
unusually large, and Boulenger has received a giant from
Damascus, which measured 125 mm., or nearly 5 inches. The
females are larger than the males.

The variations in colour are not only local but also in-
dividual, moreover the colours are changeable. The ground-tint
ranges from dull brown through olive to bright green, the dark
spots being more or. less pronounced and numerous; the light
vertebral line is olive-yellowish, bright green, or altogether absent.

Those which inhabit waters with plentiful vegetation, like
water-lilies and other luxuriant plants, are generally prettier and
more vividly coloured than those which live in swamps and
ponds with dark mud, or where the prevailing vegetation has
a sombre aspect. Cold and dull, warm and sunny days also
influence the water-frogs, and those which have been kept in a
dark tank look very different from the bright assembly which
had been put in some weeks before.

Various attempts have been made at subdividing E, esculenta
of Linnaeus into sub-species, and Boulenger has now, after the
attentive study of an enormous material, arranged them in four
principal and recognisable races. The chief differences are the
relative length of the femur to the tibia and the size of the
metatarsal tubercles.

1. Var. ridihunda, Pallas. — The right and left heels overlap
each other when the thighs are stretched out at right angles to
the vertebral column, and the tibia is closely folded up against
tlie thighs. When stretched forwards, the heel reaches the eye
or even the tip of the snout. The inner metatarsal tubercle is
feebly developed, very small and blunt ; the outer tubercle is
absent.

That part of the thighs which is concealed by the legs when
tlie animal is at rest is whitish or pale greenish, marbled with
dark olive, or bronze, or of the latter colour with or without
small light spots. Xo trace of yellow is ever to be detected
on that region, nor at the axillae or on the groin. The vocal
sacs are strongly pigmented with black, when inflated they are
pale grey. The iris is a mixture of black and gold.

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RANINAE 265

This form or race has the widest distribution, namely, all
over Europe with the exception of England, the northern half of
France, the Ehine countries, Denmark, and Italy. Southwards
it extends from France through Spain and Portugal into the
Sahara, eastwards into Turkestan. It attains a larger size than
the others, but only in certain localities in various countries,
where circumstances favour its development. Eastern countries
produce the largest of all ; those of the Volga are said to be very
large. German physiological laboratories prefer those from the
Danube, from Bohemia, and from the lakes and broad expansions
of the Spree, to specimens from other localities.

2. Var. typica {escvlenta, Linnaeus). — The heels just meet, but
do not overlap. The inner metatarsal tubercle is strong, com-
pressed, and prominent. A small outer tubercle is present. The
heel reaches to the eye or a little further ; the hinder surface of
the thighs is " marbled with black, usually with more or less
bright yellow pigment " in the living specimens ; the vocal sacs
are white or feebly pigmented. This race inclines to rather
more green than the others, the males especially are often dark
grass-green, with scarcely any markings. The vertebral stripe
is then yellowish, and the lateral stripes almost golden. The
range extends over the whole of Central Europe and the kingdom
of Italy. Its northern limit is the southern end of Sweden. In
the greater portion of Germany, Poland, and Austria it overlaps
the var. ridihunda, with which it does not seem to pair, owing to
a difference in the time of spawning ; the var. typica being about a
fortnight later, and beginning to spawn wlien the other has finished.

3. Var. lessonae, Camerano. — Except that the inner tubercle
is stronger, while the outer one is near the vanishing point, and
that the fourth toe is proportionally longer, this variety is really
not distinguishable from the typical form, and Bouleiiger himself
confesses that the distinction is arbitrary. Tlie var. lessonae
seems to have a rather sporadic distribution. It has been found
in Piedmont and other parts of Italy, in Hungary and Transyl-
vania, near Vienna, Halle, Upper Bavaria, on the Rhine, near
Brussels, Paris, and what is of especial interest to us, in a few
places in the eastern counties of England.

According to Boulenger's " Notes on the Edible Frog in
England," ^ the individuals of R csculenta which live in Foulmire

1 P.Z.S. 1884, p. .^j73.

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266 ANURA

CHAP.

Fen in Cambridgeshire, near Stow Bedon, and between The t ford
and Scoulton in Norfolk, and are generally supposed to have
been introduced from France, belong to the Italian form of var.
lesso7iae. " It used to be found in Cambridgeshire, in Foulmire
Fen, where it was discovered in 1844 ; and Bell ^ assures us that
his father, who was a native of Cambridgeshire, liad noticed
the presence of these frogs many years before at Whaddon and
Fouhuire, where they were known from their loud croak as
' Whaddon organs * and ' Dutch nightingales/ The species was
afterwards rediscovered in Norfolk, between Thetford and Scoulton,
where it is now still very abundant, and from inquiries made by
Lord Walsingham, must have existed for the last seventy (80)
years at least. These frogs belong to the var. lessonae, and differ
widely (by the much stronger inner metatarsal tubercle) from
those found in a few other places in Norfolk, which are un-
doubtedly the descendants of a number imported from France .
and Belgium in 1837, 1841, and 1842, and turned loose in th^
Fens at Foulden and in the neighbourhood. ... Within the last
ten years large numbers of all the three forms have been imported
from Brussels, Berlin, and Italy, and liberated in various localities
in West Surrey and Hampshire. Berlin specimens of the var.
ridihunda have also been introduced in Bedfordshire, and Italian
ones in Oxfordshire." ^

Leaving aside the question whether the so-called var. lessonae
is merely sporadically developed out of the typical form, the
inquiry of the possible origin of the English specimens of the var.
lessonae is of special interest. Have they been introduced, as has
been suggested, from Lombardy, or are they the last lingering
descendants of native English frogs ? The suggestion as to their
Italian origin has naturally lost in value since similar specimens
have been found in Belgium and near Paris ; but we must remember
that there existed considerable intercourse between East Anglia
and the monks of Lombardy, who, to mention only one instance,
came regularly to tlie old Priory of Chesterton, near Cambridge,
in order to collect their rents. If the frogs were introduced by
them for culinary purposes into various suitable localities their'
descendants would remain as local as they, and as the undoubtedly
introduced French typical specimens actually are. On the other

^ BrUiih JiepHles, 2nd ed. 1849, p. 110.

- Boulenger, "Tailless Batracb. of Europe," pt. ii. p. 287, Bay Society, 1897.

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RANINAE 267

hand, if we assume the lessonae specimens to be the last living
descendants of English natives, it is inconceivable why they
should now be restricted to that eastern comer while there are
hundreds of other suitable places in England and Wales which, if
on the Continent, would be perfect paradises for Water-frogs. The
same vegetation, the same insects, the same climate, and — an
enormous advantage to the frogs — no storks.

These English specimens are "olive-brown or bronzy-brown
above, with black spots, strongly marked on the flanks, where a
light longitudinal area remains unspotted ; glandular folds lighter;
the sides of the head and the ground colour of the flanks are
sometimes green ; tympanum chestnut-brown ; a pale yellow or
pale green vertebral line, frequently edged with black ; the dark
cross-bands on the limbs usually very irregular, sometimes absent ;
lower surfaces more or less profusely spotted with blackish ; iris
golden. Length of a male from Stow Bedon, 64 mm. or 2c^
inches; of a female, 78 mm. or 3 inches."^

4. Var. chinerms, Osb. — Distinguished by short glandular
folds along the back, in addition to the long dorso-lateral pair.
The metatarsal tubercle is large and shovel-shaped. Distribution
from Corea and Japan to Siam.

All these Water-frogs are decidedly aquatic. They make short

excursions on land when their homes are dried up, but as a rule

they remain in the lake, pond, river, morass, or ditch in which

they were born. Their favourite resorts are the broad floating

leaves of water-plants, for instance water-lilies, or a prominent

stone, a tussock of grass, or the banks of their homes, where

they sit motionless, basking for hours in a half- erect, alert

position, watching for insects and other small fry, which are

secured by a jump, and then lapped up. Sunshine is sure

to bring them out, and on our approach they make straight

for the water, either by one tremendous leap or with quick

bounds, but without the slightest hesitation or stopping on the

way. With folded arms they take a header, swim, with the

arms still folded, for some distance under water, and conceal

, themselves in the mud, between stones, or in the vegetation. We

/ perhaps have not seen them at all, whilst their watchful eyes and

i keen ears have noticed our approach, and the pond might appear

. iminhabited if we had not heard the plumping noise. If we

\ ' Boulenger, op. cit. y. 278.

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268

ANURA

CHAP.

^>P^

keep quite still, and they have not been disturbed previously, one
after another will wriggle out of the mud, rise slowly to the
surface under cover of the plants, and, without causing a ripple,
rise just enough for the prominent eyes and the nose to clear the
surface. Then one scrambles partly on to a leaf, but the sight
of the huge human figure strikes him as uncanny, as it certainly
does not belong to the scenery, and he doubles back, the broadly-
webbed feet making a little splash.
But another appears, jumps on to a
leaf in the middle, or at the farther end
of the pond, settles down, and utters a
little jcroak, somewhat like " ooaar,"
and soon the whole company appear
one after another, each, taking up its
favourite position. After all, their ob-
serving powers cannot be very great.
If we ourselves. keep still we may wield
a rod and fish for them. There is no
need of a hook, a piece of red cloth
tied to the end of the line and
skimmed over the water causes a lively
commotion. The new bait is noticed
at once, and arouses their curiosity;
several jump at it, and the one which
swallows the bait can be lifted out
before it has time to let go. However,
this is after all poor sport ; the game
is too eager. When a boy I have often
ciiught them with a noose of slender
wire at the end of a long hazel rod.
They do not mind the rod at all, their
attention being fixed on the person ;
they allow the noose to be slipped over their heads, and a sudden
jerk secures the ciiptive. In this way they can be singled out
individually. Old frogs are more wary and experienced than
the younger members; they take up safer positions, and by
their sudden plunges give the alarm.

The males are great musicians, singing for sheer enjoyment
not only during the pairing time, but throughout the months
of June and July. "Warm moonlit nights are the favourite times

Fig. 61. — Rana esculenta. x 1.
Three stui^es of the movement
of the tongue.

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RANINAE 269

for the concert, which takes place in the water, beginning at
sunset, and continuing until the early dawn. A few individuals
here and there utter a single note, " gwarr, 00-arr," or " coarx,"
but these are only preliminaries. The precentor — the country-
folk in Xorth Germany firmly believe that in each pond one old
male holds the dignified position of choir-master — begins with a
sharp-sounding " brekeke," and this is the signal for all the others
to chime in with the same notes, varied with all sorts of other
sounds, bass, tenor, and alto, each performer filling its resounding
vocal sacs to bursting size, and these bags then look as if they
acted as floats. When there are several hundred of these
sociable creatures, the din is continuous, and
may be heard more than a mile off. There
can be too much of this, just as there can be
too many nightingales ; and a well-stocked
pond in the neighbourhood may become a
perfect nuisance. There are accounts of
servants having been employed in the
Middle Ages for the sole purpose of keep- Fio. 52.— iia/ta esatUnta.

., .1 I T_ i.- 4.1 J Male with inflated ex-

mg the noise down by beating the pond, temai vocal sacs. xi.
throwing stones into the water, or otherwise
disturbing the frogs. Sometimes more vigorous and lasting
measures seem to have been taken ; the monks exorcised them
in order not to be disturbed in their vigils. Near the former
monastery of Chorin, in the province of Brandenburg, the frogs
have still the reputation of keeping very quiet on account of
some powerful abbot who threatened them with awful con-
sequences if they did not forego their concerts.

The length of life which these frogs can attain is quite
unknown. They do not reach maturity until the fourth or fifth
year, but this is long before they stop growing, and it is no
exaggeration to say that few, if any, frogs die of old age, since
they have so many enemies. The stork is their king in the
fable, and his daily visits to his realm strike dire distress
amongst his subjects, which soon learn to know his conspicuous
white and black garb, and seek imperfect safety at the bottom
of shallow ponds and ditches, not too deep for the long-legged
and long-billed despot. Numbers are taken by birds of prey ;
snakes and tortoises hunt them up in the water, and they are
good bait for pike and other voracious fishes. The specific

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2 70 ANURA CHAP.

name esculenta needs no comment, and this species is as much a
martyr to science as the brown Grass-frog. The destroyers of
tadpoles and young frogs are unlimited. In their turn the
frogs themselves, especially the old ones, are very rapacious, and
eat any living creature they can master, — insects, worms and
snails, other frogs, especially the brown kind, and the young
brood of fishes.

Recently caught Water-frogs are wild beyond description,
much more so than the Grass-frog, but even they calm down
after some time, learn to know their keeper, and allow him to
handle them without trying to commit suicide. by jumping on
to, into, and down anything. However, they do not thrive well
in captivity, and it is rare that they can be induced to breed,
unless their enforced new home affords them ample freedom,
and plenty of water and fresh air.

The Water-frogs appear in Grermany rather late in the year,
not before the middle of April, first the younger, then the adult
members. In Southern Europe they show themselves earlier,
and still further south they do not hibernate at all. The breed-
ing season begins in Germany towards the end of May and
continues well into June, the var. ridihunda beginning mostly a
fortnight earlier. The male clasps the female under the arms,
throwing its own round her breast, the nuptial grey excrescences
on his inner fingers pressing against her skin, the palms being
turned outwards. The embrace does not last long, rarely
extending over a few days. The eggs, to the astonishing number
of 5000 to 10,000 in full-grown specimens, are expelled in
several masses, which sink down and remain at the bottom.
The eggs measure only 1*5 mm. and are yellowish-grey above,
pale yellow below ; their gelatinous cover swells to 7-8 mm. in
width. The embryo escapes on the fifth or sixth day as a very
small larva, in which, however, the mouth, eyes, and beginnings
of the external gills are already discernible. At the age of two
weeks the gills have shrunk away, the left-sided " spiracle " is
completed, and the well -tailed tadpoles, olive brown above,
yellowish white below, still hang with their suckers on to plants
and stones, or lie at the bottom, nibbling away at any rotting
animal matter or scraping off the green algae.

It may here be mentioned that small tadpoles of any kind
can with advantage be used as cleaners of delicate and small

L*^V.

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RANINAE 271

skeletons. The object is put into a vessel, and the tadpoles
will soon nibble and rasp away all the edible portions, leaving
the skeletal framework beautifully cleaned. But they require
attention lest they rasp away the cartilage.

The tadpole stage lasts three to four months ; but cold, absence
of sunshine, and scarcity of food delay the metamorphosis well into
the end of summer, or force them to hibernate in the unfinished
condition. They are very gregarious, and when the tadpoles of
several families combine, they make imposing shows. By the
time that their hind-limbs begin to sprout, they frequently
combine into large shoals, and instead of always feeding they
swim about in their tens of thousands, all moving in the same
direction, and making almost regular evolutions. Mill-ponds
with steep banks are good places for watching these peculiar
habits. The tadpoles reach a considerable size, the total length
averaging 2^ inches, or some 60 mm. the tail taking up |^ of
the whole length. Specimens which measure more than 3 inches
are rare. The baby-frogs hop on land while still provided with
a stumpy tail ; when this is resorbed the little creature is
scarcely half-an-inch long, and for the rest of the available
season leads a rather more terrestrial life than ever after.

Ex Africa semper aliquid novi I Quite recently Boulenger
has received a consignment of Anura from the French Congo,
amongst which were several new, remarkable genera, notably
Trichohatrachus and Gampsosteonyx. Both are true Eanidae.
PupU vertical, with vomerine teeth. Omosternum with a bony
style. The outer metatarsals are bound together. In Trichu-
batrachvs robustus the toes are webbed, and both sexes have the
flanks and corresponding portions of the thighs covered with
numerous darkly pigmented, filamentous, cutaneous excrescences ;
these are several millimeters in length, giving the flanks and
thighs a " hairy '* appearance. Mr. F. F. Laidlaw has examined
these structures. Their most remarkable feature is the presence
in them of a great number of ordinary flask-shaped cutaneous
glands, whilst such glands are scarce on the surrounding skin.
They diflfer in no way from those seen in sections of the skin of
the Common Frog. The fibrous connective tissue is dense and
vascular; the pigment - cells are most plentiful at the base.
Contrary to expectation no nerve-endings were found in these

filamenta

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272 ANURA CHAP.

Gampsosteonyx has free toes. The terminal joints of the
digits stand out beyond the skin, and end in sharp, bony claws,
like those of a cat.

Sub-Fam. 3. Dendrobatinae. — About one dozen arboreal
little frogs have been separated from the Baninae proper on
account of the entire absence of teeth. This mere loss of teeth,
and the geographical distribution suggest that these frogs do
not form a natural group, but have been developed independently
from other Eanidae, the Neotropical Dendrohates from some like-
wise Neotropical genus like Prostherapis, the Malagasy Mantella
from an African form like Megalixalus,

The sacral diapophyses are cylindrical. The omo- and
meta-sternum are well developed. The fingers and toes are free,
their terminal phalanges are T-shaped and carry regular, round,
adhesive discs. The tympanum is distinct, although sometimes,
in Dendrohates, very small. The pupil is horizontal.

Dendrohates. — The tongue is elongate, entire and free behind.
The omosternum has a weak, semi-ossified style, but the meta-
sternum remains cartilaginous. The males have a subgular
vocal sac. Seven closely-allied species inhabit tropical America.

D. tinctorius. — This pretty little species, scarcely 1^ inch
in length, is quite smooth, varies much in coloration, and forms
local races to a certain extent. Some are quite bkck, others are
grey above, black on the sides and under parts ; or they are grey
with large black patches. A fourth variety is black above with
several white or pink longitudinal stripes, while the \mder parts
are grey, spotted with black. In others, again, the ground-
colour is black, with white stripes and spots above, marbled
below. But this enumeration does not exhaust the list, since
living specimens are sometimes much more conspicuously
coloured, some being black with large patches of saturated
yellow on the head and back, while the limbs are orange red
and black. This species has a wide range, from Panama to
Ecuador and to the mouth of the Amazon. It owes its specific
name to the peculiar use made by man of the strongly poisonous
secretion of the tiny glands of the otherwise smooth skin.
Other species are doubtless employed in the same way. The
poison is mainly used for " dyeing " the green Amazon-parrots.
This is done as follows : — The green and blue feathers on the
head and neck, or other parts, according to the fancy of the

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RANIDAE — DENDROBATINAE

273

operator, are plucked out, and these places are rubbed with the
poison, often simply with the living frog, certainly not with its
blood, as is sometimes asserted. This operation may be repeated
when the new, young feathers begin to bud. The result is that
these appear yellow instead of green, and since the Brazilians,
and to a certain extent the Portuguese, are rather 'partial to
these artificially -produced freaks or "contrafeitos" as they
call them, the industry is kept up. That the poison is also
used for arrows has been mentioned on p. 38.

Fig. 53. — DendrdbcUea tinctorius^ three colour- variations, x 1.

D, trimttatus, chiefly in Northern Brazil, has the first finger
slightly longer than the second. It likewise varies considerably
in its coloration, being either quite black, or spotted with white
and brown, or with a whitish forehead and several white patches
on the back and hind - limbs. D, typographus of Central
America is vermilion red, with small dark marks on the bjwik ;
the legs are black.

The various species of Dendrohates take remarkable care of
their young. D. hraccatus lives in Brazil in " varzeas," i.e.
moist but waterless places, and carries its tadpoles on its back,
to which they are attached by a peculiar secretion. The same is
said to be true of D. trivittatus, which sits down in a drying-up
puddle, lets the little tadpoles, when they are only 6-7 mm. long,
fasten themselves on, and conveys them to a safer locality, where
the water is calculated not to evaporate before -the metamorphosis
is completed.

VOL. VIII T

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274 ANURA CHAP. VI

Mantdla. — Both omo- and meta-sternum possess a bony style.
The tongue is free and distinctly mitred or cut out behind.
The skin is very granular. Several species, in Madagascar,
were formerly put into the same genus as the American forms,
until Boulenger established the genus Mantdla for them. The
coloration is strikingly pretty. M. rfiadagascariensis is a rare
instance of diflference in colour between the two sexes. The
male is bluish black, with light blue spots on the belly, while
the thighs and the inner sides of the legs are beautifully red.
The female is deep black, with a light green spot at the base and
in front of the limbs ; the rest is coloured like the male.

Cardioglossa gracilis, quite recently discovered at the Gaboon,
has likewise to be added to the Dendrobatinae, on account of the
absence of teeth. It is a small, slender, arboreal frog, bearing
an unmistakable resemblance to the other genera by its general
appearance and conspicuous, contrasting coloration of black and
white.

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PART II
REPTILIA

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" Gada uno es como Dios le hizo,
y aun peor muchas vezes. "

" We are all as God made us
and many even worse. '*

Sancho Panza,

Don Quixote.

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CHAPTER VII

REPTILIA

DEnNITION AND CHARACTERS POSITION OF THE CLASS REPTILIA

IN THE PHYLUM VERTEBRATA CLASSIHCATION SKULL AND

VERTEBRAE.

The recent Reptiles comprise, broadly speaking, the Crocodiles,
Tortoises, Lizards, and Snakes. They are the only Vertebrates
which are cold-blooded, breathe by lungs, and have a median
occipital condyle. Another equally sufficient diagnosis is the
following: — Tetrapoda, with a median occipital condyle, with
nucleated red blood -corpuscles, and with complete right and
left functional aortic arches. A still shorter diagnosis is : —
Monocondylia with a scaly skin.

If our diagnosis is to include the fossil Reptiles we have not
only to discard the characters drawn from the soft parts as
unavailable, but we are forced to treat the condition of the
occipital condyle with caution, since there exist, or must have
existed, transitional stages between Reptiles and Amphibia and
Mammals ; and the winged class Pterosauria does not permit us
to use the wings as a differential character for the Birds. In
fact, while the Reptilia are sufficiently separated from the
Amphibia by their absolutely gastrocentrous vertebrae, it is
difficult to distinguish them as a class from the Birds ; hence the
term Sauropsida, which is intended to indicate the close relation-
ship of the Reptiles to the Birds in opposition to the Mammalia,
and to the Ichthyopsida or Amphibia and Fishes. However, the
Reptilia take up a very central position in the evolution of the
main classes of the Vertebrata. On the one hand, there is not
the slightest doubt that they are evolved from some branch of

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2 78 REPTILIA CHAP.

the Stegocephali, whilst on the other hand the Reptiles, prob-
ably through some branch of the Theromorpha, have given rise
to the Mammals; some other Beptilian branch, at present
unknown, has blossomed out into the Birds.

Principal Oharacters of the Beptilia.

1. The vertebrae are gastrocentroiia.

2. The skull articulates with the atlas by one condyle, which is formed

mainly by the basioccipital.

3. The mandible consLsts of many pieces and articulates with the cranium

through the quadrate bones.

4. There is an auditory columellar apparatus fitting into the fenestra

ovalis.

5. The hmbs are of the tetrapodous, pentadactyle type.

6. There is an intracranial hypoglossal nerve.

7. The ribs form a true sternum.

8. The ilio-sacral connexion is post-acetabular.

9. The skin is covered (a) with scales, but (6) neither with feathers nor

with hairs ; and there is a great paucity of glanda

10. Reptiles are poikilothermoiia

11. The red blood -corpuscles are nucleated, biconvex, and oval.

12. The heart is divided into two atria and an imperfectly divided

ventricle. It has no conus, but semilunar valves exist at the base
of the tripartite aortic trunk.

13. The right and left aortic arch are complete and remain functional

14. Respiration is effected by lungs ; and gills are entirely absent, even

during embryonic life.

15. Lateral sense-organs are absent

16. The kidneys have no nephrostomes. Each kidney has one separate

ureter.

17. There is always a typical cloaca.

18. The eggs are meroblastic.

19. Fertilisation is internal, and is effected, with the single exception of

Sphenodon, by means of male copulatory organa

20. An amnion and an allantois are formed during development
Numbers 1, 2, 6, 7, 8, 14, 16, 18, 20 separate the Reptiles from

the Ami)hibia. Cf. also pp. 4 and 5.
Numbers 9 (6), 10, 12, and 13 separate them from the Birds and

Mammals.
Numl)ers 3, 8, and 11 separate them from the Mammak.

The evolation of the classification of the Reptiles has

to a certain extent been already treated on pp. 7-9. For a
long time only Chelonia or Tortoises, Ophidia or Snakes, and
Saurii were recognised as their principal divisions. Then the
Crocodiles were separated from the Lizards ; later the Coeciliae
were removed from the Snakes and referred to the Amphibia,

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vii CHARACTERS — CLASSIFICATION 2/9

and ultimately Sjphenodon was recognised as deserving a separate
position, equal in rank to the other groups. Stannius showed
that the Crocodiles and Tortoises are relatively near allies in
opposition to the likewise closely allied Lizards and Snakes
(^Sjphenodon was then unknown), and he expressed this by the
term Monimostylica, or creatures with fixed quadrate bones, for
the former, and Streptostylica, creatures with movable quadrates,
for the latter combination. The fossil Eeptiles were hardly
allowed proper places in the system. In various zoological text-
books they were, or are even now, treated as inconvenient, out-
lying, or supernumerary members. A long time elapsed before,
thanks to the labours of H. von Meyer, Owen, Huxley, Marsh,
Cope, Zittel, and Seeley, it was recognised that the extinct groups
form the preponderant mass of Eeptiles, and that it is the recent
groups which, in spite of the bewildering number of species of
Lizards and Snakes, are the comparatively few and much-reduced
members of a once flourishing class. With the exception of the
Lizards and Snakes, which are on the ascending branch, the
modern Sjphenodon, the Crocodiles and the Tortoises are a mere
fraction, comprising a few survivals of richly-developed groups,
while all the others, the overwhelming majority, have died out.
The classification adopted in this volume is as follows : —

Class Eeptilia.

Sub-Class I. Proreptilia.
„ II. Proeaaria.

Orders : Microeauri, Prosauri.
„ III. Theromorpha. Orders: Pareiasauri, Theriodontia,

Anomodontia, Placodontia.
,, IV. Chelonia. Orders : Athecae, Thecophora.

„ V. Dinosauria. Orders : Sauropoda, Theropoda, Orthopoda,

Ceratopsia.
„ VL Crocodilia. Orders : Pseudosiichia, Parasuchia, Eusuchia.

„ VII. Plesiosauria. Orders : Nothosauri, Plesiosauri.

„ VIII. Ichthyosauria.
„ IX- Pteroeauria.

„ X. Pythonomorpha. Orders : Dolichosauri, Mosasauri.

„ XI. Sauria. Orders ; Lacertilia, Ophidia.

The eleven principal groups are here called " sub-classes " to
emphasise the undeniable fact that these Keptilian groups are of
much greater morphological value than those which are most
generally called " Orders " in the Mammalia, that class which we
consider as the standard or model of classificatory units. The

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Fl«. 54. — Diagrams of skulls, showing especially the composition of the bony arches of
the orbito-temporal region.

A, C, D, E, Theromorpha. A, Elgin in, p. 305 ; C, Cynognathus, p. 306 ; D,
(jirrdoniaj ]). 310 ; E, hinpuxion, p. 310.

B, O, Prosauria. B, Sphenodon, p. 294 ; O, PalaeoIiaUeriay i>. 291.
F, Crocodilia, p. 434.

H, I, K, Chelonia, p. 316. H, Chdydra, p. 338 ; I, Chrysemys, p. 346 : K,
Cistvdo, p. 361.

E^ Epiotic ; /', frontal ; JTy infratem]X)ral fossa ; J, jugal, shaded vertically ;
Z, lacrymal ; My maxillary ; iN", nasal groove ; iVa, nasal bone ; 0, orbit ;
P, parietal ; Po, poster bital, dotted ; J% post-frontal ; Pm^ premaxillary ;
Pr, prefrontal ; Ptgj pterygoid ; Q, quadrate ; Qj\ quadrato-jugal ; ^Sh, supra -
occipital ; Sq, squamosal, shaded obliquely ; St (in B-E), supratemporal fossa ;
^ (in A), Supratemporal bone.

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^ !^r

Fig. 55. — Diagrams of skulls, showing especially the composition of the bony arches of
the orbito-temporal region.

L, Ptthonomorpha. Clidastes, p. 490.

H, N, O, Lacertiua, p. 496. M, VarayiiiSy p. 543 ; N, UromastiXf p. 524 : 0,

fjocertaj p. 550.
P, IcHTHTOSAURiA, p. 479. /chthi/osaunis, p. 488.
Q, Pterosauria, p. 484. Dimorphudon, p. 486.

B, Ates, generalised, for comparison.

8, Mammaua, generalised, for comparison.
T Ophidia, p. 581.

C, Condyle of mandible ; Col, columella craiiii ; t\ frontal ; /, interparietal or
pineal foramen ; J. Ay Inner angle of mandible ; J, jugal, shaded vertically ;
Z, laerymal ; J/, maxillary ; N^ nasal groove ; Sa, nasal bone ; 0, orbit ;
Oi, preorbital fossa ; P, parietal ; /y, postfrontal ; Pvh, premaxillary ; Pr, pre-
frontal ; Ptg^ pterygoid; Q, quadrate ; C^", quadrato -jugal ; »Sr/, squamosal,
shaded obliquely ; St, supratemporal bone.

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282

REPTILIA

CHAP.

families cannot well be changed, and terms like super-families
and super-orders are sometimes resorted to by those who do not
like to look stern facts in the face.

The sequence of the groups, although arranged as much as
possible in ascending order, is of necessity as unnatural as that of
the maps in an atlas. We cannot yet construct a satisfactorj-
phyletic tree of the Reptiles. The Proreptilia connect them witli
the Amphibia. Next follow the Prosauria with Sphenodon among
the Prosauri as the key to most other groups. Then follow
the Theromorpha, and it is probable that from various branches
of these have arisen the Chelonia, Dinosauria, Crocodilia, and
Plesiosauria. The descent of the Ichthyosauria is very pro-
blematic. The same applies to the Pterosauria and to the
Pythonomorpha, but it is possible that they, together with the
Sauria, are connected with the Prosauria.

With all reserve these hypothetical afl&nities may be expressed

by the following diagram : —

Dphidia

Ichthyosauria Plesiosauria

Chelonia

Sauria

irosaana
Pythonomorpha

Prosauria

Proreptilia

The eleven sub -classes of the Reptilia present so many
important differences that it is not advisable to give here a further
general account of their structure. The diagrammatic figures
A to T on pp. 280, 281, representing various types of skulls,
are intended to explain their chief modifications, all referable to
Proreptilian arid to certain Theromorphous conditions. One of
the most important features is that the mandible, which is always
composed of many pieces (cf Fig. 142, p. 550), is invariably
carried by the quadrate bone. Diagrams of the generalised
skulls of a Bird and a Mammal have been added for comparison.

As mentioned on p. 278 the vertebrae of the E^ptilia and
those of all other Amniota are gastrocentrous ; that is to say

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VII

VERTEBRAL COLUMN

283

the centra or bodies of the vertebrae are formed by the pairs
of interventralia, while the basiventralia are reduced, persisting

-^^^

Fig.

56. — Composition of vertebrae of Reptiles, illustrated by the first and second
cervical vertebrae. (1) Atlas (first cervical) and axis (second) vertebra of Ci-oco-
dilu8. (2) Atlas and axis of MetriorhynchuSy a Jurassic Crocodile. (3) Analysis
of the first two cervical vertebrae of a Crocodile ; 2, second basiventral complex
or "intercentrum" continued upwards into the meniscus or intervertebral pad.
(4) Diagram of the fundamental composition of a Reptilian vertebra ; compare this
and (6) with Fig. 1 (8 and 9) on p. 13. (5) The first three cervical vertebrae of
Sphenodon, (6) Trunk- vertebrae of Kryops^ a Permian Proreptil^ ; typitffelly temno-
spondylous ; cp, articular facet of the capitulum of a rib. (7) The complete atlas
of an adult TrUmyx hurum ; the second basiventral (intercentrum) is attached to the
posterior end of the first centrum, which, not being fused with the second centrum,
• is not yet an odontoid process. (8) The complete atlas of an adult Trionyx gawje-
ticus ; still typically temnospondylous. (9) The first and second cervical vertebrae
of an adult Platemys. (10) The complete atlas of a Chelys Jimbriata, Az, Anterior
zygapophysis ; B,D, basidorsal ; B, F, basiventral ; C,, Ca, C3, first, second, and third
centra, formed by the interventralia ; Cp^, C/7^ articular facets of the capitular
portions of the first and second ribs ; /. V, interventral ; N\^ N^, N^, first, second,
and third neural arch, formed by basidorsalia {B.D) ; Od, odontoid process = first
ceutnim ; P&, posterior zygapophysis ; /?i, R^ ribs ; Sp, detached spinous process
of the first neural arch : tj, t^ tulxjrcular attachments of the first and second ribs ;
1, 2, 3, 4, "intercentra " = basiventrals ; /, 77, 777, position of the exit of the first,
second, and third spinal nerves.

either as so-called intercentra or wedge -bones, or as inter-
vertebral pads, or disappearing altogether. At the earlier
stages of development the gastrocentrous vertebrae behave in the

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284 REPTILIA CHAP. VII

same way as that described on p. 12 (Fig. 1), except that
the interdorsal elements are suppressed from the beginning. Ii
the remaining three pairs of constituent elements of each verte-
bra (the basidorsalia, forming the neural arch ; the interventralia,
forming the body or centrum ; and the basiventralia) remain
separate, the vertebrae are called temnospondylcyus (t6/lm/g), I cut,
oTTovivKo^;, a vertebra). If the neural arches and the centra are
suturally united or are fused with each other, the vertebrae are
called stereospondylcms (ore/aed?, solid). In many Anmiota
the atlas or first vertebra remains in a relatively primitive,
embryonic condition, and is temnospondylous but for the usual
modification that its centrum becomes attached to that of the
second vertebra, and forms the odontoid process of the latter.
The composition of gastrocentrous vertebrae (cf. p. 282) is best
illustrated by the first and second cervical vertebrae of the
Crocodile (Fig. 56, 3, p. 283).

Concerning geographical distribution, even a cursory study
shows that the sub-classes have come into existence at very
different geological periods, and have each followed their own
lines of dispersal.

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CHAPTER VIII

proreptiua prosauria theromorpha

Sub-Class L—PROREPTILIA,

Permian Temnospondj/lous Reptiles with well-developed limbs
and girdles of the terrestrial type.

The two genera Eryops and Cricotus of the North-American
Permian formation had until recently^ been relegated to the
Stegocephali. By grouping them and their nearest allies
together as Proreptilia it is intended to indicate that they are
the lowest known Beptiles and that they probably link this class
to the Amphibia. The superficial resemblance of their tri- or
bi-partite vertebrae, and their occurrence in the Lower Permian,
have caused the error of classing them with the Stegocephali, but
the composition of their typically gastrocentrous vertebrae leaves
no doubt as to their affinities. After all, we feel certain that
Septiles have arisen from Stegocephalous Amphibia, and it is in
the Lower Permian, exactly where these debatable creatures lived
side by side with Stegocephali, undoubtedly likewise temnospon-
dylous, that the change from Amphibia into Eeptiles seems to have
taken place. Both are referable to Amphibia with quadripartite
vertebrae. The condition of the occipital condyles determines
nothing. This greatly exaggerated character has lost in import-
ance since we have known the condylar modifications of the Thero-
morpha ; moreover, Cricotvs itself seems to have possessed a single
condyle. We should even expect the Proreptilia to present
many Stegocephalous inheritances, for instance the condition of
the skull roofed in by dermal bones, a ventral dermal armour, a
very complete pectoral arch still without a sternum, and only one
sacral vertebra.

* Phil, Trans, clxxxvii. 1896, B. p. 23.

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2 86 PROREPTIUA

CHAP.

Until more genera are better known than they are now, it is
premature to divide the present sub-class into ordera

Eryops, with several species in Texas and New Mexico. E.
megacephalns is the most abundant and the largest species, its
broad and flattened skull measuring more than 18 inches in length
and 12 in width. With the exception of the nostrils and the
small orbits, the skull is entirely encased in bone, with a rough,
pitted surface, but without any distinguishable sutures. The
absence of mucous canals, so common in the Stegocephali, is
worthy of note. The quadrates extend obliquely outwards and
backwards, so that the joint with the mandible lies in a plane
behind the occiput. The mandibles are devoid of a projecting
angular process. The teeth are numerous,
small, and pointed. The vertebrae are
typically temnospondylous, consisting each
of three pairs of separately ossified pieces,
which, although closely packed together.
Fig. 57.— Trunk vertebrae of are not suturally connected. The neural
^()p* (cf. Fig. 66, 4, p. arches possess high spinous processes, they
of thecapituiumofarib. articulate by short and broad zygapo-
physes and are, with their triangular
bases, wedged in between the two ventral pieces, the posterior of
which (the united interventralia) is in broader contact with the
neural arch and lies behind it ; the anterior piece (the united
basiventrals) appear as typical, but large, interoentra, and bear on
their posterior, dorsal margin the facets for the ribs. The latter
are short, but are broad at their proximal ends, which are not
bifurcated ; they extend their articulation from the " intercentra "
upon the short lateral processes of the neural arches. The tail is
short and ends in a pointed coccyx, owing to fusion of the last
vertebrae.

The pubes and ischia are heavy, the former flattened and
broadened out. The limbs are of an almost ideal pentadactyloid
type ; strongly developed for terrestrial locomotion. The ulna
possesses a large olecranon. The carpus consists of ten separate
pieces, ulnare, intermedium, radiale, two centralia and five distal
carpalia. The latter support only four metacarpals and fingers,
the second finger being completely abolished, an explanation
suggested by Cope and corroborated by Emery.^

^ Anat. Anz, xix. 1897, p. 201.

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PROREPTILIA 287

CricotvSy with several species in Texas and Illinois. C. hetero-
clitus was perhaps 10 feet long and probably aquatic. The
skull has a long, narrow, depressed snout, the margins over-
hanging those of the lower jaw ; its surface is encased in dermal
bones, most of which still show sutures, so that for instance
postfrontals, postorbitals, supratemporals and squamosals can be
distinguished ; all these are in contact with the long parietals
and with the quadrato-jugal arch, covering the temporal region ;
but the supratemporals have a free projecting border, like the
squamosals of the crocodiles. According to Cope's description
the basioccipital is connected with the first vertebra by an
undivided discoid " intercentrum," probably the true centrum, while
the first basiventral mass, which would be, if independent, the
first true intercentrum, is more probably connected with the first
neural arch, thus constituting the ring of the atlas.

The vertebrae are still temnospondylous, but no longer
tripartite. The neural arch is fused with the interventralia into
one mass, which carries the capitula and tubercula of the ribs,
while the united basiventrals still remain as separate intercentral
wedges. In the tail these wedges carry chevron-bones, and are
enlarged into thick almost complete discs, or rather rings, while
the whole vertebral column is still perforated, as also in Eryops,
by the chorda dorsalis. The tail is long. The digits are devoid
of claws.

Eemains of dermal armour exist on the throat in the shape
of several large gular plates, while the whole belly is covered
with many closely packed bony scales, which are arranged in
chevron-shaped transverse rows.

Probably several other genera of American Permian and also
of European Permian strata will, when better known and criti-
cally examined, have to be referred to the Proreptilia. Thus for
instance the European Melosaurvs may have afl&nities with
Eryops, while Diplovertebron of Bohemia seems to be allied to
Cricotus. The diflSculty of division will lie with those Lower
Permian Amphibia which, like Archegosaurus; Euchirosaurus,
Actinodon, possess tripartite vertebrae, which at first sight are
strikingly like those of Eryops, But the tail-vertebrae permit
of no mistake, and since these are quadripartite in Archegosaurus,
Chelydosaurtis, and Sphenosaurus, these genera are safely to be
classed with the Amphibia, unless, indeed, for mere argument's

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288 PROSAURIA

sake, it be assumed that the intercentral discs of JHplovertebroyi
and Cricotvs are formed by the fusion of Amphibian interdorsals
with interventrals. Anyhow, simply to state that the tripartite
vertebrae of Eryops are the same as those of AcHjwdon, would be
as convincing as saying that the English and French flags are
essentially the same, both containing the same colours, but one
is white, red, and blue, the other blue, white, and red. Tripartite
Amphibian vertebrae are composed of basidorsals + basiventrals +
interdorsals, those of Septiles are made up of basidorsals +
basiventrals + interventrals. (Cf. Fig. 56, p. 283, and Fig. 1,
p. 13.)

SuB'Class IL— PROSAURIA,

Mostly extinct Reptiles, with deeply amphicoelous hut stereo-
spondylous vertebrae, with viovable chevron-hones in the tail and
frequently with intercentra in the trunk, Sphenodon, the only
recent genus, has no copulatory organs.

Order I. mOBOSAUBI.

Extinct, small Reptiles, mostly Carboniferous and Permian, with
dermal armour on the dorsal and ventral side and with bifurcated
ribs.

We retain this term of Dawson's for those small, newt-
shaped, chiefly Permian reptiles, which are allied to Hylonomtis,
after elimination of contemporary forms like Keraterpeton and
Urocordylus, which belong to the Branchiosaurian order of the
Stegocephali. Until recently ^ all these creatures had been
classed with the Stegocephali. The Microsauri in the present
restricted sense reveal themselves, however, as reptiles by the
movable chevron-bones in their tail, their broad neurocentral
sutures, the possession of two sacral vertebrae {Petrohates),
the bifurcated ribs which always articulate with the centra
(most clearly shown in Orthocosta), and the possession of five
fingers and toes.

Considering the age of these little creatures and their low
position in the reptilian scale — in fact, they stand almost as low

* Phil. Trans, clxxxvii. 1896, B. p. 23.

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MICROSAURI 289

as the Proreptilia — it is not to be wondered at that they still
retain a number of amphibian features. The skull is encased in
dermal bones as in the Stegocephali, and the dermal armour of
the trunk and tail is composed of many bony, sculptured scales,
which cover back, sides, and under surface. The middle rows on
the back are the largest, while the scales on the belly are
arranged in transverse rows, which imbricate and converge
obliquely headwards. Special gular plates seem to be absent.
The skull has an interparietal foramen. The jaws and the palate
are furnished with small, simple teeth, and there is a large
parasphenoid bone, an eminently amphibian character. The
occipital condylar articulation is supposed to be double. The
centra of the vertebrae are deeply amphicoelous, elongated, and
constricted in the middle, just like those of the Aistopoda
and Branchiosauri. The dorsal spinous processes are strongly
developed, and with the zygapophyses are very reptilian. Trans-
verse processes are absent or very short, the tubercular portions
of the ribs articulating with the centra, the capitula mostly
intervertebrally, in any case close to the anterior end of the centra.
The tail - vertebrae possess very typical, movable chevrons,
placed intervertebrally, and bear an extraordinary resemblance
to those of Geckos. The ribs are long and slender, but there is
no sternum. The fore- and hind-limbs are pentadactyle, in
opposition to the invariably four-fingered Stegocephali. The
shoulder-girdle consists of scapulae, coracoids, clavicles, cleithra,
and a T-shaped interclavicle. The pelvis also resembles that of
certain Stegocephali by the separately ossified, somewhat disc-
shaped, flat ischia and pubes, which seem to have been joined
together by cartilage into one broad mass.

Hylonomus, Dawson's type of Microsauri, was found in the
Coal-measures of Nova Scotia, within decayed tree-stumps. Closely
aUied, if not identical, but much better known is Hyloylesion, e.g.
H. longicostatum of the uppermost Permian of Nyrschan in
Bohemia. Total length under 4 inches ; eyes with bony sclerotic
rings ; neck short. The truly Permian genera Dawsonia, Melan-
erpeion, Orthocosta, and Seeleya are allied forms, the last scarcely
one inch in length, but well preserved. Peirohates of the
Triassic Lower Red Sandstone of Saxony has an arrangement of
the ventral dermal armour closely resembling abdominal ribs.

VOL. VIII u

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290 PROSAURIA

Order IL PEOBAUBL

Mostly extinct, chiefly Permian and Triassic, terrestrial, un-
armoured reptiles with deeply biconcave vertebrae, numerous
intercentra and chevron-bones, fixed quadrates, complete penta-
dactyle limbs and shoulder - girdle, entepicondylar foramina,
acrodont teeth, and many small abdominal ossifications.

The Prosauri differ from the Microsauri, vrith which they
are closely allied, by the more advanced solidification of the
vertebrae, the reduction of the tubercular portions of the libs,
the presence of an entepicondylar foramen in the humerus, and the
loss of the dermal ossifications on the upper surface.

Their ancestors are the Microsauri, whilst they themselves
seem to be very near the root whence have sprung most, if not
all, other main branches of the reptiles, notably Crocodilia, Dino-
sauria, and Sauria. In fact the Prosauri, although apparently-
few in number, seem to represent the central stem of the
reptilian tree. Only one of them is still surviving, the famous
Sphenodon, now represented by a single species in New Zealand.

Sub-Order 1. ProtorosanrL — The ventral half of the pelvis
seems to have formed one broad, contmuous mass of cartilage in
which the pubic bones are represented by a pair of oval, rather
disc-shaped ossifications, while the ischia are more elongated.
The pelvis consequently still bears a great resemblance to that of
the Microsauri, and thereby also to the Stegocephalous condition,
but the ilium seems to be attached to more than two vertebrae.
The vertebrae are deeply biconcave, perhaps even with a persistent
continuous chorda. The neural arches bear high, laterally com-
pressed spines, but no diapophysial or lateral processes, the ribs
being placed mostly intervertebrally and having lost their tuber-
cular portions. The ribs are continued to about the sixth
caudal vertebra. Intercentral wedges exist in an imbroken series
between all the vertebrae from the atlas to the tail, where they
are represented by movable chevrons. A costal sternum seems
to be absent, unless it was quite cartilaginous. The shoulder-
girdle is complete, consisting of a long interclavicle, clavicles,
disc-shaped coracoids, and scapulae ; but there are no cleithra, and
no indication of precoracoids or even notches in the coracoids. The
fore- and hind-limbs are complete and primitive, with five digits.
The abdomen is protected by numerous oat-shaped little ossifica-

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VIII PROSAURI — PROTOROSAURI * 29 1

tions, which are arranged in many transverse or rather chevron-
shaped rows, still greatly resembling the condition prevailing
in the Microsauri, except that they have sunk deeper into
the skin, being no longer directly covered by the scales. The
skull, being no longer completely encased by bones, and possessing
now wide supra- and infra-temporal fossae, appears at first sight
much like that of a generalised lizard, except that it possesses
three very conspicuous and distinct arcades in the temporal region :
namely, the orbito-squamosal bridge across the temporal fossa,
formed by the postorbital and squamosal ; the arch formed by
the squamosal with the postero-lateral buttress of the parietal ;
and the infratemporal arch or jugal bridge. The jugal itself is
long, connecting the quadrato-jugal with the maxillary and
lacrymal, and sending up an ascending process to the postorbital
bone, thus taking a considerable share in the formation of the
orbit. The quadrato-jugal is small, apparently fused with the
quadrate, which itself is firmly overlaid by the squamosal The
quadrates are further fixed by being buttressed by the pterygoids,
which rest upon short basisphenoid processes and extend far
forwards, meeting the vomers and separating the palatines. The
premaxillae are short, the nares small and terminal, the nasal
bones are large. There is a small interparietal foramen. The
teeth are acrodont and pointed, forming unbroken series on the
premaxillaries, maxillaries, palatines and dentaries, and there are
scattered little teeth on the vomers.

Pcdaeohatteria longieaudata from the Lower Red Sandstone of
Saxony. Total length about 18 inches, with six cervical, twenty
trunk, three or four sacral, and about fifty caudal vertebrae. The
teeth are ankylosed with the supporting bones. The five fingers
have 2, 3, 4, 5, 3 phalanges respectively. For the skull see Fig.
54, G, p. 280. Telerpeton elginense from the Triassic sandstone
of Scotland, and perhaps Saurosternon of the South African
Karroo sandstone seem to be allied.

Protorosaurus {nrp&ro^ =s first, &pa = spring, or dawn, not
Proterosaurus) apparently several species, e.g. P. liricki in the
Upper Permian (marl-slate and magnesian limestone) of Thuringia
and Durham. About 4 or 5 feet long, and in its general appear-
ance rather like a Monitor -lizard, with about eight cei-vical
vertebrae, most of which carry slender backwardly-pointing ribs,
sixteen long-ribbed trunk-vertebrae, followed by three or four

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292 PROSAURIA CHAP. VIII

sacrals and more than thirty caudals, some of which have bifur-
cated spinous processes.

Sub-Order 2. Bhjmchocephali — The ventral pelvic tones
resemble those of lizards and enclose a wide pubo-ischiadic foi*amen.
There are only two sacral vertebrae. The abdominal ribs are
closely packed, each transverse set consisting of only three rod-
shaped pieces instead of many small oat-shaped nodules. The
intercentra are sometimes suppressed in the trunk-region.

Rhyrvchoscmrus from the Upper Trias of Warwickshire and
Shropshire, and Hyperodapedon of the same age, found at Elgin,
in Warwickshire, and also in Central India, are rather large, H,
gordoni measuring 6 feet in length. Both have a short, broad,
and stout cranium, and curved down, toothless premaxillae,
hence the name Rhynchocephali ; the nares are confluent ; the
teeth are numerous and small, and are liable to be worn down
so that the animals ultimately bite with the edges of the jaws,
to which the teeth are ankylosed. The premaxillaries of
Hhynchosaurus are curved downwards over a slightly upcurved,
likewise toothless process of the mandibles, which form a strong
symphysis. All the teeth are very small, absent, or minute on the
mandibles, forming one series on the maxillae, several rows on
the vomers and especially on the palatines, which latter remain
separated from each other. Hyperodcq^edon seems to have lost
the intercentra; its vertebrae are solid, those of the neck are
opisthocoelous. The interparietal foramen is likewise abolished.
The hook-shaped end of the curved-down premaxillae fits into a
bifurcation of the mandibles in front of their stout sjTnphysis.
The teeth are similar to those of the other genus. Whilst these,
the earliest known genera of Rhynchocephali, are already in
various ways rather specialised, e.g. the hooked beak and the loss
of the intercentra, the two following fossil genera, although of
much later date, namely Upper Triassic, are more closely allied to
the recent Sphenodoiu

Homocosaurus x>ulcheUii8 and other species in Germany are only
6 to 8 inches long. The vertebral column consists of twenty-
three presacral and many caudal vertebrae. The first five
cervicals are devoid of ril)s. Intercentra are restricted to the
neck and the anterior portion of the tail. The mandibles are
not fused together. The nares are divided by a bony septum.
Each premaxillary has one rather broad tooth. The teeth of

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294 PROSAURIA

the maxillaries and mandibles are triangular, much worn down
in front. The ribs are devoid of uncinate processes. Closely
allied but larger is Sauranodon of France, which has lost the
upper teeth and uses the sharp margins of the jaws instead.

Fleurosaurus of Germany and France, about 5 feet in length,
is remarkable for the shortness of its still pentadactyle extremities,
for its short neck, and very long tail ; — an interesting parallel to
what has happened in many genera of recent lizards.

Sphenodon s. Hatteria is the. sole surviving member of the
whole group of Prosauria, and is represented by one species only,
S. punctatuw,, in New Zealand. As the last living witness of
bygone ages this primitive, almost ideally generalised type of
reptiles, this " living fossil," deserves a detailed description.

Total length of very large male specimens up to two feet and
a half; in general appearance like many a stoutly built lizard.
The general colour of the skin is dark olive-green with small
• white or yellowish specks on the sides. A series of slightly
erectile spines of yellowish colour extends from the top of the
head to the end of the tail, but is interrupted on the neck ; they
are cutaneous, covered with a thin sheath of horn. The under
surface is covered with numerous scales, arranged in transverse
rows ; the rest of the body is rather granular. The tail is thick,
slightly compressed laterally. The eye is large, dark brown, with
a vertical pupil.

Those who are satisfied with superficial resemblances still
group this creature with the lizards, but it reveals itself as a
primitive reptile or Prosaurian by the following characters, every
one of which distinguishes it from the lizards : — The temporal
region is bridged by three bony arcades. The large vomers,
palatines, and pterygoids form a broad bony roof to the mouth ;
the large quadrates are firmly fixed by the pterygoids, squamosals,
lateral occipital bones, and by the jugal bridge. The vertebrae
possess an unbroken series of intercentral wedge-bones. There
is an elaborate system of abdominal ribs. The humerus has an
entepicondylar foramen, and there is also, in contradistinction to
the fossil Ehynchocephalia, an ectepicondylar foramen for the
passage of the radial nerve. The carpus still has the primitive
number of ten bones, all of which remain separate, including the
intermedium. Of soft parts are to be mentioned above all the
entire absence of external copulatory organs, ^phen/)don being the

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l»ROSAURI — RHYNCHOCEPHALI

295

only recent reptile which is devoid of them ; a most primitive
condition, sufficient by itself to separate this creature from all the
other living reptiles.

The supratempord bridge is formed by the squamosal and

fTT*R

Pmx

Fig. 59. — A, Dorsal ; B, ventral ; C, left-sided view of the skull of Sphenotlon. x j|.
Coly colamella amis ; C<md^ occipital condyle ; E. /*, ectopterygoid ; t\ frontal ; Jng^
jugal ; McLX^ maxillary ; iVo, nasal ; iVo, anterior nasal opening ; Pal^ palatine ;
Par, parietal ; PmXy premaxillary ; Prf^ prefrontal ; Pt'f, postfrontal and post-
orbital ; Ptg, pterygoid or endopterygoid ; Q, quadrate and quadrato-jugal ; *S^,
squamosal ; Fo, vomer. See also Fig. 54, B, p. 280.

postorbital (Fig. 59, C, Pt.f), the latter being continued forwards
and fused with the postfrontal (A, P//). The postorbital joins
the ascending branch of the jugal, both together forming the
hinder border of the orbit ; this is bordered below chiefly by

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296 PROSAURIA

the maxillary, which is long, while the anterior process of the
jugal is much reduced. There is no pre-orbital fosseu The nares
are terminal and lateral, well separated by the premaxillaries.
The posterior temporal bridge is formed by the squamosal and
parietal, the bridge extending laterally over the quadrate and
enclosing a wide space between itself and the buttress-like expan-
sion of the lateral occipital bone. The space enclosed between
this occipital buttress, the quadrate, and the pterygoid support
of the latter is likewise very large ; it is of course the ca\dty
of the middle ear, and as such is crossed by the columellar chain
of the ear. The infratemporal bridge or jugal arch is formed by
the jugal, which joins the descending process of the squamosal,
and by the quadrato-jugal, which is small and fused with the
quadrate. The latter is consequently very firmly fixed.

The teeth are acrodont, ankylosed in one series with the
supporting bones, triangular and much worn down in older
specimens. Originally there seem to be several in the premaxilla,
but the adult bite with the somewhat curved-down portions of
the premaxillaries themselves, or with what remains of the fused
bases of the original teeth, which then, together with the bone,
look like one pair of large chisel-shaped incisors. The lateral
edges of the palatines likewise carry teeth, those of the mandibles
fit into the long slit-like space between the palatine and the
maxillary teeth. Young specimens have a few small teeth on
the vomers, which are large, and separate the long choanae from
each other. The pterygoids form an anterior symphysis, posteriorly
they rest upon short processes of the basisphenoid and send short
flanges to the quadrates.

The vertebral column is very primitive. The atlas is still
typically temnospondylous. The first intercentrum or fused pair
of basiventrals is broad and thick, and forms the ventral half of
the atlas-ring, which articulates with the first centrum and with
the second intercentrum. The irregularly shaped neural arches
remain separate from each other and from the centrum; they
carry on the dorsal side a pair of disconnected supradorsals, the
so-called pro-atlas. The second intercentrum is fused with the
first and second centrum. The second to ninth intercentra
have low median ridges or knobs, and are as a rule more
firmly attached to the cranial ends of the centra. Those of the
trunk are small. From the third or fourth caudal vertebra

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M^^

». 60.— The

first three

cervical vertebrae of

Sphenodon.

1, 2, 3, 4,

Intercentra

; c, . C„

centra; Ny

iVj, neural

arches.

viii RHYNCHOCEPHALI 297

backwards they appear as chevrons, articulating more with
the vertebra in front than with the one behind. The bases
of the right and left chevrons are frequently fused across,
so that the caudal canal is completely surrounded by bone, a
feature common in Dinosaurs. Every intercentrum, be it a pair
of chevrons, or an unpaired nodule, or cres-
cent, extends dorsalwards into a fibro-
cartilaginous ring which surrounds the
chorda. The centra of the vertebrae are
deeply amphicoelous, the cavity being filled
throughout life by the chorda ; but the
middle of the centra is solid. Most of the
caudal vertebrae are transversely divided
into two parts, the posterior of which
carries the greater share of the arches ;
they resemble in this respect those of lizards,
and the lost tail is likewise reproduced. The first three ribs are
represented by bands of connective tissue. The first is attached
to the side of the first intercentrum ; the second arises from the
second intercentrum, and forms a small tubercle on the side of
the second centrum ; the third behaves similarly. The vertebral
arteries and lateral strands of the sympathetic nerve-chain pass
through these double basal attachments of the reduced ribs.
The other ribs are osseous; they possess short capitula which
retain their partly intercentral attachment, while the short
tubercula are carried by low processes of the centra, not of the
neural arches. Already in the thoracic region both capitulum
and tuberculum merge into one facet, at first dumb-bell shaped,
further towards the tail oval, gradually shifted backwards and
dorsalwards upon the middle of the centrum, until the facet
reaches and ultimately lies right across the neuro-central suture.
The first few caudal vertebrae also possess ribs, which are how-
ever very short and fuse with the diapophyses, immediately below
which lies the neuro-central suture.

The whole column consists of twenty -five presacral, two
sacral, and about thirty caudal vertebrae. Some of the thoracic
ribs have cartilaginous uncinate processes. Three or four pairs
of ribs join a typical sternum, into the antero-lateral portion of
which are let in the coracoids. The sternum is raised into a
low median crest which fuses with the posterior branch of the

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298 PROSAURIA ciiAi'.

T-shaped interclavicle, while the lateral branches of the latter
fuse with the clavicles. The coracoids are broad and entire,
still without fenestrae or notches indicative of precoracoida
The parasternum is very elaborate ; it extends from the sternum
to the pubic bones, and consists of about twenty-four transverse
rows, each of which is composed of a median and two lateral
splint-bones. They are irregularly shaped, partly with imbri-
cating hooks, and are firmly attached to, in fact still connected
with, the deeper portions of the cutaneous scales of the belly*
The three pairs of pelvic bones are fused together at the
acetabulum. Pubes and ischia each form one symphysis, and
these are connected with each other by partly ossified cartilage
and ligaments, so that the original cordiform foramen is
divided into a pair of ovals. The lateral processes of the pubes
are thick, but very short. The ischia have postero- lateral
processes. There is also a mostly cartilaginous, unpaired
hypo-ischium.

The fore- and hind-limbs are still primitive in structure ;
both pentadactyle. The carpus consists of ten, sometimes eleven
pieces, according to the single or double nature of the central
element. The proximal series is formed by the radiale, inter-
median, and ulnare, with a pisiform. The ulna and radius remain
separate. The humerus has the usual ectepicondylar in addition
to the entepicondylar foramen common to all the Prosauri and
Theromorpha. The hind-limbs are typically plantigrade.

The tail is capable of regeneration, as in many lizards.

The development of this reptile has recently been studied
and described by Howes,^ who quotes the literature bearing
upon the whole subject.

A good account of the occurrence and habits of the " Tuatera "
has been given by Newman.^ The Maoris call it " ruatara,"
" tuatete," or " tuatara," the latter meaning " having spines."
Formerly common on the main islands of New Zealand, they are
now apparently restricted to some of the islets in the Bay of Plenty,
North Island. Bush -fires, wild pigs, dogs and cats, reptile-
eating Maori tribes, and the advance of civilisation, have swept
them away except on some of the small uninhabited islands,
difficult of access, where they dig burrows, into which they
retreat at the slightest sign of danger. They sleep during the

^ Traw. Zool. Soc. xv. - Trans. N. Zealand Inst, x. 1878, p. 222.

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RHYNCHOCEPHALI 299

greater part of the day, are very fond of lying in the water, and
they can remain below for hours without breathing. They live
strictly upon animals, but these are only taken when alive and
moving about. The kind of food seems to vary according to
the custom or fancy of the individuals. Sir W. L. BuUer observed
that some of his captives stubbornly refused to eat until
one day, rather accidentally, minnows were ofifered. Others eat
insects and worms ; those which live near the seashore not im-
probably eat also crustaceans. From November to January they
lay about ten eggs — white, hard-shelled, long and oval — about
28 mm. long, in holes in the sand, where they can be warmed by
the sun. They are as a rule lazy in their movements. The usual
pace is a slow crawl, the belly and tail trailing on the ground,
but when chasing prey they lift the whole trunk off the ground.
After running, or rather " wobbling " three or four yards, they
grow weary and stop. They cannot jump the smallest obstacle.

Von Haast ^ has carefully examined their habittitions on the
Chicken Islands. The Tuatara excavates its own hole, and this
is shared sociably by various kinds of Petrels. The entrance to
the chamber is generally 4 or 6 inches in diameter, and the
passage leading into the inner chaml^er is 2 to 3 feet long,
first descending and then ascending again. The chamber itself
is one foot and a half long, by one foot wide and 6 inches
high, lined with grass and leaves. The petrel lives usually on
the left side, the Tuatara on the right side of the inner chamber.
Whilst very tolerant of the bird with its egg and young, it does
not allow another of its own kind to live in the same hole,
which it is ready to defend by lying in such a manner that the
head is placed where the passage widens out into the chamber.
On putting one's hand or a stick into the bun'ow the Tuatara
bites at them furiously. They can run very fast, and defend
themselves with great pluck against dog or man by biting or
scratching. As soon as the sun has set they leave their holes
to seek food. During the night, and especially during the
pairing season, they croak or grunt.

The eggs, having been deposited during the Southern summer,
from November to January or February, in holes on a sunny
and sandy spot, contain nearly ripe embryos in the following
August. They are, however, not hatched until about thirteen
^ Trans, N. Zealand Inst. xiv. 1881, p. 276 : cf. also Rcischek, 0^). «'/. xiv. ji. 274.

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300 THEROMORPHA

months old. In the meantime they seem to undergo a
kind of aestivation. The nasal chambers become blocked
with proliferating epithelium, which is resorbed shortly before
hatching.

I have kept half-a-dozen specimens in a green-house for
several years, and have come to the conclusion that they are
dull, not companionable creatures, in spite of their imposing,
rather noble appearance when, with their heads erect, they
calmly look about with their large, quiet eyes. Each dug its
own hole in the hard ground underneath and between large
stones. At dusk they sat in front of the holes or walked
leisurely to the pan with the earthworms which formed their
principal food. Meat they did not touch, but they killed and
chewed up lizards and blind-worms. Sometimes they soaked
themselves for many hours in the shallow, warm water. The
skin is shed in flakes. I never found them basking in the sun,
and the pineal eye, still so well developed in these strange
creatures, caused them no distress when bright light was thrown
upon it. They grew tame enough not to run away when found
roaming about at night, but they did not like being handled,
and they inflicted the most painful bites when taken up care-
lessly. The biggest, a male, was rather quarrelsome, grunted
much, and worried the others.

Sub-Class IIL— THEROMORPHA.

The Theromorpha comprise a great number of extraordinary,
extinct reptiles, which as a group had a wide range in space
and time. The earliest known occm* in the Lower Red Sand-
stone of Thuringia and Bohemia, and in the middle Permian
strata of Eussia. The majority have been found in strata
transitional between the Permian and the Triassic age, notably
in tlie Karroo sandstone of South Africa and in corresponding
levels of North America. Closely allied to them are those of
the Triassic sandstone of Elgin in Scotland, and of India.
They seem to have died out with the Muschelkalk or Middle
Trias.

The various genera exhibit such a diversity of structure,
shape, and size, and many are still so imperfectly known, that

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STRUCTURE 30I

any diagnosis is liable to be faulty, even assuming that they are
a homogeneous group. To avoid confusion, we characterise the
Theromorpha as Reptiles with a firmly fixed quadrate, a single
temporal arch, an interparietal foramen, and a pelvis in which
the puhes and ischiaform one stout, ventral synqihysis.

The dentition is most abnormal, and pennits the division of
the Theromorpha into two or three main groups. In the Pareia-
sauri the teeth of the upper and lower jaws form rather even
series of nearly equal size ; smaller teeth are carried by the
palatal bones. In the Theriodontia the teeth are differentiated
in a tridy Mammalian fashion into incisors, prominent canines,
and multicuspid or tubercular molars. Each tooth, and this
applies to all Theromorj)ha, is implanted in a separate alveolus ;
Tritylodon only seems to have double-rooted molars. The lower
canines cross in front of the upper, just as in Mammals. In
PlacodxLS, which probably belongs to this assembly, the teeth are
few in numbers, very broad and flat, especially those of the
palate. In Dicynodon and Gordonia the teeth are restricted to
a pair of conical, sometimes very large, tusk-like upper canines,
and in Oudenodon the whole mouth is toothless.

The configuration of the skull shows two main types. In
the Pareiasauri it is completely roofed in by dermal bones, the
only holes on the surface being the nostrils, orbits, and the
interparietal foramen.

The most striking feature of the second type of skull is the
tendency to form an almost Mammalian zygomatic arch by the
junction of the much elongated squamosal with the jugal bone, both
abutting against a downward process of the i)ostfrontal bone.
The skull shows a pair of wide supratemporal foramina bordered
by the parietals, squamosals, and postfrontals. The com|)osition
of the temporal arch varies considerably in detail, and in Cyno-
f/nuthus crateronotus at least there is a small hole within the
arch, between the squamosal and jugal, probably the last
remnant of the otherwise absent infratemporal foramen. Except
in the roofed-in skulls of Pareiasauriis and Elginia there is no
separate quadrato-jugal element. The quadrate is firmly fixed by
the overlapping squamosal, and the whole pedicle for the support
of the mandible is rather elongated, and either stands vertically
or slants forwards. The mandible itself is compound. The
pterygoids extend backwards so as to approach or reach the

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302 THEROMORPHA chap.

distal portion of the quadrate; separate ectopterygoids do uot
seem to be developed. The shoulder-girdle consists on either
side of a large scapula, which is mostly directed obliquely back-
wards, and is fused with the coracoid ; a precoracoid is present or
at least indicated by a notch or foramen ; it is usually fused with
the other bones. At least some genera possess a T-shaped inter-
clavicle and clavicles; Pareiasaurus possesses also a pair of
cleithra.

The pelvis is in every respect constructed upon the Mam-
malian plan. The three constituent parts meet at the ace-
tabulum, and the ventral bones, pubes and ischia, form one broad
symphysis, leaving two, sometimes very small, obturator-foramina.
The ilium is attached to one to five sacral vertebrae, and since
the whole pelvis slants obliquely downwards and backwards, this
sacral attachment is distinctly pre-acetabular, perhaps most
markedly so in Dicynodon, The limbs are mostly stout, humerus
and femur with strong crests ; the feet are thoroughly plantigrade,
with five fingers and toes. The details of the carpus and tarsus are
not well enough known to permit of generalisation, but there is
a tendency to form a heel, and to develop the cruro- tarsal
joint into the chief joint of the hind feet. The vertebrae are
amphicoelous, sometimes with rather thin-walled centra, so that
in these crises the chorda was continuous. Intercentral wedges,
or basiventral elements, are frequent in the cervical and caudal
regions. Most of the ribs, especially those of the neck, have a
tuberculuui attached to the neural arch, and a distinct capitu-
lum which articulates either with the centrum or with the
intercentrum, or lastly, if the latter is absent, between two
centra. The axis and atlas vertebrae are united.

The occipital condyle exhibits every stage between the single
median knob {Pareiasaurus) formed almost entirely by the
basioccipital bone, a triple condyle (Dicynodon) to which both
lateral and the basioccipital bones contribute, and a kidney-
shaped or double condyle (Cynognathus) from which the middle
or basioccipital portion is more or less withdrawn.

Dermal bony armour reached an extraordinary development on
the head of Pareisaurus and Elginia ; whether other parts of the
body were protected is doubtful, but the flattened tops of
the neural spines of Pareiasaurus suggest that they carried
bony scutes. Abdominal protective ossifications are unknown.

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RELATION TO MAMMALS 303

Mauy of the Theromorpha ^ reached a considerable size, massive
skulls of one foot in length being not uncommon. , The tail
was comparatively short.

The many resemblances of these strange creatures to
Mammals have naturally suggested that the Mammalia have
sprung from some such Theromorpha or " beast-shaped " animals.
The resemblances are chiefly the dentition, the zygomatic arch,
the pelvis, the cruro-tarsal joint, the scapula which is sometimes
possessed of a spine, and the occasionally double occipital con-
dyle. The general shape of the skull of Cynognathus is indeed
strikingly like that of a Carnivorous Mammal, and the shape of the
whole body suggests rather a Mammal than a reptile ; and when
we have to deal with the fragmentary skulls of Tritylodon (cf. p.
309) it is, indeed, difficult to decide to which of the two classes
such a creature belonga But the Theromorpha possess a
number of important characters by which they reveal themselves
at once as reptiles: (1) the large and fixed quadrate bone, which
is still the sole support of the lower jaw; (2) the compound
mandible, which is composed of at least an articular, dentary,
angular, supra-angular, and splenial element; (3) the inter-
parietal foramen ; (4) the possession of prefrontal and post-
frontal bones, sometimes also postorbital, supratemporal, and
quadrato-jugal bones. Of course, any of these ancestral
bones may be lost, and the interparietal hole may be closed as in
tortoises and crocodiles. We can also imagine that the quadrate
may be relieved of its jaw-bearing function and become loosened,
but this is not easy, considering the strong development of
the squamoso-quadrate pedicle. Those Theromorpha in which the
quadrate itself is small, whilst the squamosal reaches down, or
at least approaches the mandible, as in D icy nod on and Gordonia,
are so hopelessly pledged, or specialised in other directions, that
it is impossible to connect them ancestrally with Mammals.

However, it is beyond reasonable question that the Mammals
have sprung from some reptilian stock (the attempts to derive

^ Cope, the inventor of this most appropriate name, soon changed it, un-
necessarily, into Theromora (/iwp6f= sluggish), perhaps in order not to emphasise
too much their possible Mammalian affinities ; while others rashly called them
Sanro-Mammalia. For detailed illustrations of Tlieromorpha reference should be
made to Owen, British Fossil Reptiles^ 4to, London, 1849-55, and to numerous
papers by Seeley, Phil. Trans, 178 (1887), 186 (1895), and by E. T. Newton in
Phil. Trans. 184 (1893), 185 (1894).

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304 THEROMORPHA

them from Amphibia, without the intervention of Reptiles,
are as gratuitous as they have proved futile), and the Thero-
morpha undoubtedly comprise creatures which of all animals
approach nearest to Mammals, and coincide with them in most
important features. But we have not yet found a single
Theromorph which can claim to be a direct ancestor of
Mammals. Since the latter occur already in the Trias, we have
to look for their reptilian forefathers at least in the Lower
Permian, and this naturally excludes all the known forms. The
filling up of this gap is but a question of time.

The ancestry of the Theromorpha themselves is also
shrouded in mystery. Attempts have been made to connect
them with the Permian Frotorosaurus, Falaeohatteria, and Eryops.
On the other hand, some retain various Stegocephalous reminis-
cences {e.g. the roofed-in condition of the skull by membrane-
bones, amongst which, besides others, supratemporals and post-
orbitals can be recognised ; occurrence of cleithra in Pareia-
saurus ; distinct epiotic bones in Elginia). Although they
have died out as a group, they have perhaps given rise to
several side-branches, one of which (leaving aside the question of
Mammalian origin) seems to have flourished as the Dinosauria.

We divide the Theromorpha into four orders, which ai'e, how-
ever, liable to run into each other, and it is reasonably to be
hoped that many forms may be discovered which will connect
not only these provisional orders with each other, but also with
other sub-classes.

Order L PAREIASAURL

Cranium completely roofed in by membrane-bones. The only
foramina are the nostrils, orbits, and the interparietal foramen.
The teeth are comparatively small, and stand in even series in
both jaws.

PareiasauruSy several species from the Karroo sandstone of
South Africa. P. haini was an extremely clumsy brute, of most
uncouth appearance, standing between 2 and 3 feet high, and
measuring with the short tail nearly 8 feet in length. The
skull is very massive, 18 inches long and slightly broader,
with a rugose, deeply pitted surface. The teeth are thickly
enamelled, serrated at the margin, with many pointed cusps ; those

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PAREIASAURI 305

of the vomer, palatines, and pterygoids are recurved and arranged
in several longitudinal rows. There is a small incisive foramen
in the premaxilla ; the choanae lie within the pterygoids. The
palate has a pair of large lateral vacuities. Between the squa-
mosal and quadrate is a small foramen, as in Belodon and
Sphenodon. The nares are terminal, bordered behind by the
nasals, and divided by the premaxillaries. The occipital condyle
is a single knob, but the lateral occipital bones also partake in
its formation. The shoulder-girdle is strong. The scapula
slants backwards, is broad, and possesses a longitudinal spine, an
almost exclusively Mammalian character. The scapula, coracoid
and precoracoid are fused together, and are united ventrally
with those of the other side. There is a T-shaped interclavicle,
a pair of clavicles, and a pair of slender, long cleithra, which
extend along the upper anterior margin of the scapulae. The
humerus possesses enormous crests. The broad ilium is
attached to two, or perhaps three, sacral ribs. The acetabulum
is closed. The pubes and ischia are united into one broad
mass of bone, and the obturator-foramina seem to be just large
enough to permit of the passage of the nerve. Both fore- and
hind-limbs are plantigrade and five-toed. The tibia articulates
with one large bone, which is supposed to represent the united
astragalus and calcaneum, the latter being without an indication
of a prominent heel, although there is a tendency to develop the
crurotarsal into the chief joint. The number of vertebrae amounts
to eighteen presacrals, eight to ten of which are cerviculs. There
are two or three sacral and about twenty-four mostly shortened
caudal vertebrae. The latter possess intercentral wedges and
chevron-bones ; wedges occur also between the cervical and some
thoracic vertebrae. Some of the posterior cervical ribs are very
peculiar — straight, broadened out, turned backwards, partly over-
lapped by one another, and 1 8 inches long, recalling the first two
ribs of the crocodiles. Sternum and abdominal ribs are unknown.
Elginia mirabilis, — The skull (Fig. 54, A, p. 280) — nothing else
is known — indicates one of the most remarkable reptiles hitherto
found on this side of the Atlantic. It was discovered in the
Eed Sandstone of Elgin (Lower Trias). The skull reminds us in
its general shape and by its spikes and horns of the little American
Iguanoid lizard, FJifv/nosoma. The length of the cranium is
about 6 inches, the distance between the tips of the two largest

VOL. VIII X

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306 THEROMORPHA

horns measures 9 inches. The teeth are small and resemble
those of an Iguana in their shape and finely serrated edges,
indicating herbivorous habits, but there are also several rows of
smaller teeth on the palate, the configuration of which is not
unlike that of Sphenodon. The top and sides of the skull,
except the interparietal foramen, the orbits, and nostrils, are com-
pletely encased by rugose, pitted, dermal bones, most of them
with strange, horn-like spikes. In the encasement of the
temporal region can be discerned a postfrontal, parietal and squa-
mosal, a conically projecting epiotic, a postorbital and supra-
temporal, a jugal and a quadra to-jugal, which latter almost
completely covers the quadrate bone. The interparietal foramen
lies far forwards, almost on a level with the orbits. The nostrils
are terminal, surrounded by the short nasals, the maxillaries
and the premaxillaries, which latter divide them.

Order n. THERIODONTIA.

The cranium is not roofed in, but shows a pair of large
supratemporal fossae, bordered below by the zygoma, which is
formed mainly by the squamoso-jugal bridge, and is shut off from
the orbit by the postfrontal joining the bridge. The teeth are
differentiated into incisors, canines, and molars (Fig. 54, C, p.
280). The lower canines close in front of the upper.

Cynognathas, Karroo formation of South Africa. C, cratero-
not us has a skull about 16 inches long, looking like that of
a ferocious Carnivore ; there are four .incisors, huge canines,
and nine molars, the latter with serrated edges and anterior
and posterior cusps. The wide supratemporal fossa is bordered
and closed behind by the broad lateral extension of the parietal,
which joins a similar extension of the squamosal bone. The
latter is very long, extending to the postfrontal and to a
bone which, bordering the orbit posteriorly, is either an upward
branch of the jugal, or a postorbital bone ; the latter inter-
pretation is made probable by the occurrence of a suture with
the jugal in C, 2)latyce2)s. The jugal bone is very long, begin-
ning at the quadrate, running along the squamosal, and forming
the lower border of the orbit.

The number of vertebrae is large, there being as many as
twenty-nine presacrals, six of which belong to the cervical region.

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THERIODONTIA 307

The atlas is fused with the axis ; most of the thoracic ribs
articulate partly upon the intercentra. The lumbar ribs are
very peculiar ; they are much expanded horizontally, and overlap
each other, forming thereby intercostal foramina. The broad
ilium is attached to three or four sacral ribs. The acetabulum
is closed. The ventral side of the pelvis shows a broad
symphysis and has a pair of obturator-foramina. The scapula
is large, directed backwards, and shows a distinct, very Mam-
malian spine ; it is fused with the coracoid and precoracoid.

The occipital condyle of C, platyceps is kidney-shaped, with
the concavity directed upwards ; in C. herryi it is separated into
two distinct knobs, the middle, basioccipital portion being
apparently wanting. The mandible possesses a long coronoid
process which ascends obliquely into the temporal fossa.

Aelurosaurus, LycosauruSy Galesaurus, and many others,
likewise of the Karroo formation. In the first genus the
splenial bones help to form the symphysis of the lower jaw ; teeth
are also found on the palate, in opposition to Lycosaurus. This
has a skull 6 inches in length ; the dental formula on either side
is L ^, c. \f m. ^ ; the molars are slender, conical, and recurved.
Galesaurus seems to have been rather small, the low, triangular
skull measuring only 2 to 3 inches in length, with four or five
sharply pointed incisors, prominent canines and four or five small
multicuspid or deeply serrated little molars.

Endothiodony with several species from the Karroo formation,
is of uncertain systematic position, only imperfect skulls being
known. The animals must have been large and bulky, the
skulls being very massive and at least one foot in length. The
premaxillaries and the maxillaries are toothless, their alveolar
borders forming cutting, prominent edges. The same applies to
the very strong lower jaw; but there is a pair of tooth-like
stout projections in the upper and lower jaws in the place of
canine teeth. True, enamelled, small, apparently conical or low
and perhaps blunt teeth occur on either side in one or three
longitudinal series upon the palate, and in corresponding positions
on the inner sides of the two halves of the lower jaw. It is
doubtful if the upper teeth are carried by the palatines or by the
broadened inner flanges of the maxillaries. The choanae seem to
lie between the pterygoids and the palatines, incompletely roofed
in by ventral extensions of the latter towards the middle line.

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308 THEROMORPHA chap.

Direct affinity of Endothiodon (ivioOiy within) with Placodas
is unlikely; the same applies to the Dicynodontia, although
the restriction of the teeth to the palate seems to point as
much to the former genus as do the toothless cutting edges
of the jaws to the forms like Oudenodon,

Other Theriodont reptiles have heen described from the upper
Permian of Eussia, for instance Deuterosaurus and Brithopus,
but the determination rests upon insufficient fragments. North
America has yielded many strange Theromorphous fossils, some
of which may belong to the Theriodont order, while others seem
to be intermediate between this and the other orders, jyiadeeies
of Texas, for instance, seems to be a Theriodont creature; while in
Empedias molaris, with a skull about 8 inches in length, the
teeth form an uninterrupted series without distinct canine tusks,
and the incisors are distinguished from the molars only by the
transversely broadened shape of the latter. Very small teeth are
arranged along the median line of the vomer and united palatine
bones. In Clepsydrops, Dimetrodon, and Naosauriis of Texas the
teeth are differentiated into incisors, canines, and molara, although
not so regularly as in the typical Theriodont forms described
above, one or more pairs of teeth being enlarged into canine-like
tusks. In the latter two genera the spinous processes of the
thoracic vertebrae are enormously elongated, standing up vertically
to a height of 2 feet, while the centra of the vertebrae measure
only one inch in diameter. In Naosaurus claviger these upright
spines carry on either side half a dozen transverse projections.
Stereorkadiis of the Permian of France is typically Theriodont in
the structure of its shoulder-girdle, humerus, and pelvis, but the
dentition is composed of ^ incisors, no canines, and ^^ pointed
miliars.

Tlic following genera have been placed by Seeley in the family
Gomphognatliidae. MicTogomj)hodon, with broader and less pro-
minently multicuspid teeth than those of the typical Theriodonts,
seems to lead to Gompho(jnat]ius,\i\\\{i\\ has the following dentition:
i. j|, c. |-, m. ||, with a long diastema between the canines and
molars, some of wliich latter are nearly as broad as they are long,
and liave comparatively low tubercles on tlie crowns. Tlie skull
is remarkably like that of a Carnivorous Mammal. There are
incisive foramina behind the premaxilla. The maxillaries and
palatines form a unit<'d palatal roof, and behind them open the

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ANOMODONTIA 309

choanae. The occipital condyle is kidney-shaped. The mandible
is most extraordinary, approaching that of the Mammalian,
especially the Marsupial type, except that it is still composed
of several pieces. The articular facet for the mandible is borne
by an outward or lateral projection, while the bulk of the
posterior half of the jaw projects inwards like a broad flange,
undoubtedly recalling the so-called inner inverted angle of the
Marsupial jaw. The coronoid process is large and extends
far into the temporal fossa. Nearly the whole skeleton of
Microgomphodon is known ; the lumbar ribs are broadened and
overlap as in CynognatMis, and the mandible is typically
compound, so that there is no doubt about the afl&nities of this
genus with the Theriodontia. It throws light upon Gompho-
gnathus and the three likewise Soutli African genera Diademudon,
Trirachiodon and Tritglodon, which are all known from imperfect
skulls only. Their teeth are restricted to the jaws, the molars
have flat, multitubercular crowns and bear an extraordinary
resemblance to those of Mammals. Some of the molars of
Tritylodon are said even to possess two roots, but this point,
absolutely unique in Eeptiles, but common in Mammals, is not
certain. The few upper incisors of Tritylodon are rather large,
chisel -shaped, and extend like those of the Eodent-type back
into the maxillaries; canines are absent, leaving a diastema.
Trirachiodon has prominent canines, the five upper molars are
multitubercular, rather flat, and much broader transversely than
in the longitudinal direction. Still, even these creatures, with
skulls of the size of that of a small fox, possessed distinct pre-
frontal and postfrontal bones, and are, at least in this respect,
typical Eeptiles.

Order ni ANOMODONTIA.

The cranivmi is not roofed in. The pedicle for the suspension
of the lower jaw is much elongated, slants slightly forwards, and is
composed of the long quadrate, which is laterally overgrown by
the squamosal bone. The teeth are restricted to a pair of strong,
tusk-like canines, or they are altogether absent. The margins
of the upper and especially those of the lower jaw are trenchant,
and were possibly furnished with a thick homy armature like
those of tortoises.

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310 THEROMORPHA

DicyTwdon, with many species from the Karroo formation of
South Africa, reached formidable dimensions. The thick, curved
skull is in size and outline not unlike that of a large lion, hence
D, leoniceps, D. tigriceps, etc. The zygomatic arch is almost
mammalian, except that the posterior boimdary of the orbit is
formed by a distinct postfrontal bone. The nostrils are lateral.
The canine tusks (Fig. 54, E, p. 280) are very large. The
choanae open behind the rhomboid vomer and between the sepa-
rated palatine bones, which are posteriorly confluent with the
medially united pterygoids. The latter send out flat extensions,
along the lateral side of the palatines ; these extensions reach
the maxillaries and probably represent the ectopterygoids. The
occipital condyle is distinctly triple, being equally composed of
the basi- and latero-occipital bones.

The three bones of the shoulder-girdle meet at the glenoid
fossa ; the scapula has the indication of a spine. The pelvis is
stout, attached to four or five vertebrae, converting the latter into
a very Mammalian -like sacrum, the position of which lies
distinctly in front of the acetabulum. The latter is closed,
composed by the three pelvic bones. The pubes and ischia
are fused together, leaving only a very small obturator-foramen.
The limbs are plantigrade and pentadactyle, very stout; the
humerus and femur have enormous crests.

Oudenodoiiy of which several species have been described, is so
much like Dicynodon, except for the complete absence of teeth,
that it has been suggested that these skulls belong to females of
this genus. This view is strengthened by the fact that tusk-
like canines exist, or are absent in some of the species which
have been described as Cistecephalus, a genus closely allied to
Dicynodon. The latter, which, like Oudenodon and Ci^te^ephalvs,
occurred in Africa, extended also into India, D, orientalis ha\ing
been found in the Panchet formation of Bengal, of transitional
age between the Permian and Triassic epoeha Oudenodon
rugosus, on the other hand, has been described from the Ural.

Gordonia and Geikia, of the New Ked Sandstone of Elgin, are
known from their skulls only, but these are so well preserved
that there is no doubt about their close relationship to the
typical South African Dicynodontia. The skull of Gardonia is
about 7 inches long and 4 inches high. The canines (Fig. 54, D,
p. 280) are reduced to short, but thick, conical tusks. The most

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viii PLACODONTIA 3 1 I

i-einarkable feature is the very elongated squamoso-jugal arch, which
arises moreover from the dorsal end of the long squamoso-quadrate
pedicle. The two wide and long temporal fossae are dorsally
divided by narrow parietal crests. There is a distinct interparietal
bone, and the usual interparietal foramen. The choanae are
united and lie within the palatines, which themselves are united;
the large lateral palatal foramina are otherwise enclosed by
the pterygoids,' quadrates, and laterally by the squamoso-jugal
arch.

Order IV. PLACODONTIA.

These are the latest and last members of the Theromorpha,
unfortunately known from skulls only, from the Muschelkalk or
Middle Trias of Germany and Russia. The skull of Flacodns
gigas is about one foot long, rather high and triangular owing
to the lateral expansion of the temporal arches, which diverge
posteriorly. The squamoso-jugal arch is very broad, and most
of the posterior border of the orbit is formed by the large
postorbital bone. The maxillary bone seems to extend back to
beyond the level of the orbits. The choanae lie behind the pre-
maxillaries. The palatines and pterygoids are fused in the
middle line, forming a broad bony palate, which, owing to the
broad, posteriorly extended wings of the pterygoids, much re-
sembles that of the crocodiles. The teeth are very remarkable.
There are two or three stout, conical, or chisel-like teeth in each
premaxillary bone, and three to five broad and fiat maxillary
teeth ; three pairs of huge, broad, and quite flat teeth aie crowded
together and fill up the whole vomerine and palatine portion of
the palate. These crushing teeth indicate that Placodns probably
lived upon hard-shelled molluscs, and this would be in conformity
with its occurrence in the Muschelkalk, which is a strictly marine
deposit and full of shells. Another closely allied genus is Ci/a-
wodus, one species of which is known from Russia. The teeth
are fewer in number and not so large as those of Flacodiis.

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CHAPTER IX

CHELONIA ATHECAE THECOPHORA

Sub-Class IV.— CHELONIA,

There is no mistaking a tortoise. The shell and the horn-
covered toothless jaws separate them from all other four-footed
creatures.

They may be described as terrestrial or aquatic, pentadactyle
reptiles, with walking limbs or with paddles ; ribs with capitular
portions only, two sacral vertebrae, humerus with entepicondylar
foramen, pubes and ischia forming symphyses, quadrate bones
fixed, jaws without teeth, but with cutting horny sheaths.
Trunk encased in a bony shell, composed of numerous dorsal and
ventral dermal bones, forming a carapace and a plastron, which
may or may not be covered with horny shields. Copulatory
organ unpaired, cloacal opening more longitudinal than round,
never transverse. Oviparous.

It is customary to distinguish the marine, paddle-limbed kinds
as TvTtleSy the others as Land- and Water-tortoises,

Tortoises occur already in the Trias. They reached their
greatest development towards the end of the Mesozoic and in
the earlier Tertiary periods. They are now comparatively
reduced in the number of families and genera, although they
are still represented by about 200 species. The sub-class as a
wliole is cosmopolitan, but does not occur in the colder regions.

Their origin is quite unknown. Of recent groups only the
Crocodilia and tlie Khynchocephalia come into consideration.
Combination of these groups with the Chelonia leads to some
unknown forms whence also the Theromorpha • have arisen.
Palaeontology does not lielp us, all the leading, main groups of
Chelonia liaving been in existence in the earlier Mesozoic ages ,

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CHAP. IX CHELONIA CLASSIFICATION 3 I 3

and Palaeozoic Chelonia are still unknown. We can, however,
to a certain extent, reconstruct an ideal primordial Chelouian
by assigning to it all the ancestral characters actually observed
in recent and fossil kinds, and by reducing to simpler conditions
those features which we know to be more or less exaggerated
specialisations. It is reasonable to assume that originally each
metamere, except those of the anterior half of tlie neck and the
posterior half of the tail, carried a transverse series of dermal
plates, covered with horny shields, while the trunk, according to
the greater bulk of the body, increased in size, converging towards
the root of the neck and tail. By concentration, reduction of
the number, and increase in the size of some of the remaining
plates and shields, the skull assumed its characteristic box -like
shape, the neck and tail becoming at the same time free. Chelonia
are without doubt descendants of terrestrial, or at least semi-
aquatic reptiles, and the marine paddled forms subsequently
developed from terrestrial kinds.

Olassiflcation of Chelonia. — After many vicissitudes it was
recognised that the Chelonia cannot naturally be divided
according to the modification of their feet. Tlie Tkionychoidea
were clearly separated from the rest by Stannius in 1854.
Cope, in 1870, was the first to emphasise the important
character of the mode in which the neck is either bent sidewards
(Pleurodika) or withdrawn in an S-shaped curve in a vertical
plane (Cryptodira) ; and he also separated Sphargis as Athecae
from all the other Chelonians, for which DoUo in 1886 proposed
the term Thecophora. The division of the latter into recog-
nisable families, based upon reliable, chiefly internal, skeletal,
characters, has been effected by Boulenger ; ^ and his classification
has been adopted in the present volume, after intercalation of the
more important fossil forms. The relationships between these
various families may perhaps be indicatt^d as follows : —

'Athecae . ... Sphargidae

/-p. ,. fPelomedusidae

Chelonia { \ Chelydidae — Carettoclielydidae

(Chelydridae — Dermatemydidae —
Cinostemidae
ty # y//n/u, n I* -. pij^^jyg^^^ernidae

[Testudinidae — Chelonidae
, Trionychoidea Trionychidae

* Cat, Chelonians, Brit. Mus. 1889.

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314 CHELONIA

The guiding taxonomic characters are fully mentioned at the
head of the difiFerent families, and are mostly internal The
following "key," adapted from Boulenger, and based upon ex-
ternal characters, is preferable for practical purposes.

For the i^osition and names of the horny shields see Fig. 61 on
p. 315.

Shell covered with homy shields.

Digits distinct, with 5 or 4 claws.

Pectoral shields separated from the marginals by infra-
marginals.

Tail long and crested. Plastron small and cruciform.
North America . . Chelydridae, p. 338.

Tail long, covered with rings of shields.

Plastron large. Indo-China Platysternidfie^ p. 345.
Tail short North and f Dermatemydidae, p. 341.
Central America . \Cino8temtdaef p. 342.
Pectoral shields in contact with the marginals.

Plastral shields 11 or 12, without an intergular.
Neck retractile in an S-shaped

vertical curve . . Testxidinidae, p. 345.

Plastral shields 13, an intergular being present.
Neck bending sideways uxidt^r f Ghelydidae, p. 399.
the shell . . \Pelomedu8idae, p. 390.

Limbs paddle-shaped, with one or two claws . Chelonidae, p. 378.
Shell without horny shields, covered with soft, leathery skin.
Digits distinct, broadly webbed, but with only

three claws ..... Trianychoidea, p. 404.
Limbs paddle -shaped.

Shell composed of regular series of

bony plates. Two claws . Carettochelydidae^ p. 404.

Shell composed of very many small plates arranged like

mosaic. No claw:* . . . Sphargidae, p. 333.

The vertebrae are, sometimes in the various regions of the
same individual, amplii-, opistho- or pro-coelous, or even biconvex.
Traces of the chorda remain longest in the middle of the centra,
lutercentra occur regularly on the first two or three cer^dcals,
and then again in tlie tail as paired or unpaired nodules, or as
short chevrons. The latter occasionally fuse with the caudal
end of their centra. Intercentral discs of fibrous cartilage occur
regularly in the neck and tail. The ribs develop originally in
the same transverse level with these discs, and frequently the
anterior thoracic vertebrae retain this intercentral or intervertebral
position throughout life. Farther back they often show a gradual
change from the intercentral to a more central and ultimately

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SKELETON

315

remarkable to a purely neural attachment. In all the Chelonia
the ribs are devoid of the tubercular portion.

The cervical vertebrae have no ribs, except mere traces in the
shape of small nodules. On the tail the ribs are often large, and,
when fused with their neural supports, look like transverse
processes ; the whole arrangement exactly resembles that of
Crocodilia. The first pair of thoracic ribs, those borne by the

Fig. 61.— Various plastra and their horny shields. 1, Tesliido iberu ; 2, Macrcudemmys
Ummincki; 3, Cinostemvm odtrratuni ; 4, Sterfiothaerus nigrvanji ; 5, Chelmiina
longicdlis ; 6, Ch^lone mydas. a or an^ Aual shield ; aftrf, abdominal shield ; / or
fe/n, femoral ; g or gui, gular, unpaired in Fig. 3 ; A or At/m, humeral shield ; i or
int.g, intergular ; im, iufra-raarginals ; m, marginals ; p ori><?f/, pectoral; a-, in Fig.
1, inguinal shield constituting, with the axillary xx, the last trace of infra-
marginals.

ninth vertebra, are peculiar. Tliey arise from the anterior
portion of the centrum, are much reduced, sometimes to mere
threads of bone, and lean against the anterior rim of the second
pair of ribs, in many cases without reaching the carapace. The
next following ribs, those of the tenth to the sixteenth vertebra,
are intimately involved in the formation of the first to seventh
castal plates. The ribs of the two sacral vertebrae sometimes
remain quite distinct throughout life, just toucliing the upper

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3l6 CHELONIA

ends of the iliac bones ; but since these find a much more effective
support in the shell, the distal ends of the sacral vertebrae
fuse with the eighth, or so-called last, pair of costal plates.

The neural arch of the ninth vertebra rests upon its
centrum; but the neural arches of the other trunk -vertebrae,
although long, rest upon two centra; retaining, like the ribs,
their original intercentral position ; and in most cases the
neuro-central sutures remain throughout life. The atlas and
the last cervical vertebra deserve special attention. In many
tortoises, e.g. Trionyx, Clemmys, Testudo, the three constituent
parts of the atlas, namely, the neural arch, the centrum, and
the intercentrum or first pair of united basiventralia, do not
ankylose, but remain loosely connected ; and the first centrum,
instead of forming an odontoid process, remains movably attached
to the second centrum, although it sometimes carries, and fuses
w^ith, the second intercentral piece. In other tortoises, e.g.
Platemys and Chelys, however, all the parts of the atlas co-ossify
and form a complete, solid vertebra which
articulates by a concavo-convex joint with
the centrum of the second vertebra. The
normal number of cervical vertebrae is
eight in all Chelonians. The first spinal
nerve issues between occiput and atlas,
all the others behind the neural arches of
^:iia.f;r^; '^ml^y. their vertebrae. The last, or eighth cer-
huTuiiu The second basi- vical, owiug to the retractility of the neck,

ventral (white) ia attached a ^ -, a, • • i. -j, trsi. 't^v.

to the posterior end of the f^rms elaborate jomts ; Its centre fits with
first centrum, which, not a knob into a cup of the ninth, and its

being fused with the second , , i» t i j

centrum, is not yet an post-zygapophyscs form broad, curved
odontoid process. 2, The articulating coucavc facets for the recep-

complete atlas of an adult . „ . t^ .Z.

Trionyx gangeticns, still tiou 01 the anterior zygapophyscs of the
typically temnospondyious. fi^ed ninth vertebra. In the Trionychidae

3, The first and second cer- •'

vical vertebrae of an adult the zygapophyses are most elaborate, and
p/^/.my.. 4 The complete ^^ ^^^^^ articulate with the ninth ver-

atlas of a Cheiys Jirnbnata. ^

tebra, while the centra do not join, but
remain, or rather become, separated by partial resorption. In
the Chelonidae, in conformity with the non-retractile and short
neck, all the cervical joints are much reduced.

The skull (cf. Fig. 54, H, I, K, p. 280) agrees fundamentaUy
with that of Sphenodon and of the Crocodilia, but it is

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SKULL

317

characterised by several special features. There are no ecto-
pterygoids or ossa transversa ; no lacrynial bones, no inter-
parietal or pineal foramen ; the vomer is unpaired and^ the
nasal bones are mostly absent, unless they are fused with the
prefrontals. The premaxillae are very small. The single vomer
forms a septum between the choanae ; and these are, except in
Sphargis, ventrally roofed over by wings sent out by the palatines.
The latter form a continuous bony roof to the mouth with 'the
pterygoids, and these diverge posteriorly, being connected suturally
with the quadrates, lateral and basi-occipital bones, and with
the unpaired basi-sphenoid, which appears between the basi-

FiQ. 63.— Skull of Chelone mydus.
A, from the left side ; in B, the
postfrontal and squamosal bones
have been removed, and the broad
expansions of the jiigal, quadrato-
jugal, parietal, and quadrate bones
have been reduced in order to re-
duce the skull to more primitive
conditions. f\ Frontal ; J, jugal ;
L.Of lateral occipital ; Mx, maxil-
lary ; Op, opisthotic ; Pal, palatine ;
Par, parietal ; Pr/, prefrontal ;
Pro, pro-otic ; Pt.f, postfrontal ;
Ptg^ pterygoid ; Q, quadrate ; Qj,
quadrato -jugal ; S.o, supra -occi-
pital ; Sq, squamosal.

occipital and the diverging pterygoids, but is in most cases to a
great extent overlapped by the latter. The occipital condyle is
distinctly triple ; the basi-occipital sometimes helps to border the
foramen magnum. The supra-occipital sends out a long vertical
blade, directed backwards and generally projecting far over the
neck, for the attachment of the powerful cranio-cervical muscles.
The quadrate is very peculiar. Firmly attached, and hemmed
in on nearly all sides by the neighbouring bones, it stands nearly
vertically and forms a broad articulating surface for tlie mandible.
Its posterior side shows either a transverse, horizontal groove, in
which lies the columella auris, or the groove is transformed into
a more or less closed canal. Moreover, the hinder lateral margin
of the quadrate forms most of the tympanic frame ; its margins
being curved backwards, leaving in the Cryptodira^ however, a

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3 I 8 CHELONIA

wide notch behind ; in the Pleurodira this part of the quadrate
is transformed into a trumpet, the wide rim of which, forming a
complete ring, carries the tympanic membrane. The tympanic
cavity thus formed often leads into a deep recess which extends
beneath the squamosal towards the opisthotic and bears some
resemblance to the intricate tympanic recesses which pervade
that region of the Crocodilian skull.

Dorsally the quadrate is broadly overlaid by the squamosiil,
which frequently forms an arch with the parietal Anteriorly
the quadrate is connected through a variably sized quadrato-
jugal with the jugal ; and this, by joining the maxilla and post-
frontal, helps normally to form the posterior rim of the orbit.
All the bones which border the temporal fossa vary much in
extent in the different groups of Chelonia. The extremes are
represented by Cistudo and Geoemyda, in which the bony infra-
temporal arch is absent, owing to the loss of the quadrato-jugal ;
and on the other hand by the Chelonidae and by Sphargis, in
which the whole temporal region is covered over by an additional
** false cranial" roof. This roof is produced chiefly by lateral
wing -like expansions of the parietal and postfrontal lx)nes,
which meet the likewise much expanded jugal, quadrato-jugal,
and squamosal bones. In the lower diagram of Fig. 63 (Cheloni
mydas) the squamosal has been removed, and the other bones
have been reduced to tlieir normal, or rather primitive condition,
for comparison with the external view of the complete skull of
the same animal. The lower diagram shows also the connexion
of the pterygoid with a descending process of the parietal ; this
column, paired of course, usually contains a separate bone, the
epipterygoid, the portion between Ptg and Par.

The hyoidean apparatus is well developed, and sometimes
assumes large dimensions, especially in Chelys. The two pairs
of " horns " are the first and second branchial arches, whilst the
hyoid arclies are reduced to a pair of small, frequently only
cartilaginous, nodules attached near the anterior corners of the
basis linguae, which generally fuses with the os entoglossum in
the tip of the tongue.

The pectoral arch consists of a pair of long coracoids sloping
obliquely backwards, the distal cartilages of which scarcely
touch each otlier in the middle line, and the scapulae. The
upper end of the scapula fretiuently touches the inside of the

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SKELETON

319

first costal plate, protected by a cartilaginous pad. Near the
glenoid cavity arises a long process (PC in Fig. 65), placed
transversely and approaching its fellow. The distal end is con-
nected with that of the coracoid by a fibro-cartilaginous band. The
homology of this scapular process is not quite clear. The band
just mentioned favours the idea that the process represents
the precoracoid, but its being an outgrowth from the scapula
suggests that it is merely the much enlarged acromion. It
certainly does not represent the clavicle, which forms part of
the plastron : and this is not in contact with the shoulder-girdle
at all.

Fio. 64. — Diagram of the skeleton of Testudo elephantopus, after removal of the left
' half of the carapace. The plastron is roughly indicated by a section through the
mitldle line. Fe, Femur, foreshortened ; Fi, fibula ; //, humerus ; 7/, ilium ; Is,
ischium ; P.P., pubis ; R, radius ; Scup, scapula ; Th, tibia ; w, ulna ; 3, third
cervical vertebra ; 1, 3, 5, first, third, and fifth fingers ; Xlir, thirteenth (fifth
thoracic) vertebra.

The pelvis is strong. Ilium, pubis, and ischium meet at the
acetabulum. The dorsal end of the ilium is generally broad-
ened, and is attached to one or both sacral vertebrae, but it is
also in contact with the superimposed last costal plate. This
additional connexion often becomes predominant and the sacral
vertebrae are partly or completely relieved of the iliac support,
fusing in this case more or less with the costal plates. The
pubes have strong lateral processes, directed obliquely forwards
and downwards. The pubes and the ischia, which latter are
much smaller, form broad symphyses, and these are connected
with each other by a longitudinal cartilaginous band {Chelone,
Trionyx) ; or the connecting bridge is broad and quite ossified
{Tesiudo)y forming in the latter case two roundish obturator-
foramina. Cartilage frequently remains at the anterior end of

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320

CHELONIA

the pubic symphysis, and a smaller, longer, and narrow piece of
cartilage extends sometimes backwards from the ischiadic sym-
physis, as the so-called h}^-ischium. In the Pleurodira the
ends of the ilia, and those of the lateral processes of the piibes,
are much broadened and firmly ankylosed with the posterior
costal plates and with the xiphiplastron respectively.

MalZ.

Fio. 65. — Ventral view of the bony shell of Chelone mydas^ the Greeu Turtle, after
removal of the plastron (Fig. 66). The costal plates are marked by cross lines to
distinguish them from the ribs. C, coracoid ; />, femur ; Fi^ fibula ; //, humerus ;
J/a.l-J/rt.l2, marginal plates, some of which are fused together ; A'm, nuchal plate :
PC, " precoracoid " ; /?, radius ; Sc, scapula ; 7, T", first and fifth digits ; /X, Ninth
vertebra or first thoracic.

The limbs are typically pentadactyle and complete, and are
most primitive in water-tortoises, e.g. Chehjdra and Emys, in
which the carpus consists of the typical ten . separate elements,
including the pisiform. In Testudo the centrale is fused with
the intermedium, and the first three distal carpals are also fused
together. In the marine turtles the limbs are transformed into
paddles, but all the bones retain their independence ; the pisiform

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CARAPACE

321

and the first metacarpal are enlarged and flattened, thereby
giving additional width to the paddle. The tarsus remains less
primitive ; the centrale and the proximal elements have a
tendency to fuse together, most com-
pletely in land-tortoises; the fifth
distal carpal is enlarged, and stands
out hook-like from the rest. The
number of the phalanges of the
fingers and toes varies slightly. It
is noteworthy that none of the
Chelonia possess more than three
phalanges. The three middle fingers
and toes have mostly three phal-
anges; the poUex and hallux have
always two; the number of phal-
anges of the fifth finger varies from
three to one, of the fifth toe from
two to none. The greatest reduc-
tion occurs in Testudo and its allied
genera of typical land - tortoises,
Homopus, PyxiSy and Cinixys, the
formula for the fingers being
2, 2, 2, 2, 2 or 1, and 2, 2, 2", 2, 0
for the toes. In Felomedusa all the fingers possess two phalanges
only, owing to fusion of the first and second phalanges with each
other.

The shell, which is the most characteristic feature of the
Chelonia, consists of the dorsal "carapace" and the ventral
" plastron." Each is composed of a considerable number of bony
plates which arise as ossifications of nearly the whole thickness
of the cutis, only a thin layer of subcutaneous connective tissue
remaining soft and lining the inside of the shelL We restrict
ourselves to a description of the shell of the Thecophora, leaving
the discussion of the peculiar shell of Sphargis to p. 336 f. Very
young tortoises are still soft, and the plates which are beginning
to ossify are not yet suturally united. The plastron (Figs. 66 and
67) consists of the paired epi-, hyo-, hypo-, and xiphi-plastral
plates, and the unpaired endo-plastral plate. The latter is homo-

* It should be noted that the homy pieces of the carapace are termed "shields "
and the bony pieces "plates."

VOL. VIII Y

FiQ. 66. — The bones composing the
plastron of Chelone mydas. On
the right side the position of the
covering horny shields^ is indi-
cated by dotted lines, a, Anal
horny shield ; a5, abdominal ; /,
femoral ; ^, gular ; A, humeral ;
i^jintergular ; im, infra-marginals ;
p, pectoral.

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322

CHELONIA

CHAP.

logous with the interclavicle, the epi-plastra are homologoufl
with the clavicles of other Reptiles, while the other pieces
are genetically derived from, and are further modifications
of, the so-called abdominal ribs of the Crocodilia and Prosauria.
These plastral plates are never in direct contact with the

FiQ. 67. — Bony shell of Testudo ibercu A, Ventral; B, dorsal; G, left-side view. In
B, and on the right half of A, the position of the homy shields is indicated by
dotted lines. The underlying bony plates are marked by strong lines. In B the
1st neural and costal plates, the 4th neural, costal, and 6th marginal plates, and
the 7th neural plate are .shaded. 1, 4, 6, First, fourth, and sixth neural plate ; Mt
in C fifth left maiiginal plate ; i\^u, nuchal plate.

shoulder-girdle or with any other parts of the internal skeleton.
In the young of all tortoises, and in the adult of the Chelonidae
and Trionychidae, the several plastral plates enclose large,
irregularly-shaped fontanelles. These are more or less filled up
in the other groups ; and in the Testudinidae especially the
whole plastron forms one continuous mass. The navel is situ-
ated between the hyo- and hypo-plastrals. Both these pairs are
broader than the others, and are connected with the carapace by

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CARAPACE

323

means of several marginals. The connecting region is called the
bridge. In several tortoises, e.g. Emys, the connexion with the
marginals is formed by ligaments only and remains movable.
In others, transverse, more or less perfect hinges are formed
across the plastron. A rather imperfect joint between the
hypo- and xiphi-plastrals develops with age in Testudo ibera.
In Cistudo and Cyclemys a very eflfective hinge lies below the
hyo- and hypo-plastrals, just in front of the bridge; and the
anterior and posterior lobes of the plastron can be closed

JVeunz/PleUe

CotCal
Shield

•Epidcim

>CuU8.

Fio. 68. — ^A, Diagrammatic transverse section through the shell of Testudo, On the
right side the homy shields have .been removed, on the left are shown the neural,
costal, marginal, and pectoral shields. The bony dermal plates are dotted. Cap^
Capitular portion of rib ; Sp.C^ position of spinal cord. B, Vertical section through
part of the shell, magnified and diagrammatic B, Bony layer of the cutis ; l,
leathery layer of the cutis ; m, cells of the Malpighian layer ; p, star-shaped pig-
ment-cells ; 8C, stratum comeum, composing the homy shields.

against the inner rim of the box, fitting tightly in Cistudo,
In Pyxis the front lobe only is movable.

The carapace is composed of one median series, a right and
left lateral series of costal plates, and a series of marginals which
surround the whole. The median series consists of one large nuchal
plate, normally eight neurals and one to three supracaudal plates.
The characteristic feature of the neural plates is that they are
firmly fused with the broadened neural spinous processes of the
underlying vertebrae. The nuchal plate lies in front of the
first thoracic or ninth vertebra; it overlies the last cervical
vertebrae, with the eighth of which it is connected by ligament
only ; but the posterior corner of the plate often fuses with the
spine of the ninth vertebra. In the Chelydridae, and still

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324 CHELONIA chap.

more in the Trionychidae, the nuchal sends out a pair of long
rib-like processes, which either extend to below some of the
neighbouring marginals, or their ends overlap those of the ribs
of the second thoracic vertebra (e.g. Trionyx), or, lastly, they
are in turn overlapped by the first costal plates (e.g. Cyclanorhis),
Such rib-like processes are also present, well developed in the
young, shorter in the adult, in the Dermatemydidae and Cino-
sternidae. It is possible that the nuchal plate represents the
fused neural of the eighth and the costal plates of the ninth
vertebrae. An indication of the compound nature of the nuchal
may be found in the fact that two nuchals have been described
in Chelydrojpsis caririata, a Miocene relation of Chelydra, Some-
what similar modifications have taken place in the post-sacral
region. The one to three supreicaudal plates are, namely,
neurals which have lost their connexion with, or perhaps have
never been fused with, the spinous processes of the movable tail-
vertebrae. The number of neural plates is mostly eight, but
there are sometimes individually nine or ten, the gradual
suppression taking place first in the sacral region. When such
a plate is suppressed the neighbouring costal plates usually close
up and meet in the median line. In Cistudo, for instance, there
are only seven normal neurals, the eighth pair of costals meet,
and the original eighth neural is transformed into a supracaudaL
In Cinosternum the sixth to eighth costals meet, separating the
one supracai;dal widely from the remaining five neurals. The
meeting of the last pair of costals, with co-ordinate reduction of
the neurals to seven, is almost universal in the Pleui'odira ; and
this tendency is carried out to an extreme in the Brazilian
Flatemys and in the Australian Chelodina and its aUies, in
whicli all the costals meet in the middle line, and the neurals
are completely suppressed. Every stage intermediate between
complete neurals {Sternothaerus) and interrupted, vestigial, and
vanished neiu-als, is still represented by some genus. This pro-
cess takes place independently, both in America and in Australia,
and is one of the most recently introduced modifications.

The costal plates arise, like the neurals, independently in the
cutis, but they soon come into contact with the underlying
cartilage of the ribs, which are long enough to reach the
marginals. The ribs flatten, become surrounded by the growing
membrane -bone of the plates, and the cartilage of the ribs,

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IX CARAPACE 325

instead of ossifying, undergoes a process of calcification. Ulti-
mately this is more or less absorbed, its place is taken by the
dermal bone, which forms so to speak a cast of the rib, pre-
serving in many cases the shape of the vanished rib, only the
capitular portions of which remain unaffected. The number of
costal plates is very constant, namely eight on each side, but
some fossils have nine or ten, and there are still individual
variations in recent forms, indicative of that number. In a
large Chrysemys concinna I find the last pair of costals clearly
composed of at least two pairs, and this same specimen has nine
distinct neural plates.

The marginal plates are originally paired, almost always
eleven pairs, very rarely ten or twelve ; an unpaired posterior
plate, the pygal, is always present, and is probably the result of
fusion. In the Chelonidae large fenestrae remain between the
costal and marginal plates, only covered by leathery unossified
cutis, and of course by the homy shields. In the Indian
fresh-water genus Batagur similar windows are gradually filled
up with age, and the horny shields become extremely thin and
almost confluent. On the other hand, in Testudo polt/phemus,
the bony shell, always very thin, becomes still thinner with
age and finally fenestrated by absorption.

Great reduction has taken place in the carapace of the
Trionychidae. The American species of Trionyx have only
seven pairs of costal plates ; in Cyclanorbis the neurals are
reduced to two. The whole dorsal shell is much smaller than
the body, and marginal plates are absent or merely vestigial.
It is doubtful if the ossifications in the posterior half of the
marginal flap of some genera are homologous with true marginals.

Externally the whole shell is covered, except in the IViony-
chidae,m Sphargis and Carettochelys,viit\\ horny, epidermal shields.
These are phylogenetically older tlian the dermal plates, and
they do not correspond with them either in numbers or in
position, although there exists a general resemblance in their
arrangement. On the plastron we distinguish an unpaired or
paired gular, and a pair of gular, humeral, pectoral, abdominal,
femoral, and anal shields (Fig. 66). Sometimes there are also
intergulars, paired in Macroclevimys and Chelys, unpaired in
Chelone ; in many of the Pleurodira an unpaired intergular lies
behind the gulars.

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326 CHELONIA chap, ix

The carapace of most Chelonians is covered with five neural,
four pairs of costal and twelve pairs of marginal shields, the last of
which often forms an unpaired pygaL In front of the first neural
lies the nuchal shield, very variable in size, often absent. The
Chelydridae, Dermatemydidae, Platystemidae, and Cinosternidae
possess moreover several inframarginals, intercalated on the bridge
between the marginal and some of the plastral shields. In many of
the other families these inframarginals are restricted to the anterior
and posterior comers of the bridge, as the so-called axillaries and
inguinals, mostly small and variable. Lastly, Macroclemmys has
several small supramarginals.

There are consequently eleven longitudinal rows of shields
in all ; by elimination of the supra- and infra-marginals they
are reduced to seven rows. It is absolutely certain that the
number of transverse rows also was originally much greater than
it is now. The mode of reduction of the number of the neural
and costal shields has been studied in Thalassochelys caretta (cf.
p. 388.) The accompanying illustration (Fig. 69) shows some of
the main stages actually observed in the reduction of these shields.
The chief point is that certain shields are squeezed out, or sup-
pressed by their enlarging neighbours. The ultimate result is
the formation of fewer, but larger shielda

Each shield grows individually as follows. Every year, or
rather during every periodic>ally recurring period of growth, the
area of the Malpighian layer belonging to each shield increases
peripherally in size, and at the same time produces a new layer
of horn. The original little shield, with which the tortoise is born,
remains for years, often throughout life, as the so-called " areola ; "
it increases in thickness owing to the new layer of horn added
from below, and peripherally the increase in size is indicated by
the overlapping concentric rings. Each ring represents a year's
growth, at least in tortoises which live in temperate zones, where
hibernation means a complete suspension of growth. It is not
known if the same applies to tropical species, which grow either
throughout the year, or which undergo one or more periods of
rest. The areola does not remain central ; the growth is uneven.
With age the oldest layers of the areola are frequently rubbed ofiP,
and the areola then appears enlarged. For the first dozen years or
so the annual rings can be easily followed, but when the creature
approaches maturity each shield adds very little to its growth,

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///«

Fro. 69. — Diagrams illustrating the progressive reduction of the horny shields in various
Chelonians. The shields, the fate of which it is desired to follow, are indicated by
distinctive shading. I. Hypothetical, primitive stage. Eight neural (including the
nuchal) and eight costal shields. Both neurals and costals lie in the same trans-
verse planes. II. -VI I. Successive stages in the reduction and suppression of various
shields, observed in specimens of Tfiahtssocheli/s, the normal contiition of which is
represented by VII. VIII. Six neurals and only four costals. The normal condi-
tion of Chelone. IX. The nuchal shield has become very small and the resulting
gap has been filled up by an enlargement of the first pair of marginals. This is the
. normal condition of most Cryptodirous tortoises. X. The first marginals meet in
front and the nuchal is either suppressed (Xa), e.g. in several si^eciea of Testudo^ or
it is surrounded by the marginals (X6), e.g, in StenwOuterris. (From Willey's
^ool. Results, 1899.)

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328 CHELONIA

and the rings become very fine, crowded and irregular. Only by
careful counting and comparison of the rings on the costals,
marginals, and plastrals, can a reliable average be arrived at In
some tortoises, e.g. Chrysemys, the whole outer layer of the shields
peels oflf periodically; only a thin smooth layer like mica or tracing-
paper remains, of course without any indication of rings. The
pigment is formed in the Malpighian layer, but it frequently
diffuses into the homy shields themselves, notably in Chelone
imhricata, which yields the beautiful " tortoise-shell." The colour
of the pigment is either black, yellow, or red, with resulting
combinations. The green colour, often so beautiful in baby-
specimens of Chrysemys, is optical, produced, axjcording to Agassiz,
by a network of black pigment, spread over a layer of yellow
oil.

Horny scales, sometimes forming spines, and covering a nodule
of dermal ossification, are also common on other parts of the
skin, especially on the limbs of land-tortoises, and also on the tail
of Chelydra, Sometimes the end of the tail is protected by a claw-
like nail, for instance in Pyxis. In some of the gigantic land-
tortoises, and in Cheloiie my das, this nail assimies large dimensions,
and several of the terminal caudal vertebreie are fused together
into a regular urostyle. In some subfossil specimens of Mauritian
tortoises, these ankylosed complexes are 12 cm. long and more
than 5 cm. broad !

Before leaving the description of the shell, it is worth while
to draw attention to the enormous correlative changes in other
organs produced by this case. Nearly the whole organism has been
altered. The hard, firm carapace has partly rendered the supporting
functions of the vertebral column unnecessary or impossible. In
many tortoises, especially in the large land-tortoises, the vertebrae
and the capitular portions of the ribs are reduced to mere bony out-
lines ; the reduction to thin paper-like bony lamellae proceeds with
age. The iliac bones find a better support in the costal plates ;
the contact with the sacral ribs is given up, and these ribs fuse
partly with the costal plates, or they are absorbed. The whole
mass of muscles of the trunk is completely lost in the region of
the shell, but traces of them exist in young specimens. Neck,
limbs, and tail can in most cases be withdrawn and hidden in
the shelL When this is not possible it is due to secondary
changes. The neck is withdrawn either by being tucked away

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IX REGENERATION — SENSE-ORGANS 329

sideways (Pleurodira ^), or by being bent in an S-shaped curve in
a vertical plane. In a left-sided profile-view of the animal, the
head represents the tail of the S. The neck is withdrawn by
long muscles, which are inserted into the ventral side of the
middle of the neck, and extend in the shape of vertical ribbons
far back into the shell, arising from the centra of some of the
middle or even more posterior thoracic vertebrae.

Lastly, a few remarks on the partial regeneration, or the
mending of injuries to the shell If part of the horny covering
is badly bruised, torn off, or rubbed through, or if part of the
shell is crushed, the underlying portion of the bony plate
becomes necrotic, and the horny covering also dies so far as its
Malpighian layer is destroyed. Soon, however, the uninjured
Malpighian cells, around the margin of the wound, multiply,
grow into and beneath the injured portion of the bone, and form a
new horny layer, casting off the necrotic portion. After several
months the deficiency is patched up ; new bone has grown in the
deeper remaining strata of the cutis, and the outside is covered
by a continuous horny layer, without, however, reproducing
the original concentric moulding of the shields. In badly
crushed shells sometimes almost one-third of the whole shell is
thus cast off and mended within one or two years. The re-
generation of the forcibly stripped-off shields of Chelone imhricata
is described on p. 386. Bitten-off tails and limbs, rather
frequent occurrences in water-tortoises, are of course not repro-
duced, but the wounds are healed and covered again with scaly
skin.

Sense-organs. — The eye is by far the best developed sense-
organ. It is comparatively smalL The pupil is round. The
iris is mostly dark in terrestrial forms, while in water-tortoises it
is often brightly coloured, for instance pale yellow in Chelodina,
greenish and mottled with black, pale grey, brown, etc., in
various species of Chrysemys. Cistudo presents a curious
sexual dimorphism ; the males have red, the females brown,
eyes. The sclerotic wall contains a ring of numerous small
ossified plates. There is no trace of a pecten. The eye is pro-
tected externally by the two lids and the nictitating membrane.
In some water-tortoises, notably in Chelodina, the lower lid is
transparent. Lacrymal and Harderian glands are present.

^ vXevpdvy side ; ^ct/wj, neck.

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330 CHELONIA CHAP.

The SENSE OF HEARING is apparently not very acute, although
tortoises and turtles are frightened by noise, and can distinguish
sounds ; otherwise they would have no voice, which is very tiny
and piping in most tortoises during the pairing season. In most
water-tortoises the tympanic membrane is thin and quite exposed ;
in land-tortoises it is often thick and covered by the ordinaiy skin ;
lastly, in Chelone the tympanic cavity is filled with a plug of
the much-thickened skin, possibly in adaptation to the water-
pressure when these creatures dive to considerable depths. The
ossicular chain is mostly reduced to a long, bony, columellar rod.

The SENSE OF SMELL is Well developed. All Chelonians care-
fully smell their food, in the air as well as under water. The
individual predilection shown by many species for different kinds
of animal and vegetable food, — since they are, for instance, able
to distinguish between the various sorts of cabbage, cauliflower,
sprouts, etc., — proves that they possess a considerable amoimt of
smell and taste.

Tortoises have a fine sense of touch ; even the slightest tap
on the shell is noticed, and the skin of the soft parts is extremely
sensitive. Tickling of the sides of the tail, or of the hinder
surface of a thigh, produces ridicidous scratching actions of the
same or of the opposite foot.

The digestive apparatus is simple. Only a few peculiarities
need be mentioned. The tongue is mostly broad and soft ; it
cannot be protruded. The oesophagus of the Chelonidae is covered
with many conical projections pointing towards the stomach. The
latter is simple, except in Sphargis. The intestine is devoid of a
caecum, but the difference between the small intestine and the
rectum is very marked and often abrupt. The cloaca is very
roomy. It contains the large copulatory organ, which is impaired,
grooved on its dorsal side, and is altogether constructed like that
of the Crocodilia. The large bladder opens ventrally into the
urodaeum, a recess of the cloaca ; near its base open the urinary
and genital ducts. Many water- tortoises possess also a pair of
lateral thin -walled sacs, the so-called anal sacs, dorso- lateral
diverticula of the walls of the urodaeum. These sacs, which
have highly vascularised walls, are incessantly filled and emptied
with water through the vent, and act as important respiratoiy
organs. Wlien such a water-tortoise, for instance an Emys or a
Clemmys, is suddenly taken out of the water, it squirts out a

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IX DIGESTION — RESPIRATION— EGGS 3 3 I

stream of this water, which is not, as is generally supposed, the
urine fix>m the bladder.

The mode of respiration is interesting. The lungs are very
complicated, highly - developed, spongy structures. They are
attached by their whole dorsal surface to the inner lining of the
shell. As they cannot expand through their own initiative,
and since the shell has made costal and abdominal expansion
impossible, the tortoise has to resort to other means of producing
the necessary vacuum. This is done partly by the neck and the
limbs, which act like pistons in being drawn in and out ; partly
by the greatly developed hyoidean apparatus, by which, when
the neck is stretched out, the throat is alternately inflated and
emptied, the air being swallowed, or pumped into tlie lungs.
Additional respiration, besides that of the anal sacs mentioned
above, is effected in various aquatic tortoises by slightly vas-
cularised recesses of the pharyngeal region. Most Chelonians
can exist for a very long time without breathing; sulky individuals
remain for hours or days under water. Cistudo can shut itself
up for an equally long time. Nevertheless this and other land-
tortoises easily get drowned.

All Chelonians lay white eggs, round or oval, according to
their kind, but the shape of the eggs of one set sometimes varies
within the greatest limits. The shell varies from a parchment-
like, flexible, scarcely calcareous cover to a hard, well-polished
case. As a rule the eggs, imbedded in the ground, are hatched
after a few months, but in some of the northern kinds, e.g. Emys
orbicularis, the hatching is deferred until the next spring, the
embryo's development being arrested during the winter. How
such eggs, buried a few inches only below the surface, withstand
the often very severe North German and Eussian winter is a
mystery. Whilst the plastron is generally flat, it is more or less
concave in the males of many species, notably in Testudo,
Cistudo, and Emys,

The general conclusions which can be drawn from the present
geographical distribution of the Chelonia are as few and unsatis-
factory as those applying to the Crocodilia, since all the main
groups of Chelonians, and many more extinct families, occurred
together in bygone ages in the same countries, for instance in
Europe. The marine forms are naturally cosmopolitan, but the
Testudinidae are likewise cosmopolitan, except in the Australian

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332

CHELONIA

CHAP.

region. The Chelydridae, now restricted to North and Central
America, occurred formerly also in Europa The Pleurodira, in
Mesozoic times plentiful in Europe, India, and North America,
are now restricted to South America, Australia, and Africa ; the

'i^^ ,..y\ .v"^ ■■•"-v.

v»"

9fiS DERMATCMYOAE.

i=v/mj TESTUDINIDAE. '//^^ CINOSTERNIOAE. 5^i!^ CHELYDRIDAE. ^ PLATYSTERNIDAE.

Fig. 70. — Geographical distribution of Cryptodirous tortoises.

Pelomedusidae to Africa, Madagascar, and South America ; the
Chelydidae to South America and Australia. In the latter
country all the Chelonians belong to the Chelydidae. The Triony-
choidea, occurring since the Cretaceous epoch in North America,
in Early and Mid-Tertiary times in Europe, are now restricted to

^^ ftLOMCDUSIDAE. Illl III CHELYDIDAE.

Fio. 71. — Geographical distribution of Pleurodirous tortoises.

North America, Asia, and Africa. The country richest in Chelo-
nians is America ; North and Central America together possessing
representatives of all the families except the Pleurodira, and
these we know to have died out there. The Dermatemydidae,

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IX GEOGRAPHICAL DISTRIBUTION ATHECAE 333

Cinosternidae, and Chelydiidae are now restricted to the Nearctic
sub-region (including Central America). Poorest in genera and
species, all of them Chelydidae, is the Australian region, where no
fossils of other families have yet been discovered. Europe, with its

^ />

■■ TRIONYOHIDAE. ^M CHELYDIDAE.

Fig. 72. — Geographical distribution of Trionychidae and Chelydidae.

few Testudinidae, does not come into consideration ; Asia has at
least Testudinidae and Trionychidae, and in addition the solitary
Platysternum in Indo-China, representative of a family whose
affinities with the Chelydridae again proclaim the validity of the
Periarctic region.

Order I. ATHECAE.

The vertebrae and ribs are not fused with, but are free from,
the carapace, which consists of numerous small polygonal plates
and is covered with leathery skin without any epidermal shields.
The limbs are transformed into paddles. The neck is not retrac-
tile. Marine,

Fam. Sphargidae. — Sphargis s. Dermatochelys coriacea, the
Leathery Turtle or Luth, is the only recent species and is the
largest of all recent Chelonians. The biggest specimen in the
national collection is about six feet and a half long, from the nose
to the end of the shell, which latter is about four feet long ; such
a specimen may weigh half a ton. Agassiz, however, says that
he has seen some " weighing over a ton." The general colour is
dark brown, either uniform or with yellow spots. The Leathery
Turtle has a wide distribution, ranging over all the inter-
tropical seas, but it is rare everywhere ; least so perhaps in the

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334

CHELONIA

CHAP.

Western Atlantic from Florida to Brazil and in the Indian Ocean.
According to Agassiz it breeds regularly every year in the
spring on the Bahamas, on the Tortugas, and on the coast of
Brazil, depositing its many eggs on the sandy shore like other
turtles. Accidentally it visits the northern coast up to Long
Island, and specimens, perhaps carried with the Gulf Stream, have
been caught on the coasts of Europe, for instance off Dorsetshire.
One was caught near Nantes in 1729, and is said to have made

buTTtWOITTHjo

Fig. 73. — Sphargis coriaceay the " Leathery Turtle, '
and dorsal views, x 1.

young specimeus, ventral

a terrible noise when being killed. This is perhaps the reason
why Merrem in 1820 invented the generic name Sphargis,
supposed to be derived from <T<f>apay€(o (I make a noise). It has
also been recorded from the Mediterranean. It seems to be
entirely carnivorous, living upon Molluscs, Crustacea, and fish.
The flesh is supposed to be imwholesome. It is a very curious
fact that of this rare species only large specimens, besides a very
few baby-turtles, are known or preserved in collections, while
individuals of intermediate size, say from four inches to thi-ee
feet in length, have never been recorded. If it were not for the
fact that they are still known to breed, it would look as if the

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IX ATHEC AE — SPHARGIDAE 335

species were dying out. Perhaps they are very shy, leading a
-pelsLgic life, diving at the least sign of danger, and coming near
the land only for the sake of breeding.

The structure of Sphargis is so peculiar in many respects that
it deserves a somewhat full account. The neuro-central sutures
I)er8ist on all the vertebrae. The eight cervicals are short. All
the ten trunk-vertebrae carry ribs, and these, with the exception
of the last, articulate between the centra and with the neural
arches ; the first and tenth ribs are short, the others are long
and flattened, but not broad, with wide spaces between them.
The tail is short, although it consists of about twenty vertebrae ;
these are devoid of chevrons.

The skull superficially resembles that of Chelone, chiefly owing
to the completely roofed-in temporal region. The supraoccipital
crest is rather short, covered completely by the parietala, the
posterior margin of which is rounded ofiF instead of forming, as in
the Chelonidae, a long projecting triangular crest with the supra-
occipital. The parietals are in broad contact with the post-
frontals, posteriorly they are just reached by the squamosals.
The quadrato-jugal is small, separated from the post-frontal by the
meeting of the squamosal with the jugal. The quadrate is
notched behind, and it separates the opisthotic from the
squamosal The basisphenoid is large and broad, extending far
forwards so as to separate the pterygoids widely from each other
except in their anterior portions, which, instead of sending a
lateral arm to the jugal and maxillary, as in Chelone, are widely
separated from these bones by the palatines. The choanae lie on
either side of the anterior half of the vomer, and are not roofed
over by ventral vomero-palatine wings.

The limbs and their girdles are essentially like those of the
Chelonidae, but are not derivable from them. The most re-
markable feature is the shell. The dorsal and ventral halves are
directly continuous, forming one unbroken case all round, which
is composed of many hundreds of little bony plates, irregularly
polygonal, fitting closely into each other with their sutural edges,
and giving the shell a beautiful mosaic appearance. On the
dorsal side are a median row and three pairs of lateral rows of
larger plates, and these form seven longitudinal blunt ridges
which all converge towards the triangularly pointed tail-end of
the shell. The ridges are not so much produced by thickened

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336 CHELONIA CHAP.

or spine-like edges of the plates, but by the right and left halves
of the plates being actually bent at an angle. This is most con-
spicuous at the sides of the shell where it passes into the ventral
portion. The latter has two pairs of lateral and one median
ridge. The whole shell has consequently twelve ridges. The
mosaic plates are deeply imbedded in the cutis, being externally
as well as internally covered or lined with dense leathery skin.
The epiderm is thin, and shows no indications of horny scales.
In young specimens the whole shell is soft and very imperfectly
ossified, later on it is quite rigid, although comparatively thin.
It is nowhere in contact with the internal skeleton, except by
a nuchal bone, which by a descending process articulates with the
neural arch of the eighth cervical vertebra.

The affinities of the Sphargidae and their position in the
system are still debatable. Whilst some authorities, eg. Cope,
DoUo, and Boulenger look upon Sphargis as the sole remnant
of a primitive group in opposition to all the other recent
Chelonia, Baur considered it the most specialised descendant of
the Chelonidae. Dames agreed with him. Van Bemmelen has
modified this view in so far as he regards Sphargis as the most
specialised Chelonian, but considers the differences between it and
the Chelonidae great enough to conclude that both Sphargidae
and Chelonidae represent two independent, partly parallel,
branches which have arisen from two different groups of
terrestrial tortoises. Case,^ from the study of Protostega and
other fossil forms, tends towards Baur's view. He believes
that Sphargis is the culminating form of a branch which
through Psephophorus and with Eosphargis has sprung from some
creature like Lytolomay which at the same time is the starting-
point of another branch which culminates in the genera
Thalassochelys and Chelone, while lastly a third branch contains
Protostega^ ProtosphargiSy and Psevdosphargis, In other words, he
considers them all Chelonidae. If he is right we have of course
no business to sepamte Sphargis with its fossil allies from the
rest of the Chelonia as " Athecae."

However, Case has not proved his point. It is easy enough
to understand that the characters of the cranium and plastron
of Sphargis are in a condition which by partial reduction can be
derived from that of typical Chelonidae. The structure of the

^ Jour II. Morph. xv. 1897, p. 21.

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SPHARGIDAE 337

cervical vertebrae, the absence of the marginal plates and the
peculiar articulation of the nuchal with the last cervical vertebra
can be explained as convergent analogies, just like the paddles of
Carettochelys. But the shell of Sjphargis is fundamentally different
from and not homologous with that of the others. Cope was
therefore quite justified in distinguishing the Sphargidae as
" Athecae " in opposition to the others which DoUo later on, by
contrast, named " Thecophora." Unfortunate names, since both
groups are undeniably in possession of a drjKri or shell. Both
authors meant, however, by Theca the epidermal shields, but even
this distinction is rendered invalid by Carettochelys,

The most reasonable explanation has been suggested by Hay.^
The mosaic polygonal components of the shell of Sphargis are,
so to speak, an earlier generation of osteodermal plates than the
later generation of longer and broader bony plates which in the
Thecophora come into contact, and fuse with, the neural arches
and ribs. The osteoderms of Sphargis belong to the same
category as the dermal ossifications in the scutes of Crocodilia,
whilst the plates of the carapace and plastron of the Thecophora
belong to the category of the abdominal ribs. Sphargis has the
first kind in its peculiar shell, the second kind in the deeper
lying plastron and in its neural plate. But it has lost, or
perhaps had never developed, the horny shields. The only
difficulty is, however, the presence of a plastron and of a typical
neural plate in Sphargis, This difficulty is not very serious.
The plastron is a very old institution. It occurs together with
the more superficial osteoderms in Caiman, and the nuchal plate
may be the oldest of all dorsals. We can scarcely imagine that
the direct ancestors of Sphargis had developed both kinds of
shells, and that comparatively recently the inner shell of the
carapace was lost, leaving only the nuchal plate. Fossils do not
support such an assumption. Undoubted ancestral forms of
Sphargis are very rare. Psephophorus of the Oligocene and Mio-
cene of Europe had a continuous mosaic shell much resembling
that of Sphargis ; Eosphargis is represented by a well-preserved
skull from the London clay. Then follows a wide gap until we
come to Fsephoderma of the Ehaetic, or Upper Trias of Bavaria ;
the large fragment of whose dorsal shell is composed of about 200
mosaic pieces. If this fragment really formed part of the shell

1 Atner. Natural, xxxii. 1898, p. 929.
VOL. VIII Z

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338 CHELONIA

of a Chelonian, its age would speak greatly in favour of the
Athecae being a very primitive and independent group.

Order II. THEOOPHORA.

Thoracic vertebrae and ribs united with a series of median or
neural and a paired series of lateral or costal plates. Parietal s
prolonged downwards, meeting the pterygoids directly or by intn--
position of an epipterygoid.

Sub -Order 1. Or]rptodira. — The carapace is covered with
horny shields, llie n^ck, if retractile, bends in an S-shaped cur re
in a vertical plane. The pelvis is not fiised with the shell.

Fam. 1. Chelydridae. — The plastron is small and cross-shaped
(Fig. 61, 2, p. 315) ; the bridge is very narrow, and the displa<ied
abdominal shields are widely separated from the marginals by a
few irregularly shaped inframarginals. The tail is long. The
limbs, neck, and head are so stout that they cannot be completely
withdrawn into the shell. Snout with a powerful hooked beak.
American ; only two genera, each with one species.

The temporal region is roofed very incompletely and only
anteriorly by the expanded parietals and postfrontals, which form
a long suture. The plastron consists of nine bony plates, a small
entoplastron being present ; there are lacunae in the middle line,
the plates meeting imperfectly, and the horny abdominal shields
are likewise separated by* soft skin. The carapace has a nuchal
with long rib-like processes which underlie the marginals ; the
neural plates form a continuous series. There are twenty-three
marginal plates. The pubic and ischiadic symphyses remain
separate, enclosing one large heart-shaped foramen. The five
fingers and toes are webbed and are protected by claws except the
outer toe, the nail of which is usually suppressed.

Chelydra serpentina, the Snapping Turtle, attains a large size,
namely, a shell-length of more than one foot, and a total length
from the nose to the tip of the tail of more than three feet.
Its range extends from the Canadian lakes east of the Rocky
Mountains, through the United States and Central America.
The carapace of young specimens has three very marked series
of keels, which gradually disappear with age, until in very old
individuals the shell becomes quite smooth. The skin is very
warty, especially on the neck, and there is a pair of minute

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IX THECOPHORA — CRYPTODIRA CHELYDRIDAE 339

barbels on the chin. The tail carries three series of originally
triangular homy crests, which with age are transformed into
blunt knobs. The general colour of this rather ugly creature is
olive, mottled with dark brown above and with yellowish below.

According to Holbrook the Snapping Turtle .is found in
stagnant pools, or in streams where the waters are of sluggish
motion. Generally they prefer deep water, and live at the bottom
of rivers ; at times, however, they approach the surface, above
which they elevate the tip of their pointed snout, all other parts
being concealed; and in this way they float slowly with the
current, but if disturbed they descend speedily to the bottom.
They are extremely voracious, feeding on fish, reptiles, or any
animal substance that falls in their way. They take the hook
readily, whatever may be the bait, though most attracted by
pieces of fish ; in this way many are caught for the market. It
is, however, necessary to have strong hooks and tackle, otherwise
they would be broken, for the animal puts forth great strength
in his struggles to escape, both with his firm jaws and by bring-
ing his anterior extremities across the line. When caught they
always give out an odour of musk, which in very old animals is
sometimes disagreeably strong.

Occasionally the Snapping Turtle leaves the water, and is
seen on the banks of rivers or in meadows, even at a distance
from its accustomed element. On land his motions are
awkward ; he walks slowly, with his head, neck, and long tail
extended, elevating himself on his legs like the Alligator, which
at that time he greatly resembles in his motions; like the
Alligator also, after having walked a short distance, he falls down
to rest for a few moments, and then proceeds on his journey. In
captivity they prefer dark places, and are exceedingly ferocious ;
they will seize upon and bite severely anything that is offered
them, and their grasp upon the object with their strong jaws is
most tenacious.

The Snapping Turtles, or " Snappers," are feared on account
of the ferocious bites which they inflict, and they are hated
iDecause of the destruction of valuable fish and water-fowl. They
in turn atone for this damage by being eaten, especially the
younger half-grown individuals, the flesh of the older ones being
too much tainted with the odour of musk. The round eggs,
which are laid to the number of twenty to thirty in the summer

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34Q CHELONIA

(in the Northern States about June), are likewise good to eat.
The first act of the young creature on leaving the shell is said to
be snapping and biting. In captivity they are often very sulky,
and refuse food stubbornly for many months, perhaps for a whole
year, and apparently without much harm to themselves, since
they lie quietly in the distant corner of the tank, now and then
slowly rising to the surface to breathe. Fresh-water algae grow
on the shell and in the mud which settles on it, and since this

Fiu. 74. — Macroclemmys temmincki, " Alligator Turtle. " x J.

happens also in the wild state, they are rendered as inconspicuous
as old rotten logs. In order to attract fishes they protrude a
pair of worm -like, pale pink filaments from the tip of the
tongue.

Macroclemmys temviiTicki, the " Alligator Turtle." — In size and
general appearance much like the other Snapping Turtle, but the
dorsal shields have each a strong and prominent keel, and these
three series increase in size with age. The costal shields are
separated from the marginals by an additional series of about four
supramarginals, well shown in the illustration. The shields of

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IX CHELYDRIDAE DERMATEMYDIDAE 34 1

the cross-shaped plastron are subject to much individual variation,
small shields being frequently intercalated, or rather retained,
between the usual ones, especially between the pectorals and
abdominals, in the gular region, and on the narrow bridge, where
the inframarginals number one to three or even more. This species
inhabits, broadly speaking, the whole basin of the Mississippi and
Missouri rivers.

This beast is as vicious as the other Snapping Turtle.
According to Agassiz it does not withdraw its head and limbs on
the approach of danger, but resorts to more active defence. It
raises itself upon the legs and tail, highest behind, opens the
mouth widely, and throwing out the head quickly as far as the
long neck will allow, snaps the jaws forcibly upon the assailant,
at the same time throwing the body forward so powerfully as
often to come down to the ground when it has missed its object.

It lives mostly in the water, but makes considerable journeys
overland. Both in the water and on dry land the limbs move
nearly perpendicularly, and the body is raised high. On dry
land a considerable part of the weight of the body is borne by
the long, strong tail.

" They are as ferocious as the wildest beast of prey, but the
slowness of their motions, their inability to repeat the attack
immediately, their awkwardness in attempting to recover their
balance when they have missed their object, their haggard look,
and the hideous appearance of their gaping mouth, constitute at
such times a picture as ludicrous as it is fearful and revolting.
Their strength is truly wonderful. I have seen a large specimen
bite off a piece of a plank more than an inch thick. They take
hold of a stick with such tenacity that they may be carried for a
considerable distance suspended to it free above the ground.
Fishes and young ducks are their ordinary prey. They lay from
twenty to forty or more round eggs only about the size of a small
walnut in holes which they dig in sloping banks not far from the
water " (Agassiz).

Fam. 2. Dermatemydidae* — The pectoral shields are widely
separated from the marginals by inframarginals, the gular shields
are very small or absent, and the tail is extremely short. Only
two or three genera, with three or four species in Central America.

The plastron is composed of nine plates. In Dermatemys
mawi it is large, firmly joined to the carapace, covered with

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342 CHELONIA chap.

eleven or more shields, and there are four inframarginals ; in
Staurotypus salvini of Mexico the plastron is cruciform, with the
anterior lobe movable, covered with seven or more shields, accord-
ing to the fusion of the anal shields and the presence or absence
of the gulars ; there are only two inframarginals. The pubic and
ischiadic symphyses remain separate ; the temporal fossa remains
widely open, the postfrontals scarcely touching the parietals.
There are 23 marginal shields in Staurotypus, 25 in Dermutemys,
including the unpaired nuchal. The nuchal plate has a pair of
rib-like processes like those of the Chelydridae, but some of the
posterior costal plates, sometimes only one pair, meet in the
middle line, overlying or supporting the corresponding neural
plates. The shell of these aquatic tortoises is rather flat, more
or less keeled, especially in young specimens, and in the fully
adult condition is about one foot in length.

Fam. 3. Oinostemidae, represented by the single genus
Cuwsternum, with about ten species in North and Central
America, and one in Guiana. Closely allied to the two previous
families, with which it agrees by the separation of the pubic and
ischiadic symphyses, the presence of an ento-plastral plate, the
possession of inframarginal shields (Fig. 61, 8, p. 315), the widely
open temporal fossae, and the rib-like pair of processes to the
nuchal plate. It agrees with the Dermatemydidae in the inter-
ruption of the neural plates by the meeting of several pairs of
the costal plates. Tliere are 23 marginal shields ; five or four
shields, according to the presence or absence of the gular on the
plastron, and in some species these plastral shields become, with
age, more and more separated from each other by soft skin (see
Fig. 75). The shape and size of the plastron differ considerably
in the various species ; in most of them, e.g. in C. jyennsylvanicnyn
and C. leucostoynum, but not in C, odoratum, the anterior and
posterior lobes are movable, with transverse soft hinges, so that
the animal can completely close its shell. The skin of the legs
and neck is so baggy and loose that these parts slip in, the skin
rolling off, when the creature withdraws into its shell. They
lay only a few — from three to five — elliptical eggs, which have a
shining, glazed, and thick, but very brittle shell.

Cinostei-num odoratum, the Mud-Turtle, or Stinkpot Terrapin,
so called on account of the disagreeable smell which exudes from
the inguinal glands. The head is disproportionately large, with

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CINOSTERNIDAE

343

the snout rather compressed laterally, and pointed underneath,
with several short barbels. The neck is long and slender. The
ciirapace of the young is keeled, each of the neural shields being

Fio. 75. — Cinosternum odoralwii, young specimens, x |.
A and B, males ; C, female.

raised in the middle line ; but in full-grown specimens the shell
l)ecomes quite smooth and rounded The horny shields of the
plastron are relatively largest in the young, but they soon leave
ever-increasing spaces between them, which are then filled with
soft skin only, which thinly covers the underlying bone. Tlie

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344 CHELONIA

fore- and hind-limbs, especially the latter, are extensively webbed,
and are provided with five short claws. The general coloiu:
of the shell is horny brown, either uniform or with darker spots
or streaks. The neck and limbs are mottled brown. The only
ornamental colouring is a pair of clear yellow broad lines on
each side of the head, and a similar streak on each side of the
lower jaw. On the chin and upper throat are two pairs of small
tentacles. The tail of the male is of about the length of the
hind-limbs, while that of the female is so short that its tip
scarcely reaches beyond the hinder margin of the carapace.
Length of the shell of full-grown specimens between four and
five inches. Very young specimens have a rather droll appear-
ance, owing to the long and slender neck with the large head,
and the humpy back.

This species is common in the eastern half of North America,
from Canada to Texas. It is mainly aquatic, and is one of the
dullest and shyest species. My own specimens spend most of
their time in the water, invariably in the darkest corners, pre-
ferably under a stone or a log, and they do not leave their hiding
places until dark, in searcli of worms, meat, and all sorts of
animal food. For months I could never induce them to take
food from a stick, or even to eat in my presence, and it was not
until after many weeks that one of them at last protruded its
head far enough to exhibit the yellow stripes. When taken out
of the water they draw in their heads, just allowing the vicious
little eyes to be visible, and opening the sharp-edged mouth
widely to bite deliberately and furiously at the unwary finger.
Some spent the winter in the water, in the greenhouses, feeding
as usual, others crept on land, hiding under moss, half buried in
the soil, where they slept for several months, but with inter-
ruptions in order to soak and to drink. When spring is well
advanced they prefer the water for their regular sojourn. Some
which had been sent over from New York arrived in a deplorably
dried-up condition, the skin being quite flabby and shrivelled,
but after a few hours' soaking they came round, and increased
considerably in weight, the limbs and neck becoming turgid.

C. peyinsylvanicum of Eastern North America has a larger,
more oval plastron. The head is not so strikingly large as in the
other species and, like the neck, is brown with yellowish spots, and
often has streaks on the sides. The tail of the male ends in a

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IX PLATYSTERNIDAE — TESTUDINIDAE 345

nail-like horny point. The lobes of the plastron are well hinged
in the adult.

C. leucostomum of Central America is larger, with a shell-length
of six inches. The plastron is not at all cruciform, but has a
broad bridge, and fills the box, moreover it has an anterior and a
posterior hinge, so that the box can be completely closed. Hence
the vernacular name of the Box-Terrapin.

Fam. 4. Platystemidae, represented by the single species
Platysterniim megacephalum in Burma, Siam, and Southern China.

The pectoral shields are widely separated from the marginals
by inframarginals, the plastron is large, oblong, not cruciform,
and the tail is long.

The plastron consists of nine plates, and is covered with six pairs
of shields, the most anterior of which are the broad gulars. The
nuchal plate has no rib-like processes. The neurals form a con-
tinuous series, and there are twenty-three marginal scutes. The
temporal fossae are completely roofed over, owing to the long sutures
formed by the parietals with the postfrontals, moreover the post-
frontals expand laterally so much that they posteriorly come into
broad contact with the quadra to- jugals and squamosals, anteriorly
with the maxillaries, so that the jugals are completely surrounded
by bones, and are shut off from the orbits and from the temporal
fossae. This is a unique arrangement, found nowhere else in
Tortoises. The pubic and ischiadic symphyses are connected
with each other by ligaments only.

The general appearance of this water-tortoise is rather curious,
since the carapace is much depressed, looking, especially in
younger specimens, as if it had been crushed in. The head, pro-
vided with very strong hooked jaws, is strikingly heavy and
large, and is covered above with one single large shield. The tail
is longer than the shell, which, in full-grown specimens, reaches
about six inches in length ; it is, throughout its length, covered
with rings of squarish shields. A large specimen measures 14
inches in total length, of which only five fall to the shell.

Fam. 6. Testudinidae. — The shell is always covered with
well-developed horny shields. Those which form the plastral
bridge are in direct contact with the marginals. The plastron is
composed of nine bones. The digits have four or five claws.
The neck is completely retractile. The skull is devoid of parieto-
squamosal arches.

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34^ CHELONIA

This large family is cosmopolitan, with the exception of the
Australian and the adjoining Austro-Malajan countries. It
contains genera which form a continuous gradation between
absolutely terrestrial and thoroughly aquatic tortoises ; and many
are truly amphibious. As a general rule the typically terrestrial
kinds have a more curved or arched shell, the digits are short,
the eggs are more oval or round, and they are chiefly herbivorous :
the essentially aquatic kinds have a flatter or depressed shell,
webbed feet, with longer, often slender claws, the eggs are more
cylindrical, and they live on animal diet. About 20 genera,
with more than 110 species, are recognised by Boulenger, but
their essential characters are nearly all internal, and therefore
of no avail for the determination of live or entire specimens.

Chnjsemys. — One of the most typical and widely distributed
genera of American Terrapins or water-tortoises. The carapace
is flat ; the plastron is quite immovable, with a strongly developed
bridge. Feet well webbed. Tail short. Skull with a broad,
complete, lateral, temporal arch. About one dozen species, mostly
in the eastern half of the United States, but the whole genus
ranges from Canada to Argentina.

Most of the young Chrysemys are very pretty, the ground-colour
of the upper shields being green, variegated with yellowish-brown
or blackish markings, which often form exquisitely delicate
patterns, either concentrical {Ch. co?ici7i7ia, Ch. ruhriventris), or
more longitudinal {Ch. elegans), or apparently quite irregular.
The ground-colour of the plastron is yellow, but the various
species are best distinguished, at least in very young individuals,
by the arrangement of the dark brown spots and patches. There
are, for instance, several pairs of bold lateral and several median
patches in Ch. ruhriveyitris ; five pairs of ocellated spots in Cli.
eleyans ; only small median patches, where four plastral shields
meet, in Ch. coiicinna ; while the plastron of Ch. jiicta is uniformly
yellow.

These water-tortoises are very lively and shy, most so perhaps
Ch. jplcta, which is very quick and active. The food varies, often
according to individual fancy. Most of them eat fish. Ch. picta
is partial to insects, but it also takes worms. Some of my
specimens refused meat for a long time, but ultimately they
became so fond of it and of worms, that they came out of the
pond to take the food from the fingers ; those in the Zoological

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TESTUDINIDAE 347

Gardens of London have developed a taste for biscuits. One of
my largest Ch. concinna fasted deliberately for eight months,
refusing worms, insects, meat, and frogs, only occasionally sniffing
at the food, until it was tempted with whitebait, which it took
greedily. It refused, however, smelts and pieces of soles, but
after another month it condescended to take meat regularly.
Very young individuals live chiefly on flies, which they watch
for near the surface of the water ; and they are fond of smooth
caterpillars, maggots, the larvae of humble-bees, and similar soft
creatures. They all spend most of their time in the water,
preferably floating near the surface, hidden between weeds ; and
they are fond of basking. Some of them spend the night in the
water, lying motionless on the bottom, with heads and limbs
turned in. Others prefer hiding under mos& Those species,
which, like Ch, concinna and Ch, picta, are common in the North,
are of course perfectly hardy. For the winter they dig them-
selves holes in the banks near the water, and they do not come
out again until the spring is well advanced. The eggs are hard-
shelled, mostly long and oval, and they are hatched before the
end of the summer. The larger species of Terrapin are eaten.

Ch. picta (Fig. 76), the "Painted Terrapin," of the Eastern
United States, e.g. of New York and Long Island, is easily re-
cognised by the much depressed shell, which is absolutely smooth,
and without a trace of a keel. The colour above is dark olive-brown
or blackish, with broad yellow bands across the anterior ends of the
neural and costal shields. Three or four of these transverse bands
are very conspicuous. The marginals are red, with more or less
concentric black and yellow markings. The pretty red colour,
with some black stripes, extends over the bridge, but the plastron
itself is uniformly yellow. The soft parts are likewise prettily
marked, the ground-colour is black-brown, with delicate bright
yellow and red stripes on the sides of the neck, limbs, and tail.
The stripes are originally yellow, but they develop an orange or
red line in the middle, so that each red stripe is ultimately
narrowly edged with yellow; or the yellow and red stripes
alternate, for instance on the tail, which is short, narrow, and
pointed. The head is further adorned with a pair of con-
spicuous bright yellow patches behind the eyes, and a smaller pair
on the occiput. The black and yellow stripes run across the gape
of the mouth, some of the lines even looking as if they had been

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348

CHELONIA

painted across. The nuchal shield is elongated and very narrow,
its anterior edge and that of the neighbouring marginals are finely
serrated. Very young individuals are at once recognised by the
prominent longitudinal median stripe of bright orange extending
over the nuchal and neural shields ; the yellow transverse Imnds
are still absent ; they appear when the longitudinal line vanishes.
The " Painted Terrapin " is one of the few species of which,
thanks to L. Agassiz,^ complete data of growth from the new

^„n«f^-|»i

Flo. 76. — Chryscmys jji'dOy " Painted Terrapin.

H^qDi'Tt/d.,

born to old age are known. During the first six or seven years
the rate of growth is so uniform that numerous specimens
collected at the same time are readily arranged in sets of the
same age, simply by the differences they show in their size. The
successive lines of growth on the shields indicate the number of
years. After the seventh year the age is much more difficult to
distinguish in those tortoises, which, like Ch, picta, have a
perfectly smooth epidermis. This smoothness is due to the fact
that the shields undergo a process of moulting. An upper, quite

* Contrihutions to the Xat. Hist, of the U.S.A.j Boston, 1857.

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TESTUDINIDAE

349

transpareut layer of each shield peels off completely like a piece
of mica. I have been able to confirm Agassiz' statement on Ch,
concinmi in their third and fourth springs, and on a number of
adult Ch. picta. The latter were not allowed to hibernate, being
kept in a warm tank ; they peeled completely during the late
autumn, and then the red and yellow colours underlying the
newly formed shields appeared very vividly ; others moult at
midsummer.

Growth of Ch, picta, after Agassiz.

Year.

Length of

Breadth of

Height of

Length of

carapace.

box.

tail.

inilliin.

Miillim.

millim.

Second . . . , 26-5

16-5

Third

39-5

17-5

Fourth .

21-5

20-6

Fifth

23o

21-5

Sixth

23-5

Seventh .

26-5

Eighth ( 6 )

72-5

27-5

Ninth ( 6 )

27-6

Tenth ( 6 )

Eleventh ( 6 )

28-5

Fourteenth ( 6 )

74-5

28-5

TAventy-fifth ( 9 )

121

Old 9 .

129

Very old 9

168

113

The size of the eggs varies considerably, from 26 by 17 to 30
by 1 6 millimeters ; sometimes they are perfectly round, 1 7 mm.
in diameter.

Ch. concinna. — The specific character by which this Terrapin
may be easily recognised is a pair of orange - red broad
streaks, which extend from above the eye to the sides of the
neck. The general colour is olive-brown above, variegated with
yellowish dark -edged lines, which, together with numerous
rugosities, radiate from the middle field of each shield. The
plastron is yellow, often with blackish symmetrical patches, and
sometimes these become confluent and preponderant. Very young
specimens are extremely pretty, the ground-colour of the carapace
being green, each shield with darker, somewhat concentric
markings, most conspicuous and regular on the upper surface of
the marginals, where the marks of the adjoining shields form
one pattern-system across the dividing lines. The plastron is
either uniform yellow or has a few pairs of blackish spots

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350

CHELONIA

CHAP.

which stand so closely together that they form almost median
patches.

The carapace is rough. The horny shields become very thin
with age. The anterior margin of the small nuchal and the
neighbouring marginals is faintly serrated. The posterior
marginals form slight notches or indentations between their
edges. The plastron is almost square behind. The edges of the

FiQ. 77. — Chrysemya concinnuy in its
third summer, x 1.

Fig. 78. — Chrysemys coticinncL, in its
third summer, x 1.

The

jaws are nearly smooth, without hook and receiving-notch,
tail is short.

This species inhabits the South-Eastern States of North
America, from Missoiu:i and North Carolina to the Gulf of
Mexico. Very large female specimens have a shell sixteen inches
in length. The eggs measure from 33 by 25 to 39 by 25 miii.
or about 1 .^ inch in the long diameter.

Emys. — The plastron is movably united to the carapace hy
ligament, and in the adult has a slightly flexible hinge across the
middle, between the hyo- and hypo-plastral plates and the pectoral
and abdominal shields. The plastron is large, but does not quite
close the box. Besides the small nuchal there are twelve piiw

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TESTUDINIDAE 35 1

of marginal shields. The head is covered with smooth skin ;
the temporal arch is complete. The limbs are extensively
webbed. The tail of the very young is nearly as long as the
shell, but it becomes relatively shorter with age, being reduced in
the males to about two-thirds, in the females to half the length
of the shell. Only two species are found in Europe, the other,
E. blandingi, in Canada and north-eastern U.S.A.

H, orbicularis s. enropaea s. hitaria, the European Pond-
tortoise. — The shape and coloration of the shell change likewise
much with age. In the very young the shell is round, and the
shields are rough and slightly keeled, uniform dark brown above,
black below, with a yellow spot on each marginal and plastral
shield. When half grown the dorsal shields become quite
smooth, and are striated or spotted, with yellow upon a dark
ground. The head, limbs, and tail are dark, with yellow or light
brown spots and small dots. In very old specimens all these
yellow marks disappear on the shell, which then becomes uniform
lirown or almost black. The coloration is subject to much local
and individual variation, and there are two main types, the
spotted and the radiate. It is difficult to say which of the two
is the prettier. One male which I caught in the Alemtejo was
very beautiful. The shell was almost black with a greenish shine
when in the water, and had many bright yellow and whitish
spots. In the radiate type the yellow is sometimes pre-
ponderant, so that each shield becomes a study of delicately
painted yellow, brown, and blackish lines radiating from the
centre. This variety seems to prevail in the south of Spain,
decidedly so in the Marismas, also in Northern Italy, whence
most of the European markets are supplied. The largest shell in
the British Museum is 19 cm. = 7^ inches long. Fischer
Sigwart received one from Naples which was about 9 inches
long, and this seems to have been kept as a pet, since its shell
had been gilt. Specimens about 5 inches in length may be con-
sidered as fully adult. There are very few reliable obsen-ations
on the growth of individuals. One of F. Sigwart's grew in
eleven years only about 2*5 cm. = 1 inch, when its shell was
13*4 cm. = 5^ inches long — total weight of the tortoise 491
grammes, about 1 lb. One of my own grew fi:om 11 to 13" 2 cm.
shell-length, and 8*3 to 10*6 cm. in width within eight years,
but this was one of the specimens which, living in a greenhouse.

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352 CHELONIA

did not hibernate. This European pond-tortoise is now restricted
to Southern and Middle Europe, extending eastwards towards St.
Petersburg and into Asia Minor, southwards into Algeria.
Formerly it had a much wider range, having been found in
post-glacial deposits in Southern Sweden, Denmark, the Nether-
lands, and in East Anglia. Specimens have been found in
the peat of the fens of Norfolk and Cambridgeshire, con-
temporary with bones of the Beaver, Roe-deer, and Pelican.
The same applies to North Grermany, where its gradual dis-
appearance from the western and central parts is obvious.
Except in Central France it is now practically unknown to the
west of the Elbe river. The country between the Elbe and Oder
is now debatable ground, Emys being exceedingly rare. Some
fifty years ago this seems to have been different, to judge from
the fact that farmers were rather fond of keeping a tortoise in the
water-troughs of the cattle to keep the water free from worms and
other impurities. Hence arose a silly superstitious custom. It
was considered equally conducive to the health of the pigs to keej)
a tortoise in the foul tub into which all the dish-water and
kitchen-refuse — as potato-peels, sour milk, etc., — were collected
before the mess was given to the pigs.

A specimen is still occasionally caught in the Havel and
Spree rivers. I myself have heard of one or two in the back-
waters of the Oder near Frankfurt, but they are vanishing, and
it is difficult to say exactly why. The universal lowering of the
water-level owing to better drainage cannot quite account for it,
since there are thousands of suitable ponds, swamps, and back-
waters left. In Poland and in Eastern Prussia the tortoise is
still common.

This creature lives on a strictly animal diet. Worms, insects,
frogs, fishes form its main sustenance. Fishes are regularly stalked.
The tortoise watches its opportunity, slowly it half crawls, half
swims along the bottom, rises imperceptibly by a few gentle move-
ments of the widely spread-out webbed feet, then opens its sharp
cutting jaws wide, and makes a grab at the belly of the fish.
Frogs are most easily stalked when they sit upon a floating leaf.
The tortoise rises from below, and often waits with the nostrils
and eyes just above the water and close to the frog. After a
while it sinks, and rises again, this time actually touching the
toes of the non-suspecting frog, smelling at them and deliberately

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VOL. VIII

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354 CHELONIA

biting with a sideward turn of the head. What the jaws have
got hold of is not allowed to escape again. The tortoise holds on
and tears the prey to pieces with the sharp-clawed fingers. This
takes a long time, only the scraped-off flesh and the intestines
being eaten. The skeleton remains and sinks to the bottom,
while in the case of a fish, the air-bladder floats away on the
surface, and remains there as one of the surest signs of the exist-
ence of tortoises in that locality. The bones are cleaned with
wonderful neatness. Some of my grass-snakes shared this fate,
their backbones, with the hundreds of pairs of ribs, being picked
or rather scraped clean, scarcely less well than if they had been
prepared for a museum.

As a rule the prey must be in motion to be seized, unless tlie
tortoise has watched it before, and even then the latter prefers to
smell it before biting. In captivity they soon learn to eat meat,
and they become very tame, but in their native haunts they an?
extremely shy and cautious. Fond of basking upon a stone or
on the banks, with the four limbs sprawling, or with the hind-
limbs stretched backwards, and with the webs spread out so as to
offer as large a surface as possible to the rays of the sun, they lie
motionless for hours and appear fast asleep. But the slightest
noise, or any other sign of our approach, is sufficient to send them
plumping into the water, and to make them scuttle along with
unsuspected agility. Nothing but the audible plump of the flat
body and the widening rings of the disturbed water indicate their
presence. After a long time of waiting we give it up, and turn
away. That very instant we see a little ripple, caused by the
withdrawing of the tortoise, which had come to the surface and
had been watching us, with only the nose and eyes peeping out of
the water, the rest being concealed between the floating vegetation.
Apparently they cannot see us well with their eyes still under
water, owing to the difference of refraction, otherwise they would
not peep out and then at once turn back. It is certainly not for
the want of air, since they can remain below for many hours
without breathing.

Although they generally feed in the water, they come on land
when tame and hungry enough to take the offered food. Some-
times they make long migrations, perhaps because their old home
is dried up or does not yield food enough. They hibernate during
the cold season, buried in the mud, and they do not appear until

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TESTUDINIDAE 3 5 $

the spring is well advanced. During the pairing season, on warm
spring nights, they emit short piping sounds, and when they
have found each other, the couple swim about together. The
white, hard-shelled, long, oval eggs, averaging 25 to 15 mm.,
and about ten in number, are laid on land. This is a very
laborious and curious business. The female having selected a
suitable spot, not loose sand, but rather hard soil free from grass
and other dense vegetation, prepares the ground by moistening it
from the bladder and the anal water-sacs. Then it stiffens the
tail and bores a hole with it, moving the tail but not the body. The
hind-limbs then scoop out the hole, the broad feet moving alternately
and heaping up the soil on the side, imtil the hole is about five
inches deep, that is as far as the hind legs will reach. The eggs
are laid at the bottom in one layer, divided and distributed by
the feet. Lastly, the soil is put in again, and the tortoise, by
repeatedly raising its body and falling down, stamps the soil firm
and flat, roughens the surface a little with its claws, and leaves
the nest to its fate. Nothing but an accident leads to its dis-
covery. The young are hatched, according to locality and thie
kind of season, either in the same autumn or not until the next
spring. Eggs laid in a garden at Kieff, in Eussia, were hatched
eleven months later. This implies hibernation of the embryo
within the egg, and this is probably the usual course of events,
resembling the conditions of the development of Sphenodon (cf
p. 299). The pretty little creatures, scarcely larger than a shilling-
piece, are exceedingly difficult to rear. They require a tank
with green vegetation, stones to bask on and to hide under, and
also dry ground and moss for a change. They eat flies, tiny
worms, tadpoles, etc., greedily enough, but for some occult
reasons they do less well than many another kind of water-
tortoise. Miss Durham has, however, succeeded in rearing one,
which is now in its foui^th year; the shell is 2 inches long,
and each shield shows three annual rings around the areola.
This specimen spent the winters in an unheated room under moss,
not in the water.

E. UaTidingij the North American species, has a more elongated
and decidedly higher carapace than its smaller European relation.
The carapace is dull black with many pale yellowish spots ; the
plastron is yellow, with a large dark patch on the outer and hinder
corner of each shield. The head is dark brown above, bright

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356

CHELONIA

yellow below and on the throat, a contrast which gives this
tortoise a striking appearance. This species is extremely voracious,
becomes easily tame, and spends a great part of the day on land,
hiding under grass to avoid great heat, and withdrawing into the
water for the night.

Clem my s. — The plastron is immovably united with the
carapace, and is devoid of any transverse hinge. The skull has
a complete bony temporal arch. This genus, consisting of eight
species, is otherwise very much like Emys, and is truly Periaretic.

C. Icprosa s. siffris (Fig. 79). — The upper jaw has a median
notch for the reception of the upturned point of the lower jaw ;

Fig. 80.— Skull of Clemmys lejyrosa, x f . A, dorsal view ; By ftvm the left side ; /'.
frontal ; J, jugal ; J/, maxillary ; Par^ parietal ; Pr./, prefrontal ; /*//, postlrontal,
Qy quadrate, Qj, quailrato-jugal ; *S^, squamosal.

the cutting edges of the powerful beak are smooth. The shell is
flat and long-oval, nowhere serrated. The plastron does not quit^
fill the box. In the young the shell is nearly round, and the
horny shields form three series of keels, of which the lateral pair
disappear early ; the shields are olive-brown, each with an orange
spot or streak ; the plastron is dark brown, with a yellowish
margin. The adult looks very different. The shell has become
much more oval, with the greatest width behind the bridge. The
long shields are smooth, and in elderly specimens are without
any trace of the original connective rings of growth. The general
colour of the shell is uniform pale olive-grey, inclining to yellow
on the plastron. The ground-colour of the soft parts is olive-
grey, but the sides of the head are adorned with orange -red or
yellow marks, the patch between the eye and ear and three or

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IX TESTUDINIDAE 3 57

four stripes on the neck being especially conspicuous. The limbs
have pale yellowish streaks. All these markings are, however,
subject to much individual variation. While, for instance, the
half-grown creatures are distinctly agreeably coloured, often with
a rich brown, nicely sculptured shell, and with conspicuous orange
and yellow marks on the skin, the very old ones become rather
ugly, the prevailing colour varying more and more into dull
uniform pale olive-grey.

The " Iberian Water - tortoise " is typical of the Iberian
Peninsula, and extends through Morocco and Algeria far into
North- Western Africa. Unknown to the north of the Cantabrian
range, decidedly scarcer than its cousin Emys in the northern
half of the Peninsula, it becomes common in the south. In the
Alemtejo, in the lower parts of Andalucia and in Morocco, there
is scarcely a pool, stream, or river in which it is not found,
feeding on any living thing it can master, although fishes and
frogs are its principal prey. When the streams and watercourses
run dry, during the hot and dry season, the tortoises crowd to-
gether into the remaining pools, which soon become stagnant
and filthy. But even these havens of refuge are not of lasting
avail. They are soon cleared of anything edible, and the stink-
ing water becomes dirtier and hotter day by day. Ultimately
the tortoises leave the pool to hide under ledges of rocks, where
they aestivate for months. Tliis life in the muddy, slimy pools
renders these tortoises peculiarly liable to the attacks of a certain
fresh-water alga, which enters through the cracks in the horny
shields and then flourishes in the Malpighian layer, and even in
the underlying bone itself. This becomes gangrenous in patches,
and the whole shell assumes a leprous appearance, hence the
specific name of leprosa. Everything combines in favour of this
destructive little alga. The tortoise, covered with mud, basks in
the hot sun, the horny shields become brittle and crack, often
peeling oflF in thin flakes. But those happy individuals which
inhabit permanent rivers, or pools which do not dry \\\), are, and
remain, as clean as other water-tortoises.

C. leprosa has a most disagreeable, offensive smell, something
like concentrated essence of fish, due to the secretion of a
pair of large glands situated beneath the skin of the inguinal
region, and opening behind the bridge. Freshly caught specimens
stink horribly, but when they have be(;c)me accustomed to being

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358 CHELONIA

handled, they no longer void these glands. They always with-
draw into the water for the night, and the cold season is spent in
the mud. Their time of propagation is still somewhat douhtfuL
Very young tortoises are met with in the Peninsula in March,
when they are already in the rivers. Those which I imported
in the summer and autumn invariably dug their nests and laid
their long, oval eggs (28 to 33 mm. long) in the month of
November, pairing having taken place some two or three months
previously. The mode of making the nest is exactly the same as
that described for Emys, As most of my specimens were kept in a
greenhouse with a permanent current of warm water through
their tanks, they never hibernated, nor did they pass through a
torpid time in the summer, but they showed an in^esistible love
for the hot-water pipes, huddling together by the dozen, so that
the pipes had to be screened off to prevent the creatures from
getting burnt. Until this precaution was taken, they heat«d
themselves so much that the shields and even the bones of the
plastron were injured. The artificial warm temperature and the
complete suppression of seasonal rest had no bad influence, most
of the tortoises living with undiminished appetite for more than
twelve years, but the sexual period became disturbed, pairinu'
occurring ultimately at all times of the year. The eagerness of
the males, however, had a peculiar evil secondary influence upon
the females. The male tries to fasten on to its mate by bitinj,'
into the collar-like fold of the neck into which the head is with-
drawn, and this repeated irritation produces sores and swellings,
which latter in their turn prevent the female from wiping the
eyes with the ])ack of the fore-limbs, a habit common to most, if
not all, tortoises. Ultimately the eyes fester, and the tortoise,
becoming practically blind, falls off its feed, leaves the water,
which makes matters worse, and is very difficult to cure.

In other res^jects they are very hardy, and tliey .stand
acclimatisation in England perfectly. Some, thriving in a deep
concreted pond, passed through the very severe winters of several
years ago, hiding in the mud below the ice, and appeared in
the spring in perfect health. They can also successfully pass the
winter under moss and a heap of loose garden-rubbish.

C. ca.ynca is closely allied to C. lejrrosa, which it represents in
the Balkan Peninsula and in Asia Minor. It differs from the
south-western species chiefly by having the cutting edges of the

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TESTUDINIDAE 359

upper jaw finely denticulated, and by its prettier coloration,
each shield being ornamented with yellowish streaks which form
a kind of 00 on the costals, and a ring on the marginals. The
plastron is black in the young, with yellow and black patches in
the adult. The head and sides of the neck are striped with
yellow lines, narrowly edged with black, and the rest of the soft
parts is marbled dark olive and yellow. A few other species
occur in China, Japan, and North America.

Clemmys insculjita, one of the American species, ranging from
Maine to Pennsylvania and New Jersey, is easily recognised by
the peculiar reddish-brown and brick-dust colour of the soft parts.
The strongly keeled, posteriorly eraarginate carapace is reddish
brown, with radiating yellow lines. Each shield is delicately
sculptured. The plastron, which is notched behind, is yellow,
with a large black patch on the outer corner of each shield.
Length of a full-grown specimen 8 inches. They frequent the
rivers and ponds, but are also very fond of leaving the water,
sometimes remaining for months in dry places.

Malacoclemmys of North America, with three species only, is
closely allied to Clemmys, from which it differs chiefly by the
very broad alveolar surface of the upper jaw, and by the more for-
ward position of the entoplastron, this being placed anteriorly to
the humero-pectoral suture. We mention this genus since one
of its species, M, terrapin, is so extensively eaten in the Eastern
United States. The shell is oval, slightly emarginate behind,
obtusely carinated along the middle line. The upper parts of
the shell are brown or greenish, with dark concentric lines ; the
marginals are yellow below, each with a ring of dark grey, and
forming a peculiarly up-turned rim. The plastron is yellowish,
either with concentric stripes and dusky lines or uniform yellow.
But it is the colour of the soft parts which gives this otherwise
dull-looking creature its delicately pretty appearance. The skin
is, namely, greenish white with countless small black dots. The
males remain much smaller than the females, and have the con-
centric stripes more pronounced. This species, the choicest of
the edible Terrapins, frequents the salt marshes of the east coast
of North America, from Ehode Island to the Gulf of Mexico,
being most abundant around Charleston.

The following is a condensed account of an article wliicli
appeared in the New York Sun, 18th September 1898, the data

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360 CHELONIA

of which were supplied by the manager of the terrapin-farm at
Beaulieu, Georgia. The continued hunting and the unfailing
demand for them are making them very scarce, so that enter-
prising men have established terrapin-farms or " crawls " for the
keeping and breeding of terrapina The " crawls " in question
are near the river. The larger is 310 by 60 feet, and is
divided into three compartments for three sizes. The smaller
"crawl" is for the babies, and is 100 by 8 feet. Through both
" crawls " runs a ditch connected with the river and making a
circuit of the farm. The bottom of the " crawls " is on a level
with the low tide, and is covered with a layer of mud al)Out six
inches deep. Into this the terrapins burrow in the winter. The
average population of terrapins is about 40,000, one half
"bulls" and the other half "heifers." The latter are much
better eating, and grow to a much larger size, namely, eight
inches on the plastron, while the " bulls " rarely grow over five
inches long. When a female reaches six to eight inches it is
called a " count." Those between five-and-a-half to six inches
long are known as " two-for-threes," while those from five to five-
and - a - half inches are known as " halves." They are fed
exclusively on shrimps and crabs on account of the flavour,
although they will eat almost anything. The 40,000 consume
on an average twenty bushels of crustaceans a day. They are
quite indifferent to cold. The manager saw some placed in a
block of ice and frozen ftist to it ; after four or five days they were
chopped out, thawed, and were soon as lively as ever. The
statement that it takes these terrapins only seven years to attain
full commercial growth is surprising, and is probably an under-
estimate. At the end of the large " crawl " is a board to enable
the females to creep into a sand-pit, where they lay the eggs
from April to June, eight to twelve forming a set. It is necessary
to get the babies away from their parents as soon as they hatch,
else they will be eaten. The young must not be exposed to the
cold. The old ones have a large amount of curiosity. The best
way of catching them is for two men to go out in a boat with a
net. They row c^irefully along until they come to a likely spot.
Then one man raps several times sharj^ly on the boat with a
stick, and if there are any terrapin about they will come to the
surface just as fast as they can get there to see what is going on,
and the other man scoops them up with a little net. Another

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TESTUDINIDAE 36 1

way, used in the salt marshes, is for the negroes to go tramping
through the mud and water. If they pass any terrapin these
will rise out of the mud to see what the disturbance is. The
captives are then fattened in the " crawl." When the men go
in to feed them they whistle, and terrapin from all over the
" crawl," thousands of them, come swimming through the water,
piling over each other in their efforts to get close to the man
with the shrimps and crabs.

Cistudo, — The plastron, without forming a bridge, is connected
with the carapace by ligaments, and is divided into two movable
lobes, the transverse hinge being so perfect that the box can be
completely closed after head, legs, and tail have been withdrawn.
The nuchal shield is very small ; the first four neurals are large
and broad, the fifth much broader than long. There are twelve
pairs of marginal shields. The carapace is high and arched.
The digits are almost completely free. The tail is very short.
The skull is without a bony temporal arch, the quadrato-jugal
and the jugal being absent. Only two species, in North
America.

(7. Carolina of the Eastern United States is a very interesting
species. Closely allied by its internal structure to the water-
tortoises, it has become absolutely terrestrial ; and the shape of
the head, the convex shell which is coloured black and yellow or
orange-brown, and the short webless fingers are all terrestrial
features. But the rather long toes, provided with long and sharp
claws, the broad and flat feet, enlarged by a broad fold of skin on
the outer margin, the long oval eggs, the smooth covering of the
head, and the preponderant animal diet, still proclaim the aquatic
relationship of this tortoise. It is in fact a genus which has
changed habits and features from aquatic to terrestrial life. The
head is covered with a smooth skin, and the upper beak, especially
in old specimens, is strongly and broadly hooked. The eyes of
the males are red, those of the females are brown. The plastron
of the males is concave, that of the females is flat. Liirge
females reach a length of nearly six inches. The young are
nearly round, with high, arched back and prominent keels. The
keels of the middle line remain a long time, but they gradually
flatten down with age, being prominent only at their posterior
ends. Each dorsal shield is originally nicely sculptured, with
a well-marked areola and concentric rings. Very old individuals

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362 CHELONIA CHAP.

become much flatter on the top of the shell, but the sides remain
steep, so that the whole shell roughly resembles a somewhat '
oblong box with the corners rounded oflf, and the whole upper
surface rubbed down quite smooth. The variations of colour
are almost endless, and they occur in the same localities. I
liave a number of all ages from Long Island, near Xew York.
The half-grown are beautifully reddish or orange-brown with
dark patches, median keels prominent, plastron uniform black-
brown. In others the dark -brown prevails over the lighter
markings, which are yellower and more spotted or dotted than
patched. Some of the oldest, with quite smooth shells, are
black, with small, round, light yellow spots. Others are vermi-
culated or striped with yellow and black. The soft parts ysltv
to the same extent, some showing on the neck a lieautiful
intricate pattern of yellow, reddish and brown, while in others
these colours are arranged more or less in longitudinal stripes.

These " Box-tortoises " are often caught in the States and kept
as pets in the gardens, and their owners mark them by cuttini:
their initials into the plastron. These marks heal up and
widen in time like letters cut into the bark of a tree. One of
my specimens, certainly a very old one to judge from his hooked
beak, perfectly smooth and flat shell, and from the condition of
the marginals, which have the edges rubbed down quite smooth
and rounded off, has two initials and the date 1837 on its
plastron. Of course there is no proof that the date had been
cut in that year, more than sixty-three years ago, but it was
done a long time ago. The scars on those parts of the shell
which touch the ground are almost effaced, and the letters and
figures have become somewhat distorted owing to the usual
unequal, not concentrical, peripheral growth. Moreover, this
tortoise must have been already adult, although not quite fully
grown, since the marks are large and were evidently put in such
a size and position as to fit the available space. I may mention
that this record tortoise was, when I got it, not kept in confine-
ment, but had been picked up at large.

These Box-tortoises become very tame. Although fond of
drinking quantities of water in long and slowly repeated draughts,
they do not go into the water, and if they fall in accidentally
they are liable to get drowned. They enjoy a mixed diet, but
animal food predominates, consisting chiefly of snails, the shells of

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TESTUDINIDAE 363

which are passed, slugs, earthworms, maggots, and soft caterpillars.
Their fondness for slugs is all the more remarkable since scarcely
any other Vertebrate eats these slimy, sticky molluscs ; but a Box-
tortoise will make a meal of two or more fat specimens of the
1)lack slug Arion, and it will eat dozens of small slugs. It
first deliberately smells the prey, turns the head sidewards and
gives a bite, whereupon first the intestines and then the rest
are eaten. The slime is later on scraped off with the fore-
limbs, or the head is rubbed against the grass. The favourite
time of feeding is towards dusk or in the early dewy morning,
and they are especially lively during a soft, warm rain. They
also relish various kinds of fungi and fruit, for instance half-
rotten bananas. Close observation of their habits gives us
indications as to how the change from carnivorous to herbivorous
ha])it8 may have taken place. Accidentally many a blade of
grass is bitten off and swallowed together with the molluscs, also
])its of rotten wood and moss, and their excrements are often full
of such more or less digested matter. They are not very fond
of basking, although they love warmth, creeping into the grass,
where they make a shallow form hy moving the shell backwards
and forwards. During the cooler nights they frequently retire
into a hole or under a log of wood. They require to hibernate. If
kept in a warm house they become restless in the autumn, refuse
food, drink and feed again after some weeks, but are liable to die
during the winter. If they can find a cool place they bury
themselves and sleep for several months. If left out of doors
they dig into the ground, creep into a hole, at the bottom of
which they half bury themselves, or they hide under a heap of
garden-rubbish well out of the reach of frost. Warm April days
bring them out, and the first recjuirement is a drink.

When walking about in search of food they assume a curious
attitude, with the shell well above the ground, the long neck
stretched out and raised high. Their temper varies individually.
Some become tame readily and lose all shyness, and creep up to
their friend to take food from his fingers. Others are decidedly
shy and sulky, withdrawing with a hiss into the shell, which in
some specimens shuts almost hermetically all round, and they do
not come out until all imaginary danger is past. One of my
males sulked thus for several months, at least we never saw any-
thing of it except the closed shell, but it did not starve itself.

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364

CHELONIA

Propagation takes place in the summer, the long oval hard-
shelled eggs being laid in June and July.

The TYPICAL Land-tortoises are easily recognised by their feet.
The digits are short, have not more than two joints, and are
without any trace of webs ; the metacarpals are scarcely longer
than broad. The hind-feet are club-footed. The skin on the
anterior side of the fore -limbs is covered with strong homy
scales, frequently with dermal ossifications. The plastron is
united suturally by a broad bridge with the usually strongly
arched carapace. The skull has complete postorbital and

temporal arches. The top of the head
is covered with shields. The tail is
short. There are only a few recent
genera, modifications of the central and
typical genus Testudo. The latter is
cosmopolitan in the warmer temperate
and tropical regions, except in the
Australian and Austro-Malayan countries.
Cinyxis (Fig. 82) with a few species
in Tropical Africa from the Gambia and
from Abyssinia to the Equator is re-
markable for the unique modification of
its ciirapace, the posterior portion of
Fig. 81. — Skull of Testudo which is movable, the hinge passing
tileoSap^^^^^ between the seventh and eighth marginal

and the fourth and fifth costal plates,
externally behind the seventh marginal

In the

Jl/, maxillary ; Op^ Opis-
thotic ; Pr.f, prefrontal ;
/'r.o, prootic ; Pt.f, post-

froutai ; y, (juadrate ; s.o, and the secoud costal shields.

siipra-occipital. . _ __ « , , , , , . . •

middle of the back the hmge is im-
perfect, the parts being merely flexible enough to permit the
posterior half of the box to be closed. The head is covered
with shields.

O. helliana, of Northern Tropica.1 Africa, has a small nuchal
shield, and the margin of the carapace is smooth. Length of
shell up to seven or eight inches. C. homeana, of West Africa,
has likewise a small nuchal shield, but the posterior portion of
the carapace descends vertically, and the marginals are strongly
reverted and serrated. C. enmi (Fig. 82), also from West Afriai,
has no nuchal shield ; the marginals are reverted and serrated,
but the posterior part of the carapace is sloping, and the anterior

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TESTUDINIDAE 365

portion of the plastron is strongly forked in front, and projects
beyond the anterior border of the carapace. This peculiar
creature reaches a length of nine inches. When withdrawn
within the shell, which is closed behind and depressed in front,
with the jagged edges of the plastron and the anterior marginals
protecting the drawn-in head, it has a very quaint appearance.
It lives entirely on fruit and other vegetable matter, and is said
to prefer to lie in the water, while C. helliana is supposed to be
entirely terrestrial.

Fig. 82.- Cinyxis erosa. x J.

Fijxis arachnoideSy of Madagascar, a small land-tortoise, only
four inches in length, has an immovable carapace, but the front
lobe of the plastron is hinged.

Testudo. — The plastron is immovable, except that in old indi-
viduals of some species, e.g. T. ibera, the hinder lobe develops a
transverse flexible hinge. They have existed since the Oligocene
of North America and Europe ; and are now represented by nearly
forty species in all the tropical and warmer temperate countries
excepting the Austro-Malayan and Australian region. Typically
terrestrial, herbivorous and frugivorous, although occasionally
varying their diet with worms, molluscs, and insects. The eggs
are hard-shelled, mostly less oval than those of the aquatic and
semi-aquatic tortoises. The males generally remain smaller than
the females, have a slightly longer tail, and have a concave
instead of a flat plastron. Most land-tortoises hibernate in the
ground during the cool and cold seasons, or they aestivate during
the hot and dry months of tropical countries, but this is not an
invariable rule.

T. graeca, the common " Greek Tortoise." The shell is very
convex, without keels, and has a smooth, not serrated margin.

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366 CHELONIA

The nuchal shield is narrow. The fifth or last neural shield is
much broader than the others. The supracaudal is usually
divided in the median line, so that this is really the last pair of
marginals. The plastron is notched behind ; the axillary and
inguinal shields are smalL The scales on the anterior surface of
the fore-limbs are small, and form from half-a-dozen to ten longi-
tudinal rows. The hinder surface of the thigh is quite smooth.
The tip of the tail ends in a conical, homy spur. The colora-
tion of the shell varies somewhat, but the ground-colour is
yellow, each shield with a dark brown centre and irregular
patches or confluent spots towards the margin. The plastron
has an irregular, broad black border. The soft parts are grey-
yellowish. Some specimens are rather pale, almost lemon
yellow with little black; others incline towards orange with
more or less black. The middle fields of the shields of young
specimens are granular, although this area is rubbed smooth
with age ; but the rest shows clearly marked concentric lines of
growth. The eyes are dark, with a brown or bluish tinge,
sometimes inclining to dark grey in very old specimens.

Full-grown females have a shell six inches in length. This
species inhabits the northern half of the Balkan Peninsula, parts
of Asia Minor and Syria, Italy, and most of the islands of the
Mediterranean, from the Grecian Archipelago to the Balearic
Islands.

T. ihera is closely allied to T, graeca, from which it difiers
chiefly in the following points. The last pair of marginal
shields are fused into an unpaired supracaudal, the median line
of division being almost obliterated. The fifth neural shield is
not broader, and generally a little narrower than the others.
The posterior lobe of the plastron develops with age a transverse
ligamentous hinge, and is thus rendered slightly movable,
especially in the females. The posterior margin of the carapace
is slightly expanded in old specimens. The scales of the
fore-limb are large and imbricating, and form only four or h\e
longitudinal rows. On the middle of the exposed posterior
surface of the thighs the skin carries a strong, conical, horny
tubercle. The coloration is much like that of T. graeca, except
that the yellow of the young inclines to pale olive. Some
specimens are uniform brownisli. This species reaches a much
larger size than T. ijracca, old females often measuring eight inches.

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TESTUDINIDAE 367

rarely more than nine inches in length. Its home is Morocco
and Asia Minor, extending into Persia. It also occurs in certain
parts of Southern Andalucia, where it breeds regularly, for instance,
in the sandy pine-forests of the Marismas, near the mouth of the
Guadalquivir. Whether it has been introduced from Morocco, or
is indigenous, is an open question. Its specific name refers to
its Iberian home.

T. marginata is worth mentioning, since it is the Greek
tortoise, although not that of the European markets, which are
supplied by the other two species. 2\ marginata is restricted
to Greece proper, where it is the only land-tortoise. It is less
closely allied to T. graeca than to T. ihera, of which it may be
called an exaggerated form. The posterior margin of the
carapace is much expanded or flanged, and serrated. The
supracaudal is undivided, the posterior lobe of the plastron
is movable, but the large conical spur on the thighs is absent.
The dorsal shields of adult specimens are black with a small
yellowish patch ; the ventral shields are yellowish, each with a
large black triangular patch. The British Museum possesses a
shell 28 cm. = 11 inches in length.

The habits of these Moorish and Greek tortoises are very
much alike, and since they enjoy the distinction of frequently
being kept as pets in gardens, where they are allowed to look
after themselves, a great many incidental and odd observations
have been made on them. They are essentially vegetable feeders,
but their taste varies individually and with the season, also ac-
cording to the vegetation of the country they happen to come
from. Most of them enjoy juicy plants, for instance, lettuce and
cabbage; the flowers of the dandelion attract them not merely
by their bright colour ; clover is also a favourite food, and an
enclosure of grass-land with clover in it is soon cleared of the
latter ; grass is also taken, in default of anything better. Some
of my specimens gradually bite large holes into gourds and
pumpkins ; and in Morocco I found them in the autumn feeding
entirely on the terribly astringent green fruits of the dwarf
palm Chamaerops humilis. The hirger specimens bolted the
fruit with the stones, passing the latter. In close captivity
they often learn to take and to like bread soaked in milk or
water. They drink slowly and at length, but scarcely ever when
they have succulent food. There is one thing which they do

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368 CHELONIA CHAP.

not eat, namely, " black beetles," although they are warranted to
do so by the men who hawk them in the streets. Worms,
slugs, etc. are often mentioned as part of their occasional diet,
but I am not aware that any of the hundreds which I have
watched have taken such creatures, in spite of every opportunity.
Their habits are very regular. They learn to know the geo-
graphy of their domain thoroughly, and the spot selected for
sleeping will be resorted to over and over again, be it under-
neath some broad leaves, under a bushy fir-tree, between a
cluster of wallflowers, or between some tussocks, or even in an
almost bare corner, the attractions of which are not at all
obvious. Although their mental capacities cannot possibly be
called brilliant, they soon learn to distinguish between different
persons, and they will come up to be fed ; but their memory for
localities is surprising. Here is only one instance. A tortoise
which had been put into an outhouse for hibernation was six
months later taken to its usual large enclosure, and in the
afternoon it tucked itself away on the top of a mound under
precisely the same low bush where it used to sleep during the
previous autumn. It could not see that spot from where it had
been put down, and it did not meander about during the day,
but after having enjoyed the warm sun it made straight for its
favourite place. Dr. Girtanner of St. Gallen in Switzerland
testifies to their appreciation of music. When the town-band
began to play on the square adjoining his garden, all his
tortoises crept as fast as possible towards the fence and remained
there motionless with heads and necks erect. When the piece
was finished they moved about, but when the next number
began they were again spellbound. This he has observed, not
on one but on many occasions. That they can hear, although
tlieir ears are not visible, but covered by the ordinary skin, is
obvious enough from the fact that during the pairing season
they emit feeble piping sounds.

They are extremely fond of basking in the hot sun, some-
times allowing themselves to be almost baked in it, but then
ai^ain at other times they seem to be anxious to seek the shade.
They rise late and go to bed early, l>eing absolutely diurnal.
In the summer they leave their quarters when the sun is well
up, making for a sunny spot to graze. Then they lie still and
bask, unless a shower causes them to retreat under shelter.

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IX TESTUDINIDAE 369

After some hours' rest they feed again, and in the afternoon,
long before sunset, they go to bed. Some winters in England
are of course much more severe than any which these tortoises
experience in their native countries. Still they manage to sur-
vive them, provided they find a place which they can burrow
into, deep enough to be out of the reach of frost ; and if there is
a heap of mould, rotting weeds, and leaves, they are probably
safe. Sometimes they are restless, coming out again in unusually
mild winters without, however, taking food. If they appear too
early in the spring, they run the risk of terrible colds on
prolonged wet and cold days, but in the autumn they are
hardier, and can stand several degrees of dry frost.

The pairing season begins in May, but lasts far into the
summer. In Morocco I found them pairing as late as the
month of September. The preliminaries extend over many
days. The male becomes unusually active, makes a piping
sound, runs after the female, draws in its head, and knocks
with its shell against that of the female. This is repeated
many times, until the female is excited enough to raise itself
upon its hind-limbs. The eggs, only two to four in number,
are laid several weeks later, and are buried in the ground.
They are roundish-oval, hard -shelled, and vary according to the
size of the female. Those of T. graeca measure on the average
30 by 24 mm.; those of a large specimen of T. ihera 32 to
36 by 30 mm. The newly - hatched little creatures are still
quite flexible, and apparently soon bury themselves before be-
ginning their active life in the ensuing spring.

The age which these tortoises can reach is quite unknown,
but there are reliable data of individuals having been kept for
many years. Rumpf^ kept two T, graeca in his garden at
Frankfort-on-the-Main, and let them hibernate in a box with
hay in the cellar. One lived 33, the other 23 years. The most
famous specimen of T. ihera is " Gilbert White's Tortoise," '^ which
had been kept for more than 40 years before it came into his
possession. It used to bury itself in November and to come out
in April. It died in 1794, having reached an age of fifty-four
plus an unknown number of years, since there is no record of its
size when it came to England. The same applies to every other
specimen which has been, and is being, observed as a pet. My

^ Zool. Garten. 1892, p. 260. '^ Natural History of Selhome.

VOL. VIII 2 B

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370 CHELONIA CHAP,

largest Morocco female, which has a shell 7 inches long, shows
at least 25 concentric rings of growth on the shields; the
last half-dozen rings are very narrow, while some of those of the
central area have been rubbed down. This creature is not im-
probably 30 years old. A small female, which is only 5|- inches
long, has already 14 rings on its still perfect shields. Lastly, a
little one, only 4 inches long, shows 7 rings. They grow fastest
when they are about 6 to 7 inches long, and they then seem to
be at their prime. White's tortoise, now enshrined in the
National Collection, was unusually large, the shell measuring 25
cm., or nearly 10 inches; around the much-enlarged, rubbed-
down areola of each shield are about 30 very narrow rings.

T, horsfieldi is easily recognised by its possessing only four
claws on the fore- and hind-limbs. It is closely allied to the
species last mentioned, which it seems to represent in the
sandy districts of Transcaspia and the Kirghiz Steppes to
Afghanistan.

T. eleganSy the " Starred Tortoise " of the southern half of
India and Ceylon, is easily recognised by the very convex cara-
pace without a nuchal shield, and by the beautiful markings of
the other shields, each of which has a yellow areola, whence radiate
yellow streaks upon a" black ground. Moreover, the dorsal shields
often form humps. It reaches the length of one foot. Old
specimens lose the beautiful yellow radiation, owing to a con-
siderable amount of peeling off of the homy layers.

The habits have been carefully watched by Captain Thomas
Hutton,^ who gives the following account. The tortoises live in
the grassy jungle at the base of the hills, but owing to their colour
being so blended with the rocky natiu'e of the ground, they are
with difficulty distinguished. Moreover, they remain concealed
beneath shrubs or grass during the heat of the day. In the
rainy season they are most active, wandering al)out all day,
feeding and })airing. At the approach of the cold weather they
selecit a sheltered spot and conceal themselves by thriLsting their
shell into some thick tuft of grass, remaining there in a sort of
lethargic, but not torpid, inactivity until the hot season, at
which time they remain concealed only during the heat of the
day, coming out about sunset to feed.

During the hot season Hutton's captives often soaked them-

' J. Asiat. Soc. BiHfjal, vi. 1837, p. 689.

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IX TESTUDINIDAE 37 1

selves in water, and they drank a great deal. Copulation
lasted about ten minutes ; the females received the males from
the end of June to the middle of October. On the 11th of
November a female dug a pit at the root of a tuft of grass, having
previously watered the spot, then digging with the hind-limbs
alternately, and continuing to water the soil In two hours she
had made a hole six inches deep and four wide ; she then laid
four pure white eggs, each about 1^ inches or 45 mm. long,
and filled the hole again with the prepared mud, pressing it well
in with the feet and with the weight of the body. The whole
operation took four hours. From December to the beginning of
February these tortoises were listless, they then took water and
some lucerne, but did not come out again until the middle of April,
well in the hot season. Both males and females wrestled in a
curious way. One confronted the other, with the head and fore-
limbs drawn into the shell, and with the hind-limbs planted
firmly on the ground, and in this manner shoving against each
other in any narrow space. Sometimes, if one succeeded in
placing its shell beneath the other, he tilted his adversary over
on his back, from which position he had great difficulty in rcr
covering himself.

T. ;polyjphemit8y the " Gopher Tortoise " of the south-eastern
States of North America, is one of the few American species. It
is characterised by the shape of the front lobe of the plastron,
which is bent upwards, and extends beyond the carapace. The
nuchal shield is present, not narrow; the supracaudal is un-
divided. The shell is much depressed, and flattened along the
vertebral region, with rounded margins. The fore - limbs are
armed with very strong claws. The general colour is very dark
brown above, inclining to black ; brownish yellow below, with
blackish patches. The length of the shell is about one foot, or
even eighteen inches.

The Gopher is interesting for its habits, which are described
by Agassiz, Schnee, and others. Its domicile consists of an
excavation, the mouth of which is just sufficient to admit the
animal, the burrow running in an oblique direction to the depth
of about four feet. The whole passage is sometimes more than
two yards long. It expands from the entrance, and ends in a
roomy space, sometimes with a few branches of fir trees which
have been dragged in either for food or as a lining. The burrow

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372 CHELOMA chap.

is inhabited by one pair only. When the dew is on the grass,
or after rain, they emerge in search of food, which consists of
grass, succulent vegetables, fruit, etc They also eat the gum
that exudes from trees, especially the resin of the pine. The
eggs are laid in June, not in their domicile, but in a separate
cavity near the entrance ; a set consists of five eggs, almost round,
and very large, namely, 40 mm., or more than one inch and a
half in diameter. To capture the Gopher a deep hole is dug at
the mouth of their home, into which they fall as they emerge for
food. In Southern Texeis and neighbouring parts of Mexico they
are represented by a smaller and lighter coloured species.

jT. tahulata, widely spread over Tropical South America, whence
it is often brought over as a curiosity, reaches a large size,
specimens nearly two feet in length being not uncommon. The
shell is flat on the top, and is very elongated, without a nuchal, but
with an undivided supracaudal shield. The carapace is very dark
brown or black, each shield with a yellow or orange centre ; the
pleistron is brown and yellow, the dark colour being mostly con-
fined to the middle portion. The ground-colour of the skin of
the limbs is blackish, but the scales are orange or red. The head
is yellow and black. This species inhabits the forests, and lives
chiefly on the fruits of trees ; in captivity they are said to take
bread soaked in milk or water, lemons, apples, bananas, cabbage,
gourds, and also meat, at least the males.

Gigantic Land-tortoises differ from the others in no essential
points except their large size. The term gigantic is, however,
applied to many of them by courtesy only, since they do not
exceed the dimensions of large Turtles. A truly gigantic species,
1\ atlas, has left its remains in the Sivalik Hills of late Miocene
or early Pliocene date. The skull is between seven and eight
inches long, and is well preserved, but the correctness of the
dimensions of the specimen, as it now stands, restored in the
National Collection, is open to doubt. The shell was probably not
more than six feet long. Miocene and Pliocene Europe was also
inhabited by large tortoises, with shells about four feet long, e.g.
T, 'perpigniana, whose bony plates are one inch thick ; others
have been found in North America. Such large tortoises are
now restricted to two widely separated regions of the world,
namely the Galapagos Islands (which have received their name
from these creatures, galdpatjo being one of the Spanish terms for

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IX TESTUDINIDAE 373

tortoise), and the islands in the Western Indian Ocean, namely
the Mascarenes (Bourbon, Mauritius, and Rodriguez), the Comoros,
Aldabra, the Amirantes, and the Seychelles. When they became
extinct in Madagascar is not known, but T, graiulidieri was a
very large species of apparently very recent date. Of the other
islands the Comoros only were inhabited by man, the others were
devoid of any but small and harmless Mammals. It was on these
peaceful islands that large tortoises lived in incredible numbers,
and, like the Dodo of Mauritius and the Solitaire of Rodriguez,
grew to a size far beyond that of their less favourably placed
continental relations. The same applies to the tortoises of the Gala-
pagos Islands. Plenty of food, a congenial equable climate, and
absence of enemies enabled them to enjoy existence to the fullest
extent. There was nothing for them to do but to thrive, to feed,
to propagate, to grow, and to vary. At least there was iiotliing
to check variation within reasonable limits. Scattered over the
many islands, they were prevented from inter-breeding, and thus
it has come to pass that not only every group of islands, but in
the case of the Galapagos almost every island, has or had its own
particular kind, be these called varieties, races, forms, or species.

There are four features of si)ecial interest. First, these tortoises
grow to a large size, and there are no small species on any of
these islands. Secondly, they vary much individually. Thirdly,
each island or group of islands has developed its own kind.
Lastly, there is the widely si)read tendency to reduce the thickness
of the bony plates of the carapace, in spite of its size. In some
cases, notably T, vosmaeri of Rodriguez, the bony shell is reduced
to apparently the utmost limit compatible with mechanical safety.
The horny shields are, or were, however, well developed, sometimes
much more so than in other recent land-tortoises. Whatever were
the original reasons for the development of a strong shell in
tortoises, they cannot have prevailed in these islands.

Where did all these tortoises come from, and how did they
get to these oceanic islands ? Accidental transport or migration
are out of the question. Land-tortoises are drowned within a
few hours. Moreover, there are none of their kind on the con-
tinents of Africa, Asia, and South America, although they had a
much wider distribution in past geological ages. Consequently
we have to assume that they are descendants of tortoises once
populating the land which, except the islands, lies now below

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374 CHELONIA CHAP.

the western Indian ocean. The existence of this, "Lemuria" or
" Gondwana," came to an end in Mid-Tertiary times. The large
tortoises on the remaining continents died out — ^in any case they
are gone, while those which lived on, or retreated to, what became
the present islands, survived and flourished.

The tortoises were not left in peace with the advent of man,
who found that they were good to eat. They were first ex-
terminated on the Mascarene Islands. In 1759 four small
vessels were specially appointed for the service of bringing
tortoises from Rodriguez to Mauritius ; one vessel carried a
cargo of 6000 ; and altogether more than 30,000 were imported
into Mauritius within the space of eighteen months. Dr. Gunther
very properly remarks that many of these tortoises must have
been small-sized specimens, and that many of them were probably
used for provisioning passing Government vessels. Anyhow an
inter-insular traffic was carried on, and there are records of
superfluous tortoises having been turned loose, at the end of the
voyage, in distant islands, even in Java. Importation and
exchange of choice specimens, by way of presents, seems also to
have taken place. All this makes it now actually impossible
to trace the original habitat of the few surviving specimens
with anything like certainty. At the beginning of this century
the large tortoises had been nearly cleared off most of the islands,
and at the present time only the south island of Aldabra enjoys
the reputation of still possessing some really indigenous tortoises.
The few survivors on the otlier islands are said to have been intro-
duced. The small stock at Aldabra is now under Government
protection. Representatives of various si>ecie8 will linger on for
a little time to come, when they are kept as pets on some tropical
islands, but those which have been brought to Europe are of course
doomed.

We can mention only a few of the large tortoises which
have become famous, not to say historical. A fascinating resume
of the whole complicated question has been given by Dr.
Giinther.^

Testudo gigantea s. elephantina s. hololissa s. iwnderosa,
originally confined to the North Island of Aldabra, where this

^ Presidential Address. Proc. Linn. Soc. 1898. See also Giinther, Giganik
Land- Tortoises, Brit. Mus. London, 1877 ; Gadow, Trans. Zool. Soc. xiii. 1894,
p. 313 ; Rothschild, Novit. Zool., several notes.

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TESTUDINIDAE 375

kind has been completely exterminated, is now still to be found
in the Seychelles in considerable numbers, introduced there by
planters, and kept in a state of semi-domestication. A very
large specimen was received by the Hon. Walter Kothschild, at
Tring, in 1893. In 1897 its shell measured 40 1 inches in
length, 5 2^ over the curve, and 5 0 inches across the curve trans-
versely ; it weighed 358 lbs. The measurements taken in
previous years are unfortunately not free from mistakes. " When-
ever the temperature was over 60" F. this tortoise had a free run
of 350 acres of grass park, and when the temperature showed
permanently below 58°, it was kept- in an orchid house from
September to June. When at liberty in the park it lived
entirely on grass, but in the hothouse it fed on carrots, cabbage,
lettuce, and several other vegetables " ; it was also very fond of
rotten fruit. To this species belongs the large tortoise which
has been living at St. Helena for more than the last hundred
years.

T, daudini is the species of the South Island of Aldabra.
Voeltzkow, in 1895, succeeded in carrying off seven specimens.
He gives the following description : — The island is an atoll, cut
through in three places, with a greatest length of about twenty
miles. The chief hindrance in the search for the tortoises is the
impenetrability of the island. The soil consists entirely of sharp
water-worn corals, with their points uppermost, while the whole
is covered with such thick masses of low scrub, that a way has to
be cut with an axe, so that an extended search over a large area
is out of the question. To land on the outside is dangerous, on
account of the heavy surf ; while landing from the inside of the
atoll is much hindered by the dense thickets of mangrove trees.
As drinking water, and that very bad, is only found in one
place, rainwater has to be collected from the natural hollows, and
carried along in tanks. Thousands of mosquitoes prevent one
remaining over night in those places which the tortoises frequent.
Then at last, when one has discovered, by a stroke of luck, one
of these creatui-es, in the thick scrub, where they hide during the
heat of the day, the real hard work begins, namely, the convey-
ance of the beast. Six reached Europe alive, two of them were
sent to Frankfort, and the four others to Hamburg. Mr.
Rothschild received a male of T. daudini, which, until its
recent death, was the largest living tortoise known. The length

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376

CHELONIA

CHAP.

of its shell was 55 inches, or 67^^ inches over the curve ; total
weight 560 lbs. This specimen had a chequered career.
Although its original home must have been the Aldabra atoll,
it had been known for many years on Egmont Island, one of the
Chagos Islands. According to tradition, it had been thert
some 150 years, but the first settlement on that island was

Fk;. 83. — Tr.stvdo (hit(fiiii {ixhow) and 7. abingdoni (below). x t,V,.

formed from Mauritius only at the beginning of this century.
The owner of the tortoise, M. Antelme, took it to Mauritius,
whence it came to England. On the Egmont Island it used
to bury itself for six months in the ground without eating
anything.

T. svmeirei. — This kind is supposed to have been the species
peculiar to the Seychelles. In 17G6 five large tortoises were
l)rought from the Seychelles to Mauritius by Chevalier Mariou
de Tresne. Of these only three were alive in 1898, two iu

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IX TESTUDINIDAE 377

Mauritius and one in London ; the latter specimen soon died in
the Zoological Gardens, One of the two survivors, the last of
their race, is famous. It was kept at Port Louis, and when
Mauritius became a British possession in 1810, the tortoise was
especially mentioned and taken over. It still lives there in the
grounds of the barracks of the garrison. According to the
proverbial oldest inhabitants it had in 1810 already reached
its present size, namely, a shell-length of about 40 inches with a *
greatest circumference of 259 cm. = 8 feet 6 inches. Total
weight 160 kilo = about 358 lbs. When walking it stands
6 3 '5 cm. = 25*4 inches high, with the plastron about 15 cm.
or 6 inches above the ground, and it can then carry with ease
two full-grown men on its back. This old male is now nearly
blind, but is otherwise of regular habits and in good health.
Although it has been known for nearly 150 years it had to
wait for its scientific name until the year 1892.

Another famous individual is the Colombo tortoise. It is
supposed to have come to Colombo from the Seychelles in 1798.
It died in 1897. To judge from photographs, this specimen, a
male, may possibly belong to T. sumeirei, in spite of the very flat
shell, which is 53^ inches in length.

Leaving aside the remains of sub-fossil tortoises, e.g, the thin-
shelled jT. vosmaeri of Rodriguez, and several kinds which have
been dug out in the Mare-aux-songes of Mauritius, one of which
had a markedly forked and prolonged anterior plastral lobe,
rather resembling that of the Pliocene Sivalik 2\ atlas, we now
turn to the tortoises of the Galapagos Islands. They existed in
enormous numbers towards the end of the seventeenth century,
when Dampier visited those islands. Hundreds were exported
and scattered early in the nineteenth century. When the islands
became a penal settlement of Ecuador, the introduction of con-
victs and pigs proved detrimental to them, but Darwin found
them still present in 1835 on most of the islands. His cla.ssical
account of these old giants is to be found in the Voyage of the.
Beagle. They lived on the succulent cactus plants, leaves of
trees, berries, and a kind of TJsnea, a lichen pendant from the
trees. They collected regularly at certain pools and springs,
leading to which were regular well-trodden paths, formed by
the coming and going of the tortoises. He calculated that they
could walk a distcmce of about four miles in one day. During

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378 CHELONIA

the time of propagation the males emit a hoarse bark, which can
be heard a hundred yards off The round eggs measure about
5 cm. or 2 inches in diameter, and are laid in the month of
October, about one dozen making a set.

Nearly every island had apparently its own kind. They are
all remarkable for their small head and the length of their neck,
which is decidedly longer and more slender than that of the
Eastern tortoises. The most peculiar looking are or were T.
ephippium and T, abingdoni, the shell of which is extremely
thin, with large lacunae in the osseous plates. The profile of the
shell is somewhat saddle-shaped, with the horny shields partly
concave and turned upwards at the sides. The general colour of
these and the other Galapagos tortoises is black. T. ephippium
still survives on Duncan Island. Of T. dejyJiantoptis s. vicina
Baur collected twenty-one specimens in 1893 on Albemarle Island.
Some of them are still comparatively young, only 1 6 inches long.
A large one was killed, and, being hard up for water, Baur and
his companions drank the five cups full of fluid contained in the
pericardial sac ; they found it most refreshing, and tasting some-
what like the white of an egg. One monster is said to have
measured 56 inches over the curve, with a skull 7*12 inches in
lengtli. Mr. Kothschild received one of this kind alive — a
much -travelled specimen. It came to England from Sydney,
whether it had been brought in 1880 from Eotuma Island,
north of the Fiji group. There it had probably been left with
others by Captain Porter, who, on his voyage from the Galapagos
in 1813, distributed several young tortoises from his stock
among the chiefs, and permitted a great many to escape into
the bushes and among the grass. The shell of this specimen
measured 49^ inches in length, 56 over the curve.

Fam. 6. Ghelonidae {Turtles). — The limbs are paddle-shaped,
and the shell is covered with horny shields. Only two recent
genera, with three species, widely distributed in the seas.

The neck is short and incompletely retractile. The temporal
region of the skull is completely roofed over above and laterally
by the parietals, postfrontals, squamosals, quadrate -jugals and
jugals. All these bones are much expanded, and form the
additional or false roof. The parietals are especially large, and are
in broad contact with the squamosals. Nasals are absent. The
nares are bordered by the small premaxillaries, the maxillaries.

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379

and the prefrontals. The choanae are enclosed by the palatines,
which are separated by the vomer, and are posteriorly in broad
contact with the pterygoids. The latter are connected with
descending processes of the parietals by epipterygoids. The fora-
men magnum is bounded not only by the supra-occipital and the
lateral occipitals, but also by the basi-occipital. For the skeleton
see Fig. 65, p. 320. The pubic and ischiadic symphyses are con-
nected by a narrow cartilaginous band. The pubis has a large,
broad, lateral process, but the ischium is devoid of such a process.

Fig. 84. — Skull of Thalassochdys caretta ; cf. also Fig. 63, p. 317. A^ Dorsal view ;
By ventral view ; F^ frontal ; Jg^ jugal ; Mx, maxillary ; Op, opisthotic ; P,
parietal ; Pal^ palatine ; /*r/, prefrontal ; Pt.fy pastfrontal ; Ptg^ pterygoid ; Q,
quadrate ; Quadra articular surface of quadrate ; Qj^ quadrato- jugal ; S.o^ supra-
occipital ; Sq, squamosal.

The paddles of the fore- and hind-limbs are produced by an
elongation of the metacarpal and metatarsal bones and of most of
the phalanges, and these have no condyles ; most of the carpal
and tarsal elements are flattened, and additional width is given
to the hands by the much enlarged pisiform bone. The number
of phalanges of the five fingers is 2, 3, 3, 2, 2 ; that of the five
toes, 2, 3, 3, 3, 2.

The carapace is heart-shaped and very flat. The nuchal plate
has no rib-like processes. The eight neurals form a continuous
series, and the short tail is covered by two or three pygal plates
besides the unpaired last marginal. The number of all the

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380 CHELONIA CHAP,

marginals is 23, sometimes 25 individually. The plastron (Fig,
66, p. 321) is composed of the usual nine plates, which, however,
remain entirely free from the marginals, and are only loosely
connected with each other, enclosing a very large unossified
space. The horny shields covering the plastron number 13, and
there is a series of about 5 inframarginals (Fig. 61, 6, p. 315),
There are normally 12 pairs of marginal shields, a nuchal,
5 neural, and 5 or 7 costal shields. Whilst the number of
these dorsal shields is pretty constant in Chelone, it is subject to
an astonishing amount of individual variation in Thaldssochelys.

The Chelonidae are a highly specialised ofifshoot of the
Cryptodira adapted to marine life. Fundamentally they agree
most with the Testudinidae, paradoxical as this may appear at
first sight. There is nothing primitive about them except the
complete series of inframarginal shields. Fossil forerunners of
marine turtle-like creatures appear in the Upper Jurassic deposits
of Europe and North America. The numerous genera have been
grouped together as Thalassemydidae and Chelonemydidae. They
are more or less intermediate between Chelonidae and ^//ly^-like
Chelonidae, the carapace being not too much flattened and
broadened out, the fontanelles between the ribs are mostly small,
the plastral bones are still broad, enclose a smaller ossified space,
and there is still a bony bridge in most cases. The paddle-shape
of the limbs is less pronounced, and sometimes only indicated.
In some forms, especially Lytoloma, from the Upper Cretaceous and
Eocene of North America and Europe, the anterior portion of the
skull is much longer than in the Chelonidae, the vomer and the
premaxillaries are elongated, and the anterior portion of the roof
of the mouth, with the corresponding parts of the lower jaw,
seems to have carried crushing pads. Some of the best-known
Upper Jurassic genera are Eurysternuvi and Idwchelys ; Fleaiochelys
from the Purbeck and Wealden ; AUopleAiron hofmanni from the
Upper Cretaceous of Belgium approaches Chelone. by the large
fontanelles between the small marginal and the short costal
plates. True Chelonidae are very rare and imperfect in the
Mid-Tertiary strata, but both recent genera seem to have existed
since Pliocene times.

The few recent Chelonidae are entirely marine, going on land
only in order to deposit their eggs in the sands of unfrequented
shores. Their distribution, in conformity with their oceanic life,

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IX CHELONIDAE 38 1

is . almost cosmopolitan within the warmer zones, but not a few
find their way far into the temperate seas. They are all eagerly
hunted by man either for food or for the sake of the tortoise-
shell.

Cheloiie, — With only four pairs of costal shields. Carapace
with large persisting fontanelles between the costal and marginal
plates. Two species.

Ch. mydas (the " Green or Edible Turtle ") has when adult a
nearly smooth shell, all the shields being juxtaposed, fitting closely
into each other, and becoming quite smooth with age. The
neural shields of younger specimens have a feeble keel. The
twenty-five shields which surround the carapace form a smooth, or
but indistinctly serrated rim. The head is covered with one pair
of prefrontal shields, the others are small. The horny beaks of
the upper and lower jaws have denticulated outer edges, those of
the upper jaw having two pairs of strong denticulated ridges.
The limbs have generally only one claw, namely on the first
digit. This claw, although sometimes curved and thick, and
more than an inch in length, is blunt. The general colour is
olive or brown above, with yellowish spots or blotches ; the
under parts are pale yellowish. This species attains a large
size, with a length of shell of nearly four feet, but the usual
length of full-grown specimens is three feet, and these weigh,
when in good condition, more than three hundredweight. Their
home is in the Atlantic, Indian, and Pacific Oceans, but there
are certain regions in which they are more common than in
others. Famous centres are the Island of Ascension, the West
Indies, and the coast of Mosquito, at least for commercial
purposes. As they require sandy, easily accesgible beaches for
the deposition of their eggs, they congregate in certain parts of the
world more than in others, and being strictly vegetable feeders,
they are naturally bound to the coasts, although they are some-
times met with far out at sea. Their chief food consists of
algae, and of Zostera marina, the edible " Dulce," which grows
plentifully in the lagoons of the coast of Florida. When they
have eaten their fill, they are said to chop off more of these
plants, and roll them, together with the adherent mud, into balls
of the size of a head, and these balls, receding with the tide,
are followed by the Turtles.

Whilst in the water they are caught in various ways, with

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382 CHELONIA

nets or harpoons. In some parts of the world the natives follow
them in a boat, and when they espy a turtle crawling along the
bottom, a man, attached to a rope, dives in, clasps it, and is
brought up by his companions together with his prey. Turtles
are fond of basking asleep, floating on the surface, and they are
then hai'pooned from a stealthily approaching boat. The most
original mode of catching them is that used by the natives of
Torres Straits, Madagascar, and Cuba. The turtle-fishers go out
in the boat to a spot frequented by grazing turtles ; a long
string is tied to the tail of a fish, Hcheneis, a member of the
Mackerel family, and the.Ucheneis, anxious to get away to pro-
tective shelter, makes for a turtle, and attaches itself to the
turtle's plastron by means of the large sucking apparatus on
the top of its head and neck-region. The men are guided by
the string, and the turtle is gently coaxed up towards the surface
or followed into shallow water, where it is either harpooned or
dived for. It is curious that this use of the Echeneis exists in
such widely separated parts of the world, the natives of which
cannot have any knowledge of each other. These modes of
catching turtles are sportsman-like, but the greatest and most
wanton destruction is practised at their breeding places. In
conformity with the wide distribution of these creatures, the time
of breeding is not the same everywhere. In the West Indian
region, and in the Straits of Malacca, it falls within the period of
April to June ; on the coast of West Africa it occurs from
September to January. The females come to their breeding
places from afar, reconnoitre the beach carefully, are extremely
wary and shy, taking alarm at the slightest disturbance, and at
last crawl on land. Well out of the reach of the tide the female
scoops out a hole in the sand, deposits about one hundred or
more of its round, rather parchment-shelled eggs, covers the nest
carefully, obliterating all traces of the dug-out sand, and makes
again for the sea ])y another route. At least they are said to
make a sort of circuitous route so that nobody can tell the
position of the nest, which may be anywhere beneath the broad
trail left by the heavy creature on its way from and back to the
sea. The nest is discovered by probing the sand with sticks.
The time of incubation is not known, but according to Agassiz,
lasts at least seven weeks.

The " turning " of turtles is a cruel and wanton operation,

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384 CHELONIA

since frequently many more are turned over and left to perish
than are taken away. Men lying in ambush watch the beast,
or they approach the lonely sandy shore by boat, and rush the
helpless creatures when these are surprised in sufficient numbers.
It takes several men to lift a full-grown specimen. It is there-
fore necessary to secure them by turning them over with poles or
by their flippers, lest they should crawl away. On board ship
they are either put into tanks or tied with ropes on deck, covered
with a moistened cloth ; and occasionally a piece of bread, soaked
in sea-water, is thrust into the parched mouth. In London thej
are kept in large tanks, often in considerable numbers, but since
they take no food in captivity, or rather because it is difficult to
supply them with the right sort, they are not kept long. After
the head has been cut oiff, the body is suspended for a day or two,
in order to drain it of the blood. It is not only the meat and
the fat which are used for the making of the famous soup, but
also the thick and dense layer of subcutaneous tissue which lines
the inside of the shell.

Tennent describes a revolting spectacle exhibited in the
markets of Jaffna, in Ceylon. The flesh of the turtles is sold
piecemeal by the Tamil fishermen, while the animals are
still alive. At certain seasons, says the same authority, the
flesh of turtle on the south-west coast of Ceylon is usually
avoided as poisonous, but some lamentable instances are recorded
of neglect of this, and consequent sickness, followed by coma and
death. In the Gulf of Manaar specimens are frequently found
between four and five feet in length ; and on one occasion, in
riding along the seashore north of Putlam, he saw a man in
charge of some sheep, resting under the shade of a turtle shell,
which he had erected on sticks to protect him from the sun.
In connexion with this curious sight, Tennent quotes Aelian's
statements, copied by him from Megasthenes* Indica Frag, lix. 31,
that in the Indian ocean turtles occur which measure fifteen ells,
so that not a few people may find ample shelter beneath a single
shell.

Ck. imhricata (" Hawksbill Turtle "). — The number of shields
covering the carapace is the same as in Ch. mt/das, but they
strongly imbricate, or overlap eiich other from before backwards,
until the animal is very old, when the shields become juxtaposed
In young specimens, under one foot in length, each of the neural

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CHELONIDAE

385

and costal shields is strongly keeled, the three rows of keels
converging towards the posterior end of the shell. The neural
series of keels is almost continuous, and remains longest, even in
half-grown specimens. The twelve pairs of marginal shields form
at first a strongly serrated sharp edge ; the serrations disappear
gradually on the front portion, but remain on the posterior half
of the shelL The homy covers of the jaws form a hooked beak,
with sharp but smooth or feebly denticulated margins. The fore-

Fia. 86. — Chelone imbricata (" HowksbUl Turtle "), young. x k

and hind- flippers have two claws. The young are pale brown
above, blackish below; the shell of the adult is beautifully
marbled with yellow on a rich dark-brown ground ; the plastron
is yellow. The shields and scales of the head and limbs are dark
brown, with yellow margins. The top of the head is covered by a
large unpaired frontal and a pair of prefrontal or interorbital shields.
This Turtle does not reach the size of the green or edible kind ;
the largest shell on record is in the National Collection, and
measures 85 cm. = 34 inches in length. They range over all the
tropical and subtropical seas. They are apparently strictly car-
VOL. VIII 2 c

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386 CHELONIA

nivorous, living upon fish and moUudcs, the shells of which they
crunch. Although not eaten, they are much persecuted on account
of their shells, the horny shields of which are the " tortoise-shell "
of commerce. A large specimen yields up to 8 lbs. Few of the
shields are, however, thick enough to be manufactured into the
larger articles which art and fashion delight in, but if heated in
oil, or boiled, they can be welded together under pressure, and be
given any desired shape. In genuine articles of Oriental manu-
facture these welds can generally be detected, or their compound
nature is indicated by the beautiful pattern, which is too regular
in the imitations now common. Even the shavings and
leavings can be welded and moulded into large pieces. The
stripping of the shields has been described by Sir E. Tennent
" If taken from the animal after death and decomposition, the
colour of the shell becomes clouded and milky, and hence the
cruel expedient is resorted to of seizing the turtles as they repair
to the shore to deposit their eggs, and suspending them over fires
till heat makes the plates on the dorsal shields start from the bone
of the carapace, after which the creature is permitted to escape to
the water. At Celebes, where the finest tortoise-shell is exported to
China, the natives kill the turtles by* blows on the head, and
immerse the shell in boiling water to detach the shields. Dry
heat is only resorted to by the unskilful, who frequently destroy
the tortoise-shell in the operation." The cruel process described
above is resorted to " for economy's sake," the Singhalese believing
that such maltreated turtles regenerate the shields, to be caught
and shipped again. Since none of them are actually re-caught
in the mutilated condition, this is looked upon as a proof of the
correctness of the treatment. It is more likely that they die.

New shields can be reproduced only if the imderlying Mal-
pighian layer of cells (if. Fig. 68, B, p. 323) is not killed by the
roasting. However, Dr. Charles Hose, with his long experience
in Borneo, is positive that numerous individuals are there caught
which have imperfectly mended shells, the ghields of which do not
imbricate, are thin, and almost worthless.

It is commonly believed that the same individuals return
again and again to the same spot for laying. This is very
likely the case. Tennent mentions that in the year 1826 a
Hawksbill was taken near Hambangtotte, which bore a ring
attached to one of its fins, that had been placed there by a Dutdi

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IX CHELONIDAE 387

officer thirty years before, with a view of establishing the fact
of these recurring visits to the same beach. The same homing
instinct has been observed in some females of the Green Turtle,
which, having been brought from the Tortugas Keys to Key
West off the south end of Florida, escaped, and were, a few days
later, re-caught at the Tortugas. On the other hand, experi-
ments made with turtles at Ascension are said to have had no
result.

Thalassochelys, with five pairs of costal shields. The carapace
is completely ossified in the adult, leaving no fontanelles between
the ribs and the marginals.

Th. caretta (the " Loggerhead Turtle"). — The shields of the cara-
pace imbricate only in young specimens, in the adult they become
smooth and juxtaposed. The margin is serrated posteriorly.
The carapace of the young has three strong keels. The inter-
gular shield is very small or absent. The marginals, including
the nuchal, usually number 23, rarely 25. The large head is
armed with hooked jaws, the crushing surface of the horny upper
beak has a median prominent ridge. The top of the head has
a pair of shields in front of the unpaired frontal. The flippers
of the young have claws on the first and second digits ; in the
adult usually only that of the first digit remains. The general
colour of the shell is uniform brown above, yellowish below.
Very young specimens are uniform dark brown or blackish above
and below.

Large individuals have a shell about three feet and a half in
length. The Loggerhead is carnivorous, and is commercially of
no value. Its habits seem to be the same as those of the other
Turtles, but it has a much wider distribution. Besides all the
tropical and intertropical seas, it inhabits the Mediterranean,
and is an accidental visitor to the western coasts of Europe,
especially Portugal and the Bay of Bisciiy. It has been caught
several times on the coast of Belgium, and an old female con-
taining 1150 eggs was captured in 1894 on the Dutch coast.
In 1861 one was caught near Penman, on the coast of Banffshire,
and a second in the completely laud-locked Loch Lomond.^
It has been more frequently recorded from the coast of Devon
and Cornwall.

The most interesting feature of the Loggerhead is the
' Xotes Leyden Mus. xvi. 1895, p. 211.

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388 CHELONIA CHAP.

astonishing variability in the number of the homy shields of
the carapace. The normal number of shields of the carapace,
leaving out the marginals and counting the nuchal as the first
neural, is 6 neurals and 5 pairs of costals, in all 16. The
greatest number of dorsal shields observed is 8 neurals and 8
pairs of costals, in all 24. Many of the intermediate combina-
tions have been observed, there being, for instance, specimens
with 8 neurals and 16, 14, 13, 12, or 11 costals, the latter not
being always in pairs, but unequal on the right and left sides;
or there are 7 neurals with 20 to 16 costals, or 6 neurals with
20, 19, 18, 17, or 16 costals. The interesting fact in con-
nexion with these variations is, moreover, that some of the shields
are much smaller than the others, sometimes mere vestiges in
all stages of gradual suppression, and that the abnormalities are
much more common in babies and small specimens than in
adults. The importance of these " orthogenetic " variations has
been discussed on p. 326.

Sab-Order 2. Pleurodira. — Neck bending laterally and tucked
aioaj/ in the niche formed between the anterior portion of the cara-
pace and plastron. Pelvis arikylosed to the shell, the broadened
tops of the ilia to the carapace, the distal ends of the pubes and
ischia to the plastron.

Freshwater tortoises, almost entirely carnivorous, inhabiting
South America, Australia, Africa, and Madagascar. Fossil forms
are known from the Jurassic epoch onwards.

Owing to the strong connexion of the iliac bones with the
costal plates the sacrum has become practically abolished, the
sacral ribs being reduced to one pair (the posterior of the
original two pairs) or being absent. The centra of the cervical
vertebra articulate by cup and ball joints. The formation of
the temporal region of the skull varies considerably in the three
families, some genera lacking the complete zygomatic arch, while
others have a narrow parieto-squamosal arch bridging over the
temporal fossa, or the latter is completely roofed over by the
laterally expanded parietal, which meets the jugal and quadrato-
jugal. The quadrate is always trumpet-shaped ; the rim of the
tympanum is complete, but the posterior part of the trumpet
remains open. The basisphenoid, pterygoids, and palatines form
a broad and flat roof to the mouth. The vomer is large, and
separates the palatines in the Chelydidae ; it is very much

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PLEURODIRA 389

reduced or absent in the Pelomedusidae, in which the palatines
meet. All the Chelydidae, except Chdys, have nasal bones
which remain distinct from the prefrontals. The choanae lie in
front of the palatines, divided by the vomer when this is present,
but they are not roofed in ventrally.

The ilia are solidly ankylosed in the adult with the neigh- .
bouring costal plates, mostly with the last two pairs, sometimes
also with the pygal plate. The lateral processes of the pubes
fuse with the xiphiplastra. The ischia are also attached to the
same plastral elements.

The carapace is flat and completely ossified. The nuchal
plate is always conspicuous, much larger than the neurals, and
these are often reduced by being encroached upon by the eight
pairs of costal plates, which then meet in the dorsal line. In
Sternothaerus all the eight neurals are present and form a
continuous row. In most of the other genera they are reduced
to seven, the last being squeezed out. In Rhinemys they are
reduced to the second, third and fourth and an isolated fifth, and
in Hydraspis they are all gone. The pygal plate is always, even
in SternothaeruSy separated from the last neural by the eighth
pair of costals. The marginals number 23, but in Carettochelys
only 21.

The carapace is covered with horny shields, except in CarettO'
chelys. The nuchal is absent in the Pelomedusidae and in a
few Chelydidae {Mseya and a few species of Emydura). In
Hydromedusa the nuchal is shut in by the anterior marginals,
simulating a sixth neural. The plastron is composed of the
usual nine elements, but the Pelomedusidae possess an additional
pair, the meso-plastra, inserted between the hyo- and hypo-plastra.
The bridge is strong, connected with the carapace by suture.
In SternothaeTUS the front lobe of the plastron is movable. The
intergular shield is always present ; it is terminal, forming part of
the front margin, except in Chelodina, where this shield, although
large, is shut in behind the gulars (cf. Fig. 61, 4 and 5, p. 315).

Although the Pleurodira are a peculiarly specialised group,
one of the oldest Chelonian fossils known seems to belong to
them. ProganochelySy represented by a complete shell, nearly
2 feet long, has been found in the Upper Keuper Sandstone of
Wurtemberg. Flesiochelys, of the Upper Jurassic of Switzerland,
has eight neural and three supracaudal plates, but is without the

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390 CHELONIA

ischiadic plastral ankylosis. Pleurosternum, of the English and
Continental Purbeck beds, has meso-plastral plates like the recent
Pelomedusidae. Rhinochelys, of the Cambridge Greensand, has
a broad parieto-postfrontal roof, and large nasal bone& Forms
like PodocnemiSy now restricted to South America, occur in
the Eocene of Europe. One of the most aberrant Chelonians
is Miolania, from the Plistocene of Queensland and from Lord
Howe's Island, remarkable for its huge size and the thick
armour on the head and tail ; the head especially carries large
paired projections, one pair of which extends horizontally like
powerful horns, recalling the queer Theromorphous Elginia,

We divide the recent Pleurodira into three families, of which
that of Carettochelys stands apart by its paddle-shaped limbs
and the absence of horny shields. The Pelomedusidae and
Chelydidae are closely allied. The former are not Australian, and
are externally distinguished by the absence of a nuchal shield-

Fam. 1. Pelomedusidae. — Neck completely retractile within
the shell. Carapace without a nuchal shield. The plastron is
composed of eleven plates,, there being besides the unpaired endo-
plastron a pair of meso-plastra, situated between the hyo- and
hypo-plastra ; but these meso-plastra meet in the middle line
in Sternothaerus only, while in Podocnemis and Pelomedusa they
are restricted to small pieces on the bridge, widely separated
from each other by the usual hyo- and hypo-plastral suture. A
nuchal shield is absent; there are twenty-four marginal and
thirteen plastral shields, inclusive of the conspicuous intergular.
The temporal fossa is widely open, except in Podocnemis, where
it is partly roofed in by the meeting of the much-expanded
quadrato-jugal with the parietal. The palatine bones are in
median contact, not separated by the vomer. Nasal bones
being absent, the large i^refrontals meet in the middle line.
The second cervical vertebra is biconvex.

This family is now represented by only three genera, with
about fifteen species in Africa, Madagascar, and South America.

Sternothaerus. — Skull without a bony supratemporal roof.
Meso-plastra large, extending right across the plastron. Anterior
lobe of the plastron movable, the hinge passing between the
hyo- and meso-plastral plates, and between the pectoral and
abdominal shields. Fore- and hind-limbs with five short digits
and claws. Several species in tropical and southern Africa, and

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PLEURODIRA — PELOMEDUSIDAE 39 1

in Madagascar. S. derbianus in West Africa, from the Gambia
to Angola, is the largest species, with a shell nearly one foot in
length.

Pelomedusa, — Skull with a slender parieto-squamosal arch.
Meso-plastra small and lateral. Plastron without a hinge.
Fore- and hind-limbs with five very short digits and five claws.
Top of the head with one pair of shields between the eyes, and
with a large interparietal and a pair of parietals behind.

P. galeata, the only species, occurs in Madagascar and nearly the
whole of Africa south of the Sahara, from the Cape to Abyssinia,
and in the Sinaitic peninsula. The shell, less than one foot in
length, is much depressed and is obtusely keeled ; brown above
with black spots ; brownish-yellow below. The short and broad
head is coloured like the rest, without ornamentation. In Somali-
land this species sleeps hidden on land during the dry seasons,
from July to the end of September, and from January to March,
and appears at once after the rains have set in.

Podocnemis. — With a supratemporal roof formed by the
junction of the parietal with the quadrato-jugal. Meso-plastra
small and lateral. Fore- and hind-limbs broadly webbed, with
five and four claws respectively. The fore-arms and the outer
edges of the hind-feet with several conspicuous shields, hence the
generic name. Head with an interparietal, two parietals, and a
narrow unpaired shield between the eyes. The tail is very
short. The carapace is flat and broad, strongly serrated on the
posterior margin. Chin with one or two short barbels. Several
species in South America, chiefly in the basin of the Amazon, and
one in Madagascar.

P. expansa. — Very common in Tropical South America, east
of the Andes. The female, which is much larger than the male,
has a shell nearly three feet in length. Olive -brown above
with darker patches ; yellowish below. With a few yellow
spots above and behind the eyes, and on the parietal region.
The " Arrau " turtle is of great commercial importance on
account of the eggs, which are periodically collected in enormous
quantities, chiefly for the oil. This is either eaten, like the
eggs themselves, or used for burning in lamps, or as an addition
to tar. The turtles are likewise eaten by man and beast.
Thousands of the little creatures are snapped up by Jabiru
storks, alligators, and fishes ; the adults fall an easy prey

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392 CTIELONIA

to the prowling jaguar, which turns them over on to their backs
and neatly cleans out the flesh with its sharp and powerful
claws.

Fertilisation takes place in the water, the eggs are deposited
on land, in sand-banks, the female digging a hole about two feet
deep and covering up the numerous soft-shelled eggs with sand.
The time of deposition is the early hours of the morning, but
the season depends upon the beginning of the principal rains,
since the young are hatched shortly before the torrential rains.
This season differs considerably in the various countries. The
hatching takes about forty days ; the eggs are consequently laid
in the Amazon countries during the months of September to
November, in the Orinoco district in March. This species lives
in the pools of the inundated forests, and when these are dried
up, the animals retire into the rivers themselves. Their food
consists mainly of the fruit dropping down from the trees.

Bates, in his delightful book, llie Naturalist on the River
Amazon, gives the following lively and exhaustive account of
his experience with these turtles : —

" I accompanied Cardozo in many wanderings on the Solimoes,
during which we visited the ' praias ' (sand islands), the turtle
pools in the forests, and the by-streams and lakes of the great
desert river. His object was mainly to superintend the business
of digging up turtle eggs on the sandbanks, having been elected
commandant for the year by the municipal council of Ega, of
the 'praia real' of Shimuni, the one lying nearest to Egiu
There are four of these royal praias within the Ega district, a
distance of 150 miles from the town, all of which are visited
annually by the Ega people for the purpose of collecting eggs
and extracting oil from their yolks. Each has its commander,
whose business is to make arrangements for securing to every
inhabitant an equal chance in the egg harvest, by placing
sentinels to protect the turtles whilst laying, and so forth. The
pregnant turtles descend from the interior pools to the main
river in July and August, before the outlets dry up, and there
seek in countless swarms their favourite sand-islands ; for it is
only a few praias that are selected by them out of the great
number existing. The young animals remain in the pools
throughout the dry season. These breeding places of turtles
then lie 20 to 30 or more feet above the level of the river.

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PELOMEDUSIDAE 393

and are accessible only by cutting roads through the dense
forest. . . .

" We found the two sentinels lodged in a corner of the praia,
where it commences at the foot of the towering forest-wall of
the island, having built for themselves a little rancho with poles
and palm-leaves.. .Great precautions are obliged to be taken to
avoid disturbing the sensitive turtles, who, previous to crawling
ashore to lay, assemble in great shoals off the sand-bank. The
men, during this time, take care not to show themselves, and
warn off any fisherman who wishes to pass near the place. . . .

''' T rose from my hammock by daylight, shivering with cold ; a
praia, on account of the great radiation of heiit in the night
from the sand, being towards the dawn the coldest place that
can be found in this climate. Cardozo and the men were
already up watching the turtles. The sentinels had erected for
this purpose a stage about fifty feet high, on a tall tree near
their station, the ascent to which was by a roughly made ladder
of woody lianas. They are enabled, by observing tlie turtles
from their watch-tower, to ascertain the date of successive
deposits of eggs, and thus guide the commandant in fixing the
time for the general invitation to the Ega people.

" The turtles lay their eggs by night, leaving the whaler, when
nothing disturbs them, in vast crowds, and crawling to the
central and highest part of the praia. These places are, of
course, the last to go under water when, in unusually wet
seasons, the river rises before the eggs are hatched by the heat
of the sand. . . . The hours between midnight and dawn are
the busiest. The turtles exaivate with their broad webbed paws
deep holes in the fine sand ; the first-comer, in each ease, making
a pit about three feet deep, laying its eggs (about 120 in
number), and covering them with sand ; the next making its
deposit at the top of that of its predecessor, and so on until
every pit is full. The whole body of turtles frequenting a praia
does not finish laying in less than fourteen or fifteen days, even
when there is no interruption. When all have done, the area
(called by the Brazilians ' taboleiro ') over which they have exca-
vated is distinguishable from the rest of the praia only by sign^
of the sand having been a little disturl)ed.

" I mounted the sentinel's stage just in time to see the turtles
retreating to the water on the opposite side of the sand-bank.

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394 CHELONIA CHAP.

after having laid their eggs. The sight was well worth the
trouble of ascending the shaky ladder. They were about a mile
off, but the surface of the sands was blackened with the multi-
tudes which were waddling towards the river; the margin of
the praia was rather steep, and they all seemed to tumble
head first down the declivity into the water. . . . Placards were
posted up on the church doors at Ega, annoimcing that the
excavation on Shimuni would commence on the 1 7th of October,
and on Catua, sixty miles below Shimuni, on the 25th. By the
morning of the 17th some 400 persons were assembled on
the borders of the sand -bank, each family having erected a
rude temporary shed of poles and palm-leaves to protect them-
selves from the sun and rain. Large copper kettles to prepare
the oil, and hundreds of red earthenware jars, were scattered
about on the sand.

"The excavation of the taboleiro, collecting the eggs, and
purifying the oil, occupied four days. All was done on a
system established by the old Portuguese governors, probiibly
more than a century ago. The commandant first took down
the names of all the masters of households, with the number of
persons each intended to employ in digging ; he then exacted a
payment of 140 reis (about 4d.) a head towards defraying the
expense of sentinels. The whole were then allowed to go to
the taboleiro. They ranged themselves round the circle, each
person armed with a paddle, to be used as a spade, and then all
began simultaneously to dig on a signal being given — the roll of
drums — by order of the commandant. It was an animating
sight to behold the wide circle of rival diggers throwing up
clouds of sand in their energetic labours, and working gradually
towards the centre of the ring. A little rest was taken during
the great heat of mid^day, and in the evening the eggs were
carried to the huts in baskets. By the end of the second day
the taboleiro was exhausted; large mounds of eggs, some of
them four to five feet in height, were then seen by the side of
each hut, the produce of the labour of the family.

" In the hurry of digging, some of the deeper nests are passed
over; to find these out, the people go about provided with a
long steel or wooden probe, the presence of the eggs being
discoverable by the ease with which the spit enters the sand.
When no more eggs are to be found, the mashing process begins.

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IX PELOMEDUSIDAE 395

The egg, it may be here mentioned, has a flexible or leathery
sheU ; it is quite round, and somewhat larger than a hen's egg.
The whole heap is thrown into an empty canoe and mashed
with wooden prongs ; but sometimes naked Indians and children
jump into the mass and tread it down, besmearing themselves
with yolk, and making about as filthy a scene as can well be
imagined. This being finished, water is poured into the canoe,
and the fatty mass is then left for a few hours to be heated by the
sun, on which the oil separates and rises to the surface. The
floating oil is afterwards skimmed off with long spoons, made by
tying large mussel-shells to the end of rods, and purified over
the fire in copper kettles.

" The destruction of turtle eggs every year by these proceed-
ings is enormous. At least 6000 jars, holding each three
gallons of the oil, are exported annually from the Upper
Amazons and the Madeira to Para, where it is used for lighting,
frying fish, and other purposes. It may be fairly estimated
that 2000 more jarfuls are consumed by the inhabitants of the
villages on the river. Now, it takes twelve basketfuls of eggs,
or about 6000, by the wasteful process followed, to make one
jar of oiL The total number of eggs annually destroyed
amounts, therefore, to 48 millions. As each turtle lays about
120, it follows that the yearly ofiBpring of 400,000 turtles is
thus annihilated. A vast number, nevertheless, remain unde-
tected; and these would probably be sufficient to keep the
turtle population of these rivers up to the mark, if the people
did not follow the wasteful practice of lying in wait for the
newly -hatched young, and collecting them by thousands for
eating; their tender flesh, and the remains of yolk in their
entrails, being considered a great delicacy. The chief natural
enemies of the turtle are vultures and alligators, which devour
the newly-hatched young as they descend in shoals to the water.
These must have destroyed an immensely greater number before
the European settlers began to appropriate the eggs than they
do now. It is almost doubtful if this natural persecution did
not act as effectively in checking the increase of the turtle as
the artificial destruction now does. If we are to believe the
tradition of the Indians, however, it had not this result ; for
they say that formerly the waters teemed as thickly with
turtles as the air does now with mosquitoes. The universal

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396 CHELONIA

opinion of the settlers on the Upper Amazon is, that the turtle
has very greatly decreased in numbers, and is still annuaUy
decreasing.

"The principal object of another expedition was to search
certain pools in the forest for young turtle. We started from
the praia at sunrise on the 7th of October in two canoes, con-
taining twenty-three persons, nineteen of whom were Indians.
The pool covered an area of about four or five acres, and was
closely hemmed in by the forest, which, in picturesque variety
and grouping of trees and foliage, exceeded almost everything
I had yet witnessed. The margins for some distance were
swampy, and covered with large tufts of fine grass. The pool
was nowhere more than five feet deep, one foot of which was not
water, but extremely fine and soft mud.

" Cardozo and I spent an hour paddling about. The Indians
seemed to think that netting the animals, as Cardozo proposed
doing, was not lawful sport, and wished first to have an hour or
two's old-fashioned practice with their weapons. I was astonished
at the skill which they displayed in shooting turtles from little
stages made of poles and cross pieces of wood. They did not
wait for their coming to the surface to breathe, but watched for
the slight movements in the water which revealed their presence
underneath. These little tracts on the water are called the
sirird ; the instant one was perceived an arrow flew from the
bow of the nearest man, and never failed to pierce the shell of
the submerged animal. When the turtle was very distant, of
course the aim had to be taken at a considerable elevation, but
the marksmen preferred a longish range, because the arrow then
fell more perpendicularly on the shell, and entered it more
deeply.

" The arrow used in turtle-shooting has a strong lancet-shaped
steel point fitted into a peg, which enters the tip of the shaft.
The peg is secured tq the shaft by twine, being some thirty or
forty yards in length, and neatly wound round the body of the
arrow. When the missile enters the shell the peg drops out,
and the pierced animal descends with it towards the bottom,
leaving the shaft floating on the surface. This being done the
sportsman paddles in his canoe to the place, and gently draws
the animal by the twine, humouring it by giving it the rein
when it plunges, until it is brought again near the surface, when

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PELOMEDUSIDAE 397

he Strikes it with a second arrow. With the increased hold
given by the two cords he has then no difficulty in landing his
game.

" By mid-day the men had shot about a score of nearly full-
grown turtles. Cardozo then gave orders to spread the net. , . .
Three boat loads, or about eighty, were secured in about twenty
minutes. They were then taken ashore and each one secured
by the men tying the legs with thongs of bast.

" When the canoes had been twice filled we desisted after a
very hard day's work. Nearly all the ajiimals were young ones,
chiefly, according to the statement of Pedro, from three to ten
years of age; they varied from 6 to 18 inches in length, and
were very fat. Cardozo and I lived almost exclusively on
them for several months afterwards. Eoasted in the shell they
form a most appetising dish. These younger turtles never
migrate with their elders on the sinking of the waters, but
remain in the tepid pools, fattening on fallen fruits, and, accord-
ing to the natives, on the fine nutritious mud. We captured a
few full-grown mother turtles, which were known at once by the
homy skin of their breast plates being worn, telling of their
having crawled on the sand to lay eggs the previous year.
They had evidently made a mistake in not leaving the pool at
the proper time, for they were full of eggs, which, we were told,
they would, before the season was over, scatter in despair over the
swamp. We also found several male turtles, or capitaris, as they
are called by the natives. These are immensely less numerous
than the females, and are distinguishable by their much smaller
size, more circular shape, and the greater length and thickness of
their tails. Their flesh is considered imwholesome, especially to
sick people having external signs of inflammation."

The most recent account of these water tortoises is that
published by Dr. Goeldi from the MS. of Joao Martins da Silva
Continho, a former resident at Mandos on the Middle Amazon.
The " Tartaruga " (the Portuguese name for turtles) live from
January to July in the inundated, quiet backwaters of the forest-
region, feeding upon the various seeds of palms as these ripen and
drop successively ; rarely, and only when hard up, they are
carnivorous. The creatures hide under water below the trees,
when they are espied by the Indians, who dive down to a depth
of twenty and more feet to catch them in their arms. The

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398 CHELONIA

civilised Indians use a steel-pointed lance of hard wood, about
10 feet in length. A string connects the point with the shaft
around which it is wound. When stuck into the tortoise the
shaft and point part ; the string is either tied to the boat or to
a little float of light wood. In other districts an arrow with a
string is employed.

In August, when the water subsides, the tortoises return to
the rivers, and the entrance of the lagoon is closed with nets.
A number of boats with long poles drive them with much noise
towards the entrance. On their way to the rivers the tortoises
always go up-stream, and this is called the " arribaijao das
tartarugas," the ascent of the turtles. The fishermen post them-
selves at shallow spots or on sand-banks, and wait for the
creatures which come up to find a place for landing and laying.
The arrows employed are called sararaca, i.e. a thing which can
be disjointed ; they are about 4 feet long, and consist of a govio
or internodium of wood 9 inches long with a one- or two-barbed
steel point, and the shaft into which the gomo fits loosely. The
gomo is, moreover, connected with the shaft by a string made of
palm-fibres about 30 feet in length, partly wound round the
shaft, which ultimately acts as a float.

The laying takes place from the end of September into
October. Some of the parents seem to reconnoitre on land for
a few days. As a rule only females do this, and the natives
say that they are led by a " mestra!' The laying takes place
early in the morning. The number of females is so great that
they often block the way of the boats, and make a great noise
by knocking against their neighbours' shells. Each digs a hole
about 18 inches or 2 feet deep, and lays from 80 to 200 eggs.
Sometimes the laying individual is entirely buried by its neigh-
bours which are scraping their own holes.

In some districts the eggs are wanted for "manteiga"
(Portuguese for butter) ; and the turning over, or virofod of the
tortoises takes place later. In other districts they are caught
before the eggs are laid, and this barbaric and destructive custom
was formerly forbidden by the people themselves. Although the
provincial assembly tried to reinstitute the old reasonable customs,
the inspectors are often got over by bribery.

There are two ways of extracting the oil from the eggs. To*
get the thick oil used, mixed with tar, for shipbuilding, caulking.

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IX PELOMEDUSIDAE CHELYDIDAE 399

etc., the eggs are heaped up for five days and then worked. The
fluid oil for lighting is made from fresh eggs, which are put into
a boat and then trampled out with the feet. The oil is drawn
off into large earthen jars and put on the fire. Then it is rapidly
cooled. The best oil, used for frying fish, is that which is gained
from the roasted tortoises themselves. Fresh eggs are either
fried or taken with sugar, or mixed with manioca-flour and water.
The young, which are hatched in Januaiy, are likewise eaten
fried, or they are preserved in the fat of the parents.

An average tortoise yields 5 lbs. of fat, costing on the spot
two milreis. The whole full-grown animal, of one yard in length,
costs the same, and its meat is suflBcient to sustain a family
of six people for three days. To make 24 lbs. of oil requires
3000 eggs. Two or three tortoises would yield the same amount
from their fat. Consequently the destruction of the eggs causes
an enormous waste, and is after all the least economical pro-
cedure. In the year 1719, 192,000 lbs. were exported from
the Alto Amazones, representing 24,000,000 eggs. In 1700
there were still plenty of tortoises 50 leagues above the mouth
of the Para river. Now there is no assembly of more than
fifteen tortoises to be found anywhere within 300 leagues from
Para to the mouth of the Kio Negro. On the Kio Madeira, from
the mouth to the first cataract, 186 leagues distance, there are
now only two regular nesting localities. The upper Solimoes
and the Eio Yapura are still rich. Near Ega are regular
tortoise-ponds, called " curral," which yield sufficient support to
their owners; the animals are fed with manioca-flour and
leguminous plants.

Fam. 2. Ohelydidae. — The neck bends under the margin of
the carapace, but remains partly exposed. The nuchal shield
is absent except in two Northern Australian species. There are
twelve pairs of marginal shields. The plastron is composed of nine
plates, and is covered with thirteen shields, one of which is the
conspicuous intergular. The temporal region of the skull shows
great diversity. It is quite open in Chelodina, covered in by
broad expansions of the parietal bones in Flatemys, Emydurcty
and Elseya, or bridged over by a parieto-squamosal arch, which is
very slender in Rhineviys, strong in Chehjs and ffi/drasjns. The
palatine bones are separated by the vomer ; the nasals are variable,
mostly present, but the prefrontals are always small, and separated

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400

CHELONIA

by the frontals. The fifth and eighth cervical vertebrae are
biconvex.

This family, still represented by nearly thirty species, which
are divided into eight genera, is restricted to Notogaea, namely,
South America and Australia.

Chelys fimhriata, the '' Matamata," the only species of this

Fio. 87. — Skull of Chdys fimhriaia, x 1. A, Dorsal view of skull \. B, side view of skull
and hyoid apparatus. Cop^ copular piece • Fy frontal ; Jy jugal ; Z.o, lateral
occipital ; Mand, mandible ; O/j, opisthotic ; Orb^ orbit ; Patf parietal ; i*/,/, post-
froutal ; Plg^ pterygoid ; Q, quadrate ; Qjy Quadrato-jugal ; Sq^ squamosal ; I, II,
First and second branchial arch.

genus, inhabits the rivers of Guiana and Northern Brazil. Besides
the nuchal, there are seven neural plates ; the last pair of costals
form a median suture. Nasal bones are absent. The jaws are
very weak. The Matamata has a veiy peculiar appearance. The
nose is produced into a long, soft tube, at the end of which open
the tiny nostrils. The eyes are veiy small, and the orbits are
placed very near the anterior end of the skull, while the parietal

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CHELYDIDAE

401

region is broad and much elongated (Fig. 87, p. 400). The
quadrates are drawn out into trumpet-shaped tubes. The hyoid
apparatus is very large, with enormous anterior and posterior
horns. The head and neck are as long as or even longer than the
carapace, which is covered with thick, lumpy shields. The skin
of the thick neck, of the sides and under parts of the head, is
produced into many soft arborescent excrescences or fimbriae,
those of the chin and throat and the large ear-flaps being movable
at will, and probably used to attract fishes and other prey. The
tail is very short. The fore- and hind-limbs are webbed, the former

;i^9Durl)on

>v'r^^,jj^^;

Fig. SS.—ChflijsJimbriata (" Matamata •*). x ^^,

with five, the latter with four claws. Old specimens, which
reach a total length of three feet, are uniformly dark brown, and
look like a log covered with rough bark. The young are far less
ugly, with black and yellow spots on the shell, and with dark
stripes along the neck.

Very little is known about the habits of this peculiar creature.
It is said to lie submerged in the water, waiting for fishes, frogs,
or tadpoles, which are attracted by the playing motions of its
cutaneous excrescences. The jaws being so weak, and being
covered with a partly soft lip-like skin, it is probable that
they are not used for seizing the prey, but that the latter is
engulfed into the mouth with tlie inrusli of water into the throat.

VOL. vm 2 D

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402

CHELONIA

That this can be widened enormously is indicated by the
greatly developed hyoid apparatus.

Gfielodina. — The neck is long and slender, the head small and
smooth. The nuchal is terminal ; the intergular is large.
The neural plates are completely suppressed, all the eight pairs
of costal plates meeting in the middle line. The shell is very
flat. Anterior and posterior limbs entirely webbed, and with
only four claws. The tail is very short. Three species in
Australia, one in New Guinea.

Oh, longicollis reaches a shell-length of ten inches. It
inhabits Southern Australia. The illustrations make a de-
tailed description unnecessary. The colour of the dorsal shield
is uniformly dark rich brown, while the shields of the under
surface are yellow, with broad dark brown lines along the
sutures. These " long-necked Chelodines " have a striking appear-
ance, when they swim or creep about, with the neck either
stretched out straight or bent horizontally in an S-shape. The
whole creature looks neat and elegant ; the iris is pale yellow,

and gives the eye a very
intelligent expression. They
keep well in captivity, pro-
vided they are given the
choice of land and water.
My own prefer to spend
most of the day on land,
preferably under the ledge
of a stone, or perched upon
the stone itself if the latter
is in the shade, and not
too much exposed to view.
There they lie motionless, with the neck neatly tucked under
the shell, either to the right or to the left. Although the
eyelids may be closed, they can see well enough, owing to the
transparent condition of the lower lid. They feed in the water
upon soft animals, as for instance worms, smooth caterpillars, cock-
roaches or little frogs ; and they also take meat readily, provided
this is moved about. The food is invariably taken with a quick
sideward jerk of the neck and head. My specimens soon became
so tame that they left the water, and ran up to me with the
necks stretched to their fidl length, then snatching the bit of

Fio. 89. — Cfielodina longicollis.

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CHELYDIDAE

403

food, and retiring into the pond to swallow it. When left to
themselves they are rather nocturnal in their feeding habits.

Fig. 90. — Chdodina longicdlis (Australian long-necked Chelodines). x |.

Now and then they tuck themselves away for weeks without
feeding, for instance when they gb through a regular term of

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404 CHELONIA CHAP.

aestivation in the summer. The last winter they spent buried
in the moss, but occasionally, especially on bright and sunny
days, they went into the water for a few hours, chiefly to drink,
but sometimes also to take a little food.

Hydromedusa, a South American genus, htw a neck even longer
than that of Chelodina, which it much resembles externally. But
the nuchal shield, large and broad transversely, is situated behind
the anterior marginals, looking therefore like a sixth neural
shield. The neural plates form a continuous row, only the last
pair of costal plates meeting in the middle line. ff. tectifera
occurs in Southern Brazil, and in the La Plata. The shell is
dark brown above ; yellowish, with dark spots, below ; the head
and neck are olive-coloured, adorned with a broad white, black-
edged band on either side. Fore- and hind-Umbs broadly webbed,
and with four claws. Total length of the shell about eight
inches.

Fam. 3. Oarettochelydidae. — The shell is covered with soft
skin instead of horny shields. The limbs are transformed into
paddles, with elongated digits, and have only two claws. The neck
is short, and not retractile. In other respects the skeleton, not-
ably the plastron, pelvis, and skull, conform with the Pleurodirous
type. Only one species, Carettochelys insculpta, still imperfectly
known, from the Fly Eiver, New Guinea. Length of the shell
of the only complete specimen about 18 inches. This peculiar
creature seems to stand in the same relation to the typical
Pleurodira, as do the Chelonidae to the Testudinidae, except for
the complete reduction of the horny shields upon the shell, recall-
ing in this respect Sphargis and Trionyx.

Sub-Order 3. Trionychoidea. — The shell is very flat, oval,
or almost round, and is covered with soft, leathery skin instead
of with horny shields. The limbs are broadly webbed, and only
the three inner digits are provided with claws. Carnivorous,
found in the rivers of Asia, Africa, and North America.

The head and neck are completely retractile, bending by a
sigmoid curve in a vertical plane like that of the Cryptodira.
The jaws are concealed by soft, lip-like flaps, and the nose forms a
soft short proboscis. The ear is hidden. The skull, Fig. 91, is
flat, with three long posterior processes, formed by the supra-
occipital above, and the squamosals on either side. The whole
temporal region forms a wide, shallow fossa, without any indica-

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CARETTOCHELYDIDAE — TRIONYCHOIDEA

40 s

tion of being arched or bridged over. The premaxilla is extremely
small, unpaired, not even reaching the nasal cavity or the
vomer. The maxUlaries are
correspondingly enlarged,
surrounding the choanae,
which are separated by the
narrow vomer. The palatines
form a median suture, and
are joined behind by the long
basisphenoid, which separates
the long pterygoids from
each other. The quadrate
is trumpet -shaped, with a
posterior notch for the stapes.
The zygomatic arch is com-
plete, and is formed by the
quadrato-jugal and the jugal;
the latter joins the maxillary
and postfrontal, mostly reach-
ing the orbit ; in some cases
it also just meets the parie-
tal, thereby adding to the

strength of the pOStorbital Fiq. 91.— SkuW of Trionyx hurum. ^,From

- -,, « , above ; B, from the left side ; Condf occipital

arch. ihe pretrontals are condyle ; Fr, frontal ; J, Jug, jugal ; Z.o,

laVire ; nasals are absent. The ^a^^al occipital ; Max, maxillary ; Op.o,

,..^ . 1 1 1 /» opisthotic ; Par, parietal ; Pr.f, prefrontal ;

mandible is remarkable tor /vo, prootic ; Pt./, postfrontal ; Q, quad-

the ffreat development of ^**^' ^^' quadrato-jugal ; S.o, supra-occi-

o ^ pital ; *Sr?, squamosal.

the coronoid process.

The pubic and ischiadic bones enclose a large heart-shaped
foramen, and are free from the plastron ; the ilia are attached
only to the sacral ribs. The carapace is peculiar in so far as it
is very incomplete peripherally, the ribs extending considerably
beyond the costal plates, nor are they joined by marginal plates,
which are absent, unless they are represented by a few small
ossifications imbedded in the posterior marginal flap of the disc
{Emyda of India). The rim of the disc is always formed by a
horizontal, cutaneous, very flexible flap. All the dorsal plates
have a rough upper surface, vermiculated or rugose, as usual
with such dermal bones, which have lost most of or all their
homy covering, and have sunk more deeply into the skin. The

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406 CHELONIA

nuchal plate has iisually a pair of rib-like processes. The neurals
form a continuous series, except in the African CyclanorbiSy in
which they are much reduced in size, and separated by the costal
plates.

The plastron is imperfect, all its constituent nine elements
being only loosely connected with each other, and there remains
a wide median vacuity between the lateral elements. Most of
these plastral bones are reduced to splints, which, instead of
meeting by regular sutures, loosely interdigitate with their
jagged edges. In the young all these ventral elements are deeply
imbedded in the soft, leathery skin, and they do not at all
resemble in appearance those of the dorsal side. With age they
develop upon their ventral surface stronger and denser ossifica-
tions, which ultimately broaden out, sometimes beyond the
original underlying bone, and assume the characteristic venni-
culated surface -appearance. This is undoubtedly a process of
exostosis, a step towards revival of that armour which had been
much reduced ancestrally. To appreciate this condition, it is at
least suggestive that these mud-tortoises, when kept in the usual
hard-bottomed tanks, invariably become sore, the skin wearing
through where the imbedded plastral bones touch the ground.
Thus what is crammed into the short life of a captive individual,
is in the natural course of events spread over many generations,
whereby it has ceased to be pathological, and has become a com-
paratively new, tertiary, but regular feature.

It is not open to much doubt that the charadteristic features
of the Trionychoidea are not primitive but secondarj'. This is
indicated by the whole structure and behaviour of the carapace
and plastron. The softening of the whole shell, the loss of the
horny shields, the reduction of the claws, are the direct and
almost unavoidable results of life in muddy waters.

Geologically they do not seem to be very old. They appear,
already referable to the genus Trionyx, in the Upper Cretaceous
strata of North America. In the Lower and Middle Tertiary
strata many species existed in North America and in Europe,
and it is of great importance that in these species the costal
plates were much broader, and the marginal plates better developed,
than in the recent forms. Now their half-dozen genera,
with about twenty- four species, are confined to North America,
the tropical and warmer parts of Asia, and the Malay

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IX TRIONYCHOIDEA 407

Islands, and to Africa from the Nile to the Senegal and to the
Congo.

The habits of Triouychoidea have found few observers. Accord-
to L Agassiz,^ they live in the muddy bottom of shallow waters,
burying themBelves in the soft mud, with only the head, or a
small part of it, exposed. They breathe without moving the
body, by raising up the long neck and carrying the leathery snout
above water. When moving through the water they strike
horizontally with both pairs of limbs, alternating, however, the
right and left ; but when they start suddenly, the front limbs are
seen moving together towards the tip of the snout, and then
striking simultaneously backward with great power. As the
shield does not project forward, the fore-limbs usually move
beyond the shield, and as its outer edge is sharp, and the feet are
broad, their webs reach above as well as below the plane of that
edge, so that the water is driven partly over and partly under
it. When they move along the bottom, the limbs still move
horizontally, the webs striking against the water, and the inner
toes, those with the claws, against the bottom. They also bury
themselves horizontally, becoming covered by only a thin layer of
mud. They readily resort to the shell for protection. The neck
and hejid are withdrawn entirely, the loose skin rolling off from the
greater part of the neck ; and the skin of the legs also slips off, as
far as the elbows and knees. In confinement they exhibit great
quickness ; their movements are abrupt and unsteady, except when
they swim rapidly in one direction. They then dart their long
and slender neck quickly forwards or sideways and upwards, as
snakes do, and bite in the same way, striking suddenly. Their
temper is bad or even ferocious, and large specimens are quite
dangerous.

Their food consists of all sorts of aquatic animals, fish, frogs, and
molluscs,, for instance Anodonta and Paludina. According to the
different diet, many species develop a peculiar kind of dimorph-
ism, a reasonable explanation of which has been given by
Boulenger. In the young the horny coverings of the jaws are
sharp, with cutting edges, and in those specimens which keep to
a diet of fish and other soft creatures, the jaws remain in the
same condition. But in those which take to living upon molluscs,
the hard shells of which they have to crush, the horny edges are

^ CorUribuiions to the Natural History of the U.S.A., vol. i. 1857, p. 333.

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408 CHELONIA

worn down ; and broad, thick, homy, crushing pads are developed
in their stead, the supporting parts of the jaws becoming more
massive. The masticatory muscles are likewise enlarged, and a
tubercle grows upon the lower border of the jugal bone, whence
arises part of the masseter muscle.

The eggs are round, thick -shelled, but very brittle ; they are
laid in the sand above the level of the water, and this is the
chief occasion on which these tortoises creep on land.

Trionyx. — The plastron has no special cutaneous valves for
the concealment of the hind-limbs. This is the principal genus,
with the greatest number of species and the widest distribution,
the latter coinciding with that of the whole family. The upper
surface of the shell of young specimens frequently forms numerous
longitudinal ridges or series of little horny tubercles which dis-
appear with age.

T. feroxy the commonest " Soft-shelled Turtle " of the United
States. Olive above with scattered, small, round, black spots ;
young with conical, spine-like tubercles, especially on the nuchal
border and on the posterior portion of the shell, which has a
pale, black-edged border. A light, black-edged streak passes
through the eye and joins its fellow on the snout. The limbs
are olive brown, spotted and marbled with black. The under
parts of the shell are white. Very large specimens have a shell
18 inches in length and 16 inches wide. Holbrook gives the
following account of its habits : —

" A voracious, carnivorous creature. They reside most con-
stantly in the water, swim with rapidity, and choose for their
retreat holes under the banks of rivers, or under rocks ; and
not unfrequently the trunk of some huge forest tree, fallen
into the stream, affords them shelter. Sometimes they leave the
water and conceal themselves in the mud: I have frequently
seen them thus buried to the depth of 2 or 3 inches, leaving
only a small breathing hole for the long neck and narrow head,
which is occasionally thrust out, but most commonly it is
retracted so that one would pass near without observing their
habitation ; and if seen, it might easily be mistaken for the
residence of some large insect. At other times they may be seen
in numbers on rocks in shallow water, basking in the sun,
apparently asleep. They bite severely when provoked, darting
forward with great velocity the long neck and head, and not

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TRIONYCHOIDEA

409

uufrequently spring upward at the same time and make a loud
hiss. In the month of May the females seek sandy places along
the banks of the waters they inhabit to lay their eggs, generally
about sixty in number; and it is remarkable that, though their
motions are slow and difficult on dry land, yet at this season
they sometimes mount hillocks several feet high. The flesh

Fio. ^2.— Trionyx ferox (American Soft-shelled Turtle). y -^.

affords the most delicate food, surpassing that even of the
Green Turtle. The geograpliical distribution is interesting. It
inhabits the Savannah as well as all those rivers that empty
into the northern borders of the Gulf of Mexico ; it ascends up
the broad Mississippi, and is found in all its tributaries, even
to the very foot of the Eocky Mountains ; it abounds in the
chain of great northern lakes both above and below the Falls of
Niagara, and is common in the Mohawk, a tributary of the

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4IO

CHELONIA

CHAP-

Hudson river ; but it is not found in any other Atlantic stream
between that and the Savannah river, a distance of nearly 800
miles."

T. triunguis, the only African species, ranging from the
Senegal and Congo into the Nile-system, but occurring also in
Syria, is perhaps the largest of all Trionychidae, reaching a
shell -length of almost 3 feet. The adults are olive-brownish

Fig. 93. —Trionyx gangeticus (young).

above, the throat and under parts of the shell with round, white
spots separated by a dark network. The young have whitish
specks and spots.

T. gangeticus and T. hurum are the principal Indian species.
The former is the larger of the two, with a shell of more than 2
feet in length ; olive above, the young with fine black vermicula-
tions ; head with a black longitudinal streak from between the
eyes to the nape, intersected by two or three chevron-shaped black
streaks ; under parts yellowish. T. hurum is olive brown above

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TRIONYCHOIDEA 4^1

and below, in younger specimens with conspicuous, large, yellow
spots on the sides of the head. The young are ornamented with

Fig. 94. — Trionyx fonnosa (young). x 1.

two or three pairs of large round spots on the back, and the
same applies to the beautiful young of the Burmese, T. formosa.
The three genera, Cycloderma and Cyclanorhis of Tropical
Africa, and Eviyda of India, have a pair of cutaneous femoral
valves or flaps on the plastron, beneath which the hind-limbs
are withdrawn.

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CHAPTER X

DINOSAURIA CROCODILIA

Sub-Class V,— DINOSAURIA.

Mesozoic, long-tailed, toothed reptiles, with distal ischiadic sym-
physis, terrestrial limbs, large fixed quadrate banes and
bifurcated ribs.

The Dinosaurs begin and end with the Mesozoic epoch, and
have a world-wide distribution. The name, " terrible Eeptiles,"
refera to the gigantic proportions which many of them attained,
not a few of them surpassing in size and shape the fantastic
pictures of the dragons of our fables. Although these creatures
came to an end millions of years before the first man-like beings
appeared, it is reasonable to suppose that the widely-spread m}'th8
of dragons are based upon the accidentally disclosed skeletons of
these monsters.

Tlie skull is built after a plan which may be derived
from a combination of the Crocodilian and Ehynchocephalian
skulls, but the detail varies considerably in the many and
much diversified members of this large sub-class. There is
as a rule a pre-orbital foramen, which is smallest in the Omi-
thopoda. The orbit is completely encircled by bones, and
the temporal fossa is divided by a squamoso-postfrontal or
post-orbital bridge into a smaller supra-, and a much wider
infra-temporal portion, the latter being bordered below by the
jugal and quadrato-jugal, and this is firmly connected with the
quadrate by an ascending process. The quadrate is long, more
or less verticil in position, slanting either forwards or backwards,
and firmly fixed above by the squamosal, perhaps also by a supra-
temporal bone. Tlie orbit is bordered by the jugal, lacrymal,
pre- and post-frontals. The interparietal foramen seems to be

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CHAP. X DINOSAURIA — SKELETON 4 I 3

abolished. Teeth, mostly alveolar and laterally compressed, are
restricted to the dentary, maxillary, and premaxillary bones.
In the Orthopoda the latter carry no teeth, or these are restricted
to the lateral portion, leaving a wide diastema. This toothless
part plays upon a peculiar crescent-shaped bone, the so-called
predentary, which rests loosely upon the anterior ends of the
mandibular rami, which latter do not as a rule form an
osseous symphysis. The Ceratopsia possess in addition a similar
upper toothless piece, the prerostral, a kind of pre-premaxilla.
The morphological value of these extra pieces is quite obscure ;
they were in all probability provided with thick, horny pads.
The bones of the roof of the mouth recall in their arrangement
that prevailing in the Ehynchocephalia and the Parasuchia.
There are two pairs of large vacuities ; one between the maxillae,
ectopterygoids and palatines ; the other between the lattfer, the
maxillae and the usually small or slender vomers. The pterygoids
are perhaps the largest bones, and form a rather long symphysis ;
laterally and behind they abut against the quadrate, anteriorly
against the ectopterygoids and the palatines, which latter they
sometimes separate. A peculiar feature of some skulls, e.g.
CeratosauTus and Triceratops is the great size of the groove in
which the large hypophysis of the brain is lodged.

The vertebrae are very variable, amphicoelous, opisthocoelous,
nearly plain, with a' slight concavity behind, or occasionally
procoelous in the anterior region of the tail. Besides the usual
pre- and post-zygapophyses many Sauropoda and Theropoda
possess on the posterior trunk-vertebrae additional joints, effected
by a vertical wedge, the hyposphene, which extends backwards
from between the post-zygapophyses and fits into a notch
between and below the anterior zygapophyses of the next follow-
ing vertebra. These additional articulations are analogous to
the zygosphenes and zygantra of snakes and iguanas, except that
in these Sauria the wedges are formed on the opposite, namely
the anterior ends of the vertebraa The vertebrae of the neck
and trunk are devoid of intercentra, but those of the tail carry
long chevron-bones. The number of sacral vertebrae is generally
increased to four or five. The ribs have well-developed capitula
and tubercula, and the former have the tendency to shift from the
centra or from their parapophysial processes on to the usually
much elongated diapophyses of the neural arches. This arrange-

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414 DINOSAURIA

ment, recalling the Crocodilian condition, results in an increased
capacity of the dorsal portion of the body-cavity. Intervertebral
articulation of the ribs does not occur except sometimes in the
sacral region. Abdominal ribs are rare, but they occur in some
of the Theropoda, e,g, in Gomjpsognathus,

The sternum seems to have been mainly cartilaginous, with
a pair of irregular, disc-shaped ossifications. How the coracoids
were attached is unknown ; they are small, generally with a
foramen, but the scapulae are always very strong and slant
backwards. Clavicles and interclavicles seem to be absent.

The fore-limbs are as a rule powerful, although often much
shorter than the hind -limbs, which are then enormously
developed, and in many genera of two of the main groups show
a tendency towards a semi-erect gait. Some of the Dinosaurs,
e.g. Iguanodon and Brontozoum, were absolutely bipedal. Others
seem to have hopped like Kangaroos. In correlation with this more
or less erect mode of progression the iliac bones are very strong,
much elongated horizontally, and attached to more than three,
often to five or even more, vertebrae. The pubic bones show two
main types. Each consists either of a single strong shaft, which is
connected distally with its fellow; or (Orthopoda) this main
shaft sends out, below its point of contact with the ischium, a
long process, the so-called post-pubis, which is directed down-
wards and backwards. In the latter case it runs parallel
and in close contact with the ischium. Such bifurcated pubic
bones never meet in the middle line. The ischia, on the other
hand, are always connected with each other, not so much by
fusion as by syndesmosis.

The hind-limbs exhibit all stages from a simple, plantigrade
and five-toed state to a decidedly digitigrade, four, and even three-
toed arrangement. Many genera exhibit the tendency to form an
intertarsal joint, a feature elsewhere known in birds only, where
it is typical and universal The astragalus sends up an ascending
process which tends to fuse with the anterior aspect of the distal
end of the tibia, and the calcaneum is sometimes more or less
firmly attached to the fibula. In Compsognathtis even the distal
tarsalia have begun to fuse with the metatarsalia, so that this
reptile at least has a typical intertarsal joint. The femur is
remarkable for the frequent possession of a " fourth " trochanter
on the middle of the inner aspect of the shaft, undoubtedly

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X SKELETON 4 1 5

for the insertion of the long caudi- femoral or long adductor
muscle.

Many Dinosaurs possess hollow instead of solid bones. The
vertebrae have large cavities in the Sauropoda, notably in Bronto-
saurus ; in many Theropoda, e.g. Coelurtis, Anchisaurvs, Compso-
gnathus, the limb-bones and the vertebrae are hollow, the latter
being reduced to thin-walled shells with a few inner partitions, the
bones being at the same time much swollen and enlarged. In
the Omithopoda the vertebrae are solid, but the limb-bones are
hollow. The reason of this hollowing out is not easily found.
Undoubtedly it results in a saving of material and weight,
whilst at the same time, without loss of strength, the surfaces for
the attachment of tlie necessarily powerful muscles are increased.
But Compsognathus is a small, Brontozoum a gigantic, creature.
On the other hand, the bones of the huge Stegosauri are solid.
Most probably these cavities were, as in birds, filled with air-sacs
ultimately in communication with the lungs ; and it is by no means
a baseless suggestion of Haeckel's that the Dinosaurs were warm-
blooded. Their mode of propagation can only be guessed at
from the circumstance that a rather well-preserved specimen of
Compsognathus contains in its abdomen what may possibly be
an embryo. There is nothing against the assumption that the
Dinosaurs were viviparous; on the contrary, it seems more
natural than that, for instance, an Atlantosaurus of more than
100 feet in length and many tons in weight, should have laid

Some of the herbivorous Dinosaurs, namely, the Stegosauri
and the Ceratopsia, had a dermal armour of variable extent ; the
plates were loosely imbedded in the skin, and reached their
greatest size along the middle of the back and tail, and these
crested plates were probably covered with horny scutes,
obviously weapons of defence. The Ceratopsia were armed with
a pair of huge pointed horns on the head, and a smaller one on
the nose (see Fig. 102, p. 430). It is difficult to guess the
use of the weapons of these terrestrial monsters, unless they were
employed against the equally large carnivorous Dinosaurs or in
the combats for the possession of their charming mates.

About the ancestry of the Dinosaurs we know nothing except
that their affinities lie with the Crocodilia ; but it is impossible
to derive either from the other. The oldest forms, in the

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4l6 DINOSAURIA

present state of our knowledge — those which have left their
three-toed spoors in the Trias of Connecticut — were already much
specialised by having attained to an upright bipedal gait, while
the Sauropoda, which except for their gigantic size are the most
generalised, are of comparatively recent date, none of them being
known from strata older than the Upper Jurassic. Twenty years
ago, until the discoveries of numerous kinds in the United States,
our knowledge of the whole group was very limited. There is a
widely spread notion that the birds have sprung from some Dino-
saurian stock. Huxley was the first to show clearly that birds
were an oflFshoot of the reptiles, and he said of the Dinosaurs,
especially his Ornithoscelida {Iguanodon, Scdidosaurus, Megcdo-
sauruSy Compsognathvs, and others), that they " present a large
series of modifications intermediate in structure between existing
reptiles and Aves." Baur proved to his own satisfaction that we
have to look for the ancestors of the Batitae among the herbi-
vorous Dinosaurs, especially the Ornithopodous forms, whilst the
Carinatae are descendants of the Eatitae. However, even he had
to give up this absolutely unwarrantable view.

It is easy to select a considerable number of characters
amongst the various Dinosaurs which also occur in birds, and
some of these have until a recent date been considered as peculiar
to birds. For instance, the double, bifurcated pubic bones of the
Orthopoda ; the increased number of vertebrae to which the hori-
zontally elongated ilia are attached, especially in the forms with
an upright gait, and the bipedal feature itself ; the possession of
an ascending process of the astragalus and its fusion with the
tibia in Compsognathus and Ceratosaurus among the Theropoda,
and in Ornithomimus ; the attachment of the distal tarsalia to
the metatarsalia, e.g. in Coynpsognathus, — in fact, the formation
of an intertarsal joint, a feature otherwise characteristic of,
and peculiar to, birds ; the frequent reduction of the fifth
metatarsal bone ; the backward position of the hallux and the
proximal reduction of its metatarsal in Coinpsognathus ; the
elongation and partial fusion of the functional metatarsals in
the latter genus and in Ceratosaurus ; the regular increase of the
phalangeal numbers of the first four toes from two to five in
many of the Ornithopoda ; — in short, the great resemblance
between the feet of some of the Dinosaurs and those of the birds.
However striking these arguments are, they are instances of con-

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AFFINITIES 417

vergent analogies. The upright walk, which has been assumed
and improved upon independently by members of both Theropoda
and Orthopoda, has produced the same, or nearly the same
modifications in them as in the birds.

It is easy to show that these features are mere coincidences.
The oldest bird known is Archaeojyteryx from the Upper Oolite of
Bavaria. Consequently all those Dinosaurs, which are of the
same and of later date, have to be excluded from the supposed
ancestry, and they happen to be those in which (as in Ceratosaiirus,
ComjpsognathuSy OrnithomiviuSy Iguanodon) the resemblances are
greatest. There remains only Anchisaurus of the Upper Trias,
more or less contemporary with the Brontozoum, which left its
three-toed footprints {Archaeopteryx has four well-developed toes)
with Zanclodon. Moreover, the most bird-like foot is either that
of the Theropoda, which, like Anchisaurus and Zanclodon, differ
from birds by the formation of the pelvis, or of some of the
latest Ornithopoda. What, then, is the good of selecting a
number of bird-like features from members of Dinosaurs which
we are bound to class in different groups, and which existed,
some in the lower, others in the middle, or even in the latest
Mesozoic periods ?

Lastly, the advocates of the Dinosaurian ancestry of birds
cannot have fully appreciated the enormous differences between
the wing of Archaeopter-yx and the fore-limb of any Dinosaur
with the most avian resemblances in the hind-limbs. The fore-
limbs of these reptiles are modified in a direction diametrically
opposed to that from which a bird-like wing could be developed.
The skull presents another difticulty,and here again Com2)sognat1m8,
a contemporary of Archaeopteryx, comes perhaps nearest to that of
a generalised bird's skull. The ancestors of the birds must have
combined the following characters: — Of not later than Mid-
Oolitic age, with bifui-cated pubic bones, four functional toes, elon-
gated metatarsals, complete clavicles, premaxillary teeth, and free,
not firmly fixed quadrate bones. But such creatures are not
Dinosaurs.

We divide the enormous number of Dinosaurs according to
the formation of the pelvis, that of the hind -limbs, and the
dentition, into four orders.

VOL. VIII 2 E

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41 8 DINOSAURIA

Order I. SAUROPODA.

Fuhes simjyle, with symphysis. Premaxillae with teeth.
Plantigrade.

The teeth are mostly spatulate, laterally compressed, with
sharp edges, but without serrations. Skull with a pair of large
pre-orbital fossae. The centra of the vertebrae of the trunk have
large lateral cavities. The fore- and hind-limbs are pentadactyle,
plaiitigrade, and hoofed, of the typical walking type ; the bones
of the limbs are stout and solid ; the femur is devoid of an inner
distal or fourth trochanter. The carpal and tarsal bones are free.
Herbivorous. Tlie Sauropoda comprise some of the most gigantic
terrestrial creatures which have ever existed, compared with some of
which the bulk of an elephant appears almost insignificant. Their

Fig. 95. — Skeleton of Brontosauru^ excelsu^. x y^^. (After Marsh.)

range in time extends from the Lower Oolite into the Cretaceous,
with a perhaps world-wide distribution, namely, Western Europe,
North America, Patagonia, Madagascar, and India. Although
they are, except for their size, the least specialised of all
Dinosaurs, none of the Sauropoda hitherto discovered are old
enough to claim to be the ancestors of the other Dinosaurs.

Brontosauriis excelsus of the Upper Jurassic of Wyoming was
a giant at least 60 feet long and about 10 feet high. The head
is extremely small in proportion, not so broad as the fourth of
the thirteen vertebrae of the long and flexible neck. The
trunk is comparatively short, the tail longer than the neck, and
provided with numerous chevron-bones. Most of the vertebrae
are hollow, especially the five co-ossified sacrals. The spinal canal
of the sacral region is very wide, indicating a strong sacral
swelling in conformity witli the huge posterior limbs. The
pubic bones are stronger than the ischia. The long axis of the

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SAUROPODA

419

Fig. 96. — Front view of the pelvis
of Morosauru^ grandis. x ^.
(After Marsh. ) a, First sacral
vertebra ; 6, " transverse pro-
cess " (rib) of first sacral ; iZ,
ilium ; t^, ischium ; nc, neural
canal ; pb^ pubis.

former stands almost vertically like that of elephants, and the
knee is scarcely bent in the erect position. The shoulder-girdle
consists of long scapulae, broad at the base and small, almost
square and perforated coracoids, which
latter fit into a pair of partly ossified
plates representing the sternum.

AtUintosauriis immanis of the Upper
Jurassic of Wyoming and Colorado, is
supposed to have been 115 feet long,
perhaps the biggest and bulkiest of
all animals, the femur measuring more
than 6 feet in length and 2 in width
at the upper end.

Morosaurus grandis^ of the Upper
Jurassic of Wyoming, with allied forms
in the Purbeck and Wealden of
England, reached a length of 30 feet;
in general appearance resembling Brontosaurus, but the sacrum
consists of four vertebrae only, and the ischia are bent back-
wards in their distal halves, so that their symphysis is formed
by the shafts instead of by their ends.

Ornithopsis and Cetiosaumis, likewise huge creatures, from* the
English Wealden and from the Great Oolite respectively, are
rather imperfectly known, although several species of each,
under many generic synonyms, have been described

Diplodocus longus, of the
Upper Jurassic of Colorado and
Wyoming, is almost completely
known. More than 40 feet
long, it had a head in its general
outlines not unlike that of a
horse, the skull being about two
feet long. The outer nasal
openings are confluent, elongated,
and lie far back on the top of the
skulL There is a pair of large antorbital, and a pair of smaller
lacrymal fossae. The teeth, long and slender, are restricted to
the adterior portion of the mouth, with many successors, which,
decreasing in size, lie on the inner or lingual side of the
functional tooth, like the cartridges in a repeating rifle. The

Fio. 97. — Skull of Diplodocus lonffus,
x^. (After Marsh.)

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420 DINOSAURIA chap.

functional teeth themselves are implanted in sockets. The
generic name refers to the peculiar chevron-bones, each half of
which diverges into an anterior and a posterior branch.

It is difficult to understand how these huge, long-necked
Sauropoda lived and moved about. The long neck suggests at
first sight predacious habits, but the teeth, rather feeble in
DipIodocuSy and distinctly of the plant-cutting type in other
genera, put this out of the question. The high position of the
unpaired nasal opening, and the shortened nasal bones of
Biplodocus, are features indicative of aquatic habits, but the
short-toed, plantigrade limbs are absolutely adapted to terrestrial
life, and we cannot well assume that such enormous brutes as
Atlantosaurus could possibly have ventured into swampy ground.

Order n. THEBOPODA.

Piihes simple, with symphysis. Premaxillae with teeth.
Dlgitigrade. Carnivorous.

The teeth are pointed, recurved, laterally compressed and
serrated. The nasal openings are large, lateral, and nearly
terminal. The vertebrae and the large bones of the limbs are
hollow. The fore-limbs are considerably shorter than the hind-
limbs, which are distinctly digitigrade, many of the species
having a pronouncedly upright gait. The proximal tarsalia
show a tendency to fuse with the tibia, and the astragalus
has sometimes an ascending process, by which the fusion with
the tibia is strengthened. The first and fifth metatarsals are
often reduced, while the three middle bones are elongated and
sometimes even fused with each other, so that the whole foot
assumes a striking resemblance to that of birds. The ter-
minal phalanges are protected by curved claws. Owing to
the shortness of the fore-limbs, and the often considerable
length of the hind-limbs, which are strongly bent at the knee
and the ankle-joint, these animals must have progressed some-
what like clumsy kangaroos.

The Theropoda, of which a great number of genera are now
known, from the size of a slender cat to that of an elephant,
lived from the Upper Trias to the Upper Oolite, both in Europe
and in North America.

Brontozoum giganteum, one of the oldest forms, is known

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THEROPODA 42 1

from its foot-spoors only, which, together with other three-toed
spoors in the sandstone of the Connecticut valley, were originally
described and figured by Hitchcock as Ornithichnites (txvo^ =
track, or spoor). Some of these imprints are more than a foot in
length, the right and left spoors following alternately at a dis-
tance of from four to six feet. In some cases the long trailing
tail has left a furrow behind, and the large tracks are accom-
panied or crossed by much smaller, and even by quite tiny
tracks, otherwise similar, and undoubtedly made by the young.

Anchisaurus, from the same locality, was still Sauropodous,
in so far as the metatarsals are still free, with two, three, four,
and five phalanges on the first four
toes, but the fifth metatarsal is re-
duced, carrying a vestige of only one
phalanx, and the proximal tarsal bones
are fused with the tibia and fibula

, m . 1 1 .!_ Fig. 98. — Skull of Anchisauru^

respectively. Total length some seven codums. xf (After Marsh.)

feet, of which about four belong to the ^ Nasal fossa ; 6, antorbital,

.- " c, infra- temporal, d, supra-

taiL temporal, and 0, orbital fossa ;

Zanclodon, from the Keuper of g, quadrate bone.
Wiirtemberg, about ten feet long, with pentadactyle hands and
feet. Ischia stronger than the pubic bones, which are distally
much broadened. The femur is nearly three feet long, and
possesses a fourth trochanter. The astragalus has an ascending
process, and is fused with the tibia. The toes are short, strong,
and clawed. The shoulder-girdle and fore-limb are strong, the
latter well adapted to grasping. The teeth are much compressed
laterally, with sharp, finely serrated edges. Several allied genera
have been described from the Ui)per Trias of France and England ;
others from corresponding strata of India and South Africa.

MegalosauTMSy from the Trias to the Wealden in England
and France, with other species in Colorado and India, reached a
considerable size, larger than that of any other Theropoda, the
scapula of M. hucklandi being nearly three feet long, and the
femur still longer. The hind-limbs are twice as long as the
fore-limbs. The cervical vertebrae are short, the neck being
much shorter than the tail. Hands with five fingers, feet with
four toes. Pubic bones long and slender, with a broad sym-
physis. With well-developed abdominal ribs, resembling those
of crocodiles.

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422

DINOSAURIA

Allosaurtis, from the Upper Jurassic of North America, with
only three toes. Ischia and pubes united into one sjTiiphysis.
Anterior extremities very short. Sacrum consisting of four
vertebrae. Total length of some of the larger species about
twenty feet.

Ceratosaurus nasicornis, from the Upper Jurassic of Colorado,
is about seventeen feet long. The generic and specific names
refer to the nasal bones, which are raised into an unpaired

Fig. 99. — Skeleton {^M and skull of Ceratosaurus nasicornis. (After Marsh.) a.
Nasal cavity ; 6, bony horn-supporting excrescence ; c, pre-orbital fojisa : t/, orbital
fossa.

longitudinal crest. This, by its rough surface, suggests that it
was covered by a horny sheath, or carried a horn. The large
skull, about two feet in length, is armed with strong, slightly
curved, laterally compressed, sharp teeth, unequal in size. The
pre-orbital foramen is large, bordered above by the prefrontiils,
which are raised into prominent knobs. The supratemporal
foramina are extremely small, the lateral foramina very large.
The quadrate slants backwards. The sacrum consists of five
vertebrae. The caudal vertebrae carry long and slender chevron -
bones. The pubes and ischia are long and slender, each forming
a separate symphysis at their broadened ends. The three meta-

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THEROPODA 423

tarsals are elongated and fused with each other. There seems
to have been some dermal armour in the shape of osseous plates,
which extended in one series from the occiput over the neck.

Coelurus gracilis, of the Upper Jurassic of Wyoming, and
closely allied forms in the Wealden of England, are remarkable
for the pneumaticity of the centra and processes of their ver-
tebrae, the bony parts of which are restricted to thin, hollowed-
out shells, so that the whole skeleton must have been very
light. Computed length of these imperfectly preserved creatures
*about five feet.

Hallojpus victor, of the Upper Jurassic of Colorado. Anterior
extremities very short, with only four fingers ; posterior limbs
very long and slender, especially the tibia ; the much elongated
metatarsals are separate, the first absent, the fifth much reduced,
so that the foot is tridactyle ; the calcaneum projects like a heel.
The ilium is attached to two siicral vertebrae only ; the pubes
are slender, forming a narrow symphysis, while that of the
ischia is broad. Most of the bones of this creature, which
probably progressed by hops, are hollow^ Total length about
three feet, the length of the hind-limbs being about nine inches.

Coinpsognathus longvpes, of the Upper Jurassic of Bavaria, is
one of the smallest of all the Dinosaurs. It is most remarkable
on account of its almost bird-like feet. The fibula is much
thinner and somewhat shorter than the tibia; the latter is
closely attached to, although not fused with the proximal tarsal
bones, while the distal tarsals are fused with the united and
much elongated second, third, and fourth metatarsals ; the fifth
is reduced to a short bone near the intertarsal joint ; while the
first is represented by its distal portion only, which is stowed
away on the hinder aspect of the middle of the second meta-
tarsal, and carries two phalanges. The three middle toes
consist of three, four, and four phalanges respectively. Whilst
the whole hind -limb is typically avian, the pelvis is quite
different; the pubic bones are simple, slender, and directed
forwards, forming a symphysis with their whole distal halves,
and broadening out distally into a horizontal process directed
towards the symphysis, which is likewise formed by the fusion
of the inner surfaces of the thin and rather flat iscliia. The
fore-limbs are only half the size of the hind-limbs. The
neck consists of about ten vertebrae, mostly with long and

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424 DINOSAURIA

pointed ribs. Tail long with well-developed chevrona The
skull is long and pointed, composed of thin bones, which have
lost most of the sutures; with large lateral, temporal, and
pre-orbital, but without supratemporal, foramina. Premaxillae,
maxillae, and mandible with numerous slender and rather long,
conical, alveolar teeth.

Order IH. OBTHOPODA.

Edch pubic hone consists of an aiiterior or pre -pubic and or
posterior or post-pubic branch, neither of which forms a
symphysis, Premax^illae without teeth. With a premandi-
bular predentary piece. Herbivorous,

The so-called pre-pubis is homologous with the pubis of most
recent reptiles, and with the pectineal process of birds, while the
" post-pubis " is homologous with the processus lateralis of
Chelonians and Saurians, and with the " pubis " of birds. The
right and left halves of the pubis remain widely asunder ventrally.
In many cases the post-pubis, always directed obliquely backwards,
lies closely against the shaft of the ischium, which always forms
a distal syndesmosis, or a symphysis, with its fellow. The fore-
limbs are usually very short, provided with five or four short
and strong fingers. The hind-limbs are long and strong, mostly
with three, sometimes with four functional short toes, either
plantigrade (Stegosauri) or digitigrade (Ornithopoda). Femur
with an inner distal, or fourth, trochanter. The dentition is of
the herbivorous type, restricted to the dentaries of the mandible
and to the maxillary bones, leaving the whole or the greater part of
the premaxillaries free. The additional " predentary " piece of the
mandible is possibly a calcified, but originally horny, pad. The
teeth are greatly compressed laterally, and finely serrated, but are
much ground down by use ; several rows of successional teeth lie
on the inner or lingual side. The skull is strongly built, with
large anterior nasal openings ; pre-orbital foramina very small or
absent ; orbits completely encircled by bones ; supratemporal
foramina small, lateral foramina large. Quadrate large, vertical
or slanting slightly forwards. The vertebrae are solid, not
hollow ; siicrum consisting of four, five, or more vertebrae ; ribs
bifurcated, the capitula carried either by the centra, or moved up
to the diapophyses of the neural arches ; chevron-bones numer-

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X ORTHOPODA — STEGOSAURI 425

0U8, and frequently long, especially on the anterior half of the
long and heavy tail.

Orthopoda occur from the Lias to the Upper Cretaceous,
both in Europe and in North America. The name Orthopoda,
invented by Cope in 1866, is appropriate for obvious reasons;
it comprises the Stegosauri and Ornithopoda of Marsh (1881).
The latter term is not very fortunately chosen, considering that
the whole hind-limb of the Theropodous Compsognathus is far
more ornithic than that of any three-toed Ornithopoda, in which
the tarsalia rarely fuse with the tibia and never with the meta-
tarsals. To apply the term Ornithopoda to the whole order is
quite unjustifiable, unless it is meant to apply to the strikingly
bird-like configuration of the pelvis.

Sub-Order 1. Stegosauri — The fore- and hind-feet are planti-
grade, or nearly so, the metapodials being but little elongated,
with more than three functional digits. The bones of the limbs
are solid. The ribs of the trunk are bifurcated, and are carried
by the diapophyses of the neural arches. The body, especially
the back, is protected by dermal bony plates, which are not con-
nected with the internal skeleton. •

Scelidosaurus harrisoni. One nearly complete skeleton, about
11 feet in length, from the Lias of Lyme Kegis. About twenty-
four pre-sacral vertebrae, of which six or seven belong to the neck,
four sacral and about forty caudal vertebrae. Four fingers, four
toes, with 2, 3, 4, 5 phalanges, the fifth metapodials being quite
vestigial ; the hallux and poUex are very short, so that the foot
at least is functionally tridactyle. The tarsal bones remain
separate. The head is very small Two rows of ridged bony
plates extend from the neck over the back, and converge into one
row upon the long tail ; smaller plates, arranged in many rows,
seem to have protected the sides and uuder parts. Ht/laeosaurus
and Polacanthus of the English Wealden are allied forms.

StegosauruSy with several species from the Upper Jurassic of
Colorado and Wyoming, and others, e.g. S, armatus ( = Omosanrus),
from the Kimmeridge Clay of Wiltshire in England The head
is relatively very small, and the brain is surpassed several times
in thickness by the huge sacral swelling of the spinal cord.
Teeth numerous and small. All the cervical and trunk-vertebrae
carry bifurcated ribs, those of the trunk being carried entirely by
the very high neural arches. The fore-limbs are only about half

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426

DINOSAURIA

the length of that of the hind-limbs, so that these creatures,
which were undoubtedly quadrupedal, must have had a very
peculiar gait, standing with the head, neck, and shoulders much
lower than the arched back and pelvic region. The ulna has a
strong olecranon ; the hand has four functional fingers. The pre-
acetabular portion of the ilium is much elongated ; the pre-pubic
branch stands horizontally, while the post-pubis is closely adpressed
to the ischium. The astragalus is fused with the tibia, the
calcaneum with the fibula. The foot has only three short toes,
protected, like the fingers, by hoofs. The dorsal dermal armature

Fig. 100. — Skeleton and dermal armour of Slerfosaunis unffulatus. x ^. (After Marsh.'

consists of very high, crest-like plates. aS'. ungulatus of North
America has a computed length of 28 feet, with the hind-limbs
about 7 feet long. This creature was nearly 10 feet high, when
measured from the ground to the tips of the dermal crests on the
middle of the back. These bony, laterally compressed plates are
themselves nearly 3 feet high, and are replaced, on the hinder
portion of the tail, by several pairs of pointed spikes about
2 feet in length.

Sab-Order 2. Omithopoda. — The hind-limbs are distinctly
digitigrade, usually with only three functional toes, protected by
claws. The long bones are hollow. Femur with a long fourth
trochanter. Without dermal armour-plates.

Cajivpiosaurus. — Several species, up to 10 feet in length, from

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ORTHOPOD A — ORN ITHOPOD A 427

the Upper Jurassic and the Wealden of North America and
England. Five fingers, with 2, 3, 3, 3, 2 phalanges and four
toes, with 2, 3, 4, 5 phalanges, but the hallux is much shortened
and does not touch the hard ground ; astragalus and calcaneum
separate.

Laosaurics of Colorado is a smaller form, intermediate in
structure between the former genus and Hyjysilophodon foxi from
the Wealden of the Isle of Wight. A small creature, less than
5 feet in length. Four fingers, with 2, 3, 4, 2 phalanges;
fifth metacarpal vestigial. Four toes with 2, 3, 4, 5 phalanges
and long claws. Astragalus and calcaneum separate. Post-pubis
very slender. Each premaxillary with five pointed alveolar
teeth, leaving a wide median diastema ; maxillaries with eleven,
dentaries with ten laterally compressed blade-like teeth.

Iguanodon from the Wealden of England, Belgium, and
Germany. Apparently two species, /. mantelli, about IG feet,
/. hernissarfnisis nearly 30 feet long. The preraaxiUa is
quite toothless ; the teeth of the maxillae and mandibles stand
in close series, implanted in alveolae ; they are spatulate, laterally
compressed, with finely serrated edges, and slightly curved, the
lower outwards, the upper inwards, and bear a general reseml)lance
to those of Iguana, hence the generic name. There is only one
functional set of teeth, and these are much worn down by use,
but in such a way that, owing to tlie different curvature of
the opposed teeth, the worn-down crowns form cutting, and at
the same time crushing, almost triturating surfaces, indicating
that these animals lived upon herbs. The gait of these
creatures was upright, as shown by their spoors ; the long
almost vertical ischia, which form a padded symphysis, only
slightly raised above the ground, suggest that this symphysis was
used as a true sitting support, the animal resting upon it, the
hind-limbs and the long tail. Thd latter, to judge from the lonu:
chevrons and the high neural spinous processes, must have been
furnished with strong muscles. The whole tail was undoubtedly
used as a balance during the upright position. Many of the
tendons of the dorsal spinal muscles on the back and upper half
of the tail are ossified. The post-pubic branches are very slender,
distally much reduced, and, except at the obturator- foramen,
separated from the ischia ; the pre-pubes are very strong and
broad. The femur has a fourth trochanter, a feature which

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428

DINOSAURIA

CHAP.

induced the unfortunate late Paul Albrecht to declare that
Iguanodon was a reptilian Duck ! The tarsal bones are separate.
The metatarsals and toes are reduced to three, with 3, 4, 5 phalanges
respectively, the first being a mere styliform vestige. The
anterior limbs are likewise very powerful, but are much shorter;
the hands are adapted for grasping, possibly for defence and
oflFence, as indicated by the pollex, which, although short, is
transformed into a formidable spur-like weapon, firmly fixed at
a right angle to the other four fingers, the phalanges of which
number 3, 3, 3, 4 ; the second and third fingers were protected

Fig. 101. — 'S^elQion oi Iguanodon heniissartensis. x^.^. (After Marsh.)

by hoof-like nails, the fifth finger is feeble, and stands somewhat
apart. The whole vertebral column consists of more than eighty
vertebrae, of which ten are cervical, eighteen thoracic and lumbar,
while five or six are fused into the sacrum. The cervical verte-
brae are opisthocoelous, and cairy short ribs, except the atlas,
which possesses two separate supra-dorsal pieces, which fill the
gap between it and the occiput.

Many specimens of /. heniissartensis, which is now completely
known, including even the hyoid bones, were discovered in 18 78,
in the Belgian colliery of Bernissart, between Mons and Tournai,
close to the French frontier. The bones were in a fault or crack,
filled with clay of Wealden age, about one thousand feet below
the present sea -level, and there about thirty Iguanodons, all

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ORTHOPODA — ORNITHOPODA 429

apparently adiilt, had become embedded. Five of them are now
mounted in one of the public galleries of the Brussels Museum,
of which these perfect monsters form one of the chief attractions.
Having proved to be such a valuable find, they were claimed by
the Government, on the ground that Iguanodons were not in-
cluded in the license of the Coal Mining Company. The fact
that not only /. hernissartensis, but also a few specimens of
/. mantelli, already known from England, where the large form
likewise occurs, were found in the same place, makes the specific
differences somewhat dou])tful ; they are perhaps sexual.

Claosaurus of the uppermost Cretaceous strata of Wyoming, is
one of the latest of Dinosaurs. It is nearly allied to If/uanodon,
but has only three functional fingers, the fifth being absent, whilst
the poUex is very short.

Hadrosaurus s. Dicloniiis of the same level as the preceding
genus in North America, apparently also in the Middle and
Upper Chalk of England and Belgium, has a most peculiar
spoon-shaped bill, the premaxilla and the predental bone being
spatulate and quite toothless. The teeth in the upper and lower
jaws are numerous and small, and whilst one set of teeth is
being ground down, the several successional series are already
functional. If. mirahilis has in all about 2000 teeth; the total
length of the skeleton is 38 feet, of which nearly 4 feet are
taken up by the skull ; in other respects this genus is allied to
lyuanodon.

OrnithomimuSy of tlie Upper Cretaceous of Colorado, is known
only from its fore- and hind-limbs. The fore-limbs are short,
with three fingers. The hind-limbs are very long and strikingly
bird-like. The metatarsals, of which only the second, third, and
fourth are developed, are much elongated ; the proximal half of
the third is pushed back between the second and fourth, and
imperfectly fused with them, exactly as in young birds. The
astragalus has a long ascending process, and is fused with the
tibia. The fibula is very slender, distally much reduced ; the
calcaneum is represented by a tiny nodule ; the terminal
phalanges end in pointed claws. 0. grand is must have reached
a considerable size, to judge from its middle metatarsal, which
is 60 cm. or 2 feet long. Until more is known of these
extraordinary creatures, nothing definite can be said about their
affinities. They may perhaps belong to the Theropoda.

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430 DINOSAURIA

Order IV. CERAT0P8IA.

Puhic hones simple.forming a symphysis, post-puhic branches hein^
absent. The mandible carries a toothless " pre-dcntal" and the
fused premaxillaries carry a similar, toothless, " rostral " hone.

The teeth of the upper and lower jaws are alveolar, and
have two roots. The fore-limbs are little shorter than the hind-
limbs ; pentadactyle and plantigrade, with broad hoofs. Femur
without a fourth trochanter. Limb-bones solid. The skull is
large, and remarkable for a pair of long frontal bony cores, which
probably carried large, pointed horns ; the parietal bones form a
huge, horizontally broadened out crest, which extends backwards

Fig. 102. — Skeleton of Triceratops proraus. XyV* (After Marsh.)

over the neck. Upon this cranial neck-shield follow small
dermal bony plates. These miraculous creatures flourished during
the Cretaceous epoch in Europe and in North America. Some,
for instance, the American Triceratops Jiahellatus, reached a huge
size, its skull alone measuring more than 5 feet in length,
while that of T, prorsus is, including the neck-shield, about 7
feet long. The total length of this monster, the back of which
stands about 8 feet high, is more than 20 feet. Other genera
seem to have a well-developed dermal armour, e.g, Nodosaurus
of the Middle Cretaceous period of Wyoming.

The Ceratopsia combine characters of the Sauropoda and of
the Stegosaurian Orthopoda ; in their pelvis they agree with the
former, in the development of dermal armour and a predental
bone they agree with the latter, while they differ from either by
the possession of a rostral element.

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CERATOPSIA — CROCODILIA

431

Scb-Class VL— CROCODILIA.

If we had to deal only with the recent Crocodilia the follow-
ing would be an all sufl&cient diagnosis : — Four footed, long-
tailed reptiles, with fixed quadrate bones, with teeth separately
implanted in alveolae and restricted to the upper and lower
jaws.

To define Crocodilia in general and to distinguish them from
various extinct groups we have to resort to additional characters.
The vertebrae are solid ; the ribs of the neck and thorax possess
a distinct capitulum and tuberculum ; there is a series of loose,

Fig. 103.--1, Atlas and axis of CrocodUus. 2, Atlas and axis of Metriorhj/nchus, a
Jurassic Crocodile, see p. 439. 3, Analysis of the first two cervical vertebrae of a
Crocodile. 4, Diagram of the fundamental composition of a Reptilian or other
Amniotic typically gastrocentrous vertebra. Az, Anterior zygapophysis ; AZ>,
basidorsal ; B. V, basiventral ; C^ Cj, first and second centra, formed by the
interventralia ; Cp\ Cp^, articular facets of the capitular portions of the first and
second ribs ; /. l\ interventral ; ATj, N^, first and second neural arch, formed by
the basidorsalia {B.D in 4); Od, odontoid process = first centrum ; 1% posterior
zygapophysis ; Rj, R^ ribs ; Sp, detached spinous process of the first neural arch ;
^1, t^ facets of the tubercular portions of the first and second ribs ; 1, 2, intercentra
^^basiventralia ; * (in 3), second basiventral "complex or intercentrum," continued
upwards as a meniscus or intervertebral pad ; /, //, ///, position of the exit of
the first, second, and third spinal nerves.

compound abdominal ribs; the humerus is devoid of an ent-
epicondylar foramen ; the iliac bones are broadened out and
attached to two sacral vertebrae ; the pubic bones are simple,
not bifurcated, and neither they nor the ischia are ventrally
united. The skull always has a strong, bony, quadrato-jugal
arch. The possession of a longitudinal cloacal opening and of

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432 CROCODILIA chap.

an anterior or ventral single copulatory organ can of course be
asserted of recent forms only.

In spite of these many characters common to all Crocodilia,
it is very difficult to separate the latter from the Dinosauria, the
only absolute difference lying in the ventral pelvic bonea It
is therefore most suggestive that the fore-limbs of the Mesozoic
Crocodilia are so much shorter and weaker than their hind-
limbs, a discrepancy which is not lessened before the Tertiaiy
epoch. The Mesozoic Crocodilia were almost entirely marine ;
the strongly-developed ankle-joint (indicated already by such
early forms as Aetosaurus and Mystnosaurus) must have been
inherited from some terrestrial group with digitigrade tendencies
and shortened hind-limbs. All this points to some Theropodous
Dinosaurian stock of which the Crocodilia may well form an
aquatic, further - developed branch. Loss of the pubic and
ischiadic ventral symphysis is not a serious modification. So
far as faiodern reptiles are concerned only the Chelonia and
Sphenodon are related to the Crocodilia, whilst Monitors and
other lizards resemble them only superficially. We divide them
into three Orders.

Order I. PSEUDOSUCHIA.

The few members of this peculiar group of reptiles are all
restricted to the Keuper or variegated marls, although they
seem to have had a wide distribution, some having been found
in Germany, others in New Mexico. They perhaps form ai?
early side-branch of the generalised Crocodilian stock, which died
out with the Jurassic age.

The skull is distinctly short and pointed. The premaxil-
laries are very small and are dorsally separated from each other
by the large nasals, which also keep the maxillae widely asunder.
Tlie nostrils are latero-terminal, bordered chiefly by the nasals,
below by the premaxillae and part of the maxillae. The orbit
is bordered below by the strong jugals, in front by the prefrontal,
above by a supra-orbital and a small postfrontal, behind by a
postorbital, which, firmly connected with the jugal and squamosal,
shuts oif a supratemporal foramen. There is also a lateral
temporal fossa, and a large hole enclosed by the lacrymal and
the maxillary bones. The teeth are restricted to the anterior

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PARASUCHIA 433

half of the jaws. The neck, back, and tail are covered by two
rows of large and broad, closely-jointed bony plates; smaller
plates protect the sides and the ventral surface. The vertebrae
are still unknown.

Aetomums ferratus of the Upper Keuper near Stuttgart is
the best known. One of the greatest treasures of the Stuttgart
Museum is a slab of sandstone, about 2 square yards in size, upon
which lie huddled together twenty-four individuals of various
sizes, the largest measiuring 86 cm. or 2 feet 10 inches. They
are in a beautiful state of preservation, and many of them are
in the most life-like attitudes, just as if a mass of sand had
fallen upon them and crushed them down, and as if they were
struggling to get out.

Erpetosuch'u^ and OrnithosiLchus of the Elgin sandstone seem
to be allied forms.

Order n. PARASUCHIA.

As the name implies, a collateral branch of the true Crocodilia.
They are, like the Pseudosuchia, restricted to the Keuper forma-
tion. The vertebrae are mostly biconcave, sometimes with
nearly plain, scarcely concave, central joints. The premaxillae
are very long and powerful. The nostrils lie far back, rather
near the orbits, on the top of the snout, within the anterior half
of each nasal and almost above the choanae. The latter are
situated in front of the palatine bones and are divided by a back-
wardly directed process of the vomer, which is plainly visible on
the roof of the mouth. The palatines and pterygoids leave a
wide median space between them. The pterygoids are narrow
and have three processes, the antero-lateral of which joins the
palatines and the maxillary bones (there being no separate
ectopterygoid), the inner joins the basi-occipital, and the postero-
lateral the quadrate.

The orbit is surrounded by the frontal, prefrontal, lacrymal,
postorbital and postfrontal, while the strong jugal is excluded.
The temporal region shows a lateral and a dorsal foramen ;
the latter opens backwards and above the occiput, being bordered
in front by the parietal, laterally by the squamoso-occipital
bridge.

The vertebrae are amphicoelous. The first and second

VOL. VIII 2 F

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434 CROCODILIA CHAP.

vertebrae are devoid of ribs ; the cervicals and first thoracics
carry separate capitular and tubercular processes for the attach-
ment of the ribs, while the ribs of the rest of the trunk are
carried entirely by the long diapophyses, as in the modern
Crocodiles. The dermal armour consists of two rows of broad,
dorsal, and several rows of smaller, lateral, bony plates.

Belodon is by far the best-known genus, with several species
in South Grermany and North Americ^x, some of which reached
a length of 10 feet, without ventral armour. The closely allied
Stagonolepis of the Elgin sandstone in Scotland had dorsal
and ventral armour. Other genera in the Triassic formations of
India and North America.

Order m. EUSUCHIA.

Crocodilia in the stricter sense. The premaxillae are short
and always enclose the nostrils. The choanae lie behind the pala-
tines, in recent forms even within the pterygoids. They occur
from the Liassic or Lower Jurassic period to the present time.

The direct ancestors of the Eusuchia are still unknown.
They cannot have been developed froin the Pseudosuchia, nor do
we know intermediate stages which connect them with the
Parasuchia. The nostrils, situated within the premaxiUaries,
always lie in front of the na&ils, although these sometimes
extend forwards and form a bony internasal septum fusing with
the usual cartilaginous septum. The choanae, instead of opening
immediately behind the vomer, are carried far back, owing to the
formation of a secondary bony palate. In the Jurassic Crocodiles
this roof is formed by the meeting of the palatine bones in the
medio-ventral line, and the choanae open immediately behind.
From Cretaceous times onwards this roofing is continued by the
pterygoids, which likewise form a median suture ; and the united
choanae (which may, or may not, be divided by a thin bony
septum) are pushed towards the posterior end of the pterygoids.
Since the Jurassic times there exists also a tendency to enclose the
Eustachian passages (the remnants of the first gill-clefts) by bone.
In the earlier members they were still wide slits or open grooves
on the ventral side of the basi-occipital bone. Since the Cretaceous
epoch they have been transformed into bony canals and open
through one median hole, situated between the basi-occipital and the

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EUSUCHIA 435

basisphenoid, immediately behind the posterior symphysis of the
dorsal portion of the pterygoids, which latter almost completely
cover the basisphenoid. The vomer is not visible (except in Caiman
niger), being covered by the ventral junction of the palatines
and maxillaries. The broad, lateral wings of the pterygoids
are connected by separate bones, the ectopterygoids = trans-
palatines = transverse bones, with the maxillaries, and in recent
forms also with the jugals. Thus an extensive, very firm bony
palate is produced ; and the large palatal foramina, between tlie
palatines, maxillaries, ectopterygoids and pterygoids, are closed
by the same dense mucous membrane which cover the whole
roof of the mouth.

The opisthotic and epi-otic bones fuse early with the lateral
and with the supra-occipital bones ; only the pro-otic remains
longer as a separate element, perforated anteriorly by a large
hole for the exit of the third branch of the trigeminal nerve.
The basisphenoid is scarcely visible, being covered by the
pterygoids. The presphenoid is large, continued forwards and
upwards into the usually cartilaginous interorbital septum.
Near the anterior and upper margin of the presphenoid is a
large notch on either side for the passage of the optic nerve, the
three eye-muscle nerves and the first branch of the trigeminal
nerve. There are no separate orbito-sphenoids, their place being
taken by membrane or cartilage in continuation with the inter-
orbital septum, but the alisphenoids are large, abutting upwards
against the frontals. Each prefrontal sends down a vertical
process which joins the palatine of its side.

The configuration of the snout varies much. There are two
parallel lines of development since the Jurassic epoch, namely,
long-snouted creatures, of which two still survive as Gavialis and
Tomistoma, and more broad and short-snouted members like tlie
rest of the Crocodiles and Alligators. In opposition to the
Parasuchia the elongation of the snout is effected by the
maxillarie& The length of the nasals varies much, mostly in
conformity with that of the maxillaries. As a rule they reach
the premaxillaries but not always the nasal groove. In Gavialis
they are short, far separated from the premaxillaries by the
maxillaries, which meet in the dorso-median line. The orbit is
bordered by the frontals, which at an early age fuse into an un-
paired piece, and by the prefrontal, lacrymal, jugal, and postfrontal.

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436 CROCODILIA

At a deeper level the orbit is partly divided from the lateral tem-
poral fossa by a strong column which is formed by the meeting of
a downward process of the postfrontal with an inner process of the
jugal, and an ascending process of the ectopterygoid (cf. Fig. 108,
p. 458). This arrangement adds considerably to the strength of
the skull. The lateral temporal fossa is bordered in front by
the column just described ; below by the jugal and the quadrato-
jugal, which is firmly wedged in between the jugal and quadrate ;
behind by the quadrate ; above by the postfrontal, which forms a
strong superficial bridge with the squamosal. This rests upon
and often fuses with the quadrate and an intervening transverse
wing-like extension of the lateral occipital bone. By this
squamoso-postfrontal bridge part of the original temporal fossa
is divided into the lateral one just described, and a dorsal fossa.
The latter is bordered by the postfrontal, squamosal, and united
parietals. This dorsal temporal fossa is consequently not
homologous with that of the Parasuchia, a vestige of which is
however present in many, especially in young skulls of Crocodiles,
in the shape of a narrow passage which extends backwards from
the dorsal fossa, bridged over by the junction of the parietal with
the squamosal, and bordered below by the occipitals.

The size of the upper temporal fossae stands in an inverse
ratio to that of the lateral fossae. In the older Eusuchia the
upper were the larger of the two. The temporo-mandibular
muscle which lifts or shuts the lower jaw arises from the walls
of the upper fossa, passes beneath the jugal arch, and is inserted
into the supra-angular portion of the lower jaw. In the more
recent Crocodiles this muscle is more and more superseded by
the pterygo-mandibular muscle, which, arising chiefly from the
dorsal surface of the much broadened-out pterygoid bone, fills
the widened space beween the latter and the quadrate, and is
inserted into the outer surface of the os angulare of the lower
jaw. This muscle, owing to its general disposition, is capable of
much more powerful development and leverage than the temporo-
maxillary muscle, which latter, being more reduced, allows the
dorsal fossae to be more and more closed up by the surrounding
bones.

Tlie fossae are still comparatively large in the long-snouted
genera Gavinlis and To mi stoma yVfhii^h live entirely upon fish and
scarcely chew their food, whilst these holes almost completely

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X SKULL 437

disappear in some of the Alligators, namely in the broad- and
short-snouted members, which, having a varied diet, taken from
every available group of the animal kingdom, chew their
prey.

The quadrate extends obliquely backwards, and is immovably
wedged in and partly fused with the quadrate -jugal, the
squamosal, and the lateral occipital wings. Between the latter
and the quadrate remains a slit-like canal, well visible from
behind, through which passes the continuation into the mandible
of the columellar or ossicular chain of the auditory apparatus.
Intricate passages, used as additional enlargements of the space of
the middle ear, pervade the proximal portions of the quadrate
and the roof of the cranium beneath the parietal bridges
mentioned above, the two sides communicating with each other.
The supra-occipital bone is visible from behind ; its top is covered
and partly fused with a continuation of the parietals, which
are, like the frontals, fused into an unpaired mass. The
occipital condyle is formed entirely by the basi-occipital bone, so
far as the articulating facet is concerned, but it is supported on
either side by a lamella from the lateral occipitals.

The two halves of the lower jaw form a symphysis of very
variable length. Each half is composed of six bones. (1) The
articulare, perforated in its upper, posterior, inner comer by a canal
for the reception of the siphonium, a narrow tube of connective
tissue, which connects the cavities of the middle ear with the large
empty space enclosed within the lower jaw ; (2) the angulare ;
(3) the dentary, which alone carries the teeth ; (4) the splenial,
a long splint-like bone on the surface of the inner or median side
of the jaw, of variable length ; (5) the operculare, the counter-
part of the splenial on the outer side; (6) the supra-angulare,
which forms the dorsal border of the lower jaw between the
dentary and the angulare.

The teeth, which are more or less conical or compressed
laterally, are deeply implanted in separate sockets. They are
often shed throughout life, the successors lying on tlie median
side, and with their caps partly fitting into the wide, open roots
of the teeth to be expelled. The number of teeth in the pre-
maxilla is universally five on either side in recent forms, but in a
few species, e.g. Crocodilus niloticus and C. porosns, the second pair
is lost with maturity and is not replaced. In the broad-snouted

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43 8 CROCODILIA

kinds, especially in the Alligators, most of the upper teeth overlap
laterally those of the lower jaw. In most species of Crocodiius
the overlapping is less marked and the teeth partly interlock, but
the fourth mandibular tooth, generally the strongest and longest,
is received into a lateral notch at the junction of the pre-
maxillary and maxillary. Frequently those of the longer lower
teeth which fit into pits of the upper jaw, gradually transform the
pits into holes by continued pressure upon the bone, and in old
specimens the tip of the lower tooth may even perforate and
stand out above the skin of the snout.

The vertebrae are solid, but remnants of the notochord per-
sist for a long time in the middle of the centra. These are still
amphicoelous in the Jurassic Eusuchia, and there were probably
considerable intervertebral portions of the notochord. From the
Lower Chalk onwards the vertebrae are procoelous, with the
exception of the first caudal vertebra, which has a knob at
either end, so that naturally the posterior of the two sacral verte-
brae is opisthocoelous. This peculiar formation of the first
caudal is probably correlated with the flexibility of the tail

Cartilaginous intercentral rings, pads or menisci, occur
regularly throughout the vertebral column, unless they are
abolished by fusion of adjoining vertebrae. It is most instructive
to follow the attachment of the ribs in one and the same
individual. The position of the capitulum, vertically below the
tuberculum in the neck, changes in the thorax into one in wliich
the capitulum lies anterior to the tuberculum and in the same
horizontal plane with it. Moreover, whilst on the cervical
vertebrae the capitulum is carried by the centrum (enclosing
with the tuberculum a typical transverse canal for the vertebral
artery, etc.), further back it moves its point of attachment up-
wards, lying right upon the neuro-central suture on the tenth and
eleventh vertebrae. From the twelfth vertebra backwards both
capitulum and tuberculum are carried by the transverse process
or diapophysis of the neural arch. The ribs of the five or six
lumbar vertebrae are merely vestigial or absent. The ribs of the
two sacral vertebrae are very stout, fusing in the adult with both
centrum and neural arch. Some of the anterior caudal vertebrae
also carry ribs, attached across the neuro-central suture ; long
before maturity they fuse with their vertebrae, and then look like
transverse processes. Most of the caudal vertebrae carry also a

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X VERTEBRAL COLUMN 439

pair of chevron-bones, and these are continuous with the inter-
central rings of cartilage.

The atlas and the epistropheus or axis are of supreme interest.
Crocodiles are, in fact, the only animals in which these two
vertebrae retain all their constituent hard parts in an almost un-
disturbed primitive condition (Fig. 103, 1-4). The basal piece of
the atlas-ring, the first basi ventral or intercentrum, carries a pair
of long ribs attached by their capitular portions. A small knob
near the dorsal edge of the rib occurs in many specimens, and
is the last remnant of the tubercular portion. The latter was
still complete in Jurassic Crocodiles, for instance in Metrio-
rhynchus (Fig. 103,2, t{). The first centrum joins that of the second
vertebra as its so-called odontoid process, not directly, however,
but by the intercalation of the complete second basiventral, repre-
sented by a cartilaginous disc, and by a large unpaired pyramidal
piece (Fig. 103, 3^^). This, serially homologous with the ventral
half of the atlas- ring, is the second basiventral intercentrum,
wedged in from below between the odontoid process and the
second centrum, with which it soon fuses. Moreover, it cairies
the capitulum of the second rib (2, Cp^), the tuberculum of which
is articulated with a facet of the second neural arch in Jurassic
Eusuchia (t^). In recent Crocodiles this tubercular portion is much
reduced, and, curiously enough, is attached to a knob which
belongs to the odontoid piece or first centrum. This shifting
explains the apparently anomalous condition that " the atlas of
the Crocodiles carries two pairs of ribs, the second vertebra none."
To complete the account of the atlas we have to mention the
separate unpaired piece which lies upon the two neural arches.
It is the detached neural spine, and not the remnant of a
" pro-atlas."

The first and second ribs (E^ and i?.,)* *^ ^^^st in the recent
forms, are very long and are quite movable. Those of the next
five cervical vertebrae are firmly fixed, short, and adze-shaped.
The eighth and ninth are again long, and make the transition to
the thoracic ribs, which are mostly eight in number, some with
uncinate processes. Then follow several shorter or floating ribs,
mostly two or three pairs. The next following three presacral
vertebrae carry no ribs. The two sacral and the caudal ribs
have already been mentioned.

Ab a rule the vertebral column of recent Crocodiles, Alligators,

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440 CROCODILTA

and Gavials is composed of twenty -six precaudal vertebrae
(namely, nine cervical, fifteen thoracic and lumbar, two sacral),
and about thirty-four to forty or more caudal vertebrae. Indi-
vidual variations, including lop-sided attachment of the iliac
bones, are by no means uncommon.

The sternum remains cartilaginous. It consists of an anterior
rhomboid portion, which carries the coracoids and two pairs of
ribs, and a posterior longer and narrower portion formed by the
median fusion of the next following five or six ribs. Posteriorly
the sternum bifurcates, each half carrying two or three ribs, of
which the last sometimes loses its proximal connexion, and thus
appears as a xiphisternal process. Ventrally, upon the anterior
part of the sternum lies the longitudinal, originally paired,
episternum. The shoulder -girdle consists of the coracoids and
the scapulae, which fuse with each other into one bony piece
on each side. A pre-coracoid is indicated in fossil forms by a
notch in tlie coracoid.

The space between the posterior end of the sternum and the
pubic bones is occupied by the so-called abdominal sternum, com-
posed of seven pairs of ossifications, resting upon the ventral side
of the rectus abdominis muscle. Each pair consists of two closely
apposed pieces, while the right and left remain separate in the
median line. The last pair is much stronger than the rest, is more
deeply imbedded in the rectus muscle, and is loosely connected
with the anterior margin of the two " pubic " bones.

The limbs are built upon the typical terrestrial pentadactyle
type, but were in the Jurassic species undoubtedly more adapted
to swimming locomotion. The fore-limbs were conspicuously
shorter and smaller than the hind-limbs, and it is only since
Tertiary times that the difference has decreased to a great
extent. Ulna and radius remain separate. The proximal
row of carpal bones consists now of the ulnare and radiale,
both strong and distinctly elongated. On the outer side, be-
tween ulna and ulnare, lies a pisiform bone. Upon the
radiale follows a compound bone, often imperfectly ossified
towards the median side, and consisting of the first distal
carpal, the centrale, and the intermediimi. The third, fourth,
and fifth carpals are fused into one mass. The second distal
carpal remains separate. All five fingers are present and well
developed. The number of phalanges of the pollex is two, of

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X LIMBS 44 1

the others three, four, four and three respectively. During the
embryonic development the number of phalanges of the fourth
and fifth finger increases temporarily, to as many as seven on the
fourth, to five or six on the fifth finger. Before the young
animal is hatched the numbers are reduced again, chiefly by
fusion of adjoining phalanges. This hyperphalangeal condition,
typical of Plesiosauri, Ichthyosauri, Cetacea, and several other
absolutely aquatic animals, naturally suggests the descent of the
present Crocodiles from more essentially aquatic ancestors, but
hitherto no trace of supernumerary phalanges has been found in
any Jurassic Eusuchia, nor in the Parasuchia and Pseudosuchia.

The composition of the pelvis is difficult to understand. It
consists in the adult stage of three separate bones, of which two
only partake in the formation of the acetabulum. The broad
ilium sends out two processes ; the posterior and stronger arti-
culates with the ischium, which sends out a short and stout
process towards the anterior process of the ilium, enclosing a
foramen. This process contains a separate centre of ossification,
possibly homologous with the true pubis, while each club-shaped
bone, loosely attached to it and directed forwards, generally called
the pubis of the Crocodiles, would then be equivalent to an
epipubis. Neither the *' pubes '* nor the ischia form a ventral
median symphysis.

The femur is devoid of a prominent inner trochanter. Tibia
and fibula are of almost equal strength. The tarsal elements are, '
in the adult, reduced by fusion to five bones. The fibulare is
transformed into a typically projecting, heel-shaped calcaneum,
while the intermedium is fused with the tibiale into a broad
astragalus. The first, second, and third distal tarsalia are much
reduced towards the inner side, and form one wedge-shaped, partly
cartilaginous mass. The fourth tarsale lies between the fibulare
and the fourth metatarsal, while the fifth tarsale is hook-shaped
and loosely attached to the outer side of the fourth. It has lost
its metatarsal and the rest of the fifth finger. Embryos are
hyperphalangeal, the fourth toe developing six phalanges, and
there are traces of the fifth toe. The numbers are ultimately
reduced to 2, 3, 4, 4, 0 on the five toes. The fourth toe remains
without a claw.

SkilL — The epidermal horny layer is not shed periodically nor
in pieces ; the wear and tear is made good imperceptibly. The

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442 CROCODILIA ' chap.

scales, which cover the whole body, have a hard, horny, water-
proof covering, but between them the skin is soft. Each scale of
the sides, belly, and tail, and especially those of the lower jaw,
shows a little dot or pit. At this spot the epidermis ia not
cornified or thickened, and a nerve with sensory corpuscles ends
beneath the bottom of the pit. Sometimes these pits are filled
with debris of cells, and on the lower jaw, especially on the chin,
these organs, instead of forming pits, are raised into little wart-
like prominences.

The scutes or dermal portions of the scales consist of
thickened, cutaneous connective tissue, and are more or less
extensively ossified, thus forming a proper dermal armour. In
most recent Crocodilia the armour is restricted to the back,
with occasional osseoas plates on the throat, as in Osteolaemus ;
regular although thin ossifications in the ventral scutes occur in
the Caimans only. The Crocodile and Alligator skins of com-
merce consist entirely of the tanned cutis, minus the epidermis
and the horny coverings of the scutes. In some fossil genera
the ventral armour was extensively developed, especially in
Teleosaurus, in some genera to the exclusion of dorsal ossifications.
The armour of the recent forms consists, so far as the large scutes
are concerned, of a considerable number of scutes, which are
arranged in transverse rows, each row corresponding with one
skeletal segment of the trunk proper. Mostly there is a detached
cluster of scutes on tlie back of the neck. On the trunk some of
the scutes are larger and more crested than others, and form in
their totality a variable number of longitudinal rows. The
median pair is generally the most conspicuous on the back.
Some of the more lateral rows of keeled scutes converge more and
more towards the tail, the inner rows drop out imperceptibly, and
two lateral rows combine on the middle of the tail into an un-
paired series of vertical blades. These are no longer bony, but
show more strongly developed horny sheaths; they are very
flexible, and transform the tail into an effective propelling organ.

Most of the larger scutes and the upper surface of the bones
of the skull have a peculiar gnawed-out, almost honeycombed
appearance, as is usual wherever most of the cutis itself is trans-
formed into bone or co-ossifies with underlying bone, while the
uppermost layers and the horny layer of the epidermis are much
reduced and thinned out.

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SKIN-GLANDS — TONGUE 443

All the recent Crocodilia possess two pairs of skin-glands, both
secreting musk. One pair is situated on the throat, on the inner
side of the right and left half of the lower jaw. The opening
of the gland, visible from below (see the figure of Crocodihis
niloticus, p. 461), is slit-like, and leads into a pocket, which in
large specimens is of the size of a walnut ; the bag is filled with
a smeary pale brownish substance, a concentrated essence of musk,
much prized by natives. The secretion is most active during
the rutting time, when the glands are partly everted. My young
Crocodiles and Alligators often turned them inside out, like the
finger of a glove, when they were taken up and held by force. The
other pair lies within the lips of the cloacal slit, and is not visible
from the outside. The use of these strongly scented organs,
which are possessed by both sexes, is obviously hedonic. The
sexes are probably able to follow and find each other, thanks to
the streak of scented water left behind each individual.

The tongne is flat and thick, attached by its whole under-
surface, so that it can be elevated but not protruded. It fills the
whole space between the two halves of the lower jaw behind
their symphysis. Tlie dorsal surface shows numerous irregular
polygonal fields, in the middle of most of which opens the duct of
a large mucous gland. Tactile and gustatory corpuscles are
scattered over the surface in the shape of tiny wartlike elevations.
The hinder margin of the tongue is raised into a transverse fold,
which, by meeting a similar fold from the palate, the velum
palatinum, can shut off the mouth completely from the dee'p and
wide cavity of the throat, which leads of course into the gullet.
Dorsally the choanae open into this cavity ; and since the narial
passages are transformed into long tubes, completely surrounded
by bone. Crocodiles can lie submerged in the water, with only tlie
nostrils exposed and with the mouth open, and breathe without
water entering the windpipe. The opening of the latter, the
glottis, is a longitudinal slit, protected by the laryngeal cartilages,
opened and closed by muscles. There is also a pair of membranous
folds within the glottis, which serve as vocal cords. Ventrally
below the larynx lies the cartilaginous, broad, shield-shaped hyoid;
on the sides are attached the short hyoid horns. The trachea is
long, consists of about sixty or more complete cartilaginous rings,
and divides into two short bronchi, likewise protected by complete
rings. The trachea is depressed ; its transverse diameter decreases

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444 CROCODILIA

from the glottis backwards. The IwigB have attained a high
degree of efficiency. Each lung is an oval sac, and is transformed
into a complicated system of tubes, at the end of which are the
countless honeycomb-like respiratory cells, the whole lung being
spongy. The main bronchus is continued straight down to the
posterior end of the lung, and sends off during its course regular
secondary bronchi, and these send off tertiary bronchL The
whole arrangement is very regular, the tubes coming off like rows
of organ-pipes. Each lung hangs freely in the thoracic cavity.
Besides its ventral attachment by its arteries, veins, and the
bronchus, it is connected by loose tissue with the liver and the
pericardial septum. Each half of the thoracic cavity is partitioned
off from the abdominal cavity by a strong transverse mesenteric
lamella. The partition between the lungs and the stomach is at
first simple, it then divides, to enclose the liver ; the anterior
partition passing between liver and lung to the inner surface of
the sternum ; the posterior lamella between the liver and the
stomach. Both meet on the ventral surface of the liver, and are
continued into or attached to the peculiar " diaphragmatic "
muscle. This is covered by the internal rectus muscle of the
abdomen, arising from the last pair of abdominal ribs near the
pubic bones ; it is innervated by a branch of the last precrural
nerve, and extends as a broad but thin muscular sheath (always
within and unconnected with the abdominal wall) to the ventral
posterior vein of the liver ; thence it is continued as an
aponeurosis, together with the peritoneal lamella mentioned
above, to the inner surface of the sternum. Contraction of this
singular muscle indirectly widens the pulmonary cavity, and
thereby directly aids inspiration. It acts consequently like the
diaphragm or midriff of Mammals, although it is morphologically
an entirely different muscle.

The stomach is smaller than one might expect from the fact
that large Crocodiles can eat up nearly a whole man ; but a great
deal of their prey is stowed away preliminarily in the wide gullet
until the rapid, powerful digestion, which dissolves every bone,
makes room in the stomach. This consists of a wide, some-
what globular gizzard, rather muscular, with a pair of tendinous
centres like those of birds, and a much smaller pyloric, globular,
more glandular compartment. It leads into the duodenum,
which is coiled up into a double loop, and receives at its end the

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VISCERA 445

hepatic and pancreatic ducts. The small intestine is narrow,
and is stowed away in a few irregular coils ; the rectum is wide ;
a caecum is absent.

The cloaca is peculiar. The coprodaeum and urodaeum, cf.
p. 498, are confluent, and form a wide, oval bag, closed in front
and behind by strong sphincters, and it acts normally as a urinary
receptacle. In the dorsal wall open the two ureters; a little
towards the sides, and ventrally, open the two oviducts, on the
right and left, near the base of the clitoris. Then follows a
transverse, soft, muscular fold, which shuts off this cavity from
the proctodaeum or outermost chamber. In the latter is
stowed away the rather large copulatory organ. It arises out of
the medio-ventral wall of the cloaca, and has a deep, longi-
tudinal groove on its morphologically dorsal side for the con-
duction of the sperma, the vasa deferentia opening near its
basal end. On either side of the root of this organ, in both
sexes alike, opens a peritoneal canal, wide enough in large
specimens to pass a goose-quill. The outer opening of the
cloaca forms a longitudinal slit; within it, dorso-laterally, are
the openings of the two anal musk-glands.

The kidneys are much lobed. The testes are long and oval ;
the ovaries are much elongated and flat ; and the eggs con-
tained therein in great numbers are extremely small, except
those which ripen during the time of propagation.

The vascular system has attained the highest state of
development of all reptiles. The heart is practically quadri-
locular, the partition between the right and left ventricle being
complete ; but there is still a small communication, tlie foramen
Panizzae, which lies in the middle of the wall common to both
aortae, where they leave their respective ventricles. Tlie left
aortic arch conveys all the arterialised blood out of the left
ventricle, and supplies head, neck, trunk, and tail. The right
aortic arch, coming from the right ventricle, sui)plies venous
blood, mixed with what little arterial blood it receives througli
the foramen Panizzae, to most of the viscera. On a level with
the stomach both descending aortic arches are still connected
with each other ; the left aorta supplies most of the gut ; the
right, the trunk and the kidneys.

The outer ear lies in a recess, dorsally overhung by the
lateral edge of the bony squamoso-postfrontal bridge ; and tliis

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446

CROCODILIA

carries a flap of skin, provided with muscles, to close the ear
tightly. The tympanic membrane is visible at the bottom of
the recess ; shining through it is part of that cartilage which is
homologous with the malleus of the auditory ossicular chain ;
the outward extension of the latter on its way to the mandible,
behind the joint, passes as a partly cartilaginous string through
the slit-like hole which is visible at the back of the skull,
between the quadrate and the latero-occipital wing.

The eyes have, besides the lower and upper lid, a third, the
nictitating membrane, which can be drawn over the front of the
eyeball. In the upper lid lies a cup -shaped bony plate of
variable size. The pupil contracts into a vertical slit. The iris
is greenish.

Fig. 104. — Map to illustrate the present distribution of Crocodilia.

The recent geographical distribution of the various kinds of
Crocodilia loses its mystery when we recollect that during the
Tertiary period Alligators, Crocodiles, and long-snouted Gavials
existed in Europe. The solitary species of Alligator in China is
the last living reminder of their former Periarctic distribution.
The group, taken as a whole, is otherwise now intertropical.
Crocodiles alone inhabiting the Palaeo-tropical region, together
with long -snouted forms in the Oriental sub -region, while
Alligators and Caimans, with a few Crocodiles, live in America,

They are all rapacious, doing much damage by their pre-
datory habits, and are fierce and sulky in temper. But the
danger to man differs much in different countries. While Croco-
diles are dreaded in some localities, they are in others considered
almost harmless, and men swim through the haunted waters
without hesitation. It seems as if certain old and wily indivi-
duals turn into man-eaters, just like tigers and lions.

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X DISTRIBUTION — HABITS 447

Their home is the water, in which they pass the night, their
time of hunting. The prey is either patiently watched or
stalked, and nothing falls amiss. Water-birds are seized by the
beast, which rises imperceptibly from below. Some species are
said to make use of their powerful tails for hitting the victim
and even jerking it into the mouth. The strength of their jaws
is enormous, and they do not let go what they have seized, unless,
in the case of a man, he has the presence of mind and the
opportunity to dig his fingers into the monster's eyes whilst
being dragged down.

In the morning they crawl on to sandbanks, or on to logs
of wood, which they closely resemble, in order to bask, mostly
in such a position that on the slightest alarm they can plunge
into the water. For this reason they frequently make a half
circle before they settle down to rest, with the heads turned
towards the river. There they bask all day long, apparently
fast asleep, often with gaping mouths. But their sense of
hearing and of sight is sharp, and they learn from experience,
old individuals being by far the most wary. Commercially the
skins are now of considerable value. The flesh is white, and is
tolerable eating but for the combination of fishy and musky
odour, which, although faint, is not to everybody's liking.

All the species have a voice, a kind of loud, short bark or
croak, heard at night and when angered. The female lays
several dozen or even three score white, oval, hard-shelled eggs in
the sand, well out of the reach of moisture ; and some species
construct an elaborate kind of nest. The mother watches it,
takes care of and fights for her offspring, numbers of which
fall an easy prey to large storks, fishes, and to the stronger
members of their own kind.

In the cooler countries they hibernate in the ground ; and in
hot countries, which are subject to drought, some kinds aestivate
in the hardened mud ; or they migrate. When during a pro-
longed drought on the island of Maraj6, at the mouth of the
Amazon, the swamps and lakes were dried up, the Alligators
migrated towards the nearest rivers, and many perished in the
attempt. On one farm were found 8500 dead, and at the end
of Lake Arary more than 4000. Such occurrences in bygone
times may perhaps explain the masses of bones found here and
there in a fossil state.

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448 CROCODILIA

The age to which Crocodiles can live is quite beyond calcula-
tion- They are capable of propagation long before they are
anything like half-grown, maybe at an age of little more than
ten years ; then they continue to grow perhaps for more than
one hundred years, until they die.

It is customary to divide the Eusuchia, most of which are
extinct, into a longirostral and a brevirostral section. In the former
the snout is much elongated and narrow, and the nasal bones,
although they are sometimes very long, do not reach the nasal
groove. The mandibular symphysis is very long, and is formed
not only by the dentary but also by the splenial bones. In the
brevirostral section the snout is shorter, sometimes broad and
rounded off, and the nasal bones are supposed to reach the nasal
groove, or at least to approach it very nearly; the mandibular sym-
physis is formed by the dentaries only. But these distinctions
are quite arbitrary, and there exist all kinds of intermediate
forms. For instance, in Goniopholis and I}ijplocyiwdon, which
are both undoubtedly near allies of the recent Crocodiles and
Alligators, the nasal bones are considerably removed from the
nasal groove ; and in Crocodilus cataphractus they are separated
even from the premaxilla by the medio -dorsal suture of the
maxillaries. Again, in Gonioj>liolis the mandibular symphysis
is so long that it comprises part of the splenial bonea Both
typically long- and short-snouted forms occur already in the
Upper Oolite, but in the Lower Jurassic age only long-snouted
kinds seem to have existed. The latter cannot easily be con-
nected with Belodon, one of the Parasuchia, on account of the
position of the nostrils; the mere shortening of the long
premaxillaries of Belodon would not transfer its distinctly
paired nostrils to the anterior end of the premaxilla. To
account for the position of the nasal groove in the Eusuchia,
we have to go back to a primitive condition, such, as that of
the Pseudosuchian Aetosaurus, and this consideration shows that
the Parasuchia and Eusuchia are collateral branchea

The Eusuchia have been split into many families. Zittel,
for instance, divides them into ten, some of them on insufficient
grounds, since there are too many intermediate forms ; and more,
soiuetiines quite unexpected, modifications are still being found
Several of the accepted ftimilies represent collateral or con-
vergent lines of development. There is the same tendency to

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VOL. VIII

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450 CROCODILIA

transfer the choanae further back, owing to the formation of
a solid secondary roofing in of the moijth, to transform the
amphicoelous into procoeloiis vertebrae, to reduce the supra-
temporal foramina, and to obtain a better development of the dorsal
armour, whilst that on the ventral side is gradually reduced.
Lastly, there is a tendency towards a shortening and broadening
of the snout, a condition which has reached its culmination in
the Alligators, while the Gavials are survivals of another branch.
The notches in the premaxilla, for the reception of some of the
lower teeth, have also been acquired independently. Although
the recent Crocodilia cannot now, as has been pointed out by
Boulenger, be separated into different families, no valid diagnoses
being possible owing to the existence of Tomistoma, their phylo-
geny shows them to belong to at least two heterogeneous groups.

Key to the Genera of recent Crocodilia.

I. Snout very long and slender. The mandibular symphysis extendB at

least to the fifteenth tooth, and is partly formed by the splenial bones.
a. Nasal bones very small, and widely separated from the premaxil-
laries ..... Gavialu gangeticu^, -p, 451.

h. Nasal bones long, in contact with the premaxillaries.

Tomistoma schlegdiy p. 453.

II. Snout not slender, but triangular or rounded off. The mandibular sym-

physis does not reach beyond the eighth tooth, and does not reach the
splenial bones,
a. Fourth mandibular tooth fitting into a notch in the upper jaw.

1. Without a bony nasal septum . . CroeadiltiSy p. 454.

2. Nasal bones dividing the nasal groove. Osteolaenuis^ p. 466.
h. Fourth mandibular tooth fitting into a pit in the upper jaw.

1. AVithout a bony nasal septum . . Caiman^ p. 471.

2. Nasal bones dividing the nasal groove . Alligator, p. 466.

Fam. 1. Teleosauridae, in the Lias * and Oolite of Europe ;
marine. — Snout very long and slender. Nasals widely separated
from the premaxillae by the maxillaries. Choanae at the pos-
terior end of the palatinea In front of the eye a small
sub-lacrymal foramen, Supratemporal foramina large. Vertebrae
amphicoelous. Anterior limbs scarcely half as long as the
posterior pair. The dermal armour consists of two rows of broad
scutes on the back, while the belly is protected by a shield of
numerous bony scutes, which are connected with each other by
sutures. Teeth numerous and rather slender. General appear-
ance like that of Gavials.

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X TELEOSAURIDAE GAVIALIDAE 45 1

Teleomvrvs of the Middle and Upper Oolite in England and
France. Snout very slender. Nasals narrow and short. The
under side is protected by a beautifully finished armour, consisting
of a square breast-shield of four rows of bony scutes, and a larger,
long, oval shield on the belly, with about six longitudinal and
seventeen transverse rows of scutes.

Mystriosaurvs, of the Upper Lias in France and Germany,
reached a length of 15 feet, and is characterised by an additional
series of keeled but smaller caudal plates running parallel with
the middle pairs, which are neatly sutured together.

Fam. 2. Metriorhynchidae, in the Upper Oolite of Europe ;
marine. — Nasals broad posteriorly, sometimes extending with a
pointed wedge very near the premaxillae. Without sub-lacrymal
foramina. Eyes with a ring of ossifications in the sclerotic.
Dermal armour unknown. Vertebrae and choanae like those
of the previous family. Metriorhynchvs and Geosaums.

TejSL. 3. Macrorhynchidae, in the freshwater deposits of the
Purbeck, Wealden, and Greensand of Europe. Snout long and
slender. The nasals are narrow, and so elongated that they
meet a similar long extension of the premaxillaries. Choanae
between the palatines and pterygoids. Vertebrae amphicoelous.
Dermal armour consisting of two imbricating dorsal and eight
ventral rows, e.g. Pholidosaurus of the English Wealden.

Fam. 4. Gavialidae. — Snout long and slender. The choanae
are situated entirely within the pterygoids. Vertebrae pro-
coelous. Members of this family make their first appearance
in the littoral marine deposits of the Upper Chalk of Europe and
North America ; others are common in tertiary, marine, and
freshwater deposits, whilst only two genera and species occur
now in the Oriental sub-region.

Thoracosaurus in the Upper Chalk of New Jersey and France
and Belgium is intermediate between Gavialis and I'omistoma,
The prefrontal bones are very small, while the lacrymals are
very long and surround the nasals posteriorly. The nasals them-
selves are slender, and reach the posterior likewise long and
narrow prolongations of the premaxillaries.

Gavialis, — The snout is extremely long and slender. The
mandibular symphysis is so long that it comprises a great
portion of the splenial bones, and extends backwards almost to
the level of the last teeth and to the palatal foramina. The nasal

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452

CROCODILIA

bones are very short, and are separated from the premaxillaries
by the long suture of the raaxillaries. About twenty-eight upper
and twenty-five lower tefeth on each side.

G. gangeticiiSy the only recent species, is essentially Indian,
inhabiting chiefly the basins of the Ganges, Brahmaputra,
and Indus ; it occurs also in the Mahanadi of Orissa and in

Arakan, but does not live in the
Irrawaddy, nor in the Narbada, Kistna,
and farther south. In spite of its
great size, which reaches 20 feet or
even more, it is harmless, and lives
entirely upon fish ; hence its Hindus-
tani name, gharial, meaning fish-eater,
of which the generic name is a corrup-
tion.

The nuchal and dorsal scutes form
a continuous shield, but there are two
small postoccipital scutes. General
colour, dark olive -brown above; the
young are paler, with dark markings.
The male is remarkable for several
peculiarities. The nose is very much
swollen, and can be inflated like a bag
when the nostrils are closed. In con-
nexion herewith, probably produced by
the recoil of the air in the long narial
passages towards the choanae or pos-
terior nares, there is a pair of hollow
globular swellings, in large specimens
of the size of a goose's egg. The
shell of these globes is formed by the
dorsal wings of the palatine bones
above the floor of the choanae, and
they extend forwards to the right and left of the ethmoid
almost to the vertical downward process of the prefrontals.

Although the Gharial is common enough, we know next to
nothing about its habits, and in zoological gardens it is rather
rare. A. Anderson ^ has, however, made the following observ^a-
tions. Forty eggs were dug out of the sand, where they were

1 P.Z.S. 1876, p. 2.

Fig. 106.— Skull of Oavialis gan
(/elicits (the Gharial). x |
F, frontal ; /, jugal ; Z, lac
rymal ; Mamlj mandible
3/x, maxillary ; Xa, nasal
Pary parietal ; Pinx^ premaxil
lary : Pr/, prefrontal ; PtJ\
postfrontal ; Qj\ quadrato-
jugal ; Sqj squamosal.

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GAVIALIDAE GONIOPHOLIDAE 453

lying in two tiers, twenty below and twenty above, with a foot
of sand between. The young ran with amazing rapidity the
moment they were hatched. Some of them actually bit his
fingers before he had time to remove the shell from their bodies !
The length of these new-born creatures was 15 to 16 inches,
9 of which belonged to the long and slender tail.

Several fossil species have been described from the Pliocene
deposits of the Sivalik Hills of India ; and in the same district
occurred the closely allied Bhamphostuihus crassidens, which
reached the gigantic length of about 50 feet !

Tomistoma. — The general configuration of the skull and snout
is that of Gavialis, but the nasal bones are long and reach the
premaxillaries, although not the nasal groove, thereby separating
the maxillaries. The first and fourth mandibular teeth fit into
notches of the upper jaw, while most of the others fit into pits
between the teeth of the upper jaw. About twenty upper and
eighteen lower teeth on each side.

T. schlegeii, the only species, reaches a length of 1 5 feet ;
it inhabits the rivers and swamps of Borneo, Malacca, and
Sumatra. Fossil specimens of Tomistoma have been found in
the Miocene of Malta and Sardinia. Gauicdosuchus of the
Miocene of Hungary is closely allied.

Fast 6. Atoposanridae. — The few members of this family,
Atoposaurus, Alligatorium, and Alligatm'eJlus, lived in the Upper
Oolitic period of France, and were small, about one foot in length.
The vertebrae are amphicoelous. The nasal groove is divided
by a prolongation of the nasal bones. The head is short, and in
its general shape rather like that of a lizard.

Fam. 6, Ooniopholidae, in the Purbeck and Wealden of
Europe and the corresponding level of North America. The
vertebrae are amphicoelous. The choanae are still elongated
but are situated between the palatines and pterygoids. The
premaxillaries are rather large, and each sends a broad triangular
process between the nasal and maxillary. The nasals are broad
and are well separated from the nasal groove. The splenials
help to form the mandibular symphysis.

Goniopholis, — The general configuration of the skull is rather
like that of Crocodilus vulgaris. There is a pair of deep notches
in the upper jaw for the reception of the lower canine teeth. G,
simus and G, crassidens in England and continental Europe, and

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4S4 CROCODILIA CHAP.

others in Colorado, were large-sized Crocodiles, some with a skull
2 feet in length. The dermal armour consisted of a pair of
dorsal rows, a thoracic and an abdominal shield, composed as in
the Teleosauridae of six to eight longitudinal sutured rows.

Fam. 7. OrocodiUdae. — Beginning in the Upper Cretaceous
period of Europe and North America, many forms of Crocodiles,
Alligators, and Caimans existed in the Tertiary period in America,
Europe, and India ; persisting in Europe until the Plistocene. The
vertebrae are procoelous. The choanae are completely surrounded
by the pterygoids. The nasals reach the nasal groove, except
in Crocodilus catajphractus. The orbits are larger than the small
supratemporal fossae, and always continuous superficially with
the lateral temporal fossae, the postfronto-jugal bridge not reach-
ing the surface. The dorsal armour consists of more than one
pair of longitudinal rows, while the ventral armour is much
reduced in thickness or absent.

Dij)locynodon. — Common in the Oligocene and Miocene of
Europe, e.g. D, hastingsiae. The skull resembles that of the
Alligators, but has a pair of lateral notches in the pre-
maxilla for the reception of the third, and sometimes also of the
fourth mandibular tooth. The ventral armour is still ratlier
strong.

Crocodilus. — The fourth mandibular tooth fits, as a rule, into
a notch in the upper jaw. The other teeth are more or less
interlocked with those of the other jaw. The fifth upper tooth
is the largest. The nasal bones form the posterior border of the
nasal groove, but do not extend into it as a septum. The bony
scutes of the dorsal shield are keeled, and stand closely together,
being rarely united by suture ; and they form from four to six
principal rows.

Crocodiles have occurred since the Upper Chalk in Europe ;
many species existed in the Tertiary epoch in Europe and North
America, decreasing in numbers in the Pliocene and disappearing
with the beginning of the Plistocene. About ten recent species
are known, and these have now a somewhat scattered distribu-
tion ; namely, three species in Africa, one of them extending into
Syria ; three in tropical America and the West Indian Islands ;
the rest in the Malay, Indian, and North Australian countries.

C. palustris. — The " Mugger " of India. The premaxillo-
maxillary suture is transverse, as in the Alligators. The adults

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CROCODILIDAE

455

'(\..-:J

retain the five teeth in each half of the premaxilla. The
mandibular symphysis is short, extending only to the level of
the fourth or fifth tooth. The snout is stout, rather broad ; the
top of the head is rough but without any ridges. The upper and
lower jaw each contain nineteen teeth
on either side. The nuchal scutes,
six in number, are packed closely
together, the four biggest forming a
square. Four smaller scutes are
arranged in a curved line on the
occiput. The dorsal shield is com-
posed of four, sometimes of six rows
of larger scutes, of which the central
pair is the broadest. The fingers are
webbed at the base ; the outer toes
are broadly webbed, and the outer
edge of the hind-limbs is turned into
a serrated fringe. The general colour
of the upper parts is dark olive-
brown ; the young are pale, with
black spots. The length of twelve
feet is considered a fair average size
for a large specimen.

This, the " Marsh Crocodile," has
a wide distribution. It inhabits the rivers, ponds, tanks, and
marshes of India and Ceylon, extending eastwards through
Burma and Malacca iato most of the Malay islands, westwards
into Beluchistan. This species is fre(j[uently venerated by the
Hindoos, and is kept in a kind of domesticated condition,
attended by fakirs. One of the most famous crocodile ponds,
the so-called " mugger-peer," lies in an oasis of the sandy
stretches to the north-west of Karachi. A. L. Adams has
described a visit to this pond.^

" The greater pond is about 300 yards in circumference, and
contains many little grassy islands, on which the majority of the
Crocodiles were then basking ; some were asleep on its slimy sides,
others half submerged in the muddy water, while now and tlien
a huge monster would raise himself upon his diminutive legs, and
waddling for a few paces, fall flat on his belly. Young ones,

^ JFandcrings of a Naturalist in India^ Edinburgh, 1867.

Fig. 107. — Dorsal view of the skull
of Crocodilxis palustris. x \.
The arrangement of the nuchal
scutes is shown in the upper
left-hand corner ; ii', position of
the ear-flai).

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45 6 CROCODILIA CHAP.

from a foot in length and upwards, ran nimbly along the margin
of the pond, disappearing suddenly in the turbid waters as soon
as we approached. The largest crocodile lives in a long narrow
tank separated from the others. The fakirs, and natives who
worship in the neighbouring temples, have painted his forehead
red ; they venerate the old monster, making a salaam to his
majesty whenever he shows himself above water. A handsome
young Beloochee, whose occupation it was to feed the animals,
informed us that this specimen was upwards of 200 years old,
and that by way of a ' tit-bit ' he was in the habit of devouring
the young crocodiles. During our visit this enormous brute
was asleep on the bank of his dwelling-place, and seemed quite
indifferent to our presence, although we came within a foot of
him, and even attempted to arouse him by rubbing his nose with
a leg of goat's flesh, which, however, a young one greedily seized.
Our attendant tried in vain to excite their ferocity, but beyond
a feeble attempt to snap their trenchant teeth, the animals
showed no disposition to attack us.

" A pony was wading about in the pond and feeding on the
grassy hillocks, but the crocodiles took no notice of him.

" The crocodiles dig deep in the sand, under the neighbouring
date-trees, and there deposit their eggs. Quantities of deciduous
teeth, of various sizes, were strewn along the slimy sides of the
pond.

" Strangers are expected to stand treat, not only by the fakirs
and natives, who gain a livelihood by hanging about the pond
and showing the monsters, but even the crocodiles themselves
seem to anticipate a feast, and on the arrival of a party come
out in unusual numbers. Accordingly, we had a goat slaughtered,
during which operation the brutes seemed to rouse themselves,
as if preparing for a rush. Then our guide, taking piece after
piece of the flesh, daslied it on the bank, uttering a low growling
sound, at which the whole tank became in motion, and croco-
diles, of whose existence we had been before ignorant, splashed
through the shallow water, struggling which would seize the
prize. The shore was literally covered with scaly monsters,
snapping their jaws at one another."

Sir J. Emerson Tennent^ has had many opportunities of
studpng the habits of the Marsh Crocodile. According to him
* Sketches of the Natural History of Ceylon^ London, 1861.

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CROCODILIDAE 457

it is essentially cowardly in its instincts, and hastens to conceal
itself on the approach of man. One of these creatures, which
was overtaken in the jungle by a gentleman riding on horseback,
fled to a shallow pool, and thrusting its head into the mud till
it covered up its eyes, remained motionless, in profound con-
fidence of perfect concealment.

"There is a popular belief that the crocodile is exceedingly
sensitive to tickling, and that it will relax its hold of a man if
he can only contrive to reach and rub with his hand the softer
parts of its under side. An incident of some reality in this
piece of folk-lore came under my own observation. One morning
... we came suddenly upon a crocodile asleep under some bushes
of the buflfalo-thorn, several hundred yards from the water. The
terror of the poor wretch was extreme when it awoke and found
itself discovered and completely surrounded. It was a hideous
creature, upwards of 10 feet long. ... It started to its feet
and turned round in a circle, hissing and clanking its bony
jaws, with its ugly green eye intently fixed upon us. On being
struck with a stick, it lay perfectly quiet and apparently dead.
Presently it looked cunningly round, and made a rush towards
the water, but on a second blow it lay again motionless and
feigning death. We tried to rouse it, but without effect ; pulled
its tail, slapped its back, struck its hard scales, and teased it in
every way, but all in vain ; nothing would induce it to move
till, accidentally, my son, then a boy of twelve years old, tickled
it gently under the arm, and in an instant it drew the limb close
to its side and turned to avoid a repetition of the experiment.
Again it was touched under the other arm, and the same emotion
was exhibited, the great monster twisting about like an infant to
avoid being tickled."

In the dry season, when the tanks become exhausted, the
Marsh Crocodiles have occasionally been encountered in the
jungle, wandering in search of water. During a severe drought,
in 1844, they deserted a tank near Kornegalle, and traversed
the town during the night, on their way to another reservoir in
the suburb; two or three fell into the wells; others, in their
trepidation, laid eggs in the street, and some were found en-
tangled in garden fences and killed.

Generally, however, during the extreme drought, when un-
able to procure their ordinary food from the drying up of the

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CROCODILIA

CHAP.

watercourses, they bury themselves in the mud and remain in a
state of torpor till released by the recurrence of rains.

C, porosus s. biporcatus. — The premaxillo- maxillary suture
on the palate does not form a transverse line, but is W-shaped,

and extends backwards as in the
rest of the species of Crocodiles
to be described. This Indian
species is easily recognised by
the prominent longitudinal
ridge which extends in front
of each eye, over the prefrontal
bones, and by the absence of
sub-occipital scutes. The nuchal
scutes consist of four large ones,
which form a square, and one
or two smaller scutes on each
side. The dorsal shield con-
tains four to eight principal
longitudinal rows. The digits,
webs, and the seiTated fringe of
the legs are like those of C.
pcdustris. The head and snout,
however, are distinctly longer,
and more slender in proportion,
and the adult has only four
teeth in each premaxilla. The
general colour is dark olive-
brown. Young specimens, as
usual, are much paler and are
spotted with black.

This species attains a much
larger size than the Marsh
Crocodile. Specimens of 15
to 20 feet in length are not
uncommon, and there is a record of one monster of 33 feet.
Consequently this is, both in bulk and length, undoubtedly the
largest species of recent reptiles. It is essentially an inhabitant
of tidal waters or estuaries, frequently entering salt water and
going out to sea. Herewith corresponds its wide distribution,
namely, the whole coast of the Gulf of Bengal, extending to

Fio. 108. — Dorsal view of the skull of
CrocixlUiva porosus. x about \. Col,
buttress connecting the postfrontal with
the jugal and ectopter>'goid ; F, frontal ;
•^^« jugal ; Mx^ maxillary ; iVo, nasal ;
/*, parietal ; Pwt, premaxilla ; Po/, post-
frontal ; Pry, prefrontal ; Q, quadrate ;
QJj quadrato-jugal ; R, the characteristic
ridge on the prefrontal bone ; Sq, squa-
mosal ; r, perforations in the premaxilla
caused by a pair of lower incisor teeth.

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CROCODILIDAE 459

Southern China^ and across the Malay Archipelago to the
northern coasts of Australia. Eastwards it ranges to the
Solomon Islands and even to Fiji. Curiously enough, it does
not seem to occur on the west coast of India.

According to Tennent it is ready to assail man when pressed
by hunger, and the same authority mentions the following
serio-comic incident. A man was fishing, seated on the branch
of a tree overhanging the water, and to shelter himself from the
drizzling rain he covered his head and shoulders with a bag
folded into a shape common with the natives. While in this
attitude, a leopard sprang upon him from the jungle, but
missing its aim, seized the bag and not the man, and fell with
it into the river. Here a crocodile, which had been eyeing the
angler in despair, seized the leopard as it fell, and sank with it
to the bottom.

I have had some personal experience in the bringing up of
the young of this species. Two dozen of them had come from
Ceylon when quite young, only one foot long. At first they
were very shy, and huddled together in their tank, but they
took food greedily — strips of fish and, later on, sheep's heart.
When frightened they emitted peculiar, high-pitched, half-
croaking sounds. Some of them snapped at the finger when
touched ; others were of a more gentle disposition ; the shy ones
were undoubtedly the most vicious. Within one year they
grew to 18 or 20 inches, and added much to their bulk. Then
they were transferred to a deeper and larger tank in a green-
house, in which they could roam about at liberty. In the
daytime they dozed on the margins of their pond, mostly in
such a position that, at the slightest alarm, they could plunge
kick into the water. The strongest specimen left the tank
entirely, and took up its favourite place for basking on the
stump of a tree, to reach which it had to climb up a rough wall
of stones. After three years, several had grown to the length of
three and a half, and even four feet, and had by this time
become formidable pets. Although handled frequently, they
never became tame, the only change in their behaviour being
that, instead of rushing off in a fright, and hiding for half an
hour at the bottom of the tank, they became more vicious and
confident, making for and snapping at the hand which fed them.
The nights were spent regularly in the water, either floating

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460 CROCODILIA

with just the nostrils exposed, or in search of food, frogs being
their favourite prey, while their main sustenance consisted of
" lights," with an occasional mouse, or a piece of solid meat by
way of an entrie. Small pieces were bolted. The tough " lights,"
namely lungs with the windpipe and blood-vessels, were causes of
great quarrels. Two or three would get hold of a lump of this
kind, tearing at it, and twisting and rolling over in opposite
directions. The supply of warm water came through a stout
pipe of red india-rubber, and this was an irresistible attraction
to the crocodiles. On many a morning the tube was found
twisted into a knot, one of the creatures having spent hours
in chewing it and in trying to wrench it oflE In order to aid
digestion they swallowed pebblea The most favourable
temperature of the water was SS'' F. ; if below 75° F. they
refused to eat, but a continued exposure to 60° F. did not hurt
them. When the temperature rose above 95° F. they left the
water, although means had to be taken to
prevent them from lying on the hot-water
pipes,

C, niloticus s. vulgaris, — The pre-
maxillo-maxillary suture on the palate is
W-shaped. The nasal bones form only a
small part of the posterior border of the
nasal groove. There are eighteen or nine-
teen upper and fifteen lower teeth on each
side. In old specimens some of the
anterior mandibular teeth perforate the
premaxillae, as indicated in Fig. 109, and
they even pierce through the integument
so as to be visible from above. The
Fig. 109.— Dorsal view^ho nuchals are composed of four large scutes,
skull of a very old specimen with a smaller One on each side and

of Orocodilus niloticus^ in . "ii-i jxi

whuih most of the bony somctimes One behind, and there is a row
sutures are obliterated, of smaller pieces across the occiput. The

X about yS- j 1 1 • 1 t . . . 1 . . -I

dorsal shield contains six to eight principal
longitudinal rows. The fingers are webbed at the base : the
outer toes are very broadly webbed; and there is a serrated
fringe on the outer side of the leg. The general colour of the
adult is dark olive -brown ; the young are paler, with black
spots and vermiculations. The under parts are yellowish white.

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461

The Nile Crocodile is essentially African, ranging from the
Senegal to the Cape and to Egypt. It is also very common in
Madagascar. Nothing is known about its occurrence in Arabia,
but a few specimens of
rather small size seem
still to exist in Syri«f,
in the Wadi Zerka, an
eastern tributary of
the Jordan.

Even in historical
times the Crocodile
must have been very
common in lower Egypt,
to judge from the
number of mummies
preserved by the old
Egyptians. Now it is
practically extermi-
nated, and there are
scarcely any left below
Wadi Haifa.

Such a conspicuous
and dangerous creature
has naturally always
enjoyed notoriety. It
is well described in one
of the oldest writings
of the world, the Book
of Job. " Canst thou
draw out leviathan
with an hook ? or his
tongue with a cord
which thou lettest
down ? . . . His scales
are his pride, shut up
together as with a close
no air can come between them,
they stick together, that they

FiQ. 110. — Ventral view of a young Crocodilus nUotiatSj
showing the arrangement of the bony scutes and
the two openings of the musk -glands on the lower
jaw. The upper right-hand figure shows on a larger
scale the disposition of the nuchal scutes and the
first row of dorsal scutes.

seal.

One is so near to another, that
They are joined one to another,
cannot be sundered. . . . Lay
thine hand upon him, remember the battle, do no more." Bows
and arrows, spears and clubs, are of little avail against such a

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462 CROCODILIA CHAP.

monster ; the dragging out of a hooked, full-grown specimen
requires many men and is a formidable task. Of coui-se fire-
arms have changed all this, and its invulnerability to bullets is
nonsense. It is true that a bullet sent into the head is generally
ineffective, since it is a hundred to one that the bullet does not
hit the small brain, and even if it does, the creature sinks to the
bottom and is lost to view until decomposition sets in and the
gases developing in the body cause it to float.

Herodotus has quaint stories about these crocodiles and their
worship. Amongst other stories he mentions that the bird
Trochilus, supposed to be the Fluviantcs aegyptius, a kind of
Plover, slips into the gaping mouth to pick off the leeches
which infest the reptile's gums. " In Egypt it is called
Ghampsa, but the lonians call them KOKoBpiXoi on account of the
resemblance to the lizards which live on their garden walla"
This is in fact the origin of the name crocodile, KopBvT^jo^ being
the ancient Greek for lizard and newt. With reduplication
KopKophvXo^ and by metathesis ultimately KpoKoBetXo^. The
Arabic name is ledschun.

The story about the Plover seems to be true. These birds
are sometimes seen sitting upon basking crocodiles, and since the
latter are in the habit of resting, perhaps half asleep, with the
mouth wide open, it is possible that these agile birds do pick
their teeth, and that they, being also very watchful, by their
own cry of warning and by fluttering off on the approach of
danger, give the alarm to the crocodiles and thus benefit them
in more than one way.

But the equally old story about the Ichneumon or Mongoose
is an idle invention. Mongooses are paitial to eggs, but they
certainly prefer those of hens and other birds to those of the
crocodile, which are far too hard and strong to be broken by
such a little animal. Moreover, as we shall see presently, the
eggs are far too well concealed.

The best account of the habits of these crocodiles is the one
given recently by Voeltzkow,^ who has spent a long time in
Madagascar to collect material for the study of their develop-
ment.

He says that C. nilotictts is not only the most common
reptile, but perhaps the most common vertebrate in Madagascar.

1 SUzber, Ak. Berlin. 1891, p. 115 ; 1893, p. 347.

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CROCODIUDAE 463

It occurs in every pool and river in great numbers, especially
upon the sandbanks of the Betsiboka River, where one may see
more than one hundred within one hour's paddling down stream.
The largest specimen measured by Voeltzkow was 1 3 feet long ;
the largest in the National Collection is a little less than 15
feet.

The -crocodiles are caught in various ways. The simplest
apparatus consists of two pointed sticks, which are fastened cross-
wise within the bait to which is attached a rope, and this is
made fast on the bank of the river or lake. The animal, when
it has once swallowed this spiked bait, keeps its jaws firmly
closed, so that it can be dragged out of the water. Another
method is more reliable. A long and strong rope is made into
an easily slipping noose, with an opening of about 18 inchea
The bait is attached to the upper part of the noose, while the
lower portion is kept open by a springy branch, the whole
thing being so balanced that it will float upright. When a
crocodile seizes the bait, which it does with a side jerk of the
head, the branch falls out of the noose and the latter closes
around the upper or lower jaw.

These crocodiles dig long subterranean passages of 30 to 40
feet in length ; the passage opens in the bank below the level of
the water, and gradually ascending ends in a somewhat wider
compartment, which allows the creature to turn round. Two or
three air-holes are pierced through the ceiling of the burrow, in
which bones and other remains of food are often found, so that
the natives' belief, that the crocodiles retire into these chambers
in order to devour their prey in undisturbed secrecy, appears
very probable. When suddenly disturbed or frightened they
take to these lairs, and since their position is clearly marked by
the air-holes, the natives block the passage and then dig the
animal out from above.

Eggs are laid, in Madagascar, from the end of August to the
end of September ; the number of one set varies from twenty to
thirty. They are deposited in a nest. This is in the ground,
mostly in white sand, and consists of a hollow 18 inches to
2 feet deep. The walls are rather vertical, but near the bottom
they are undermined, and here the eggs are placed. The centre
of the pit being somewhat higher, the eggs roll by themselves
into the imdermined peripheral region. The laying takes place

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464 CROCOblLIA

during the night, mostly a little before daybreak. After one
half of the eggs has been laid, they are covered up with
sand, whereupon the other half is deposited. Then the hole is
completely filled up and no visible traces are left behind ; but
the mother sleeps upon the nest and thus leads to its discovery.
The position of the nest is so chosen that it cannot be reached
by moisture from below ; the eggs are most susceptible to moisture,
a very slight amount of which causes them to turn bad.

The shape of tlie eggs of one and the same clutch varies
much, some being elliptical, others cylindrical with rounded off
ends. Their size varies from 5*5 to 9 cm. in length, and 4 to
5 cm. in width. The shell is white and glossy, thick and hard,
either roughly granular or smooth. They are hatched in about
twelve weeks.

Voeltzkow feels certain that the mother returns to the nest
at the proper time in order to dig the young ones out and to
conduct tliem to the water. To test this story he had a nest
sun'ounded with a fence ; the mother returned several times and
partly destroyed the fence, which was then replaced by a stronger
one. One day, when the young had been hatched, the nest was
found to be filled with sand, the shells and one dead little
crocodile being at the bottom of the hole. The mother had dug a
deep ditch below the fence, but had not succeded in reaching the
nest, although she had received and conducted her offspring
away. As a rule, when the young are hatched, the sand and the
shells are found to be scraped out of the nest. The mother is
probably warned by the hiccough-like sound which the young
emit while still within the unbroken shell. Voeltzkow heard them
piping from the other end of his room, the eggs being covered
with a layer of sand two feet high. The sounds were heard
when he walked past the nest, or knocked against the box.
Possibly the young hear the mother when she retires to the nest
to sleep on it, and give her warning to remove the eggs out of
the groove. However, they do not break the shell until several
days later.

The hatching is not caused by the rainy season, since it took
place a fortniglit before the first showers. The " egg-tooth " of the
newly liatched young is 0'5 to 0*75 mm. high, bicuspid, and acts
like a borer or auger. It is still visible on the tip of the upper
jaw, in front of the nose, when the creature is two w^eks old. The

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CROCODILIDAE 465

newly hatched crocodile is of an astonishing size, so that it is rather
puzzling to understand how it was stowed away in the egg.
For instance, an egg of 8 cm. length and 5 cm. width, sends
forth a crocodile 28 cm. or 11 inches in length. Even at this
early age they snap at the finger.

The egg is covered by a hard shell, within which is a thicker
outer and a thinner inner membrane. The " white " is jelly-like,
sometimes of a greenish tinge, and is so consistent that it will not
flow. ' The yolk is round, and so large that it nearly reaches the
shell-membrane in the short diameter. The yolk itself is sur-
rounded by a very thin but strong membrane.

The embryo begins to develop long before the egg is laid.
When laid the germ is about 4 mm. long and shows about twelve
somites. The cephalic bend begins at the end of the second
week, the tail grows longer and the embryo becomes curled up.
At the end of the third week it measures 10 mm. in a straight
line from brain to vent. The limbs begin to bud in the fourth
week. With the sixth week the final shape begins to reveal
itself, and is completed at the age of eight weeks ; but a third
month is necessary to ripen the embryo.

C. cataphractiis is the Common Crocodile of West Africa,
from the Senegal to the Congo. In opposition to C. niloticus
it does not enter brackish water. It is easily recognised by the
very slender snout, which rather resembles that of the Gavial ;
but the mandibular symphysis, although extending to the level
of the eighth tooth, does not reach the splenial bonea The
premaxillo-maxillary suture on the palate is not transverse, but
extends backwards. In conformity with the length of the
snout the maxillaries meet in the dorso-medial line behind the
nasal opening, thus excluding the nasals from the latter. The
nuchal scutes consist of two large pairs, almost in contact with
the dorsals, six of which form the principal longitudinal rows.
The gular and ventral scutes ossify in the adult, hence the
specific name. The fingers and toes are slightly webbed.
General colour above, dark olive-brown ; yellowish below. The
young are olive with large black spots.

The natives of the Lower Congo catch the crocodiles with two
pointed sticks tied together cross-wise, surrounded with entrails
by way of a bait. The whole is fastened to a pole or a strong
rope and thrown into the river ; and a narrow line, with a float

VOL. VIII. 2 H

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466

CROCODILIA

attached to the cross-sticks, indicates the whereabouts of the
crocodile when it has taken the bait and has sunk to the bottom.
C. johnstoni, of Northern Australia and Northern Queensland,
and (7. intermeditts, of the Orinoko, are allied to C. cataphr actus,
at least so far as the configuration of the bones of the slender
and long snout is concerned. The former is small, scarcely

reaching the length of 7 feet, while
the South American species grows to
13 feet.

(7. americanus s. acutus. — This
species, which inhabits the West
Indian Islands, being there the only
representative of the order, occurs
also in Florida, and extends through
the warmer parts of Central America
into Venezuela, Colombia, and Ecuador.
Its characteristic feature is a median
ridge or swelling on the snout. The
length and relative width of the
latter varies considerably. The
maxillaries sometimes meet dorsally,
or they remain separated by the
narrow nasals, which in this case
reach the posterior corner of the
nasal groove. The nuchal scutes
vary likewise ; there being often a
pair on the side of and
another behind the four principal
scutes, which, as usual, form a square.
A transverse row of little suboccipital
Largest size about 12 feet long.

Ptr.

Fio. 111. — Dorsal view of the skull
of Crocodilus americanus. x J.
Fj Frontal ; Jg^ jugal ; L, lacry-
mal ; Mxy maxillary ; iVo, nasal ;
P, parietal ; Pwix, premaxillary ; smaller
Pr/j prefrontal ; P{f, post-
frontal ; Qj\ qnadrato-jiigal ; Sq^
squamosal ; T, tooth -perfora-
tion.

scutes is also common.

Osteoldemus tetraspis s. frontatus, — The only species of this
genus inhabits the rivers of the west coast of Africa, from Sierra
Leone to the Ogowai It differs from Crocodilus chiefly by the
bony septum of the nasal groove, produced by forward extension
of the nasal bones. The snout is rather short and stout ; the
upper surface very rugose and deeply pitted, but without ridges.
The gular and ventral scutes are ossified, hence the generic nama
Total length about 5 feet.

Alligator. — The fourth mandibular tooth fits into a pit in

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CROCODIUDAE 467

the upper jaw, and this pit is in some adult specimens trans-
formed into a hole, the tip of the tooth appearing on the upper
surface through the perforation. Most of the other teeth of the
lower jaw are overlapped by those of the upper jaw. The number
of teeth on either side amounts to seventeen to twenty in the
upper and eighteen to twenty in the lower jaw. The nasal bones
form not only the posterior border of the nasal groove, but they
divide the latter by a median bony septum. The dorsal shield is
formed by six or eight longitudinal series of keeled bony scutes,
which, although standing close together, do not articulate with
each other. Ossification of the gular and ventral scutes is absent
or very slight.

Alligators occur in the fluviatile deposits of the age of the
Upper Chalk in Europe, w^here they did not die out until the
Pliocene age ; they are now restricted to two species, one in the
Southern States of North America, the other in China.

A. mississippiensis. — The much-depressed and broadly rounded
snout bears some resemblance to that of a pike, hence the now
discarded specifis name of lucius. The neck is protected by two
pairs of large scutes, which form a square, interrupted in the
middle line, with a pair of small scutes in front and another behind.
Of the eighteen transverse dorsal rows of scutes eight are broad
and prominent. The fingers are about half webbed, the outer
toes about two-thirds webbed. The general colour is greenish black
or dark brown above, yellowish below. Young specimens have
yellowish cross-bands on a darker brown groxmd.

The Alligator's northern limit is the mouth of the river
Neuss in North Carolina, 35"^ N. lat. From this point they
abound near the mouths of all the creeks and rivers as far south
as the Bio Grande, ascending the Mississippi to the entrance of
the Eed River in 33° 50' N. lat.

The habits and the embryology of the American Alligator
have been described by S. F. Clarke,^ who gives the following
vivid and minute account : —

" Usually one finds them in the waters of the smaller streams
and ponds, lying with only the tip of the nose and the eyes
exposed, or lying on an exposed place on the bank where the
grass and other plants are beaten down, and the black, rich mud
of the river bank is smoothed by the repeated movements of the

^ J, Morphol. V. 1891, p. 181.

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CROCODILIA

CHAP.

alligators in climbing up and down. There they bask in the
sunlight until disturbed by the hunter or the desire for food.
When aroused they make for the bottom, and I have never
waited long enough to see one return unless he were vigorously
stimulated with a long pole. They frequently dig a cave for

Fig. 112. — Skull of AUigator mississippiensis. A, Dorsal ; B, ventral ; C, lateral view.
Ag, Angalar bone of mandible ; Cd^ occipital condyle ; Ch, choanae or posterior
narial openings — the median small hole behind them indicates the position of the
opening of the Eustachian tubes ; Jg^ jugal ; Z, lacrymal ; Mx, maxillary ; No,
nostrils ; Pa^ palatine ; Pw, premaxillary ; PU pterygoid ; Q, quadrate ; T, TV, trans-
verse bone or ectopterygoid.

themselves in the bottom of the pond or stream, or in the bank
beneath the water. Oftentimes one can start them out of the
cave by using a pole, but if very obstinate, the hunters dig them
out with spades.

" As tlie water decreases in the streams and ponds with the
summer heat, the alligators travel to the larger bodies of water.

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CROCODILIDAE 469

During the breeding season, from the end of May to the beginning
of July, the males are very active, wandering about to various
ponds and rivers in search of the females. Fierce battles are
said to take place during this time between the excited males ;
and the mutilated specimens that one sees are weighty evidence
for the truth of this assertion. ... It is in the breeding season
also that their bellowing is mostly heard, and more in the night
than during the day. I have frequently heard them, while lying
in the swamps at night, when they were in ponds fully a mile
distant.

" The largest specimen I saw measured 1 2 feet in length ; and
none of the many hunters and other natives of Florida I have
met have seen any longer than 13 feet. All the hunters agree
that it is only the males that acquire the great size ; no one had
ever seen a female that measured over 8 feet, and the majority
are not over seven.

" The male has a heavier, more powerful head, and during
the breeding season especially is more brilliantly coloured. The
more brilliant colour occurs in patches and streaks on the sides
of the head and body ; it is generally a light yellow, or even
whitish, and on one large male I saw a fairly bright red spot
over each eye.

" The alligators are rapidly diminishing in numbers under the
stimulus of the high prices offered to the hunters for their hides.
Both Whites and Indians make increasing war upon them.
Several thousand skins were brought into the little station of
Fort Pierce in 1890. The pioneers and settlers always destroy
the nests and eggs, because the alligators eat their pigs ; and the
cleaned eggs and young alligators are sold by hundreds in the
curio shops farther north. As their numbers diminish in Florida
it is noticed that the Moccasin snakes increase. In Louisiana
also the alligators are disappearing ; and there the musk-rats are
at the same time increasing, and are doing much damage by
burrowing in the levees along the Mississippi. While the
alligator can make a very stout fight, I have never seen one
offer fight if there was any chance of retreat. They never offered
to molest us, even when we waded through the ponds where they
were.

" The nest of the alligator is very large, and is built by the
female. A great quantity of dead leaves and twigs, together with

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470 CROCODILIA CHAP.

much of the finely divided humus underlying them, ia scraped
together into a low mound about 3 feet high ; this varies consider-
ably in its other dimensions, being in some instances 8 feet in
diameter at the base. The nests are built on the bank of a stream
or pool, and the female digs a cave under water in the bank close
to the nest. Careful examination of the largest nest found showed
a root of a neighbouring palmetto-tree, nearly an inch in diameter,
running through it at about a foot above the ground ; there were
also roots of a grape vine growing near, which extended nearly
through the nest. This furnishes strong support to the state-
ment of many of the himters, that the nests are used for more
than one season. I could get no evidence whatever that the
nests are used more than once a year.

" The eggs are laid near the top of the nest, within 8 inches
of the surface, are four or five layers deep, and have no regular
arrangement or uniform position of their axes in relation to the
nest. The number of eggs to a nest varies from twenty to thirty,
and averages twenty-eight ; the maximum found was forty-seven.

The eggs are white, elliptical, and vary in length from 50 to
90 mm. or 2 to 3^ inches, and in the shorter diameter from 28
to 45 mm. Generally there is only slight variation in the eggs
of one nest, but occasionally a nest is found in which most of the
eggs are about the average size, while from two to five are very
much smaller.

" The shell is much rougher than that of a hen's egg, and much
thicker. The shell membrane consists of an outer and an inner
layer, in both of which the fibres are arranged spirally about
the egg, but at right angles to one another.

" The white of the egg has the consistency of a very thick
jelly, is very clear and transparent, and is so firm that the whole
egg, when perfectly fresh, may be turned out of the shell and
shell membrane, and transferred from one hand to the other
without breaking, and with but slight change of form. The
white lies mostly at either end of the shell, but extends also in a
thin layer between the yolk and the sides of the shell. The yolk
holds a median position in the egg, is spherical, of a very light
pale yellow, and so large that it almost touches the shell
membrane about the midline."

According to Holbrook the young as soon as they are dis-
engaged from the shell seek the water and shift for themselves.

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CROCODILIDAE 47 1

the parents taking no care of them, though they may remain for
some weeks in the same locality. In the spring and early
summer months, and during the time of incubation, and especially
on cloudy days or in the evening, alligators make a great noise ;
their croak is not unlike that of the bullfrog, but louder and
less prolonged. On the approach of winter they select holes in
the ground, where they remain torpid until spring. In this
state of hibernation many are dug out by the negroes, who
esteem the tail as an article of food.

A, sinensis. — The first intimation of the existence of a
Crocodilian in the Yang-tse-kiang was made by Swinhoe in
1870, but it was not until nine years later that FauveP
described the creature as A, sinensis. The same gentleman gave
also an exhaustive account of the former records of this species
in Chinese literature. According to Boulenger its nearest ally
is A. missi^ippiensis, but it approaches the Caimans by the
presence of ossifications in the ventral shields, which ossifications
are, however, wide apart from each other. There are three pairs
of large nuchal scutes in contact in the median line, besides
smaller scutes in front of the nuchals and behind the occiput.
The dorsal shield contains six rows of larger scutes. The fingers
are not webbed. The general coloiu' is greenish black above,
speckled with yellow ; greyish below. Total length only about
six feet.

Caiman. — The five species of this genus, confined to Central
America or to the East Andesian parts of South America,
resemble the Alligators in most features, but differ from them in
the following points. The nasals, although bordering the nasal
groove, do not form a bony nasal septum. The supratemporal
fossae are very small ; or closed up, as in C. trigonaius and C.
palpehrostts of Guiana. The ventral armour is composed of over-
lapping bony scutes, each of which is formed of two parts united
by a suture.

C. sclerops has the widest distribution, from Southern Mexico
to the northern half of Argentina. The upper eyelid is rugose,
although only incompletely ossified, and is often more or less
produced into a small horn. C. niger has flat upper eyelids.

According to Bates, Caimans exist in myriads in the waters
of the Upper Amazons. One species, C. trigonatus, the Jacari-
^ J. China Asiat. Soc. xiii. 1879, pp. 1-36, with Figs.

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472 CROCODILIA chap, x

tinga of the natives, reaches only six feet in length and has a
slender muzzle and a black-banded tail. Another species, C.
niger, the Jacar^-nassu or large Caiman, attains an enormous
bulk and a length of 20 feet. They migrate annually, retreating
to the flooded forests in the wet season and descending to the
main rivers in the dry season.

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CHAPTER XI

PLESIOSAURIA ICHTHYOSAURIA PTEROSAURIA PYTHONOMORPHA

Sub-Glass VIL— PLESIOSAURIA.

Mesozoic aquatic reptiles, with two pairs of pentadactyle limbs,
firmly fixed quadrate hones, single tem2)oral arches, numerous
alveolar teeth, and ribs which articulate only unth the centra
of the biconcave vertebrae.

The Plesiosauria comprise the Mesosauri, Nothosauri, and Plesio-
sauri in an ascending order of development, which concerns
especially the changes from a semi -terrestrial to an absolutely
aquatic life ; — elongation of the neck with corresponding
shortening of the tail, and the graduafl transformation of the
limbs into hyperphalangeal paddles.

The skull varies considerably in length. Seen from above it
shows the nostrils, orbits, very large supratemporal foramina,
and the interparietal hole. The nostrils lie rather far back, in
front of the orbits, between the elongated premaxillaries, short
nasals, and the usually large maxillaries. The orbits are rather
small, bordered behind by the postfrontals and postorbitals,
which two bones fuse together in the Plesiosauri. The temporal
bridge is long, and is formed by the junction of the two bones just
mentioned with the squamosal mass, which overlaps the greater
portion of the quadrate, and perhaps contains the quadrato-jugal.
The dorsal branch of the squamosal joins a corresponding diverg-
ing branch of the parietal, and completely shuts off the posterior
region of the supratemporal foramen. The interparietal hole is
small and placed far back. The palate possesses a row of teeth
on the pterygoids in Lariosaurus. The choanae open separately
between the vomers and maxillaries. . The pterygoids are very
long ; posteriorly they join the quadrates, anteriorly they extend

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474 PLESIOSAURIA

right up to the vomers, separating the palatines from each other
thereby. Palatal vacuities are absent in Nothomurus; small
and oval, between the palatines, pterygoids, ectopterygoids and
maxillaries in Lariosaurus ; still smaller in the Plesiosauri.

The vertebrae are mostly biconcave, in the Triassic genera
still perforated by the chorda, while in many Plesiosauri the
centra are solid, with almost plane articulating surfaces. The
neural arches are usually firmly sutured, or quite fused with the
centra. Intercentra are absent, except as chevrons in the tail.
Although the cervical and some of the thoracic ribs of the
Triassic genera have typical capitula and tubercula, they articu-
late exclusively upon the centra, and not upon the neural arches
also. The number of cervical vertebrae amounts to nine in
Mesosaurus ; in Lariosaurus it is increased to about twenty ; and
in some Plesiosauri to between thirty and forty. The cervical ribs
are very short, but they increase gradually towards the thorax,
which is well protected by long and strong ribs, which decrease
again very gradually, being still long in the lumbar region.
There is, properly speaking, no sacrum, because the one to four
sacral ribs remain quite separate. The tail is still long in
Lariosaurus, consisting of about forty much shortened vertebrae ;
considerably shorter than the neck in most of the Plesiosauri.
A sternum is absent, but the belly is protected by many strong
abdominal ribs, crowded together, and consisting each of a median
and two pairs of lateral pieces.

The shoulder-girdle is very strong, composed of scapulae,
very strong coracoids, clavicles, and an interclavicle. The pre-
coracoids are indicated by a process and a notch in the Triassic
genera ; in the later forms they are abolished. The coracoids
always meet in the median line, and often produce a strong
symphysis. The scapulae possess a very prominent and large
acromial process, upon which rest the dorsal or lateral ends of
the clavicles. In some Plesiosauri the shoulder-girdle has under-
gone an absolutely unique modification. The correct interpre-
tation has been given by C. W. Andrews after the examination
of exquisitely preserved specimens of Cryptoclidus from the
Oxford clay of the Middle Oolite, near Peterborough. The
dorsal portion or main shaft of the scapula is reduced to what
now looks like a dorso-lateral process, while the broad acromial
process is much elongated, and lies in Plesiosaurus upon the

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SKELETON

475

ventral surface of the clavicle; the latter and the irregularly
T-shaped interclavicle being, however, still visible from below.
In CryptocliduB the two acromial processes meet each other and form
a long ventral symphysis, which meets that of the much-enlarged
coracoids, the latter enclosing with the scapulae a pair of roundish
foramina. The clavicles ai-e not visible from below; they rest upon
the dorsal surface of the scapular symphysis, and the inter-
clavicle seems to be suppressed. Young Cryptoclidus (Fig. 1 1 3, B)
and various species of Plesiomurus show intermediate con-
ditions.

This unique arrangement is correlated with the enormous

•&d^0^

^--riil,^;;

Fig. 113.— a, Restored outlines of a PlesiotauruSf x-^; B, dorsal view of the pectoral
arch of an immature Oryptoclidus, from the middle Oolite ; C, fore-limb of a Plesio-
saurua, from the lias. A, Acromial process of scapula; C?, clavicle; Co, cora-
coid ; fff humerus ; i, carpale intennedium ; Mi to mg, first to fifth metacarpals ;
p, pisiform bone ; It, radius ; r, radial carpal ; S, scapula ; U, ulna ; n, ulnar carpal.

development of the fore-limbs, although nothing of the kind has
taken place in the Ichthyosauri, which have similar large
paddles. The limbs exhibit considerable differences in the
various groups of Plesiosauria, but they are all pentadactyle. In
the oldest, the Mesosauri and Nothosauri, the limbs are still of
the terrestrial type, although fitted for swimming; the chief
bones are still slender and elongated, and none of the five
fingers and toes have more than five phalanges, the usual
number of which seems to be 2, 3, 4, 5, 3 for the first to fifth
digits respectively. In the Plesiosauri the limbs are trans-
formed into long hyperphalangeal paddles, unfit for progression
on land, rather like those of the Ichthyosauria, with much

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476 PLESIOSAURIA chap.

shortened radius and ulna, tibia and fibula ; but the phalanges,
which increase to about ten, are always longer than broad, and
there is no indication of an increase of the number of fingers, or
of additional, lateral, phalanx-like nodulea The pelvis is very
strong ; the broad 'pubes and ischia meet in the middle line, and
they either enclose one wide undivided foramen, or the two
symphysial portions meet, and there are then two obturator-
foramina. The pubes are generally much larger, especially
broader, than the ischia; and although partaking in the
formation of the acetabulum, they do not articulate with the ilia,
at least not in Plesiosauri. The ilia are always small ; in
Plesiosauri attached to only one or two sacral ribs ; to three or
four in the Triassic genera.

Ichthyosauri and Plesiosauri were combined as " Enaliosauria "
by Conybeare. Owen recognised their fundamental diflerences,
and separated them as " Ichthyopterygia " and " Sauropterygia,"
according to the structure of the limbs. We now know that
the paddles of the Ichthyosauri bear but a superficial resemblance
to the fins of fishes, and are fundamentally referable to the
pentadactyle type, as are tlie paddles of the Plesiosauri,
although the latter retain more of the typical fe^itures of
reptilian limbs. It was soon recognised that the Nothosauri
are allied to the Plesiosaul'i, but the Mesosauri (until then
vaguely grouped with the Ehynchocephalia, or linked Math
Protorosauri as Proganosauria) have only recently^ received their
proper place in the system as members of the Plesiosauria,
which we divide into two main groups.

Order I. NOTHOSAUBL'

The limbs are of the terrestrial type; the five digits have
the usual number of phalanges, which do not exceed five. The
bones of the limbs are slender; the humerus has an entepi-
condylar foramen.

Fam. 1. Mesosauridae. — The neck contains about ten
vertebrae. The vertebrae are deeply biconcave, perforated by the
chorda dorsalis. Sacral vertebrae four in number. Clavicles strong;
interclavicle very small. Mesosaurus, the only genus, with one
species, M. tenuidens, about one foot in length, was foimd in
* Boulenger, Trans, Zool, Soc, xiv. 1898 (read Nov. 1898). * F6^ot=spurioaa.

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XI NOTHOSAURI — PLESIOSAURI 477

South Africa, probably in Triassic sandstone. Very similar
specimens are known from S^o Paolo in Brazil.

Fam. 2. Nothosanridae. — With sixteen to twenty-one cervical
and three to five sacral vertebrae. The vertebrae are biconcave.
The clavicles are strong; the interclavicle ifi much reduced.
Coracoids with distinct acromial processea

NothosauTus mirahilis, of the Muschelkalk of Germany.
Total length about ten feet. Length of head about one foot.
The teeth are very irregular. About five slender, long teeth
are implanted in each side of the premaxilla, with wide spaces
between them, similar to those of the symphysial portion of the
lower jaw. Those of the maxillaries are numerous and small,
except two large pairs in the anterior portion, on a level between
the orbits and nostrils. The upper and lower teeth overlap, or
cross each other. The palate of the long and slender skull is
quite bony, without anterior palatal or infra-orbital vacuities.

Zariosav/rus balsami, about one foot in length, from the
fresh-water deposits of the Upper Trias in Lombardy. Neck
with about twenty, tail with about forty vertebrae. Head com-
paratively shorter ; more triangular than in Nothosaurvs ; palate
with small infra-orbital vacuities. The number of the phalanges of
the fingers and toes is apparently 2, 3, 4, 4, 3 and 2, 3, 4, 5, 4.

ATiarosauriLS pumiliOy of the Muschelkalk, near Magdeburg,
and Neusticosaurus and Simosaurus of the same geological age,
are allied forms.

Order IL PLESIOSAUBL

The limbs are transformed into hyperphalangeal paddles. The
clavicles are small, and are overlapped ventrally by the strongly
developed acromial processes of the scapulae. The vertebrae are
slightly biconcave or plane. The neck consists of at least twenty
vertebrae ; those of the thoracic region have long . transverse
processes; the sacral vertebrae are mostly reduced to two or
one. Very large, massive animals.

Fam. 1. Pliosaoridae. — About twenty cervical vertebrae, with
proximally bifurcated ribs. The scapulae do not meet ventrally ;
they enclose with the coracoids a single large foramen, and are
fused with the clavicles. Fliosaurus, the principal genus, con-
tains several species of gigantic size ; for instance, F. grandis, of
the Kimmeridge clay. Upper Oolite, of England, has a skull

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478 PLESIOSAURIA chap.

nearly 5 feet long and 2 feet broad, armed with many enor-"
mous conical teeth, some of which reach one foot in length,
inclusive of the long collar and root -portion. The neck is
rather short, owing to the much condensed, disc-shaped centra
of the vertebrae. Total length of this species about 30 feet.
Other species in England and continental Europe as far as Bussia.
Fam. 2. Plesiosaaridae. — ^The neck is very long, and consists
of from twenty-eight to forty vertebrae. The scapulae do not meet
ventrally, but the symphysial portion of the coracoids meets the
clavicles and the interclavicle, the pectoral arch thus enclosing
two foramina. Chief genus Plesiosaurus, with many speciea The
head is comparatively small, the neck very long, the tail short,
although consisting of from thirty to forty vertebrae. The third
digit (Fig. 11 3, C) is the longest,and possesses nineor ten phalanges.
The abdominal ribs are very strong, and reach from the pectoral to
the pelvic girdle. Eange from the Lower Trias to the Lower
Oolite, chiefly European. P. dolichodirus and P. conybeari, the
latter reaching a total length of more than 15 feet, from the
Lower Lias, especially at Lyme Kegis.

. Fam. 3. Elasmosaaridae. — The neck is extremely long, posses-
sing from thirty-five to seventy-two vertebrae, with single-headed,
not bifurcated, ribs. The scapulae meet ventrally, and enclose with
the very broad coracoids two foramina. The tail is short. The
pisiform bone articulates with the humerus. Otherwise much re-
sembling the Plesiosauridae. Principal genus Cimoliasaurus, with
many synonyms, and many species from the Middle Oolite to the
Upper Chalk ; cosmopolitan distribution, e.g. C. cantahrigierms, of
the Greensand and Upper Chalk ; C. trocharUericiis, of the Kim-
meridge clay ; C. haasti in New Zealand ; C. australis, C. chilensis ;
others in North America. Cryptoclidns of the Middle and Upper
Oolite of Europe. Masmosaurus, of the Upper Cretaceous forma-
tion in Kansas, with a computed total length of 45 feet, of which
22 belong to the neck, with its seventy-two vertebraa

Sub-Class VIIL—ICHTHYOSAURIA,

Marine y ichale- shaped reptiles, with the anterior and posterior
limbs transformed into hyperphalangeal paddles. Restricted
to the Mesozoic age from the Trias to the Upper Chnlk,

The skull is long, owing to the elongated slender snout, which

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ICHTHYOSAURIA 479

is formed mainly by the premaxillary bones. The nostrils lie
far back, in front of tlie orbits, and are bordered by the long
nasals, the premaxillaries, a small part of the maxillaries, and
posteriorly by the large lacrymal bones. The eyes are large, and
are strengthened by a sclerotic ring composed of many closely over-
lapping bones. The orbits are very large, and are directed side-
ways so as to be scarcely visible from above. They are formed above
by the long prefrontals, which join the postfrontals ; behind by
the long postorbitals ; below by the long and slender jugals ; in
front by the lacrymals and prefrontals. The postorbito-temporal
region of the skull is short but high, and, with the exception of
the supratemporal foramen, is entirely closed in by bones, namely,
the quadrato -jugals, supratemporals, and squamosals. The
latter, with the parietals and large postfrontals, surround the
supratemporal foramina. The parietals and the small frontals
enclose the parietal foramen. The whole temporal arch conse-
quently recalls much that of the Pareiasauri and Stegocephali,
chiefly owing to the presence of conspicuous supratemporal and
postorbital bones, which, together with the quadrato-jugal, close
in the whole side without any indication of a lateral or infra-
temporal foramen. The postorbital completely separates the
jugal from the quadrato-jugal, and this almost hides the quadrate.
The occipital condyle is single. The lateral occipitals and the
supra-occipital bones retain their sutures. The pro-otic and
opisthotic bones remain separate. The latter lie between the
basi- and lateral occipitals, the squamosal, quadrate, and pterygoid.
The pterygoids, which posteriorly touch the quadrato-jugals,
basi-occipitals, opisthotics, and basisphenoid, are very long and
remain widely separated from each other ; in the space between
them appears the long ensiform presphenoid. Anteriorly they
are connected through the ectopterygoids with the maxillae, and
touch the palatines. These are likewise narrow and slender, but
touch each other in the middle line, and contain the well-
separated, slit -like choanae, laterally to which lie the elon-
gated, rather narrow, palatal vacuitiea The vomers are mostly
not visible; when they appear on the surface they are long
and narrow, and enclose the choanae between them and the
palatines.

The teeth are pointed, conical and thickly covered with enamel,
which in transverse sections forms vertical ridges, recalling

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480 * ICHTHYOSAURI A

the folds of the Labyrinthodonts. The teeth have open roots,
and are not implanted in separate alveoli, but lie in long
grooves of the premaxillaries, maxillaries, and dentals.

The vertebrae are numerous, up to 150, two-thirds of which
belong to the tail The centra are deeply biconcave and short,
not co-ossified with the neural arches, which have therefore often
broken loose. The atlas much resembles the other cervical verte-
brae in so far as its centrum is concave in front and scarcely
ankylosed with that of the second. Its basiventrals, equivalent
to the ventral half of the atlas-ring of other reptiles, thus become
an unpaired intercentral wedge, between the first centnmi and the
basis of the cranium ; the neural arches rest upon the centrum,
but remain separate from each other, or at least diverge dorsally.
The atlas carries no ribs. Intercentra occur also between the
second and third vertebrae ; they reappear in the tail as chevron-
bones. All the other vertebrae carry ribs, which gradually
increase in length towards the trunk and decrease again equally
gradually on the tail. In the neck and trunk they have separate
capitula and tubercula, which articulate upon short knobs of the
centra ; towards the tail these shift farther and farther towards
the ventral side, and ultimately unite. Although the ribs of the
trunk are so long, there is no trace of a sternum, but there are
many " abdominal ribs " crowded together, each consisting of a
middle and a pair of lateral piecea

The shoulder-girdle is very complete, but the pieces remain
separate, or at least do not co-ossify; it consists of a T-shaped inter-
clavicle, clavicles, broad coracoids touching each other in the
middle line, and short scapulae. The existence of small separate
precoracoids is doubtful The pelvis is much reduced ; the small
ilium is quite unconnected with any vertebrae ; the small pubes
and ischia form no symphyses. The fore- and hind-limbs are
very similar to each other; the posterior are, however, much
smaller. Both are transformed into highly specialised paddles.
It is of the greatest importance, as an indication that the
Ichthyosauri are descendants of a terrestrial stock, and have been
modified into what they are owing to having taken to marine
life, that in the oldest members known, the paddle-like structure
of the limbs was less advanced than in the later specie& In
Mixosaurus of the Muschelkalk of Europe the ulna and radius
are still distinctly longer than broad, and they enclose a space

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SKELETON

481

between them. They articulate with three carpal bones, the ulnare,
intermedium, and radiale, while a small pisiform bone lies on the
outer side, between the ulnare and the outer distal carpal bone.
In Ichthyosaurus^ from the Liassic period onwards, the ulna and
radius are much shortened, broader than long, and touch each other
without any intervening space ; the pisiform element is enlarged.
Lastly in Ophthalmosaurus of the Middle Oolite (but not in con-
temporary species of Ichthyosaurus) the ulna and radius are still
more reduced, and the pisiform has moved up to the humerus, so
that the latter articulates with three bones.

Fio. 114. — A, Ventral view of the shoulder-girdle and right fore-limb of an Ichthyosaurus,
from the Lias ; B, part of the fore-limb of a Mixosaurus, from the Trias ; C, part
of the fore-limb of an Opkthulmosaurus, from the Chalk. C], e^ first and second
centrale carpi ; CI, clavicle ; Co, coracoid ; H, humerus ; /, interclavicle ; i, inter-
medium carpi ; .je>, pisiform ; R^ radius ; r, radial carpal ; <&, scapula ; U^ ulna ;
u, ulnar carpal.

Other important features of these paddles are not only the
much-increased number of phalanges (sometimes up to twenty or
more), but also the increase of digits to six or more, produced
apparently by a splitting of the third finger into two series, and
by the development of additional rows of phalanx-like bones
on the outer and inner margins of the paddle. This increase of
fingers exists, for instance, in Ichthyosaurus communis, but not in
/. tenuirostris. Owing to this peculiar development of paddles the
constituent bones are extremely numerous, and from the radius
and ulna downwards they are all closely packed, and have
assumed a polygonal, often hexagonal, shape, dwindling to more or

VOL. VIII 2 I

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482 ICHTHYOSAURIA chap.

less ilatteued nodules towards the ends of the digits. These
carpal and phalangeal bones are common objects in amateurs'
collections ; they fit together by the short angular facets, while
the two flat and broader surfaces are those of the dorsal and
ventral sides.

The Ichthyosaurs lived upon fishes and cuttlefish, as is indicated
by their dentition and the shape of the snout, and proved by
the coprolites, most of which are full of fragments of bones and
ganoid scales of fishes, and of the beaks and shells of cuttlefish ;
the larger of these true coprolites (literally " petrified dung)," in
coprolite-beds, contain also an abundance of other fossils, such as
Ammonites, Terebratulae, molluscs and fish-remains ; they are
several inches long, and many of them show on the outside ring-like
impressions, undoubtedly caused by a spiral valve of the intestinal
canal. In conformity with their absolutely aquatic life the
Ichthyosaurs were viviparous. Several well-preserved adult
specimens have been found, which contain the skeletons of one
or more rather large young within the body, in exactly the
position in which such foetal creatures would lie, namely, with
the head in the pelvic region of the mother, while the rest of the
body stretches along the vertebral column towards the chest
The suggestion that these young Ichthyosaurs have been
swallowed by their cannibal elders is too idle to require serious
refutation.

Until within a few years Ichthyosaurs were always restoi^d
with a smooth and even back, but several well-preserved
specimens have come to light in Wiirtemberg which show the
complete contour of the animals, with a long, somewhat jagged
fin on the middle of the back. Since then not a few specimens
in various collections have on closer examination revealed the
same feature, except, of course, those in which the outlines of
the fin had been chiselled away in order to " improve " the look
of the slab. The fins were undoubtedly of the "adipose" kind; —
raised folds of the skin. The latter is now known to have
been covered, at least at the bases of the dorsal fins, with hard
little scales, probably osteoderms.

Many specimens are beautifully preserved, others present a
very peculiar appearance. They look, namely, like long rolls of
clay, and nobody but an expert would suspect an Ichthyosaturus
within such a log. The explanation is simple. The dead

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ICHTHYOSAURI 483

creature was rolled about by the waves of the surf on the Liassic
muddy beach until it was wrapped in a mantle of clay and
then imbedded on the shore.

The distribution of Ichthyosaurs in time and space is wide.
The earliest are found in the Middle Trias ; in the lias they are
very common, fairly frequent in the Oolites, dying out with the
Cretaceous epoch. They have left no descendants, being far too
specialised, and their origin is quite unknown. Mixosaurus, the
oldest genus, occurred in Europe, and has also been found in the
Triassic strata of Spitsbergen. Ichthyosaurus, the chief genus, is
known from the Liassic, Oolite, and Cretaceous strata of Europe,
a famous place being Lyme Regis ; and also from the Cretaceous

Fia. 115. — Restored outlines of Ichthyosaurus quadriscissus. (After Fraos.)

strata of Queensland and New Zealand. The Jurassic of
Wyoming has yielded Baptanodon.

Order I0HTHT08AUBL

The few genera are easily recognised.

MixosauruSy Triassic, with radius and ulna still elongated,
a longitudinal space occurring between them. Both jaws with
numerous uniform teeth.

Ichthyosaurus, with much shortened radius and ulna ; botli
jaws with uniform series of teeth. Many species are known, some
with four to five, others with several additional and incom-
plete rows of fingers and toes. /. trigonodo7i of the Lias in
Wiirtemberg seems to have reached the size of 30 feet, the
vertebrae showing a diameter of 9 inches, while the skull is 6
feet long. /. covimunis and /. tenuirostris are common in the
English Lias. The long-snouted /. camjyylodon, with large, spaced
teeth, occurs in the Gault of Cambridge, Dover, and France ; and

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484 PTEROSAURIA chap.

there are many others. Ophthahfiosaums, of the Upper Oolitic and
Cretaceous formations of England, had very small vestigial teeth.
Baptanodon, of the Upper Jurassic epoch of Wyoming, was
toothless, and was one of the six-toed forms.

Sub^Class IX.— PTEROSAURIA

Mesozoic reptiles with fixed quadrate bones and with the anterior
limbs transformed into wings, the enormously elongated ulnar
finger carrying a patagium.

The skull bears a superficial resemblance to that of Birds.
It articulates with the neck by a single condyle, at nearly a right
angle. The interparietal foramen is absent, but there are five
pairs of foramina on the surface of the skull, namely, the nostrils,
orbits, supra- and infra-temporal and pre-orbital foramina. Most
of the constituent bones of the cranium fuse with each other, and
the composition of the various arches is therefore difficult to
make out with certainty. The premaxillaries are fused together, and
extend dorsally backwards between the nasals, which themselves
diverge towards the prefrontals. The nostrils are bordered
chiefly by the maxillaries, nasals, and prefrontals. The orbits
are very large, mostly shut off in front from the pre-orbital
foramina by a bridge, which is formed by descending processes
of the prefrontals and ascending processes of the jugal. Above
and behind, the orbits are bordered by the frontals, postfrontals,
and possibly the quadrato-jugals. The whole temporal region is
shortened from before backwards, but heightened dorso-ventrally,
and the whole temporal fossa is divided into a supra- and infra-
temporal portion by the junction of the postfrontal with the
squamosal, the latter joining the parietal, thus closing the supra-
temporal fossa behind. This is conspicuous only in the older
forms, e.g. Dimorphodon, but is very small in Pterodactylus, and
quite abolished in Fteranodon, The infratemporal fossa is a
narrow slit, slanting obliquely upwards and backwards, between the
quadrate and the quadrato-jugal. A foramen of this kind occurs
elsewhere only in the Ehynchocephalia. The quadrate is long,
firmly fixed, and slants so far forwards that the mandibular joint
lies on a level below the middle of the orbit. The pterygoids
articulate with strong and long processes of the basisphenoid,
touch the quadrate posteriorly, enclose an interpterygoid vacuity,

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XI SKELETON 485

and extend forwards as slender bones to the vomer, separating
the palatines. The choanae are enclosed by the vomer, palatines,
and maxillaries, and they lie in dorsal recesses above the level
of the roof of the mouth. The teeth are alveolar, pointed, of
variable size, and restricted to the jaws ; in the Pteranodonts
they are absent.

The brain is known from the natural cast of ScaphognathuSy
and shows some remarkably bird-like features, especially the
width of the hemispheres, which touch the well-developed cere-
bellum, while the optic lobes lie on the sides of the cerebellum,
with a pair of appendices, the so-called flocculi, elsewhere known
in birds only.

Tlie caudal vertebrae are still amphicoelous, while the pre-
sacral vertebrae are procoelous.
Abdominal ribs are few in number
and are very thin. The true ribs
possess capitula and tubercula ;
those of the neck are very short
and directed backwards ; in the
thoracic region they are long, and
some are attached to a broad
sternum with a keel and a median
anterior process, on the sides of
which latter articulate the cora-
coids, Precoracoids and clavicles
are absent. The scapulae are long, ^"- "^-^f^'l^^nltl^^'^""^-
sabre-shaped, and turned back as

in birds ; in Pteranodon they show the unique modification of
articulating with special processes of the neural arches of several
ankylosed thoracic vertebrae.

The hand possesses only four fingers ; the four phalanges of
the ulnar finger are very much elongated for the support of the
patagium ; the other fingers remain short and are provided with
little claws. The ilia are expanded horizontally, and are firmly
attached to from three to six vertebrae, which mostly fuse together
into a sacrum. The ventral half of the pelvis consists of a pair
of broad bones, which contain a small obturator-foramen ; they
form a ventral symphysis, and are usually fused with the ilium.
These bones represent the conjoint ischia and pubes, while the
so-called pubes, a pair of flat and club-shaped bones, are excluded

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486 PTEROSAURIA chap.

from the acetabulum. The whole arrangement resembles that of
the Crocodilian pelvis. The hind-limbs are bird-like in so far as
the fibulae are reduced to splints, and attached to the proximal
halves of the long and slender tibiae. The feet contain five
separate toes with rather long metatarsals and short claws.
Many of the bones are hollow.

The Pterosauria have no relationship with the birds, in spite
of the number of apparently striking resemblances {e.g, choanae,
pre-orbital foramina, brain, scapula, fibula, cervical vertebrae),
which are, however, coincidences, cases of convergence, in con-
formity with the aerial life. The totally different plan of the
wings is sufficient to show this. On the other hand, the real
affinities of this group of flying reptiles are unknown. They
turn up " fully fledged " in the Lower Lias, and they reach their
highest specialisation in the Upper Cretaceous epoch, with which
they have died out. In fact we do not know any forms through
which to connect them with other extinct reptiles. The skull
shows some Ehynchocephalian features; the pelvis, Crocodilian
featur^ ; and this combination points back a long way.

Order PTEB08AUBL

Sub-Order 1. PterodactyU, with alveolar teeth in the upper
and lower jaws. Imperfect remains, impressions of phalanges of
the long patagial finger, are known from the Bhaetic of WUrt-
emberg. The oldest well-known genus is Dimorphodon, Lower
Lias of Lyme Regis. D. macronj/x. — Total length between 3
and 4 feet, of which the large light skull takes up about 9
inches, and the long thin tail about 2 feet. The patiigial finger
is about 20 inches, the whole wing about 28 inches long.
Rhamphorhynchus longicavdatus of the Upper Oolite of Germany
is remarkable for the long slender teetli, which are directed
forwards and separated by wide spaces from each other. The
nine or ten cervical vertebrae are elongated. B. phyllurus of the
same geological age has left impressions of the flying membranes.
They extend from the whole length of the wing and the
sides of the trunk to the thigh as far as the knee, and from the
inside of the hind-limbs to the tail The end of the tail
carries a spatulate membrane. Allied is Ornithocheirus, with
many species in the English Wealden and Greensand. Ptero-

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XI PTEROSAURI PYTHONOMORPHA 487

ddctylus, with many species from the Upper Oolite, chiefly of
Grermany. — The tail is very short, consisting of a few vertebrae
only. The seven neck- vertebrae are so much elongated that the
neck is as long as the trunk with the tail. P. longirostris
measures about 1 foot in total length, while P. sfpectahilis is
one of the smallest, only of the size of a lark. The wings,
however, measure 10 inches from tip to tip. The largest is
P, giganteibs, with a " spread " of more than 5 feet.

Sub-Order 2. Pteranodontes. — The beak is long, pointed,
toothless, and laterally compressed ; mandibular symphysis very
long. Pteranodon longiceps. — The skull has a long parieto-supra-
occipital crest, which extends far back. The supratemporal
foramina are abolished. The pre-orbital and orbital foramina are

Fig. 117. — Khamphorhxpichus muensterif x f, as restored by Marsh. (From Geikie.)

confluent. The scapulae are attached to several thoracic
vertebrae. The skull of this gigantic species has a length of
two feet and a half, and the spread of the wings measures nearly
20 feet. This, and several much smaller species, are from the
Middle Cretaceous formation of Kansas.

Sub-Class X.— PYTHONOMORPHA.

Very long-necked and long-hodied marine Cretaceous reptiles, vnth
movable qtoadrates, siTigle lateral temporal arches and pro-
coelous vertebrae ; with paddle-shaped, pentadactyle limbs ;
and with the teeth a.nkylosed to the jaws.

The skull possesses many of the essential features of the
typical lizards. The premaxillaries, frontals, and parietaLs are
fused into unpaired bones. There is an interparietal foramen.

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488 PYTHONOMORPHA

. The nostrils are dorsal, bordered by the premaxillae, nasals, pre-
frontals, and maxillaries. The quadrato-jugal arch is incomplete,
and the orbit is posteriorly confluent with the infratemporal
fossa, but a supratemporal space is shut off by the single arch,
which is composed of the postfrontal, squamosal, and supra-
temporal. The latter is interposed between, and connects the
squamosal and quadrate with the latero-posterior branch of the
parietal. There is a space between this parieto-squamosal arcade
and the epi-otic, which is fused with the lateral wing of the
lateral occipital bone. The foramen magnum is bordered by the
two supra-occipital, lateral occipital, and the unpaired basi-
occipital bones ; the condyle is triple. The quadrate is movable,
articulating with the squamosal and laterally expanded epi-otic.
There is no bony connexion of the quadrate with the jugal, which
is restricted to its anterior half, and attached to the maxillary
and lacrymal. The quadrato-jugal is absent as a separate bone ;
it is probably fused with the anterior surface of the quadrate, as
indicated by a perforation of the quadrate, resembling in this
respect the Ehynchocephalia. The vomers are long, and separate
the elongated choanae from each other. The palatines separate
the vomers from the pterygoids, which enclose a long median
vacuity and are not connected with the quadrates. The teeth
are conical, and stand near the inner margin of the jaws upon
little prominences, with which they fuse. Some genera have
teeth upon the pterygoids also.

The vertebrae are very numerous and are mostly procoelous.
They are noteworthy for the possession of an additional anterior
and a posterior pair of articulating processes on the neural
arches, homologous with the zygosphenes and zygantra of
Snakes and Iguanidae (see p. 582). Intercentra are absent,
except in the tail. The ribs have no tubercula, and articulate
with the centra of the vertebrae to which they belong.

The pectoral arch is strong. The scapulae are short and broad ;
the coracoids, fused with the precoracoids, except for a notch,
are flat and broad, and meet ventrally ; posteriorly they articulate
upon the anterior margin of the flat sternum, to the lateral
margin of which are attached several ribs. Clavicles and inter-
clavicle seem to be absent. Abdominal ribs are likewise absent.
The pelvic girdle is feeble ; the ilia, ischia, and pubes are
loosely connected with each other, the pairs of ventral elements

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XI DOLICHOSAURI MOSASAURI 489

meeting also in the middle line. The ilia are loosely attached
to two vertebrae in the Dolichosauri ; in the Mosasauri they
have lost this connexion. Both anterior and posterior limbs
are transformed into pentadactyle paddles, vdth much shortened
and broadened bones of the arms and legs. The digits are to
a certain extent hyperphalangeal, since several of them possess
five phalanges.

The Pythonomorpha are undoubtedly allied to the Sauria,
but they are certainly hot their ancestors, since typical Autosauri
occur in the Lower Chalk ; nor are the Snakes their descendants,
in spite of many convergent resemblances. We consider them
to be the marine collateral branch of the Sauria, which rapidly
developed highly specialised, often very large forms, restricted to
the Cretaceous epoch, with a wide, cosmopolitan distribution.

Order I. DOUOHOSAUBI.

This older group is characterised by the sutural symphysial
connexion of the two mandibles and by the possession of two
sacral vertebrae. The body is snake-like. Pleurodont. Doli-
chosaurus longicollis of the Lower Chalk of Kent and Sussex ;
total length about 3 feet, with about seventeen cervical vertebrae
and pleurodont teeth. Acteosaurus of Istria ; anterior extremities
distinctly shorter than the posterior pair ; tail long. Vertebrae,
like those of Dolichosaurus, with zygosphenes. PlioplcUecarpus
of the Upper Chalk of Holland has a slender interclavicle ; the
vertebrae are without zygosphenes, but those of the cervical
region possess a downwardly directed long hypapophysial process
with a separately ossified epiphysis.

Order n. MOSASAUBL

The two halves of the lower jaw are connected by ligament
and are -therefore movable as in Snakes. There are no sacral
vertebrae, the pelvis having lost its connexion with the vertebral
column. The formation of the limbs into paddles is more pro-
nounced than in the Dolichosauri.

Mosasaurus, the chief genus, so called from Mosa, the Latin
name of the river Maas, with several species from the U})per
Cretaceous strata of the Netherlands, England, and North

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490 PYTHONOMORPIIA chap, xi

America. M, camperiy from Belgimn, with a skull about 4 feet
in length, armed with many large, curved, acrodont teeth. The
vertebral column consists of about one hundred caudal and thirty-
four precaudal vertebrae, of which seven are cervical, without
zygosphenes. The total length of the type-specimen is estimated
at 25 feet.

Plateearptts of North America and New Zealand, and various
other North American genera, also contained species of large
size.

Liodon. — Premaxilla without teeth, the others nearly smooth
instead of being ridged. With a very wide distribution in
the Chalk of Europe, North America, and New Zealand. Z.
haumuriensis of New Zealand seems to have been the giant
amongst these monstrous marine creatures ; its total length has
been computed from imperfect fragments at 100 feet.

Clidaates, of the Upper Cretaceous of North America and
Europe, although not so massive, comprises the most elongated
forms. The cervical vertebrae possess long median hypapophyses
with separate epiphyses ; most of the vertebrae are much
elongated and have well-developed zygosphenes. C. tortor had
a skull nearly two feet and a half long.

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CHAPTER XII

SAURIA AUTOSAURI OR LACKRTILIA LIZARDS

Sub-Class XL — SAURIA.

Reptiles with movahle qtuidrate hones, with a transverse, external,
cloacal opening, near the posterior lateral corners of which
open the eversihle, paired {righ t and left) copulatorg organs.

The Sauria, which comprise the Autosauri or Lacertilia in
the wider sense and the Ophidia or Snakes, are the most
recently developed groups of Eeptiles. No fossils are known
from strata earlier than those of the Cretaceous epoch. Their
origin has probably to be looked for among the Prosauria, of
which Sphenodon, cf. p. 294, is the only surviving member. The
Sauria have attained their great development within the Tertiary
period. They, both Autosauri and Ophidia, are now the two
dominant Beptilian groups, and they have, so to speak, a future
before them, being apparently still on the increase in numbers
and species, but certainly not in size.

Order I. AUTOSAURI or IJlOESlTJUA— LIZARDS

Saurians which have the right and left halves of the mandibles
connected by a stUural symphysis.

The overwhelming majority possess well -developed limbs,
movable eyelids and cutaneous scales, covered by the mostly thin
and horny epidermis. But there are many kinds of Autosauri,
especially those belonging to the degraded, burrowing families,
which have lost not only one or both pairs of limbs, but even
the limb-girdles, while the eyes have become concealed beneath
the skin, and in some cases the scales have been lost, or reduced

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492 LACERTILIA

to mere vestiges. Moreover in some of these burrowing and
limbless forms the quadrate bones have become more or less
immovable.

We divide the Autosauri into three sub-orders : — I. Geckones,
p. 502; II. Lacertae, p. 513; III, Chamaeleontes, p. 567,
with about 270, 1500, and 50 species respectively.

The Autosauri are of great interest, since they exhibit a
great, almost endless variety in shape, size, and structure in
direct adaptation to their surroundings. Most of these modifi-
cations are restricted to the external organs, or rather to those
which come into direct contact with the outer world, namely the
skin, the limbs, the tail, or the tongue. The majority of the
Autosauri are terrestrial, but there are also semi-aquatic forma
There are climbing, swiftly running, and even flying forms,
while others lead a subterranean life like earthworms. Most
of them live on animal food, varying from tiny insects and
worms to Birds and Mammals, while others live upon vegetable
diet. According to this diet, the teeth and the whole digestive
tract are modified. The intestine is relatively short in the
carnivorous, long in the herbivorous species. But swiftness,
the apparatus necessary for climbing, running, and digging, the
mechanism of the tongue, the armament and the muscles of the
jaws (hence modifications of the cranial arches, etc.), stand also
in correlation with the kind of food and with the way in which
it has to be procured.

A very interesting study of the influence of the climate and
the nature of the country upon Reptiles has been made by
Boettger^ with especial reference to the Transcaspian desert-
region. The winter is there short, but very severe, and there is a
considerable amount of snowfall, while the summer is intolerably
hot. The spring arrives suddenly. Lilies and tulips, which
have been asleep for nine or ten months, sprout towards the
end of February, and a carpet of flowers covers the ground
for a short time. Then everything shrivels up during the
rainless and fierce heat of the summer, and the autumnal storms
of dust and sand kill off the last remnants of vegetation. There
are no trees, and even prickly shrubs are rare. Instead of broad
leaves the plants have grass-like blades or needles. The little
shrubs do not form coherent patches, but they are scattered

1 ZooL Oart, 1889, p. 1.

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XII ADAPTATION TO ENVIRONMENT 493

about, and around the roots of each shrub the wind accumulates
little mounds of sand and dust, a place of retreat for rodents,
lizards, snakes, and even for the female tortoises. G. Eadde's
" law of the steppe " is in full force ; — there is little change of
forms in a wide district, but all these forms are peculiar, and
they congregate socially in great numbers. Most characteristic
are those kinds of Geckos which, like Teratoscinctts, cf. p. 507,
have become inhabitants of sand instead of climbers of rocks and
trees; various kinds of Phrynocephalus, cf. p. 521, and Varanus
griseus ; the four desert-species of Lacertidae are brownish-grey
or sandy yellow, with conspicuous stripes or spots. Of snakes
are to be mentioned Eryx jacvltcs, digging in the sand, and
about ten other non-poisonous snakes. Tropidonotus is, of course,
restricted to permanently watery places, where they can get
frogs and fishes. Of poisonous snakes there is the Cobra and
Ilchis arenicola. Of Amphibia only Bufo viridis and Bana
esculenta var. ridibunda exist in suitable places, but there are
neither Tree-frogs nor Newts.

Characteristic features of these inhabitants of the desert are
the following : —

1. Velocity. The Lizards are slender. The Sand -snake,
Tephrometopon, is whip-like; even the Cobra has a relatively
narrower and longer tail than the Indian specimens, although
the number of the vertebrae and of the scales is the same. All
the desert-snakes are remarkable for the great number of their
ventral shields, two hundred and more.

2. Hard, scaly covering, for instance in Agama, Echis, Gymno-
dactylus, Teratoscincus ; the latter with its fish-like scales is
exceptional among Geckos, resembling the likewise deserticolous
Geckolepis and Homopholis of Africa.

3. Capacity for digging in the sand in order to escape
great cold, or burning heat. All the Lizards and the Tortoise,
Testudo horsfieldi, have strong claws. The snakes Typhlops and
Eryx dig with their specially modified snouts, and their tails are
very short and blunt. The Sand-viper, Echis, has the scales of
the back arranged in very oblique rows, so that it can heap
sand upon its body by wriggling, shaking, and up-and-down
motions of the body. The Agamoid Fhrynocephalus does this
by means of lateral folds of the skin.

4. Arrangements for running on sand. The lizard Eremias

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494 LACERTILIA

has very large crural shields ; Scapteira has the digits broadened
out into shovels ; others, e.g. Phrynocephcdus and Teratoscincus,
have long lateral fringes on the digits, a very rare arrangement
among Geckos, occurring elsewhere among them only in Ptenopus
and StenodactyluSy which are likewise inhabitants of the desert.

5. Protection against the everlasting, ubiquitous sand. In
the digging species the nostrils are directed upwards instead of
forwards ; in most of the snakes they are protected by compli-
cated valves, or they are reduced to small pin-holes. The eyes
of Typhlops are overhung by the head-shields. In Agamu and
Ph^ynocephalus the margins of the lids are broadened into plates
and are furnished with peculiar scales. In TeratosciJicus the
upper lid is enlarged. . The lizard Mabuia has the lower lid much
enlarged, with a transparent window in it, so that the eye can
be closed without impeding sight, an arrangement carried to the
extreme in Ablepharus, cf. p. 560. The ear-opening is either
small, or protected by fringes of scales, or it is abolished, e.g,
in Phrynocephalus,

6. Coloration. Pure green is quite absent, even in Bufo
viridis and in Rayia esculenta, since there is no green in that
country, at least not of long duration. White, with grey and
black spots, occurs only in the nocturnal Geckos. Yellow,
brownish, reddish colours are common, in adaptation to the sand.
The advantages of the carmine -red, and of the blue spots of
Phrynocephalus, and the yellow or bright red imder surface of its
tail, are unknown. Striation is of frequent occurrence among
the lizards and snakes, probably in adaptation to the dry grass
heaped up around the scattered shrubs.

Concerning the various organic systems of the Autosauri only
some of the more important features may here be mentioned.

Skeleton. — The vertebrae are precocious, with the exception
of most of the G^ckones, in which they are amphicoeloua So-
called intercentra, in the shape of unpaired nodules or wedges,
persist between most of the cervical vertebrae. In the tail
these wedges, the remnants of the basiventralia, are generally
present, frequently in the shape of chevron -bones Sometimes
they fuse with the centra of the vertebrae; occasionally the
axial or central portion of these basiventrals persists as a sort of
fibrous disc, which may calcify separately, and is interposed
between the caudal end of the centrum and the articulating

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XII SKELETON 495

knob. The caudal vertebrae of the Geckones and of most
Lacertae are liable to break across, like those of Sphenodon.
They are enabled to do this owing to a transverse split, which
makes its appearance with the ossification of the vertebral
bodies and extends later into and across the neural arch and
the various lateral processes. The split is ultimately referable
to a transverse septum of cartilage, wrongly called chordal
cartilage, which develops in the shell of the body of the vertebra,
destroys the chorda, and extends peripherally. The cells of this
septum retain throughout life their juvenile quasi-embryonic
character. When the tail is broken off — and this always happens
at such a septum — the cells of the remaining half reproduce a
new tail The latter is, however, in reality a sham tail, since
neither new centra nor arches, but only a non-segmented rod or
tube of fibro-cartilage is produced by this process of regeneration.
Reproduction of centra is precluded by the previous normal
reduction of the chorda, around which alone proper bony centra
could be formed. The regenerated tail is, however, invested
with new muscles, and with skin, but the scales often differ
considerably from those of the normal organ. Boulenger ^ has
found that the new or aberrant scaling is in some cases a
reversion to an ancestral form. This is, for instance, the case in
Psevdopus, and in the Tejoid genus Gymnophthalmus ; to a
certain extent also in Geckos and Skinks. On the other hand,
Lacertidae, Gerrhosauridae, and also Anguidae reproduce a
caudal scaling true to their type. Injured or broken-off tails
are often reproduced double, or even trifid; sometimes an
additional little tail grows out from an injured spot, anywhere
on the side of the old remaining but mended taiL

The ribs of the trunk articulate by their capitula only,
while the reduced tubercula are attached to their vertebrae by
ligaments. In the tail the capitular portion is much reduced,
while the tuberculum is much stronger and lies behind, no
longer above, the capitulum, fusing sometimes directly with the
centrum. The ribs of the poststernal region of Geckos and
Chameleons are very long, and meet each other in the middle
line, forming thin cartilaginous hoops.

The limbs are of the typical pentadactyloid type. The distal
tarsalia are often fused with the metatarsals, so that the chief

1 P.Z.S. 1888, p. 351, and 1891, p. 466.

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49^ . LACERTILIA

bending of the foot is effected by truly intertarsal joints. The
greatest modification occurs in the foot of the Chameleons, in
which the proximal tarsalia are reduced in nimiber, and form a
globe for the articulation with the tibia and fibula

The shoulder-girdle and sternum much resemble that of Spheno-
don in their completeness. The coracoids articulate with the
sternum ; the precoracoids and the basal parts of the scapulae
often send out several processes towards those of the other side,
so that several fenestrae are formed. The clavicles are complete,
but are absent in the Chameleons. The interclavicle is mostly
T-shaped. A presternum is absent, but the sternum proper is
well developed, often forming a rhomboid plate, usually carti-
laginous, often diverging backwards into xiphisternal processes.

The pelvis is attached to two vertebrae by means of several
ribs. The ischium and pubis form symphyses. The pubis
carries a well-developed lateral process, and the obturator-nerve
pierces the shaft of the pubis. Epipubic and hypo-ischial
cartilages are of frequent occurrence.

The hyoid apparatus consists of a median, styliform rod,
which extends forwards into the tongue ; it is often bifid
behind. The unpaired piece carries two pairs of horns. The
posterior of these, the first pair of branchial arches, extends
backwards along the gullet, and is very long if the tongue is very
slender and protractile. The anterior pair, the hyoid arches,
consists of two pieces on either side, one short and directed
forwards, the other long, connected with the former at a sharp
angle and continued upwards to the sides of the skull, often in
direct continuity with the columellar chain of the ear.

The modifications of the skull concern chiefly the composition
of the temporal arches, see Figs. 55, M, N, O, p. 281. The
quadrate bone is movable, but it has become fixed in various
degraded families, where the skull shows a great reduction
and concentration ; the postorbital and temporal arches, the
interorbital septum, and with it the columellae cranii are lost.
The columella cranii of the Chameleons, which is generally stated
to be absent, is really present, although in a much reduced state,
and is partly imbedded in the interorbital septum. The occipital
condyle has become bifid in Amphisbaenidae.

Burrowing and living in sand are often correlated with
partial or complete reduction or loss of the limbs and their

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XII SKELETON — SKIN 497

girdles. This loss of limbs is as a rule correlated with an
elongation of the trunk, not always at the expense of the tail,
which in such cases is much shortened. The vestiges of the
hind-limbs come to lie as near the vent as possible. This
reduction of the limbs occurred in several families which are
not directly related to each other. Moreover, it does not occur
in all the members of the family, not always in those of the
same genus, and there is a considerable amount of individual
variation. In most cases of reduction the fore-limbs disappear
before, or are smaller than, the hind-limbs. In the Amphis-
baenidae (c£ Chirotes, p. 566), and in the Tejidae the reverse
takes place. In extreme cases the reduction is so complete that
even the pectoral girdle has disappeared, leaving scarcely any
trace, e,g, in Dihamus, p. 564.

The skin is normally covered with scales, which are formed by
the cutis and have a horny epidermal coating. The latter, thin
and transparent, is shed periodically, peeling off in flakes, ex-
cept in Anguis and perhaps other snake-shaped creatures, which
shed the skin in one piece. In the Amphisbsienidae the scales
have practically disappeared. When well developed the scales
are prominent, and imbricate or overlap with their free posterior
edges ; but in many cases the scales are not " scale-like " at all,
only like little tubercles, which give the skin a granular
appearance. Frequently, for instance in the Scincidae and
Anguidae, all the scales contain " osteoderms," or ossified portions
of the cutis, and encase the whole body and tail. In other
families, e,g, Lacertidae, such osteoderms are restricted to the
scales or shields on the head, where they come into contact and
fuse with the imderlying cranial bones, and moreover roof in the'
supratemporal fossa.

The skin of the Autosauri is entirely devoid of glands. The
femoral and pre-anal pores of many families, occurring especially
in the males, are probably not glands. They are arranged in
rows on the under surface of the thighs and in front of the anal
opening. Each of these organs perforates a scale and leads into
a tubular invagination, which is lined with epidermal cells, the
proliferation of which produces a horny yellowish d(5bris, and this
fills the tube and appears above the surface in the shape of a
little cone. The use of this " excretion " is unknown ; it is
possibly hedonic.

VOL. VIII 2 K

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498 LACERTILIA chap.

Most Autosauri are capable of changing colour. In most of
them this faculty is restricted to the assumption of paler or darker
tints owing to the shifting of the colouring matter contained in
the chromatophores. In others new, often vivid colours are the
result. The mechanism is described in detail in the Chameleon
on pp. 570 and 574.

Pigment is deposited either directly in the upper strata of
the cutis, just below the Malpighian layer, or it is contained in
chromatophores. The latter are imbedded in the deeper layers
of the cutis, and send out movable contractile processes, in
which their pigmented protoplasm is conveyed towards or away
from the surface. The only colours available are black, red, yeUow,
and white, with their combinations of grey and brown. The
white pigment consists of guanin- salts. Blue and green are
structural colours, not due to pigment. The same can no longer
be said of the Ophidia, since Boulenger has observed accidentally
that green Tree-snakes (e.g. Dryophis) give the alcohol in which
they are kept the colour of green Chartreuse.

Digestive organs. — The tongue is very variably developed,
and affords good taxonomic characters. It is always furnished
with many tactile, or with gustatory, corpuscles. When the
tongue is very long and narrow it is generally forked, and in
these cases, for instance in the Varanidae, is almost entirely used
as a sensory organ. In others, especially where it is broad, it
assists in catching the food, and in the Chameleons it has
attained a most elaborate development (see p. 569).

Salivary glands are restricted to labial glands. In Ifeloderma
those of the lower jaw are transformed into • poison-glands, an
analogy to what prevails in the poisonous snakes. The intestinal
canal is longest in the herbivorous forms ; the rectum sometimes
possesses a short blind sac or caecum.

The cloaca of the Sauria is somewhat modified ; instead of
the Coprodaeum, Urodaeum, and Proctodaeimi forming three
successive chambers, the urodaeum is practically reduced to its
dorsal half, forming a dorsal recess between the two other
chambers. The Coprodaeum is constricted into several successive
chambers, and is always well shut off from the urodaeum by a
strong sphincter. The urodaeum receives the urinary excretions,
which are mostly chalky white and are rather consistent instead of
being fluid. The right and left oviducts also open into it. The vasa

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XII DIGESTIVE ORGANS — EGGS — LUNGS 499

deferentia open into the dorso-lateral portions of the walls of the
urodaeum, but the sperma is conducted by folds of the lining of
this chamber towards the bases of the copulatory organs, which,
although arising from the lateral and posterior comers of the
cloaca, where uro- and procto-daeum meet, are stowed away outside
the cloaca. These organs are always paired. The proctodaeum
or outermost cloacal chamber is shallow. Its inner opening is
round and is furnished with a sphincter, but it is surrounded and
covered by lips of the outer skin, which form a transverse slit.
This is due to the peculiar arrangement of the copulatory
organs.

Each organ consists of a tube of erectile tissue, and can be
everted like the finger of a glove. To the apex of the tube is
attached a long retractor muscle, which arises from the ventro-
lateral surfaces of the caudal vertebrae and extends a considerable
distance back. When at rest and withdrawn the organs form slight
conical, longitudinal swellings on either side of the root of the
tail, an external feature by which male specimens can generally
be distinguished. Only one organ is inserted at one time.

The majority of Autosauri lay eggs, surrounded by a white
or yellowish shell, which is either hard, for instance in Geckos,
in Lacerta viridis and L, agilis, or parchment - like, e.g. in
Chameleons and in L. vivipara. Eggs with a thin and soft
shell sometimes exhibit the paradoxical feature of increasing in size
after they have been laid. This is explained by the growth of
the embryo, which stretches the sheU and does not merely live
upon the white and yellow contents of the egg itself, but also
takes in air and moisture. Many Lizards do not lay their eggs
until they contain ripe embryos, which burst the shell shortly
after deposition. Some, for instance Lacerta vivipara, Angvis
fragiliSy and Chamaeleo pumilus, are practically viviparous. 1 he
embryos, especially those which are enclosed in hard - shelled
eggs, are provided with a sharp, calcareous " egg-tooth " on the
top of the snout.

The longs are thin-walled sacs, sometimes provided with
lateral ex-sacculations, and these reach their greatest development
in the Chameleons. The breathing is effected by the motion of
the ribs. Inflatable sacs on the throat, or on the sides of the
neck, for ornamental or sexual purposes, occur in various families.
The lungs of much-elongated, snake-shaped Lizards are generally

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500 LACERTILIA

asymmetrical ; the right being reduced in Amphisbaenidae ; the
left in other cases.

Several Autosauri, for instance the Geckos, Psammodromtis,
and various other Lacertidae have a weak voice.

The Fat-bodies are mysterious organs which are situated
beneath the skin, and extend from the inguinal region forwards
along the ventral sides of the belly. They are often of consider-
able dimensions ; largest in the spring, in both sexes, at the
time of propagation. Their colour is greyish-white or yellow,
owing to the great accumulation of fat in the meshes of the
connective tissue which composes the frame-work of these organs.
An artery enters them, breaks up into capillaries, and these com-
bine to form an efferent vein. After the time of propagation
these organs are reduced to grey or reddish flaps, consisting mainly
of very vascular connective tissue. G. W. Butler ^ has written a
long paper on their morphology. The same author ^ has investi-
gated the " sub-divisions of the body-cavity in Lizards, Crocodiles,
and Birds," with reference to peritoneal diaphragmatic structures.

The geographical distribution of the Aatosaari teaches
few, but important lessons. We have to restrict ourselves to
the principal families, leaving out those which are small and
have a limited distribution; also those which, like the few
Anelytropidae in Africa and in Mexico, are not natural groups.

The Geckones, which are probably the oldest of modern
Autosauri, are practically cosmopolitan, being absent only in the
cold and in the cooler temperate regions. They are common
even in Oceanic Islands, for instance in New Zealand and in the
Sandwich Islands. Although not at all aquatic, they are
particularly fit to be transported accidentally on or in the trunks
of floating trees, to which they cling firmly, and they can exist
without food for months. I once received a little South
American Gecko in perfect health from a grocer, who found it in
a well-closed wooden box containing canned meat, two months
after delivery of the box in Cambridge.

The Scincidae, likewise an old family, are equally cosmopolitan,
but although many exist in the islands of the Pacific they do
not occur in New Zealand. Many of the genera have a very-
wide distribution ; for instance, Lygosoma, with its one hundred
and sixty or more species, occurs in the Australian and Palaeo-

1 P.Z.S. 1889, p. 602. « KZ.S. 1889, p. 452.

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XII GEOGRAPHICAL DISTRIBUTION 501

tropical regions, and also in North and Central America, not ex-
tending, however, into South America. Mabuia, with more than
sixty species, occurs in the Palaeotropical and the Neotropical
i-egions. Whether these and other widely-distributed genera are
all natural is another question.

The Agamidae, Varanidae, Lacertidae, and the Chamaeleontes
are restricted to the Old "World. The Agamidae and Varanidae
have the widest distribution, occurring in the whole of the Old
World with the notable exception of Madagascar and New
Zealand. The Lacertidae are Palaearctic and Palaeotropical, being
however absent in Madagascar, and, broadly speaking, not extend-
ing eastwards beyond Wallace's line. It is a most suggestive
fact that most of those families of Reptiles, and even of other
Vertebrates which have a wide distribution and are apparently
debarred from transgressing Wallace's line, are also absent from
Madagascar.

The Chameleons are essentially African, with their centre of
greatest abimdance and development in Madagascar, only one or
two species occurring in Socotra, Southern Arabia, and in Ceylon
and Southern India. If they existed, or could be shown to
have existed, on the various islands in the Indian Ocean, for
instance in Mauritius and the Seychelles, the Chameleons would
be an excellent illustration of the former existence of a direct
land-connexion between Southern India and Southern Africa.

The Iguanidae are essentially American, with the remarkable
exceptions of Chalarodon and Hoplums in Madagascar, and
Brachyloph'us in the Fiji and Friendly Islands. This peculiar
distribution finds some analogies in that of Dendrobatinae (p.
272), certain Boinae (p. 601), and Centetes and Solcnodon among
Insectivora. An Iguana (/. europaea) has, however, been described
from the Eocene of France and England. The supposed relation-
ship of the Iguanidae with the Agamidae makes the problem
only more puzzling, since Agamidae are absent in Madagascar.
If we have recourse to the Zonuridae, which are confined to Africa
and Madagascar, and are supposed to be intermediate between
Anguidae and Iguanidae, then we may have ultimately to con-
clude that the Malagasy Iguanoid genera and the American
Iguanidae are a case of convergent evolution.

The Amphisbaenidae are distributed over America, including
the West Indies, Africa exclusive of Madagascar, and the

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502 LACERTILIA

Mediterranean countriea This is very puzzling, considering that
these subterranean, helpless creatures positively cannot travel.
Boulenger regards them " as a degraded type of the Tejidae, with
which they are to some extent connected by Chalcides and its
allies," i.e. genera with reduced limbs, cf. p. 562.

However, this supposed relationship with a strictly American
family does not explain the occurrence of Amphisbaenidae in
Africa. Either they are not a natural group, or they had,
as already degraded, limbless creatures, a much wider range ; and
this would imply their being a very old family, perhaps as old as
we suppose the Coecilians to be.

Anguidae occur in North and South America, in Europe and
the Mediterranean parts of North Africa, and in Trans-
Gangetic India. Their older relations, the Zonuridae, inliabit
Africa and Madagascar.

Madagascar is consequently devoid of Agamid«ie, Varanidae,
Lacertidae, Anguidae, and Amphisbaenidae, while it possesses,
besides the cosmopolitan Scincidae and Geckones, only Chameleons,
G^rrhosauridae, and Zonuridae, — all three essentially African
families, — and a few Iguanidae. This means that the Autosaurian
fauna of Madagascar is intimately related to that of Africa, and
that it possesses only old families so far as Sauria are concerned.
But since this great island was separated from its continent not
earlier than in Mid-Tertiary times, it follows that most of these
" old " families are comparatively recent.

Australia possesses only Agamidae and Varanidae besides the
ubiquitous Geckos and Skinks. Besides the latter two families it
has nothing in common either with Madagascar (an analogy with
the Anura) or with America. The Autosauri consequently do not
support the idea of a Notogaea, cf. p. 74. This again indicates
the comparatively recent age of Autosaurian families. The
marked difference which exists between the Old and the New
"World points to the same conclusion. On the other 'hand, the
Autosaiud support the idea that the Palaeotropical region is but
the tropical and therefore richer continuation of the now
impoverished Palaearctic sub-region.

Sab -Order 1. Geckones. — The typical Geckos are char-
acterised as followa Four-footed Autosauri with amphicoelous
vertebrae; skull without bony temporal arches; clavicles dilated
arul with a perforation near the ventral end; parietal hones

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XII GECKONES 50 3

separate; eyes {with few exceptions) without movable lids;
plevrodont ; tongue fleshy and hroad, slightly nicked anteriorly,
and capable of protrusion.

This definition does not apply to a few forms. In the
EvMepharinae the vertebrae have advanced to the procoelous
condition, and the parietak are fused together, while the eyes are
provided with typical, movable lids. In the Uroplatinae the
clavicles are not dilated, and the nasals are fused into one bone.
The Geckos seem to be not only a very independent but also a
very old branch of Saurians. Although fossil representatives are
unknown, the resemblance of their vertebrae to those of the
Palaeozoic Microsauri is at least remarkable. They are now
practically cosmopolitan within the wanner zones, being found in

- ^

I CECKONIOAE.

FiQ. 118. — Map showing the distribution of Geckonidae.

abundance in all intertropical countries and islands, even in New
Zealand. About two hundred and seventy species are known,
which have been subdivided into about fifty genera. The generic
differences are trivial with few exceptions, and refer mostly to
the structure of the digits.

The more important features of the vertebral column are the
absence of axial joints and the persistence and life-long growth
of the chorda dorsalis. Each vertebral centrum consists of a
cartilaginous tube, more or less calcified or ossified, with a narrow
waist and a cartilaginous septum in the middle. In the tail this
septum, which is only slightly invaded by ossification, coincides
exactly with the line of transverse division of the vertebrae into
an anterior and a posterior half. This is the level where the
tail breaks off and whence it is renewed. Between every two
successive centra lies an intercentrmn, broadest ventrally, crescent-

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504 LACERTILIA chap.

or wedge-shaped. Dorsally it is continued as fibro-cartilage, and
the whole ring acts as an articular pad instead of the joint.
Che^^^on-bones are common in the tail.

The ribs are bifurcated, but the tubercular portion is frequently
reduced. The post-thoracic ribs are usually very slender, and
so long that they meet each other in the middle line, in
this case bearing an extraordinary resemblance to the so-called
" abdominal ribs " of other reptiles.

The bony frame of the skull is slender. There is a complete
absence of bony arches spanning over the temporal fossae, or
bordering the orbit, which is open posteriorly. The upper jaw,
owing to the slender and flexible nature of the respective l)ones,
is movable upon the rest of the skull ; in this respect not unlike
the upper jaw of a duck. The dentition is pleurodont and the teeth
are minute. The eyes of the typical Geckos are peculiar. They are
covered with an absolutely transparent skin, shaped like a
watch-glass, beneath which the eye moves freely, while the true
upper and lower lids are reduced to tiny folds. The covering
" watch-glass " is probably a modification of the nictitating mem-
brane. In the Eublepharinae, however, and in the few sx)ecies
of the Geckonine genera Aelurosaurus of Borneo and Australia,
and Ptenopus of South Africa, the upper and lower lids are
present and movable. The pupil contracts mostly into a vertical
slit, except in the few diurnal kinds, e.g. Phehwma, of the islands
in the Indian Ocean, and the African Lygodactylus.

Another peculiarity of at least many Geckos is the extra-
ordinary development of the endo-lymphatic sacs of the ear,
which, being filled with the chalk-like otoconia or otolithic
crystals, perforate the skull, and are stowed away in the shape of
a pair of large bags behind the ears, or on the sides of the neck.

The skin exhibits considerable variety. It is mostly soft
above, with little granular tubercles, sometimes containing small
dermal ossifications or calcifications. The latter are most de-
veloped on the head, where they occasionally fuse with the
underlying bones. A few species of Tarentola possess supra-
orbital bones, independent remnants of such osteoderms. The
ventral surface is generally covered with small imbricating
scales, but in some genera, e.g. JIomojihoHs, such scales occur also
on the dorsal surface, reaching their highest development in
Teratoscincics (p. 507). In a few forms, notably in Ptychozoon

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GECKONES 505

(p. 512), the skin of the sides of the Ixxiy and tail is produced
into a series of lobes and flaps, the object of which seems to
assist adhesion. Many, perhaps the majority of Geckos, have ad-
hesive digits, by means of which some kinds are enabled to climb
absolutely smooth and vertical surfaces, for instance a window-
pane ; or, what is more startling, they run along the smooth,
white-washed ceiling, back downwards. The apparatus is com-
plicated in its minute detail, but is very simple in principle. The
adhesion is effected neither by sticky matter, nor in the way
described in the Anura (p. 187), but by
small and numerous vacua. The under
surface of each digit is furnished with
many transverse lamellae. The pressing
down of the foot upon a smooth surface
causes the lamellae to spread asunder and
to drive out the air; partial retraction ^W

lets them return to their original position ^'^^ "^^ > J

by virtue of their elasticity; and little ^ \ ^ i /

vacua are produced. Each lamella is ^,^y^iK >■ v ,
further beset with tiny hair-like ex- V.. . ~^ 1;""=^

crescences, which secure adpression to ^=4;,

even the slightest irregularity of surfeuie
and at the same time enhance the elas-

. ■ ., « i.1 1 mi J. !• Fig. 119. — Ptychozomi hoDuUo-

ticity of the pads. The arrangement of c^p/utium. A, Veutrai view
the lamellae and pads differs much in of the right hand. x2.

^, . -i-i • ^ ^, B, Side view of a finger to

the various genera. For instance, the show the peculiar arrange-
lamellae are either broad and entire, or 7^?"*^ ^^ t^® claw -bearing
they are divided into two parallel rows,

with or without lateral hairy fringes; or the under surface
of the digits is granular, but strongly fringed ; or the lamellae
are restricted to the dilated tips of the digits, etc. The fingers
and toes are mostly furnished with sharp, curved claws, and
these are in many cases retractile between some of the lamellae,
or into a special sheath. Those Geckos which live on sandy,
barren ground are as a rule devoid of adhesive pads, the digits
being narrow. The typically padded, adhesive digits cause a
peculiar sensation when a Gecko hangs on to one's finger, and
this feeling has perhaps given rise to the erroneous notion of
stickiness.

The tail exhibits many kinds of shape and size. Mostly

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S06 LACERTILIA GECKONES chap.

cylindrical and tapering to a point, it is leaf-like in Gymno-
ditctyltLs jplaturus of Australia ; provided with many lobes, and
used as a parachute in the Malay Ptychozoon. In Ncphmrus
asper of Eastern Australia the tail is quite short, much shorter
than the limbs, much swollen at the base, and very thin towards the
end, which carries a round knob. The tail of all Geckos is very
brittle and can be quickly regenerated, except the long rat-like tail
of the Persian Agamura. In many other desert-forms the tail is
long, slender, and laterally compressed, acting in such cases like
that of desert-forms among the Lizards.

Many Gecko§ have a voice, mostly rather feeble, and sounding
like a soft " click " or " chick " produced by our tongue. Repe-
tition of this sound resembles in some species the word " gecko."
They lay eggs, rather globular, or but slightly oval, hard-shelled,
and wliite, mostly two in number. Navltimbs eleyans of New
Zealand is said to be viviparous. The males are generally larger
than the females, and they are further distinguished by the
possession of femoral or pre-anal pores.

All Geckos feed upon animals, chiefly upon insects, 'but the
larger forms take anything they can master. With few excep-
tions they are nocturnal, which, however, does not prevent them
from occasionally baking themselves in the sun. They are
capable of changing colour, but since their ground-colour is
almost universally grey, yellow, or brown, the range of the colour-
changes is restricted to the adoption of darker or lighter hues.
The skin is shed in flakes and eaten.

Geckos are absolutely harmless; they cannot even inflict
painful bites. However, in many countries they are feared as
much or even more than the most poisonous snakes. In the
south of Spain and Portugal, for instance, where Geckos are
plentiful in and outside the houses, and are consequently objects
of daily observation, the " osga " is considered a dreadfully
poisonous creature. They become very tame, or rather confiding
in their regular habits, provided they are not molested. If
caught — and they have many enemies among other lizards and
snakes — the only safety of these defenceless and mostly small
creatures lies in their tail, which, being extremely brittle, is left
in the claws or jaws of the pursuer. The remaining stump soon
produces a new tail, in shape and size like the old one, but with
a different and simpler scaling. I knew of several specimens of

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GECKONIDAE 507

the Portuguese Platydcxtylus fdcetanvs, which, having lost their
tails in the act of being caught, were kept in a box for six weeks
without food. On their arrival in England they had each
grown a new stump nearly half an inch long !

Fam. Oeckonidae. Sab-Fam. 1. Oeckoninae. — Vertebrae
amphicoelous ; parietal bones separate ; clavicles dilated and
perforated. Hereto belong the overwhelming majority of Geckos,
only a few of which can be mentioned.

Teratoscincus scincus. — This most peculiar creature, about six
inches in length, inhabits the steppes of Turkestan and neigh-
bouring desert-r^ons of Persia. It is a thorough desert-form.
The digits are devoid of adhesive lamellae, but are granular
inferiorly and strongly fringed laterally, an arrangement which is
rare among Geckos, practically restricted to it with Ftenopvs and
Stenodactylus, which are likewise deserticolous. This is a beautiful
illustration of adaptation to the surroimdings. A Gecko, instead
of climbing rocks and trees, has lost the climbing apparatus, or
has transformed parts of it for running upon loose sand. The body
is covered with imbricating, rather large and smooth scales.
The tail is round at the base, compressed in its posterior half,
covered below and on the sides with scales like those of the
body, but on the upper side with a series of large, transverse,
nail-Hke plates. By rubbing these plates upon each other, this
Gecko produces a shrill, cricket-like noise, sitting at night in
front of his house, perhaps in order to attract grasshoppers. The
noise is made by both sexes.

Ptenopus, a Gecko of Damara Land, likewise adapted to desert-
life, produces a similar chirping noise by its throat.

Phyllodactylus is a genus of world-wide distribution, occurring
in tropical America, Africa, Madagascar, and Australia, extending
to the Norfolk Islands and to Lord Howe's Island. One species,
Ph, europaeus, occurs on the islands in the Western Mediter-
ranean. The digits are furnished with transverse lamellae, the
greater number of which are broken up into small scales forming
three longitudinal series. The ends of the digits are dilated,
with two large plates inferiorly, separated by a longitudinal
groove into which the claw is retracted. The upper parts of the
body are covered with juxtaposed scales intermixed with larger
tubercles. The abdominal scales are small and imbricating.
The cylindrical, tapering tail is slightly prehensile, covered with

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5o8

LACERTILIA — GECKONES

small scales arranged in verticils. This species is devoid of
femoral or anal pores. General colour above grey-brown, with
darker and lighter markings ; a dark streak on the side of the
head, passing through the eye. Under parts whitish. Total
lengtli up to 3 inches. The eggs are almost round, measuring
8-7 by 7 mm.

Hemidactylus, likewise a widely distributed genus, with many
species. The digits are dilated, inferiorly with two rows of

Fig. 120. — Hemidactylus turcicus (left), and Tarentola mauritanica (right).

lamellae ; the clawed joints are slender, bent at an angle, and
rising from within the extremity of the dilated portion. H.
turcicus, between 3* and 4 inches long. — The upper parts of
the body are covered with minute granules, mixed with larger
tubercles. The abdominal scales are small and slightly imbri-
cating. The male has several pre-anal pores. The tail is
covered above with minute scales and tubercles, below with a
series of large transversely dilated plates. The general colour is
white below, brown above, with darker spots, and with white
specks on many of the tubercles. This species extends from

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XII GECKONIDAE 509

Southern Portugal and Spain to Karachi. Like Phyllodactylus
and various other kinds of Geckos, the body is semi-transparent ;
80 much so indeed that the white eggs shimmer through the body-
in certain lights.

Tarentola mauritanica s. Platydactylus fatetanuB, — The digits
are strongly dilated, with undivided lamellae below, and a flat,
nail-like scute on their upper surface near the tip. Only the
third and fourth digits are clawed. Femoral or pre-anal pores
are absent. The upper parts are covered with scales and
granules, and bear several longitudinal rows of strongly keeled,
large tubercles ; the under parts have hexagonal scales. General
colour above greyish-brown, with darker or lighter markings;
with a dark streak through the eye. Total length of large males
about 6 inches. This species is one of the commonest Geckos
in the Southern Mediterranean countries. In Portugal it extends
northwards to the Douro. It has been introduced by ships into
the ports of Cette, Toulon, and Marseilles. It is easily kept in
captivity, like most Geckos indeed, provided they axe supplied
with a variety of insect-food, water in the shape of drops, and
suitable places to hide in. A female, which I had received from
Algiers in a little tin box, with a lump of meat (presumably its
food!), laid two eggs six weeks after its arrival. This was
towards the end of April. Towards the end of June in the
same year it again laid two eggs, measuring 13 x 10 mm.
Another specimen laid in June in two successive years. These
and other Geckos live very well in a greenhouse, or in a large
glass cage. They change colour most adaptively. Tliey hunt
preferably at night for insects, which are stalked and then
suddenly rushed at. Drops of water are tak^n by a lapping
motion of the tongue. For sleeping-places they selected bits of
hollow bamboo, but these had to be vacated when some tree-
frogs crept into them for the daytime, and the Geckos took to
some curved pieces of bark, on the under side of which they
slept, with their backs downwards. This is, by the way, a
favourite position of rest of most Geckos. But Stenodactylu^
guttatits of Egypt lies flat on its belly, tucks the fore-feet under
and inwards like a cat, rests the head upon them, and stretches
the hind-limbs out backwards. The little Geckos are rather
intelligent. They take no notice of a finger put against the
other side of the glass to which they happen to cling; but

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5IO LACERTIUA — GECKONES chap.

when the hand is put inside their cage and approaches them too
near, they dart off suddenly. When driven into a comer they
wriggle and wag their tails, or even raise the latter, perhaps as
an invitation to grasp it, in which case it would of course
break off. When caught, they emit feeble soimds, and attempt
to bite with the mouth widely open. During the moulting,
which takes place at least twice a year, in the spring and in the
autumn, the skin peels off in flakes ; if, as happens sometimes,
the skin upon the lamellae is not stripped off neatly, these refuse
to act, and the creatufe cannot climb until all the old skin has
been rubbed off.

In their native haunts they are very regular in their habits.
Favourite resorts of theirs are old olive trees or oak trees, the
rough and cracked bark of which affords excellent places for
hiding in. Hollow trees are of course preferred. Not a single
specimen is seen during the early hours of the morning or in the
forenoon ; but when the sun has become broiling hot, and our
own shadow passes over the stem of a tree, we become aware of
flitting little shadows which jerk over its surface. These are
Geckos which had been basking, motionless; very dark grey,
almost blackish, just like the colour of the grey bark upon which
the last wet season's moss has been scorched to a black cinder. It
is difficult to espy a Gecko whilst it is glued on to such a tree.
Only the little beady eyes betray it, watching you carefully.
Nothing appears more easy than to catch that motionless thing.
You put out your hand and it is gone ; like a flash it has moved a *
foot higher up, or down, to the right or to the left, just where you
least expected it to go, and there it clings on motionless as before.
It does not seem to run ; it glides along, dodging over to the other
side of the stem and back again. There is system in its motions, since,
taking a last leisurely look around, it gently disappears in a rent or
hole. Towards the evening, or when the shadows become longer,
the Geckos become lively. One after another appears on the
surface, upon the tree, or at the entrance of the cave, and they
all move about in their peculiar rushing jerks. Spiders, flies,
mosquitoes, moths, form the principal diet, and the hunting
goes on well into the night. Where a gecko has been seen
once it is sure to reappear the next day at the same hour.
Those which take up their abode inside a house become almost
domesticated. They are strange sights when hunting for flies,

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XII GECKONIDAE 5 I I

running up and do.wn the papered walls; but we fairly gasp
when they come to the upper corner, calmly bend over, and with
the next jerk slide along the white- washed ceiling. We are
accustomed to flies performing such feats, but at animals five
inches long, supple and fat, we are inclined to draw the line.
However, that is the way of (Jeckos, and — ^be it confessed — the
more we ponder over the mechanism of their fingers and toes,
the less we comprehend how such little vacua can support or
suspend such heavy creatures from a dry and often porous
surface.

Gecko, — The digits are strongly dilated with undivided
lamellae. All, except the pollex and hallux, have a very short
compressed terminal phalanx with a retractile claw. Males with
femoral or pre-anal pores. This Eastern genus includes some of
the largest of all Geckos.

ft stentor of the Malay countries reaches a length of 15
inches. G, verticillattLS s. verus s. giUtatus ranges from Eastern
Bengal to China and through the Indian archipelago. It grows
to about one foot in length. The head is large ; the back is
covered with small granules and about a dozen rows of large
tubercles. The tail, when intact, and the belly are covered with
scales, those of the tail being arranged in transverse rows, several
of which make up distinct rings. The upper parts of the body
are grey or yellowish with red spots and vermiculations. Accord-
ing to Theobald ^ it lays about eight hard-shelled white eggs as
big as a musket-ball, cementing them to trees, rocks, or secluded
buildings. The cry is " touk-tay," several times repeated, and
ending in a long-drawn out, diminuendo, guttural rumble. This
animal does not confine itself to insects, but eats young rats also.
Dr. Mason has seen it devour smaller species of house-lizards,
and Theobald has seen it seize a bat flying round the room, and
devour it.

Tennent^ tells the following story about one of these creatures :
" In an ofl&cer's quarter in the fort of Colombo, a Gecko had
been taught to come daily to the dinner-table, and always made
its appearance along with the dessert. The family were absent
for some months, during which the house underwent extensive
repairs, the roof having been raised, the walls stuccoed, and the

* F. Mason's Burvuiy London, 1882.
* Sketches of the Nat. Hist, of Ceylon, London, 1861.

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LACERTILIA GECKONES

ceilings whitened. It was naturally surmised that so long a
suspension of its accustomed habits would have led to the
disappearance of the little lizard ; but on the return of its
old friends, it made its entrance as usual at their first
dinner the instant the cloth was removed."

Ptychozooiu — The digits
have the same structure as
described in the genus Gecko,
but they are entirely webbed.
The extraordinary feature of
Ptychozoon is the membran-
ous expansions on the sides of
the head, body, limbs, and tail,
which are said to act as para-
chutes. P, homalocephalum y
the only species, inhabits the
Malay Islands and the Malay
Peninsula. It reaches a
length of 8 inches. A speci-
men obtained by F. H. Bauer
in Java, in the month of
November, laid two eggs a
few days after its capture.
One young was hatched in
the middle of the following
May, and two days later
another came out of the
second egg. The character-
istic folds of the skin were
already clearly discernible.

Sub-Fam. 2. Euble-
pharinae. — Differing from
the true Geckos by their
procoelous vertebrae and the fusion of the two parietal bones
into one. The eyelids are not reduced, but remain functionaL
This sub -family is undoubtedly a heterogeneous assembly, as
indicated by the very scattered distribution of its few species
(about seven), in India, West Africa, and Central America.

Sub-Fam. 3. Uroplatinae, composed of a few species of the
genus Uroplates in Madagascar. The distinctive characters of

Fig. 121. — Ptychozoon homalocephalum.

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LACERTAE 5 I 3

these otherwise typical Greckos are the fusion of the nasal bones
into one, the small size of the interclavicle, and the non-dilated
shape of the clavicles.

Neither the Eublepharinae nor the Uroplatinae are more nearly
related to other Autosauri than are the other Geckos. They are
modifications within the sub-order of the Geckones.

Sub-Order 2. Lacertae. — Autosauri with procoelous, solid
vertebrae, and with the ventral portions of the clavicles not dilated.

Cope/ discarding outer appearances as deceptive in the classi-
fication of the Lacertae, laid stress upon internal characters,
notably the presence or absence of osteoderms, the formation of
the skull, and the structure of the tongue. Boulenger ^ has
followed and improved upon Cope's arrangement, and has
elaborated the classification, which, being used by himself in the
three volumes of the Catalogue of Lizards in the British Museum,
has also been followed in the present work, with slight alterations
in the order of treatment of the families. . For our present purpose
we diagnose the families as follows, giving preference to such
characters as are most easily ascertained : —

Synopsis of the Families of Lacertae.

Fam. 1. Agamidae, Acrodont. Tongiie broad and thick. No oeteo-

dernis. Old World, p. 515.

Fam. 2. Iguanidae. Pleurodont. Tongue short and thick. No

oflteoderms. America, Madagascar, Fiji Islands,
p. 528.

Fam. 3. Xenosauridae, Pleurodont, solid teeth. Anterior part of tongue

retractile. Osteoderms on the body. Mexico,
p. 536.

Fam. 4. Zonuridae. Pleurodont Tongue short, not retractile. With

osteoderms at least upon the skull, where they
i*oof in the supratemporal fossae, African
sub-region, p. 636.

Fam. 6. Anguidae. Pleurodont, solid teeth. Anterior i)art of tongue

emarginate, retractile into the posterior por-
tion. Osteoderms on body and head, roofing
over the supratemporal fossae. Limbs mostly
reduced. America, Europe, India, p. 537.

Fam. 6. Helodermatidae. Pleurodont, lower teeth grooved, with poison-
glands. Tongue bifid. Osteoderms tiny.
Postfronto-squamosal arch al)6ent, p. 540.

1 P. Ac. Philad. 1864, p. 224, and P. Amer. Ass. xix. 1871, p. 236.

^Ann. Nat. Hist. (5) xiv. 1884, p. 117.
VOL. VIII 2 L

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LACERTILIA LACERTAE

Fam. 7. Lanthanotidae.

Fam. 8. Varanidae.

Fam. 9. Xantusiidae.

Fam. 10. Tejidae.

Fam. 11. Lacertidae.

Fam. 12. Gerrhoeauridae.

Fam. 13. Scincidae.

Pleurodont, Tongiie short and bifid. Post-
fronto-squamoeal arch absent No osteoderms.
Borneo, p. 541.

Pleurodont Tongue very long, bifid, smooth,
very protractile. No osteoderms. Postorbital
and temporal arches incomplete. Old World,
p. 542.

Pleurodont Tongue very short and scaly. No
osteoderms. Supratemporal fossa roofed over
by the cranial bones. No movable eyelids.
Central America and Cuba, p. 547.

Teeth solid, almost acrodont Tongue long,
deeply bifid, with papillae. No osteoderms.
Limbs sometimes reduced. America, p. 547.

Pleurodont Tongue long, bifid, with papillae
or folds. With osteoderms on the head.
Supratemporal fossae roofed over by the cranial
bones. Old World, p. 549.

Pleurodont Tongue long, with papillae, but
feebly nicked. With osteoderms on the head
and body, roofing over the supratemporal
fossae. African sub-region, p. 559.

Pleurodont. Tongue scaly, feebly nicked.
Osteoderms on the head and body. Liinl)8
often reduced. Cosmopolitan, p. 559.

" families " are much degraded in con-
formity with their usually subterranean life, see p. 496: —

Fam. 14. Anelytropidae. Without limbs. Body covered with scales.

Mexico and Africa, p. 564.
Venuifonn, limbless body covered with scales,
witliout osteoderms. Australasia and Nicobar
Islands, p. 564.
Without linil)s ; body covered with scales, with-
out osteoderms. California, p. 564.
Fam. 17. Amphisbaenidae. The lx>dy is covered with soft skin, forming

numerous rings ^vith mei-e vestiges of scales.
Without limbs, except Ghiroies with four-
clawed fore-limbs, p. 565.
Snake-shaped, with scales. Fore-limljs absent,
hind-linil)s appearing as a pair of scaly flapsL
Australia, p. 567.

These eighteen " families " of the Laeertae fall into four main
groups. We naturally assiune that the presence of osteoderms and
of complete cranial arches indicate more archaic conditions than
their absence, just as we conclude that limblass forms have been
evolved from creatures with fully developed limbs. We arrange
the four groups with their families as follows : —

The

following

five

Fam. 15. Dibamidae.

Fam. 16. Aniellidae.

Fam. 18. Pygopodidae.

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XII AGAMIDAE 5 1 5

Group I. Zonuridae and Anguidae assume a central position, with Iguanidae
and Agamidae as two parallel families of highest development
Aniellidae as the most degraded forms. Helodermatidae and
Lanthanotidae as rather primitive and solitary survivals.

Agamidae Iguanidae

Xenosauridae

Zonuridae — Anguidae — Helodermatidae.

Aniellidae.
Group II. Xantusiidae — Tejidae — Amphisbaenidae.
Group III. Scincidae — Gerrhosauridae — Lacertidae. — Here also Anelytro-

pidae and perhaps also Dibamidae as degraded Scincoids.
Group IV. Varanidae, which are in many respects the most highly developed
ofalL
Pygopodidae are of obscure relationship.

Fam. 1. Agamidae. — Acrodont, Old- World Lizards, with a
broad and short tongue. The teeth are usually differentiated

^M AGAMIDAE. ^

Fia. 122. — Map showing the distribution of the Agamidae.

into incisors, canines, and molars. The orbit is closed posteriorly ;
the temporal fossa is bridged over by an arch which is formed
chiefly by the squamosal and the well-developed jugal ; the post-
orbital mostly remaining small, and the postfrontal and supra-
temporal bones being either absent or not present as separate
elements. The limbs are well developed. The eye, provided
with complete eyelids, is distinctly small and has a round pupil.
The skin is devoid of osteoderms, although large and numerous
spines are often present, especially on the head and on the tail.
The Agamidae, of which about two hundred species, arranged

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5i6

LACERTAE

in about thirty genera, are known, exhibit a great diversity of
mostly flat -bodied, terrestrial and more laterally compressed,
arboreal forms. The majority are insectivorous, a few Agamas
have a mixed diet, while UromasUx and some others are chiefly,
if not entirely, frugivorous and herbivorous. They are an ex-
clusively Old- World family, avoiding the cooler parts of the
Palaearctic sub-region, and also, a very curious fact, Madagascar.
The majority live in Australia and in the Indian and Malay
countries, comparatively few in Africa, chiefly the genus Agama,
Draco (" Flying Dragon "). — The body is much depressed and

Fia. 123. — Draco volans. x |.

the sides extend as a pair of large wing-like membranes, which are
supported by five or six of the much-elongated posterior ribs, and
can be folded up like a fan. On the throat are three pointed
appendages, a short one on either side and a long one in the
middle. The tail is very long and slender, but not brittle. About
twenty species of this extraordinary genus inhabit the various
Indo-Malayan countries ; one, D. (hissumieri, occurs in Madras.
D. volans of the Malay Peninsula, Sumatra, Java, and Borneo is
about 10 inches long, 5 of which are taken up by the tail. The

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XII AGAMIDAE $17

male has a small nuchal crest. The upper parts of this pretty
creature have a metallic sheen, with small dark spots and
undulating cross-bands upon the rich brown ground-colour. The
wings are orange with black markings. The gular sac of the
male is orange, that of the female is blue.

The " Flying Dragons " use their wings as parachutes, but their
sailing powers are said to be very moderate. Certainly they do not
fly by moving the wings, but when at rest upon a branch, amidst
the luxurious vegetation and in the immediate neighbourhood of
gorgeously coloured flowers, which partly conceal them by their
likeness, they greatly resemble butterflies, especially since they
have the habit of opening and folding their pretty wings.

Ceratophora. — This exclusively Ceylonese genus is remarkable
for a flexible, erect, and pointed appendage which arises from the
top of the snout ; it is best developed in the males, vestigial or
absent in the females. Gular appendages are absent. The trunk is
crestless, slightly compressed, and covered with partly keeled scales.
The tail is slender and very long, about two-thirds of the total length
of the animal The general colour is olive-brown, with irregular
darker markings and with light streaks on the head and thighb'.
C. stoddarti and C. tennenti are about 10 inches long, the former
without, the latter with, little scales upon the rostral appendage.

Lyriocephalus, with Z. scutatus (Fig 124) of Ceylon as the
only species, is remarkable for its Chameleon-like appearance. A
splendid case of convergent evolution, but most improbably of
mimicry. The tympanum is quite hidden. The head is raised into
a pair of sharp bony edges. On the top of the nose is a thick
globular lump, recalling the genus Ceratophoray and also various
Malagasy Chameleons. The back and sides are covered with very
small granular scales, intermixed with several rows of enlarged
scales as in Chameleo pnmilus, and there is a serrated crest along
the back from neck to taiL The under parts are covered with large
keeled scales with sharp points directed backwards, especially on-
the tail. The whole body is laterally compressed. The poUex and
the fifth toe are strongly opposed to the other digits. The general
colour is greenish above, whitish below. Total length about one foot.

Calotes, with many species in India and in the Malay Islands,
is distinguished by a crest on the neck and back. Many of the
males have a gular sac. The tail is extremely long. These
lizards are remarkable for their changes of colour.

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5i8

LACERTAE

C. versicolor ranges from Afghanistan through the whole of
India to Southern China, and attains a length of 14 inches, 11
of which are taken up by the tail It possesses no gular sac, but
has a well-developed crest. The whole body and tail are covered

Fig. 124. — Lyriocephalus scutcUus,

with strongly keeled scales. When the lizard is irritated, or
swallowing its food, the head and neck become brilliant red, whilst
the usually brownish tint of the body is converted into pale yellow.
Mr. Annandale has favoured me with the following observations
on C. emmu : — " In the Malay Peninsula the Europeans misname
this lizard Chameleon. The colour-changes appear to be brought
about by emotions, although the lizard is often darker towcurds

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AGAMIDAE 5 1 9

evening than it is at mid-day. The males are very pugnacious,
and change colour as they fight. At the time of courtship a
curious performance is gone through by the male, the female
remaining concealed in the foliage hard by. He chooses some
convenient station, such as a banana leaf or the top of a fence,
and advances slowly towards the female. His colour is then
pale yellowish flesh-colour, with a conspicuous dark spot on each
of the gular pouches, which are extended to their utmost. He
stands upright, raising the fore part of the body as high as
possible, and nodding his head solemnly up and down. As he does
so, the mouth is rapidly and repeatedly opened and shut, but no
sound is emitted. When he is driven away, caught or killed, the
dark spot disappears entirely from the neck. If one male is
captured, another takes his place within a few hours."

G, ojphiomachus of Southern India and Ceylon reaches 2
feet in length, has a fold of skin in front of each shoulder, and is
generally known as the " blood-sucker " on account of the red
colour displayed during excitement on the head and neck.

C, mystaveuSy chiefly in Burma and Siam, but also in the
Nicobar Islands and in Ceylon, has a small gular sac and an
oblique fold in front of each shoulder. The specific name seems
to refer to the yellowish lips. Mason ^ gives the following vivid
account of it : —

" This is a very common species in gardens in Toung-ngoo.
A pair made their home in the mango trees near my study
window. The female blundered into the house a few days ago,
but I found her a very unattractive animal of a uniform earth-
brown colour. The male, however, is sometimes a beauty. He
may be often seen jerking his head up and down, with the head,
pouch, and whole front of the body a glowing ultramarine blue,
contrasting beautifully with the reddish brown of the hinder part
of the body and tail. From the nose to the shoulders, below the
eye, is a broad white band, which is interrupted by three reddish-
brown patches, in line with the white band, before reaching the
uniform reddish brown of the hinder part of the body. Occasion-
ally the white band below the eye assumes a brownish colour,
and the animal appears to have a brown band down each side.
He does not always, however, appear in this gay dress. While I
am writing, I see him coming down the trunk of one of the trees
^ BurTnUf Us People and Prodv^tions^ London, 1882.

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520 LACERTAE chap.

in a very faded garment. His skin suggests a bright calico
after it has been washed, whose colours succumb to soap. The
blue is there, but it is no longer the bright blue of yesterday.
It has changed to a dull light indigo colour. He runs across
the grass to the foot of another tree, and stops on the bare
ground at its base, where for a minute or more he bites with
great energy at a struggling grasshopper, and while thus exercising
himself the blue fades out from his body altogether, and his
whole body takes the colour of the brown earth on which he
stands. After tarrying a minute or two he ran up the other tree,
and the dull light blue colour seemed to return to him."

Agama with many species in Africa and Asia ; some in South-
Eastern Europe. The body is somewhat depressed. There is a
fold across the throat and a pit on either side ; the presence of a
gular sac is variable. A dorsal crest is absent or but feebly
developed. The males have pre-anal pores.

A. sanguinolenta, — The body is covered with strongly keeled
and pointed scales. On the sides of the head are a few spine-like
scales. The ear-opening is partly concealed by a fringe of spinous
scales. The males have a gular pouch. This is a typical
inhabitant of the deserts and steppes of Turkestan. Zander ^ has
observed the habits and many changes of colour of this lizard.
The usual garb is earthy brown above, with somewhat darker and
rather indistinct markings. The under parts are whitish. Some-
times the creatiu*e changes to dirty white, at other times into
blackish or grey brown. Bluish-red stripes may appear on the
sides of the body ; blue lines begin to show on the throat, and
ultimately the whole belly, originally white, may become ultra-
marine blue. When the general tone happens to be sulphurous
yellow, blue often appears on the tail and limbs. Brick red
appears on four longitudinal rows of patches on the sides of the
body. Sometimes the whole animal assumes a vinous tinge, or
it is at first greenish before turning into blue. The change
begins on the tail and limbs, extends over the head, and at
length reaches the back. Eed appears in both sexes, more
frequently in the female; blue almost entirely in the male.
Sunlight and warmth only intensify the colours. Adaptive
coloration, besides the usual sandy garb, has not been observed.
The lizards live on soil which is baked as hard as bricks, or in
1 Zool, Garten, 1895, p. 232.

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. AGAMIDAE

521

cavities of old walls, provided there is vegetation. They require
vegetable food, besides insects, grazing on grass, and having a
fondness also for Meaemhryanthemum cardiforme. Very large
males reach a total length of one foot. They are pugnacious,
especially during the time of breeding. The male inflates its
gidar sac into the size of a walnut, stands up upon its four
limbs, with its head slightly lowered and turned to one side.
Then it darts upon the foe which it has been eyeing for some
time.

Fio. 125. — Afjaina sidlio. x ^.

A, stellio is the commonest Agama in Egypt, Asia Minor, and
in some of the Grecian Islands, where the Greeks still call it
korkordilos, just as they did in the time of old Herodotus. The
Arabic name is hardun. This lizard is easily recognised by the
irregular folds on the neck, which are beset with spinous horny
scales. It grows to a length of 15 inches. The general
colour is brown, with dark patches on the back. When basking
they become abnost black ; in the breeding season the male
assumes red tints on the head and neck.

Fhri/noceph(du8. — This is a tj^pical Agamoid of the steppes
and deserts of Asia. The head is short and thick, the ear is

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522 LACERTAE chap.

hidden. The body is depressed, devoid of a dorsal crest ; on the
throat is a transverse fold but no sac.

A. Zander^ has made interesting observations upon the habits
of several speciea

Ph, helioscopus lives on hard stretches of soil, which are
absolutely bare of vegetation, the soil being baked as hard as a
paved road. The lizards live on any insects they can get hold
of, chiefly, however, upon mining ants. When chased they run
with short jerks, carrying the tail high or rolled up.

Ph. intersenpidaris occurs, in Transcaspia, on the shifting,
loose sand. It runs so fast that one scarcely sees anything but
its shadow. The tail is rolled upwards. With short jerks
it suddenly changes its direction, stops behind a few blades of
grass, or in the open, makes a- few shaking, wavy movements,
and covers itself lightly with sand. Shortly after that the top
of the head appears, the grains of sand rolling off between the
strong supraciliary ridges, and the little creature, only about 3
inches long, peeps out of its temporary hiding-place.

Ph. mystaceus, which inhabits Transcaspia and parts of
Southern Russia, often faces its aggressor, raising itself upon its
fore-limbs, curling and uncurling its tail in its excitement, and
holding its mouth widely open. The creature, which attains a
length of 9 inches, inclusive of the long tail, then assumes a
markedly changed aspect. The flaps of skin at the corners of
the mouth swell up into a half-moon-shaped transverse plate,
the hinder surface of which is covered by the outer skin, while
the front is a continuation of the rosy lining of the mouth,
which thereby appears hugely enlarged. When biting it hangs
firmly on to the finger. This frightening attitude is interest-
ing, since it occurs in a much more developed condition in the
following genus.

Chlamydosaurus kingi. — This peculiar Agamoid, which
inhabits Queensland and Northern and North-Westem Australia,
is easily recognised by the large frill-shaped dermal expansion on
either side of the neck. The two halves are confluent on the
throat. The whole frill can be erected, and is worked by the
much-elongated arches or horns of the hyoid apparatus, which
extend into the flaps of skin, somewhat like the ribs of an
umbrella. ITie specially modified hyoidean muscles spread out

1 Zool. Garten. 1895, p. 257.

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AGAMIDAE

523

and fold the frill. When this curious creature is pursued it folds
the frill and runs in a semi-erect position upon its hind-limbs,
with its fore-limbs hanging down. However, it cannot keep
up this peculiar gait for long, and it then suddenly turns to bay,
frequently at the root of a tree, which it can climb with ease.
When standing at bay it spreads out the shield to its full extent,
in the middle of which appears the widely opened mouth, which
is red inside and armed with powerful teeth. Altogether this

[D.di^t^iTL

Fio. 126. — Chlamydosaurus kingi. x^.

lizard presents a formidable aspect, and is an enemy not to be
despised, considering that it is strongly built and grows to nearly
3 feet in length. For a further accoimt of the habits and of
the mechanism of the frill see De Vis.^

Physignathus. — This is a water-loving genus, inhabiting well-
watered districts with luxurious vegetation in Australia, Papuasia,
Siam, and Cochin China. The body and the very long tail are
laterally compressed and furnished with a low, serrated crest.
Ph, lesueuri of Queensland reaches a length of about 18 inches.
The general colour is dark olive above, with darker and lighter

^ P. Linn. Soc. N.S. Jr. viii. 1883, p. 300.

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LACERTAE

CHAP.

cross-bands, and with a broad black band reaching from the eye
to the shoulder. The under parts are pale olive, with small black
dots. The throat, although devoid of a special sac, is frequently
bulged out by the hyoid apparatus, as shown in Fig. 127, taken
from a specimen in the Zoological Gardens in London.

Uromastix is a typical desert-form, inhabiting the dry and
sandy tracts of North Africa, Arabia, Syria, Persia, and North-
western India. The genus is easily recognised by the short and
thick tail, which is covered with whorls of large spinous scales,
while the much-depressed body and head are almost smooth, being

Fig. 127. — Physignathus ksiieuri: x^.

covered with very small scales. The tympanum of the ear is
quite exposed. The incisors are large, uniting in the adult into
one or two pairs of large cutting teeth, separated from the molars
by a toothless space. There is a transverse fold on the throat.
Pre-anal and femoral pores are well developed.

These " Spiny-tailed Lizards " live chiefly upon vegetable food,
leaves, grass and fruit, but they vary this diet with insects, at
least in captivity, where they become rather partial to meal-
worms. They are absolutely terrestrial and diurnal, preferring
sandy places, where they bask or rather roast themselves in the
Sim ; for the night, at the approach of rain, or on dull and
chilly days, they retire into their burrows, which they dig in

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AGAMIDAE 525

the sand or in the hard ground, unless they hide in the cracks
of rocka They have a regular mania for digging with their
strong limbs and short, curved claws. Although they love a
great amount of heat, and l)ecome stiff when cooled down to
about 16° C. = 60 F., they can stand several degrees of dry frost
without injury. During the cold season they hibernate. The
spiny tail is used for defence. The lizard lies as a rule in such
a position in its hole that the tail blocks the narrow passage ;
when touched with the hand it deals out jerky side-blows with
the tail. The bite is deUberate and very painful.

C/". hardwicki is a native of North-Western India and Beluchi-
stan, occurring especially in Sindh and Kajputana, for instance
near Delhi and Agra. This species is of a delicate sandy colour,
with dark dots or vermiculations, interspersed, occasionally, with
pale blue specks. The under parts are whitish on the tail with
a greenish hue. A distinctive and obvious mark is a large
blackish patch on the anterior side of the thigh. Total length
up to one foot.

I have several times received consignments of the Indian
Spiny-tailed Lizard through the kindness of friends, but I must
confess that they are far less easily kept than one is led to
believe from certain exaggerated accounts. They are lovely,
most interesting, and surprisingly tame creatures. I received
one lot in the month of June. They made burrows in the dry
soil, basked in the sun and on the grassy sods of their roomy
cage, and showed great curiosity. When approached, they at first
scrambled off or sank down flat, shut their eyes and feigned death.
They then opened their tiny yellow eyes a little, while others
peeped out of their retreats to see if all was safe, or attracted by
some noise. Soon they became so tame that they crawled over
my hand. But the difficulty consisted in feeding them. They
greedily lapped up drops of water. Their dung consisted of the
indigestible parts of some species of Uquisetum or Mare's tail,
mixed with fragments of beetles and ants. Lettuce, cabbage,
cauliflower, grass, the flowers of red and white clover. Mare's
tail, wheat, rice, and Indian corn were offered, but they only took
a few blades of grass and the hard Indian corn, besides meal-
worms. This is all the more astonishing since other specimens are
known to partake freely of herbaceous food None of them sur-
vived the late autumn, and most of them succumbed to a disease

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LACERTAE

known as intussusception of the gut. They certainly could not
complain of the want of heat, since the bottom of their cage was
kept permanently warm by a lamp, and in the autumn they in-
variably slept in the warmest part of the soil, avoiding the cool
regions which would have given them a chance of hibernating.

Another consignment arrived in the month of February.
None of them ate anything or survived the early summer.

U. ncanthinurus and U. spinipes are common in Algeria,

Fig. 1.28. — Uromastix acanthinurus. y.\,

Tunis, and Eg}^t, where they prefer sandy and rocky localities.
Their Arabic name is Dah. In Algeria they are sometimes
called " lezards des palmiers," perhaps because they eat dates,
besides berries, grass, and various flowers. Very large specimens
attain a length of 18 inches. Like the other species of Uromnstix
they have no voice. The African species can change colour
to a great extent. At a low temi)erature they are mostly grey
or brownish black above, dirty white below^ When it is warmer
they change to lighter shades of brown or even to orange yellow
and to green, with black or brown specks and vermiculations.

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XII AGAMIDAE 52/

A young specimen of U, acanthinurus has been observed to
gTQw within twelve months from 90 to 150 mm. in length.

Moloch, — The mouth of this peculiar-looking creature is very
small ; the lateral teeth of the upper jaw are implanted horizon-
tally and directed inwards. The body is much depressed, and, like
the short tail and head, is covered with small scales or tubercles
intermixed with large spines. M, horridus, the only species,
inhabits the sandy districts of Western and Southern Australia.
Nothing is known about its habits except that it seems to live
upon ants. Its extremely rough skin is, according to an acci-
dental observation by Dr. Willey, highly hygroscopic. He
happened to put a live specimen into a shallow dish with water,

Fig. 129. — Moloch horridus. x |;

when, to his surprise, the water was sucked up as by blotting-
paper.

Liolepis. — The body is depressed, without a crest, and is
covered with minute granular scales. The tail is long, and has
small keeled scales. There is a strong transverse gular fold, and
a fold along the side of the body. The tympanum is distinct.
Femoral, but no pre-anal, pores are present.

L. belli, the only species, about 18 inches long when full
grown, is a native of South-Eastem Asia. The general colour is
brownish, with pale black-edged spots along the back ; the sides
are marked with black and orange vertical bars ; the under parts
are orange, variegated with blue. Annandale remarks that this
is perhaps the commonest lizard on the barren stretches of sand
in Lower Siam, especially near the coast. It is exceedingly
active and timid. Though its colour is brilliant, the green and

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528 LACERTAE chaf.

grey eye-like spots which ornament its back, and the orange and
purple stripes on its sides, are not conspicuous amidst the natural
surroundings, the former harmonising with the shadows cast upon
the sand by the scanty vegetation which it supports, and the
latter being more or less concealed by the folds into which the
skin that covers the ribs naturally falls. When the male is
roughly handled and is unable to use its powerful jaws, it
flattens its body in such a way that the stripes become most
conspicuous. The female is unable to do this with such effect,
as her ribs do not seem to be so mobile and her colours are leas
bright. Ziolejyis lives in holes in the ground, which often go
down vertically for more than 2 feet before there is a bend in
their course. Each burrow generally contains a pair of these
lizards, which, according to the natives, are strictly monogamous,

Fam. 2. Iguamdae. — Pleurodont lizards with a short and
thick, non-protractile tongue ; almost entirely American, with the
remarkable exception of two genera, Hoplurus and Chalarodon
in Madagascar, and one, Bnichylopkus, in the Fiji Islands. Most
of the Iguanidae are insectivorous, but some of the most striking
forms are herhivorouSy e.g. IguaTUiy Amhlyrhi/nchus, a.nd Basilisciis.
In their general structure the Iguanidae closely resemble the
Agamidae, from which they differ chiefly by the pleurodont
dentition. The orbit is surrounded by bone, and the temporal
fossa is bridged over by an arch which is formed by the junction
of the squamosal chiefly with the postorbital, the jugal taking
as a rule less share in the arch. Dermal ossifications are absent
on the body.

There are about three hundred different species, which have
been grouped into about fifty genera, representing arboreal,
terrestrial, burrowing, semi-aquatic forms, and even one semi-
marine species. Their external appearance varies consequently
within wide limits.

Anolis is distinguished by the partial dilatation on the middle
phalanges, which carry a series of transverse adhesive lamellae.
In its general shape Anolis resembles slenderly built and long-
tailed Lacertidae, which it may be said to represent in tropical
and sub- tropical America, inclusive of the West Indian Islands.
The males have a large gular appendage, which can be distended
by the hyoid bones. Anolis is an expert climber, living in trees,
or rushing about on fences or walls of houses in search of insects ;

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IGUANIDAE

529

most species can change colour to a great extent. More than a
hundred species are known, of which we mention only one, very
common in the Southern United Statea

A, carolinends of the South-Eastern United States and of
Cuba is beautiful golden green on the whole upper surface ; the
gular sac becomes vermilion when inflated ; when flaccid, it is
white with occasional red lines and spots. The under parts are
white. In cold weather and in confinement this little lizard,
which is scarcely 6 inches in length, appears dark brown,
sometimes with a white line along the back. The changes of
colour are very sudden. They are thoroughly arboreal, leaping
from leaf to leaf like Tree-frogs.

^ ANCUIDAE. illtlllll IGUANIOAE. !'»:» ZONURIDAE.

Fio. 130. — Map showing the distributiou of Anguidae, Iguanidae, and Zonuridae.

In Anolis, Polychrus, Hoplurus, Chalarodon, Ziosawrus, and a
few others, the posterior ribs are much elongated and imbedded
in the abdominal muscles, often reaching the medioventral line, a
feature elsewhere known in the Geckos only.

Polychrus. — The body is laterally compressed, covered with
small scales, but devoid of crests. The digits are likewise com-
pressed, with keeled lamellae on the under smrffice and with
four large scales at the base of each claw. Both sexes have
femoral pores. The male possesses a small gular sac. A few
species in Tropical America.

P. marmoratus in South America, where it is often called the

Chameleon on axjcount of its power of changing colour. The tail

is nearly three times as long as the head and body, and is covered

with keeled scales. The general colour of this arboreal creature,

VOL. VIII 2 M

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530 LACERTAE chap.

which reaches a length of 18 inches, is green, but the hues are
very variable, and within a short time the creature can change
into dull brown, with or without blackish spots and bands, or
with whitish spots and black lines on the head and other parts
of the body.

Basiliscus is remarkable for the high and erectile crests which
are developed on the back and tail of the males. The toes are
bordered on the outer side with small lobes. The limbs are
long, the hind-limbs when stretched forwards reaching the tip

Fio. 131. — Basiliscus atnericanus (male).

of the snout. Several species in Central America and the
adjoining countries to the south.

B. americanus reaches the considerable length of nearly 3
feet. The male has a crest on the top of the head, and this is
produced backwards into a leathery lobe. The back is adorned
with a \exy high crest ; the folds and dark-coloured marks in
which give, in the accompanying illustration, the impression that
the crest is supported by spines. The long tail carries a similar
crest. The general colour of the " Basilisc " is green and brown
with dark cross-bars on the back. The crest of the male is
said to be red. These creatures are very common amidst the

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xiT IGUANIDAE 531

luxuriant vegetation on the banks of the rivers of the Tierra
Caliente of Mexico and in Guatemala. They lie upon the
branches of trees, preferring those which overhang the water,
into which they plunge at the slightest alarm. The high
crests, being restricted to the male sex, are not essential to
.their swimming ; they propel themselves by rapid strokes of the
fore-limbs, letting the long rudder-like tail drag behind. The
eggs, measuring 20 by 13 mm., are laid in April or May, and are
hidden in a hole at the base of a tree. About one dozen make
a set, and they are said to be hatched within a very short time.
Owing to their being strictly herbivorous, these pretty and
striking-looking creatures do not endure captivity in Europe,
unless indeed their particular food can be prociffed.

Iguana, — The body and tail are laterally compressed and are
covered with very small scales, while those on the top of the
head are large. The neck and back carry a high crest, which
is composed of separate, laterally compressed, soft spinea A
similar but lower crest borders the anterior edge of the large
gular sac, which is not dilatable. The lateral teeth are remark-
able for their finely serrated or denticulated anterior and
posterior edges. Both sexes have long rows of femoral pores.
Only two species in Tropical America, absolutely herbivorous.
Their delicate flesh is much esteemed as food.

/. tuherculata (Fig. 132), of South and Central America and the
West Indies, reaches a length of 5 to 6 feet. The general colour
of the upper parts is a mixture of green and blackish, frequently
speckled with white or yellow, and there is usually a pale band
in front of each arm. The flanks are marked with dark, light-
edged bars. The under parts are pale greenish or whitish. The
Iguanas live in the trees, and when there is danger they jump
into the water whatever the height of the tree, coming down with
violence. In going up some of the narrow, unfrequented creeks
in the Mosquito country, according to Napier Bell,^ the voyager
often encounters quite a shower of falling Iguanas, and runs some
risk of getting his neck broken. Large specimens, 6 feet long,
weigh perhaps 30 lbs. They burrow deep horizontal holes in
the sloping side of a bank. About two dozen eggs, nearly 2
inches long, are laid in a hole, where they are hatched in tne
month of May.

^ Tangweera, London, 1899.

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532

LACERTAE

CHAP.

Iguanas are often brought to the markets, either lashed length-
wise to a branch of the tree on which the specimen happened to
be surprised, or tied up with the long tendons of their own
toes.

FiQ. 132. — Iffiiana tubercidata.

Metopoceros cornutus of Hayti is closely allied to Iguana^
but the male has three conical horn-like scales on its head. The
general colour of the whole animal, which grows to more than
one yard in length, is dull black.

The following two genera, each containing one species only,
are restricted to the Galapagos Islands. Darwin ^ gives a long
and vividly written account of their habits.

Conolojphus suhcristatus, — Fully grown specimens are a yard
long. Their shape is stout, the head and fore part of the body
appearing especially heavj^ The head is covered, or rather paved,
with large cobble-stone-like scales. On the neck is a low crest
of recurved spines, while the median line of the back appears
simply serrated. All the teeth are trilobate. A gular sac is
absent. The colonition is striking. The head is lemon-yellow ;
* Voyage of the Beagle f London, 1845, chap. xvii.

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XII IGUANIDAE 533

the back is red, merging into dark brown on the flanks. The belly
is dark yellow ydth a tinge of reddish brown.

This lizard was found by Darwin on some of the Galapagos
Islands. On James' Island it was so common that the party
found it difficult to pitch their tent, on accoimt of the groimd
being undermined by the many burrows of the reptiles. They
feed during the daytime upon the succulent cactus and the leaves
of various trees. The perfectly harmless creatures are, or were,
eaten by the inhabitants.

Amhlyrhynchus cristatvs is closely allied to Gonolophus, of
which it may be said to be an aquatic modification. The top of
the blunt head is covered with low, conical, broad-based scales.
Over the neck, back, and tail extends a continuous crest of low,
recurved, spiny scales. All the teeth are trilobate. The body
and even more so the tail are laterally compressed. The general
colour is dark brown above, paler and inclining to whitish below.
Yoimger specimens have pale grey spots and blackish cross-bands
on the back and sides. Total length up to 4 feet. The
remarkable feature of this Iguanoid is its semi-marine life. It
inhabits the rocky and sandy strips of coast of most of the
Galapagos Islands, feeding upon certain kinds of algae, which it
has to dive for, since these plants grow below tide-marks.

Phrynosoma (" Horned Toads "). — The body of these little
creatures is much flattened and broadened, devoid of a dorsal
crest, but covered with larger and smaller, strongly keeled scales.
The head is bordered posteriorly by conspicuous osseous spines. The
under parts are covered with small, very regular scales. Both
sexes have a long row of pores on the under surface of the
thighs. The general colour of the upper parts is a mixture of
yellow, grey, brown, and black, the larger spiny scales causing
the animal to look as if it were sprinkled with the dried husks
of seeds, for instance those of Buckwheat. The object is
concealment, by close adaptation to the arid, sandy localities
which are the home of " Homed Toads." About one dozen
species inhabit the western half of the United States and Central
America, All the species are viviparous, almost the only
instance among Iguanidae.

Ph, cornutum has five spikes on each side of the head : one
postorbital, three temporal, and one occipital, the latter being by
far the largest. The sides of the lower jaw project in the shape

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534 LACERTAE

of prominent ledges, and are protected by a series of small
spines. The ventral scales are keeled. The under parts are
yellowish, frequently with a few brown spots. This species.

Fig. 133. — Phrynosoma cornutum ( ** Homed Toad "). x 1.

which grows to a length of 5 inches, ranges from Illinois
through Kansas and Texas to Northern Mexico.

Fh, coronatum, an inhabitant of California, has an additional
smaller spine between the two large occipitals. The scales of
the belly are quite smooth.

These peculiar-looking and interesting creatures recall some
of the extinct Dinosaurs in the curious configuration of their
head: small miniatures indeed. In order to be kept in good
health, and to be observed properly, they require, above all,
warmth, simshine to bask in, sand to burrow in, and proper food.
The latter consists of all kinds of small insects, the necessary
variety of which is best procured by making sweepings with a
butterfly-net in a meadow. Tliey take green-flies, house-flies,
ants, smooth caterpillars, small moths, meal-worms, wood-lice,
etc. The food is snapped up very quickly by a flash of the
tongue, which can be turned out, almost like that of a frog, but
only to the extent of half an inch. Water in the shape of dew,
or drops, is absolutely necessary. When in good condition, they
defaecate regularly every alternate day. They love to bask in

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XII

IGUANIDAE

535

the broiling sun, heating themflelves well through ; and in the
afternoon, long before sunset, when the sand is warmed up to
40° C, or fever-heat, they prepare to go to bed. For this they
select a dry and soft spot, and within a few minutes manage to
dig themselves in flat, literally sinking into the sand by pushing
themselves forwards, and by shovelling the sand upon their
backs with peculiar motions of the fringed sides of their flat
bodies. Sometimes the spines of the head remain sticking out,
looking like dry thorns scattered over the sand. To prevent the

:3t^^^

FiQ. 134. — Phrynosoma eoronatum (" Horned Toad "). x 1.

latter from getting into the nostrils, these are provided with
closely-fitting valves. Thus they remain concealed during the
night, and not until the sun is well up do they leave their
hiding-place, first peeping out, and then raising their head and neck,
letting the sand roll oflf between the spines. Still half con-
cealed, the back covered with little pebbles, seeds, or bits of dry
leaves, they wait for a long time before they feel lively enough
to sally forth. Although mostly slow and deliberate in their
movements, stalking about with arched back, and raised upon
the fore-limbs, they can rim fast enough for a few yards before
they stop again and nod in a ridiculous way. When they
see themselves observed, they shut their eyes and slowly sink

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536 LACERTAE chap.

down. On cool and dull days they do not appear at all, and
during part of the cooler season they require artificial heat until
they are ready to hibernate. Unless they are allowed to hiber-
nate, they will keep on feeding through the winter, but in that
case are sure to die in the following spring.

Fam. 3. Xenosauridae, with Xeriosaurus grandis in Southern
Mexico as the sole species, seems to connect the Iguanidae with
the Anguidae. According to Boulenger, its afl&nity to the
former is shown by the T-shaped interclavicle, the absence of
symmetrical bony shields on the head and of osteodermal plates
on the body. Affinity to the Anguidae is indicated first by the
short tongue, which has a narrow, feebly incised, retractile
anterior part, covered with flat papillae ; secondly, by the teeth,
which, instead of being hollow at the base, are solid ; lastly, by
the palatine bones, which are widely separated.

JT. grandis, scarcely one foot in length. The body is de-
pressed, covered above with minute granules and tubercles, below
with smooth scales. A distinct fold of skin extends from the
axilla to the groin, recalling the more strongly developed lateral
fold of some of the Anguidae.

Fam. 4. Zonuridae. — This family, comprising four genera
with about one dozen species in South and Tropical Africa, and
in Madagascar, likewise * seems to connect Iguanidae and An-
guidae. It is distinguished from the former by dermal ossifica-
tions, which roof over the supratemporal fossa ; from the latter
by the tongue, the hollow teeth, and, in Zonurus at least, by the
occurrence of dermal ossifications on the trunk and tail. The
tongue is short, villose, scarcely protractile, entire, or but feebly
nicked at the tip. The Zonuridae may therefore be defined as
pleurodont African lizards with a short tongue, and with a bony
roof to the supratemporal fossae,

Zonurus. — The whole head, back, and tail are covered with
bony scales, the horny covering of which forms very sharp spikes,
especially on the tail The body is depressed. The ear-opening
is large. South Africa, in dry and rocky localities ; one species,
Z. tropidosternum, in Madagascar.

Z. giganteus s, derhianus, with strong spikes on the occiput,
neck, and tail. Greneral colour yellowish brown. Total length
about 15 inches.

Chamaesaura of South Africa closely approaches the An-

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537

guidae by its snake-shaped body, extremely long tail, and
vestigial limbs. In Ch, aenea both pairs of limbs are still
present and pentadactyle, but are very small ; in Ch, anguina the
limbs are reduced to little styliform stumps ; and in Ch, mcLcro-
lepis they are altogether absent. The scales of the body and

Fig. 1 3.5. — Zonurus giganteus. x\.

tail are strongly keeled and imbricating, but are devoid of dermal
ossifications. Total length up to 2 feet.

Fam. 6. Angnidae. — Pleurodont lizards with osteoderms, and
with the tongue composed of two distinct portions, of which the
anterior is thin, emarginate, extensible, and retractile into the
posterior thicker portion. The supratemporal fossa is roofed in
by dermal bones. The whole body is protected by bony plates
underlying the imbricating scales. The teeth vary much in
shape, but they are always solid, the new teeth not growing into
the base of tha old ones, but between them. The limbs are in a
very unstable condition, there being in the family a general
tendency to reduce and lose the limbs. The shoulder- and
pelvic - girdle however remain, although sometimes merely
vestigial. The tail is long, very brittle, and easily reproduced.
All the Anguidae are strictly terrestrial, and live on animal diet.

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538 LACERTAE

Some Anguis, at least, are viviparous. The distribution of the
seven genera, with some forty species, is very scattered. The
majority, chiefly OerrhoTwttts, inhabit Central America, a few
occur farther north and south — two, Anguis fragUis and Fseudoptis
pallasi, in Europe, and one in the Himalayas and in Burmah.

Gerrhonotus has a pair of deep longitudinal folds, each of
which extends from the region of the neck along the side of the
body towards the tail. The four limbs are well developed and
pentadaxjtyle. . The teeth are conical Many species, mostly in
Central America. G, coerulevs has the widest range, extending
from Costa Eica to Vancouver. It is also one of the largest
species, reaching a length of more than one foot. The tail is
nearly twice as long as the rest of the body. General colour
above brown with blackish bars and spots, especially on the
more yellowish flanks; under parts whitish with a greenish
tinge, often with brown spots arranged in longitudinal rows.

Ophisaurus s. Pseudopus is closely allied to the previous
genus, being possessed of the same kind of deep lateral folds ; the
limbs are, however, reduced to a pair of tiny spikes, half concealed
at the sides of the anal cleft. The teeth are conical, and in the
adult have somewhat flattened crowns. The body and tail are
very long and snake-like, but the head is that of a typical lizard.

0, apus s. Psevdopus pallasi, the Glass-Snake of the Balkan
Peninsula, South Bussia, Asia Minor, and Morocco, grows to
more than one yard in length, of which about two-thirds belong
to the tail. The general colour is brown above, paler below.
Young specimens are olive-grey with dark brown cross^bands on
the ba^k. 0. gracilis inhabits the Eastern Himalayas and
Burmah, the others live in North America.

The "Glass-Snake" inhabits bushy localities, where it can
hide under the fallen leaves and in the sand ; it cannot climb,
and avoids the water. Its movements resemble those of a
snake, but are far less graceful, owing to the stiff armour in
which the whole body is encased. The food consists chiefly of
snails, the shells of which axe crushed, and. of mice, but
nothing comes amiss which can be mastered, namely insects,
worms, small lizards, young birds, and vipers. The prey, when
caught, is rapidly twisted round and round, or shaken until it is
giddy or stimned, whereupon the Glass-Snake proceeds to chew it
with its powerful jaws, and then to swallow it in pieces.

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Although it can bite so well, it never does so when caught, but
resorts to the much more disagreeable defence of twisting
itself around one's hand and arm, and besmearing them with its
disgustingly stinking excrements. Those who have observed
Glass-snakes praise their tameness, and the inteUigent way in
which they hunt about in search of their food. They lay eggs
under moss and leaves, and the young seem to require many
years to grow up.

AnguiSy with only one species, -4. /rai^iZis, the "Slow- worm" or
" Blind-worm," is devoid of a lateral fold. Limbs are entirely
absent. The whole body is ^ covered with smooth roundish

tfiEl-rWiT

Fio. 136. — Anguis/ragilis (the Slow- worm). x\.

scales, with a substratum of dermal ossificationa The teeth are
curved backwards, fang-shaped, and have a very faint longi-
tudinal groove on their anterior surface. The ear-opening is
very minute, more or less hidden by surrounding scales. . The
eyes are perfectly well developed, provided with movable lids, and
it does not speak well for the power of observation of most people
that this creature should generally be known as the " Blind-worm."
The whole skin is shiny, metallic, quite smooth, brown above,
blackish below. But the coloration is subject to much indi-
vidual variation. Old specimens are sometimes adorned with
blue specks. The very young are exquisitely beautiful, the
upper surface being silvery white, with a median and two more
lateral lines of deep black ; the under parts are black. The iris
is yellowish red. Very large specimens measure more than one

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540 LACERTAE

foot in length, more than half of which belongs to the tail. One
in the British Museum is 425 mm. = 17 inches long.

The Slow-worm is viviparous, i.e. the young are fully de-
veloped, and burst the transparent, soft, yellowish eggs immedi-
ately after these are laid. This takes place in the months of
August or September, about one dozen making a litter. The
little creatures are at first about one inch and a half long, and
as thin as an ordinary match. They eat the smallest of spiders
and delicate insects; later on earth-worms, which they bite
into and then suck out before devouring them. When six
weeks old and well fed they are about 3 inches long, but it
is at least four or five years before they are mature. The
little ones carefully avoid the hot sunshine, and the adults
are likewise rather partial to the shade, although strictly
diurnal. Their chief food consists of earth-worms and snails.
For the night they retire under moss, leaves, stones, or into the
ground. In the autumn the Slow-worms dig passages or burrows,
which often serve as the winter-quarters of mauy specimens, as
if there were no other place available, or rather as if the spot
selected were by far the best with regard to safety, dryness, and
warmth.

Fam. 6. Helodermatidae. — Fleurodont, poisonoiis lizards of
North America. The teeth are fang-like, recurved, with slightly
swollen bases, rather loosely attached to the inner edge of the
jaws. Each tooth has a groove on its anterior and posterior
surface, and a series of labial glands which secrete the poison open
near the bases of the teeth of the lower jaw. The skull has strong
postorbital but no postfronto-squamosal arches. The pre- and
post-frontals are in contact, separating the frontal from the orbit ;
the premaxillaries are fused into one; the nasals and frontals
remain separate. The limbs are short, but strong and well
developed. The tongue is villose, with an anterior smooth
portion, which is bifid and protractile, resembling the tongue of
the Anguidae and of Aniella, The skin of the upper surface is
granular, with many irregular bony tubercles, which give it an
ugly warty look. The under parts are covered with flat scales.^

Hdoderma, the only genus, with H. horridum in Mexico and
If, suspectum in New Mexico and Arizona, reaches about 2 feet

* For further anatomical details see Shufeldt, P.Z.S, 1890, p. 148: Boulenger,
P.Z.S, 1891, p. 109; and Stewart, P.Z.S. 1891, p. 119.

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XII

HELODERMATIDAE LANTHANOTIDAE

541

in length. The animal, stout, depressed, thick -tailed, looks
rather repulsive when it squats down in its usuial lethargic way.
The whole skin is blackish brown and yellow or orange, these two
"warning" colours being distributed unevenly, except on the
thick, peculiarly-shaped tail, where they are arranged- in alternate
rings. The specific differences are rather imaginary. The New
Mexican form is supposed to be more orange and yellow than black,
with a somewhat smoother skin and with shorter toes and tail.
The " Gila Monster " inhabits dry localities, spends most of

Fio. 137. — Hel(M.lenna suspectum (the Gila Monster), x^.

the daytime in concealment between the roots of trees, and crawls
about in the evening in search of worms, centipedes, frogs, and
the eggs of large lizards. Frogs are probably paralysed or killed
by the bite which, although not so dangerous as that of poisonous
snakes, is effective enough to produce severe symptoms even on
man, and a few cases of death of people who had been bitten are
on record. In captivity they are very partial to eggs, which they
break and then lap up. During the dry and hot season they
aestivate.

Fam. 7. Lanthanotidae. — Lanthanotus borneensis, of which
only two specimens are known, one in the Vienna Museum, the

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542

LACERTAE

CHAP.

other in the Sarawak Museum, was described by Steindachner as
the type of a distinct family, near the Helodermatidae. Boulenger,^
after examination of the Sarawak specimen by means of a sciagraph,
has come to the conclusion " that the aflinity of Lanthanotus to
the Helodermatidae is fully confirmed." The teeth of Lanthanotus
show, however, no traces of grooves ; poison-glands are probably
absent, and there are no osteoderms. The skin is covered with
wart-like tubercles, each with a homy, peeled scale. The eyes
are very small, the ears are concealed. The general colour is
reddish brown above, yellowish, with brownish bands, below.
Total length about one foot, a little more than half of which
belongs to the roundish tail.

Fam. 8. Varanidae. — Pleurodont Old- World Lizards, with a

Fig. 138. — A, Ventral, B, dorsal view of the skull of Varanua griseus. x 1. B.0, B&si-
occipital ; B.Sy basisphenoid ; Co/, columella auris or stapedial rod ; E,P, ecto-
pterygoid ; Fr, frontal ; Jug, jugal ; LaCj lacryraal ; A", nasals ; /*a/, palatine ; Par,
parietals ; Pr./, prefrontal ; Pt/, postfrontal, fused with postorbital ; Ptg, pterygoid
(endopterygoid) ; Q, quadrate ; 2Y>, turbinal ; To, vomer.

long, deeply bifid and protractile smooth tongue. They reach a
large size, and the neck is relatively much longer than that of
other lizards. The limbs are well developed. The skin is

' P.Z.S. 1899, p. 596.

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XII VARANIDAE 543

covered with very small juxtaposed scales and tubercles above,
while the ventral scales are squarish and arranged in transverse
rows. Osteoderms are entirely absent. The tail is very long,
often laterally compressed. The teeth are large and pointed,
dilated at the base. The premaxilla is unpaired and dorsally
extends backwards to the likewise impaired nasal. There is a
pair of small supra-orbital bones, easily lost dming maceration.
The orbit is open behind, the jugal being short and not meet-
ing the postfrontal; the postorbital forms a slender arch with
the supratemporaL The vomers are long and diverge posteriorly.
The palatines, pterygoids,
and ectopterygoids enclose
on either side an oval
infra - orbital foramen.
The Varanidae contain
only one genus, Varanus,
with nearly thirty species
in Africa, Southern Asia,
and Australia, but not in
Madagascar. ^* varanidae.

rarani/s.— The name ^^^- ^^^■"ofTfafvImfidlll'*^*"^^^^

of " Monitor " bestowed

upon these creatures has a curious origin, owing to a ridiculous
etymological mistake. The Arabic term for Lizard is " Ouaran " ;
this has been wrongly taken \iO mean warning lizard, hence the
Latin Monitor, one of the many synonyms of this genus, e.g.
HydrosauTus and Psammosaiirics, Many of the " Monitors " are
semi-aquatic, others inhabit dry, sandy districts, while others are
at home in well-wooded localities. They are all rapacious, taking
whatever animals they can master according to their size, which
in some species amounts to 6 or 7 feet.

V. niloticus inhabits the whole of Africa, except the north-
western part. It reaches a length of more than 5 feet. The colour
of the adult is brownish or greenish grey above, with darker reticu-
lations and yellowish ocellated spots on the back and limbs. The
under parts are yellowish with blackish cross-bands. The groimd-
colour of the yoimg is black above with yellow lines on the
head and neck, and with yellow spots on the back and limbs ; the
tail has black and yellow bars.

V. salvator ranges from Nepal to Ceylon, Cape York, and

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544 LACERTAE chap.

Southern China, inclusive of the Malay Islands and the Philip-
pines. This is the largest species, specimens of 7 feet in
length being on record The general colour is dark brown or
blackish above, with yellow spots or ocelli. The snout and chin
have transverse black lines on a lighter ground. A black band,
bordered with yellow, extends from the eye along the side of the
neck. The under parts are yellow.

Mr. Annandale has favoured me with the following observa-
tions : — ** Varanus salvator is common in Lower Siam, where it is
equally at home on land, in water, and among the branches of
trees. The eggs are laid in hollow tree-trunks. When in the
water the lizard swims beneath the surface, the legs being closely
applied to the sides, and the tail functioning both as oar and
as rudder. Their food is very varied. In the states of Patalung
and Singora, in which the Siamese practise a form of tree-burial,
these great lizards are accused, probably with justice, of eating
the flesh of the corpses in the aerial cofldns. I have disturbed a
large Monitor devouring the body of one of its ojyn species, which
had evidently been dead for some days. Another, which was
chased by some men, dropped from its mouth a small flying
squirrel (Sciuropterus) ; a third, which I dissected, had lately
swallowed a small tortoise, the hard shell of which had been
broken into innumerable fragments. The stomachs of several
others contained nothing but dung-beetles, for which Varanus
may often be seen himting, turning over the dung of elephants
and buffaloes with its fore-feet. The Malay name of these lizards
is Biawak."

According to Mason and Theobald ^ all the Varanidae and their
eggs are highly esteemed for food, and are sought for in hoUow
trees with the aid of dogs. If not wanted at once, the wretched
creature has its fore-feet bent over its back, a few of its toes are
broken and the sinews drawn out and tied into a knot, rendering
the animal helpless. The Karens, who are extravagantly fond of
the flesh, steal up the tree with a noose at the end of a bamboo,
and often noose them while leaping for the water, or catch them
in a boat which is brought under the tree. The head, the natives
say, is venomous, and they discard it altogether, but the flesh of
the other parts, which smells most odiously, is deemed preferable
to that of fowls.

* Burmaf its People and Productions, London, 1882.

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VARAMDAE 545

Sir J. G. Tennent^ gives the following account of V,
salvator : —

" The ' Kabara-goya ' of the Singhalese is partial to marshy
ground, and when disturbed upon land will take refuge in the
nearest water. From the somewhat eruptive appearance of the
yellow blotches on its scales, a closely allied species, similarly
spotted, obtained the name of Monitor exanthematicuSy and it is
curious that the native appellation of this one, Kabara, is
suggestive of the same idea. The Singhalese, on a strictly
homoeopathic principle, believe that its fat, externally applied, is
a cure for cutaneous disorders, but that taken inwardly it is
poisonous. The skilfulness of the Singhalese in their preparation
of poisons and their addiction to using them are unfortunately
notorious traits in the character of the rural population. Amongst
these preparations the one which above all others excites the
utmost dread, from the nimiber of murders attributed to its
agency, is the potent kahara-tel, a term which Europeans some-
times corrupt into cohra-tel, implying that the venom is obtained
from the hooded-snake ; whereas it professes to be extracted from
the Kabara-goya.

" In the preparation of this mysterious compound, the unfor-
tunate Kabara-goya is forced to take a painfully prominent part.
The receipt, as written down by a Kandyan, was sent to me from
Kornegalle by Mr. Morris, the civil officer of that district ; and
in dramatic arrangement it far outdoes the cauldron of Macbeth's
witches. The ingredients are extracted from venomous snakes
by making incisions in the head of these reptiles and suspending
them over a basin to collect the poison as it flows. To this,
arsenic and other drugs are added, and the whole is boiled in a
human skull, with the aid of three KaVmra-goyas, which are tied
on three sides of the fire, with their heads directed towards it,
and tormented by whips to make them hiss so that the fire may
blaze. The froth from their lips is then added to the boiling
mixture, and so soon as an oily scum rises to the surface, the
kabara-tel is complete. Before commencing the operation of pre-
paring the poison, a ccxjk has to l)e sacrificed to the demons.

" This ugly lizard is itself regarded with such aversion by the
Singhalese that if one enter a liouse or walk over the roof, it is
regarded as an omen of ill-fortune, sickness, or death ; and in

* Sketclics of the Nat, Hist, of Ceylon, London, 1861.
VOL. VIII 2 X

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546 LACERTAE

order to avert the evil, a priest is employed to go through a
rhythmical incantation."

Captain Eobinson, renowned as a hunter of tigers on foot in
the old days of muzzle-loading rifles, has told me the following
unique use to which these large lizards are put by ingenious
thieves in India. In order to be able to get over a wall too high
for climbing without assistance, the thief provides himself with a
strong lizard, ties a rope round its waist and lets the animal go,
when it at once scales the mud wall by its strong and sharp
claws, and jumps down on the other side. The weight of the
lizard, which, moreover, holds vigorously on to the groimd, and

Fig. 140. — Varanm salvator swallowing a Fowl's egg. x |.

the friction of the rope on the top of the wall, are sufficient to
help the man over !

It is a sight, never failing in its attraction to the visitors of
the Zoological Gardens in London, to see one of the big Monitors
fed with an e^^^. The lizard knows the treat well that is
in store for it. It raises itself up high in expectation, then
examines the egg with the long tongue, takes it up gingerly
and swallows it entire, crushing it by the contraction of the
muscles of its gullet. On one occasion it was given a rotten
egg which burst in its mouth, and the lizard refused for a long
time to take another.

V. gouldi is common in Australia and in New Guinea. It
reaches a lencrth of a]>out 4 feet. Its colour is brown above

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XII XANTUSIIDAE TEJIDAE 547

with yellow spots on the back and limbs, and with yellow rings
on the tail. Two yellow streaks separated by a black band
extend from the temples along the side of the neck. The under
parts are yellowish, sometimes with black spots.

Fam. 9. Xantusiidae. — Three Central American, or West-
Indian genera, with less than half-a-dozen species. Pleurodont
vnth a short tongtie and with the supratemporal fossa roofed over
by bone. The tongue is scarcely extensible, with oblique over-
lapping folds which converge towards the median liue, and with
scale-like imbricate papillae towards the tip. The skull possesses
complete postorbital and postfronto-squamosal arches, the latter
meeting the parietals and roofing over the supratemporal fossa.
The palatines are in contact with each other, and there are no
infra-orbital fossae. There are no osteoderms; the body is
covered above with small granular scales, below with larger scales.
The eyes are devoid of movable lids. The tympanum is exposed.
Femoral pores are present. Limbs and tail well developed.
Xantusia and Lepidophyma,

YdJSL 10. Tejidae. — American Lizards with a long and bifid
tongue. The greater portion of the tongue is covered with
scale-like papillae; the anterior forked and pointed ends are
smooth. The teeth are solid and implanted almost upon the edge
of the jaw, being therefore intermediate between the acrodont and
pleurodont condition. The shape of the posterior teeth shows
several modifications ; they are conical or tricuspid, or molar-like
in the adult Tejus; iuDracaena they are transformed into large, oval
crushers. The palatines are in contact anteriorly. The infra-orbital
fossae are surrounded by the palatine, pterygoid, and ectopterygoid
bones, the maxillary being excluded from the fossa, as in Varanus
(see Fig. 138, p. 542). The skull has no supra-temporal arch.
Osteoderms are absent ; the body is covered with small scales,
or the skin is simply granular above; the under surface is
covered with larger scales, generally arranged in transverse rows.

This large family, which comprises nearly forty genera with
more than a hundred species, exhibits great diversity of form.
Some are inhabitants of forests and are arboreal, while others are
strictly terrestrial, preferring hot and sandy plains, or they dwell
below the surface and are transformed into almost limbless and
blind-worm-shaped creatures. The range of the family extends
over the whole of the South American continent, over the West

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548

LACERTAE

Indian Islands, and through Central America into the warmer
parts of the United States.

I'upinambis (" Teju "). — The skin of the back is smooth,
covered with small scales; with large scales on the top of the
head. The skin on the neck is generally thrown into two ir-
regular transverse folds. The long and narrow tongue is capable
of being telescoped into a sheath at its base. The lateral teeth
are compressed and tricuspid in the young, but the later genera-
tions of teeth have obtuse crowns in ' the adult. T. teguixin is

mcgoyri|ttTTL

Fio. 141. — Tvpinavihi^ ni^jropunctatus. x ^.

>T7«^«f

the largest member of the whole family ; it reaches a length of a
yard, most of which, however, belongs to the tail. The general
colour is bluish black, with pale or whitish -yellow spots on the back,
flanks, and tail, combining into more or less transversely arranged
])ands. The limbs are black, with many and tiny yellow dots.
The ground-colour of the under parts is reddish yellow, with
irregular black bars. This species inhabits the greater part of
Soutli America, east of the Andes, from Uruguay to the West
Indies. T, nigropunHatua is confined to the Continent, and lacks
the (lark cross-bands on the belly, which is uniformly yellowish
or s])eckled with black.

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XII TEJIDAE — LACERTIDAE 549

The "Tejus" frequent forests and plantations, and are
carnivorous. Their strength and swiftness enable them to
catch all kinds of animals, from insects and worms to frogs,
snakes, mice, and birds. As thej take chickens and eggs from
the farms they are considered noxious, and they are frequently
hunted down with dogs for the sake of their flesh, which is
regarded as good to eat. They defend themselves with lashing
strokes of their long tail and with their powerful jaws. They
retire into burrows, and they deposit their hard-shelled eggs
in the ground. In captivity they can easily be kept on meat.

Dracaena guianensis of the Guianas and the basin of the
Amazon has the lateral teeth transformed into regular large
molai:s, with broad and rounded crowns. The tail is strongly
compressed, with a double, denticulated keel. It seems to be
semi-aquatic, and, to judge from the teeth, herbivorous.

Ameiva and Cnemidophorus, with many species chiefly in
tropical America, have laterally compressed hi- or tri-cuspid teeth.
The skin forms a double fold on the neck, and is covered oh the
upper surface of the body with very small scales ; those on the
ventral surface are large, and arranged in regular rows. Most of
the species are small, under one foot in length, and are extremely
pretty, very active, timid, and mainly insectivorous.

C. sexlineatus is one of the few species of Cnemidojphorus which
inhabits the southern half of North America. Like all its
relations it has the appearance of an ordinary lizard {Lacerta), The
head is dark brown. A purple or brownish band extends over the
back and tail, bordered on either side with three golden-yellow
longitudinal lines. The flanks are brown, the under parts bluish
white. The iris is golden, and the inner margins of the lids are
bordered with a narrow band of bright yellow. This species is a
very fast runner, and frequents dry and sandy places. Its total
length amounts to about 10 inches.

Fam. 11. Lacertidae. — Pleurodont Old- World Lizards, with-
out osteoderms on the body, and vAth the supratemporal regions
roofed over hy osteoderms.

The limbs are always well developed, and have five fingers
and five toes, always provided with sharp claws. The skin
covering the head forms large shields, mixed with small scales ;
most of which, especially the shields, contain dermal ossifications.
These frequently fuse with the underlying bones of the top of

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S50

LACERTAE

the skull. The latter is always well marked off from the neck.
The postorbital arch is complete. The temporal region is com-
pletely roofed over by bones dorsally, chiefly owing to the size of

Fio. 142. — Skull and lower jaw of Lacerta iHridi^. A, Dorsal view ; B, ventral view ;
C, from the left side ; D, right half of the lower jaw, from the inner side, with some
of the pleurodont teeth. ±:,P, Ectopterj'goid ; F, Fr, frontal ; jt4^, jugal ; Zac,
lacrymal ; Max, maxillary ; iV, ^Vo, nasal ; iV^j, in B, inner narial opening ; Pal,
palatine ; Par^ parietal ; Prnxy preniaxillary ; Pr.f, prefrontal ; Pt/y postorbital ;
Pt./2, postfrontal ; Ptff^ pterygoid ; Q, quadrate ; &ang, supra-angular ; Sq, squa-
mosal ; To, vomer.

the postfrontal (Fig. 142, jt^^/g), which fills the space between the
parietal and the squamoso-postorbital bridge, thus abolishing the
supra-temporal fossa. The squamosal is very small, placed be-

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XII LACERTIDAE SSI

tween the postfrontal (pt.f^), the lateral occipital and the supra-
temporal. The large jugal and the quadrate are not con-
nected with each other. The columella cranii is well developed.
The infra-orbital fossae are surrounded by the palatines, pterygoids,
ectopterygoids, and maxillaries. The palatines and pterygoids
remain separated in the middle line. The pterygoids frequently
carry little teeth. The other teeth are typically pleurodont,
hollow, slightly curved, and bi- or tri-cuspid.

The skin covering the body, the legs, and the tail is devoid of
osteoderms. The scales on the dorsal surface vary much in size,
from large, strongly keeled scales to tiny granulations. Those
of the ventral surface are large, broader than long, and are
frequently arranged in regular transverse and longitudinal rows.
The tail, generally long and pointed, is very brittle. All the
sense-organs are well developed. The tympanum is exposed.
The tongue is deeply bifurcated, narrow, flat, and covered with
scale-like papillae.

Various Lacertidae, especially some of those genera which live
and dig in the sand, have a transparent disc in the middle of the
lower eyelid, so that they can see while the eye itself is pro-
tected. This is for instance the case in some specimens of the
Indian and African Ereviias. In the Indian genus Cahrita the
transparent disc is very large, and in Ophiops, which inhabits
sandy stretches from North Africa to India, the lower eyelid is
fused with the rim of the much-reduced upper lid, and forms a
large transparent window.

The Lacertidae or True Lizards comprise nearly twenty
genera, with about one hundred species, and are typical of
the Old World, being foimd in Europe, Asia, and Africa,
but not in Madagascar nor in the Australian region. They
are most abundant in Africa. Their northern limit coincides
fairly closely with the limit of the permanently frozen
under-ground. This is indicated in the map (Fig. 143) by the
dotted line. All the Lacertidae live upon animal food, chiefly
insects, and after them worms and snails ; but the larger lizards
take what they can master, frequently other lizards, and even
younger members of their own kind. Many of them love sugar,
which they lick, and all require water. They are all terrestrial,
preferring, according to their kind, such localities as yield
them their particular food. Sunshine and warmth make a

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5 52 LACERTAE chap.

marvellous change in the same individual, which on dull, raiuy,
or cold days lies in its hole, or shows only sluggish movements.
Tlieir sense of locality is great, or rather each individual inhabits
one place, of which it knows every nook and corner, cranny, tree,
and bush. It has its favourite hole to sleep in, a stone, the
branch of a tree, or a wall to bask upon, and when disturbed or
chased it makes with unerring swiftness for a safe spot to retire
into. The same lizard, when once driven away from its own
locality, seems to lose all its presence of mind, floundei-s
about, and is comparatively easily caught. Most lizards are
extremely curious, although shy, and this state of their mind can

= LACERTIDAC.

Fia. 143. — Map showing the distribution of the Lacertidae.

be made use of by those who want to catch them without injury,
and above all without getting the animal minus the brittle tail.
This safe way of catching lizards consists in taking a thin rod
with a running noose of thread at the end, in drawing the
latter over the lizard's head, and then raising it. The little
creature does not mind the rod in the least ; on the contrary, it
watches it carefully, and often makes for the thread. The boys in
Southern Italy have improved upon and simplified this mode of
catching lizards by bending the end of a wisp of grass into a
noose, and covering the latter over w4th a thin film of saliva.
The shiny film, Uke a soap-bubble, is sure to excite the curiosity
of the creature. The late Professor Eimer ^ refers to this practice

^ Organic Evolution. Translation, London, 1890.

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XII LACERTIDAE 553

as carried out by the children of two thousand years ago,
and he sagaciously explains that the beautiful statue of the
so-called Apollo Sauroctonos represents a boy who is in the act
of noosing the little lizard on the tree.

Lacerta. — A row of enlarged scales forms a distinct collar
across the ventral half of the neck, in front of the chest. The
scales on the back are much smaller than those on the tail, which
is long, round, and pointed. The digits have smooth, tubercular
lamellae on the under surface. Femoral pores are well marked.
This genus, with about twenty species, ranges through Europe,
Northern and Western Asia, and Africa north of the Equator.

i. vivipara, the Common English Lizard, has a very wide
range, through Northern and Central Europe and Siberia to the
Amoor country and the Island of Saghalien. It occurs through-
out Great Britain, even in Ireland, where it is the only species
of reptile, occurring, for instance, in the County of Meath and in
the south-eastern counties, e.g. Waterford. It does not occur
south of the Pyrenees or south of the Alps. The supra-ocular
and the supraciliary scales are in contact with each other, not
being separated by a series of little granules. Normally there is a
single postnasal and a single anterior loreal shield. The ventral
scales are arranged in six or eight longitudinal series, of which
the second series on each side from the median ventral line is the
largest. The coloration of this species is subject to much
variation. The general colour of the adult is brown or reddish
above, with small darker and lighter spots; many specimens
have a blackish vertebral streak and a dark lateral band edged
with yellow. The under parts are orange to red in the male,
with conspicuous black spots ; yellow or pale orange in the
female, either without or with scanty black spots. The newly-
born specimens are almost black. The males are slightly smaller
than the females ; males of a total length of 6 inches, and
females 7 inches long, may be considered rather large specimens.

This lizard is, as the specific name implies, viviparous, i.e, the
six to twelve young burst the eggs immediately after they
have been laid ; sometimes the mother has to retard the laying,
in which case the young are born free. The female does not
make a nest, but simply deposits her offspring on the ground
and leaves the young to their fate. For the first few days the
little ones, which scarcely measure three-quarters of an inch in

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S 5 4 LACERTAE chap.

length, remain almost motionless between leaves or in cracks of
the ground, and they do not take any food. They grow, how-
ever, quickly, living upon the remains of the yolk which has
slipped into their body. Their first food consists of Aphides and
similar tiny insects.

The Common Lizard prefers moist localities and is very hardy.
It extends northwards to Archangel, and in the Alps it ascends
to nearly 10,000 feet above the level of the sea. However, on
the approach of the cold season, in the month of October, it
withdraws into its winter quarters, frequently in company with
many of its own kind.

i. agiliSy the Sand-Lizard, has nearly the same wide range
as L. vivipara, except that it does not go so far north and does
not extend eastwards beyond Central Siberia. It is absent in
Ireland and Scotland, while in England it is restricted to the
southern half.

The characters which distinguish the Sand-Lizard from L.
vivipara are few, although the majority of the specimens of either
kind are very distinct in their coloration, and i. agilis is strictly
oviparous, depositing its eggs in the ground, imder leaves, in
heaps of weeds and similar places. The Sand-Lizard has usually
a single postnasal and two superposed anterior loreals, the three
shields forming a triangle. The temples are covered with flat
scales, two or three of which are enlarged and in contact with
the parietals, but there is no tympanic scale.

The coloration is subject to much variation, local as well as
individual. As a rule the Sand-Lizard gives the impression of
being striped longitudinally, the striation being caused by rows
of dark and white spots and patches along the sides of the back,
flanks, and tail. In the male a more or less pronounced green,
in the female brown and grey are the prevailing ground-colours.
A typically coloured male during the breeding season is grass-
green on the sides and suffused with green on the yellow under
parts ; the sides are dotted with black, with whitish eye-spots.
The under parts are spotted with black. The adult female is
brown or grey above, with large dark brown, white-centred spots,
which are arranged in three rows on each side. The under parts
are cream-coloured, with or without black specks. The yoimg
are grey-brown above with white, l)lack-edged spots ; the under
parts are whitish. Total length of the adult up to 8 inches.*

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LACERTIDAE 555

The male is a little smaller than the female but has a relatively
longer tail, a little less than half the total length.

The Sand-lizard is easily kept in captivity, and lives for years
if allowed a variety of food and proper places to hibernate in.
It pairs in the spring, in England in May or June ; the white,
parchment-like eggs, numbering five to eight, are hatched in the
following July or August

L, viridis, the Green lizard, inhabits Southern and Middle
Europe and South- Western Asia. The general colour of this
beautiful lizard is emerald-green above, changing into greenish
yellow on the flanks and into yellow on the beUy. The throat,
especially in the males during the breeding season, is blue. The
upper parts are frequently speckled with black. The young are
brown or green above with one or two yellowish lateral stripes,
which persist in some adult females. There are usually two
superposed postnasal shields. The semilunar collar on the neck
is well pronounced, and there is usually a distinct gular fold.
The tail is often very long, especially in the males, sometimes
nearly three-quarters of the total length, which in very large
males reaches 16 or 17 inches. The females do not quite reach
this length.

The Green or Emerald-Lizard prefers rocky localities, from
the sea-level, as for instance in Jersey, up to a height of
several thousand feet. It is extremely swift and can
climb trees, which it sometimes resorts to when chased. When
hard pressed it takes tremendous leaps down to the ground,
marvellously enough without injury to body or tail, which latter
is otherwise very brittle. They pair in the spring or early
summer after much fighting between the males ; the eggs, to the
number of about ten, are whitish and are deposited a month
later. The young are hatched after another four weeks.

This beautiful lizard does not keep well in captivity, although
it becomes very tame ; it eats meal-worms, snails, earth-worms,
and insects, especially butterflies, but it sickens after the first
winter even if it has been allowed to hibernate.

In Portugal and Spain L. viridis is represented by a slightly
different kind, L. schreiberi, the chief interest of which lies in the
fact that it approaches L. ocellata in several respects. The
occipital shield is large and is usually broader than the inter-
parietal. The dorsal scales are smaller, and there are eight well-

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5 56 LACERTAE

developed rows of ventral scalSs. Instead of being uniformly
green, the upper parts are usually spotted and vermieulated with
black ; sometimes, especially in the females, the black spots have
a white ocellus in the centre. The under parts are yellowish,
with or without black spots. The throat is blue. The young
look very different. They are olive-brown above with large
yellow, or bluish-white, black-edged ocelli on the side of the head
and body.

Other forms, perhaps of sub-specific rank, approaching L.
ocellata, occur in the Balkan Peninsula, where, for instance in
Dalmatia, the typical L, viridis attains its most beautiful de-
velopment.

L, ocellata, the Eyed Lizard, inhabits Spain and Portugal,

FiCf. 144. — Lacerta ocdlata (the Eyed Lizard), x {.

extending northwards into the South of France and into the
Eiviera, southwards into Morocco and Algeria ; these southern
forms (Z. ^Jrt^^r and L, tanfjitana) approach Z. viridis. The
Eyed Lizard is green or dark olive above, with black or yellowish
dots, which are sometimes combined into a kind of network
pattern. The under parts are uniformly greenish yellow. The
sides of the body are adorned with about two dozen blue, black-
edged spots or " eyes." The intensity of the blue and the depth
of the green ground-colour vary much according to sex, time of
the year, and state of health. Males during the breeding season
are most beautiful and brilliant. The occipital shield is broad ;
there are two superposed nasal but no tympanic shields. The
supraoculars are separated from the supraciliaries by a series of
granules. The collar is well marked, but not the gular fold.

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LACERTIDAE 5 57

The dorsal scales are minute and granular ; the ventral shields
are arranged in eight or ten longitudinal rows.

The " Eyed Lizard " reaches a considerable size, especially, the
males, which develop a very strong and thick head, and are
much more robust and powerful than the more slender females.
Old males reach a length of 2 feet, two -thirds of which
length belong to the tail ; but the latter varies much, even if it
has never been broken and renewed.

The Eyed Lizard keeps extremely w^ell in captivity, and in
this respect is unlike the Green Lizard. A case has been re-
corded of its living thirteen years. This species is very intelligent.
Although at first ferociously wild and biting furiously, these
lizards soon become tame anij take food regularly. One of my
own, a half-grown male from Xorthern Spain, about one foot in
length, made its home in a little niche of the greenhouse-wall,
whence it emerged regularly to take the offered food from my
hand. It soon knew the whole place thoroughly, making use of
the creepers whilst scaling up to its retreat, jumping over certain
gaps, descending to the ground at certain spots, basking on
certain stones, invariably in the same methodical way. In the
month of October it retires into the ground on the coolest side
of the greenhouse, and although the latter is well warmed, the
lizard remains invisible until the next February or March,
when on some fine day it is rediscovered basking upon exactly
the sfime stone where it had been seen five months before. The
only drawback in connexion with keeping this kind of lizard in
company with other creatures is their voraciousness ; since large,
fully adult specimens attack and eat any other small lizard,
slow-worm, or snake they can find. They also take mice. The
eggs are often deposited in hollow trees.

L. riiuralis, the Wall -Lizard, is very common in Southern
Europe, Asia Minor, and -Northern Africa. Northward it
extends into Belgium and into South Germany. In the Iberian
Peninsula it ascends up to 5000 or 6000 feet above the level
of the sea. This graceful little creature, with an average length
of 6 to 8 inches, is easily recognised by the series of granules
between the supraocular and supraciliary scales and usually by
having only six rows of ventral scales. The great variety in colora-
tion has given rise to the establishment of many races, varieties, and
sub-species. In the typical forms the upper parts are brown or

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5S8 LACERTAE

greyish, with blackish spots or streaks, sometimes with a bronzy
greenish sheen. The under parts are white, yellow, pink, or red,
either uniform or, especially in the males, with large black spots.
The lateral rows of ventral shields are frequently blue. The
colour-varieties are almost endless. One of the most noteworthy
is that described as var. coerulea by Eimer; this, confined to
the Faraglione Rocks near Capri, is blackish above, like the rock,
and sapphire -blue below. Similarly coloured specimens, var.
lilfordi, occur on some of the rocky islets of the Balearic Isles,

The Wall-Lizard deserves its name, since in the Mediterranean
countries there is scarcely a wall on which these active lizards
do not bask or run up and down, often head downwards, in
search of insects. They are oviparous. The hibernation is
short and not very deep, since these lizards can sometimes
be seen basking on sunny winter days before their regular
appearance in the early spring.

Psamviodromus, with a few species in South- Western Europe,
notably in the Iberian Peninsula and in North- Western Africa,
has no distinct semilunar collar, but has a short fold in front of
each arm. The back is covered with large, rhombic, strongly
keeled and imbricating scales. The lateral scales pass gradually
into the ventrals, which are smooth and arranged in six longi-
tudinal rows.

P. hispanicMs is bronzy brown above, with small black and
white specks, and with one or two longitudinal streaks on each
side. The under parts are white. Total length about 5 inches.
Although also found inland, this species prefers sandy dunes,
studded with prickly and scanty vegetation. It runs very fast
and digs itself rapidly into the sand when pursued. When
caught it either utters a faint cry like " tsi-tsi," or it feigns
death. The pairing takes place in June ; half-a-dozen eggs are
laid about eighteen days later, deeply imbedded in the warm
sand, and they are hatched in eight weeks. The eggs are said
to grow^ after they have been laid from 13 by 7 mm. to 17-
20 by 10-11 mm. The newly hatched little creatures
measure about 2 inches in length, more than half of which
belongs to the tail.

P. (Trojndosaura) algirus has the same range as P, hispani-
ens, but grows to 10 inches in length, and is much more

* Fischer, Zool. Garten. 1884, p. 38.

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Xii GERRHOSAURIDAE SCINCIDAE 559

beautifully coloured. The upper parts are bronzy brown with
one or two golden, dark-edged, lateral streaks ; the under parts
are whitish ; the male has one or more blue-eyed spots above
each shoulder.

Acanthodactyliis is distinguished by the laterally fringed
digits. This genus ranges throughout Northern Africa to the
Punjab. One species, A, vulgaris^ extends into Spain and
Portugal The dorsal scales are small and almost smooth, but
those on the tail are strongly keeled; the ventrals are much
broader than long, and are arranged in eight to ten rowa The
fringes on the digits are but feebly developed in the shape of
lateral denticulations. The adults are grey-brown with faint
longitudinal stripes, and with more conspicuous black and pale
spots ; in the breeding season larger blue-eyed spots appeeu: on the
sides near the limbs. The tail is often pink, especially on the
under surface. Total length about 7 inches.

Fam. 12. Gtorrhosanridae. — Pleurodont African Lacertidae
with osteoderms on the head and body.

This family is intermediate between the Lacertidae and
the Scincidae. The tongue is constructed like that of the
Lacertidae, but is only feebly nicked anteriorly. Dermal ossifi-
cations roof over the temporal region, and femoral pores are
present. On the other hand, the osteoderms, which cover the
whole body, are in their structure and arrangement typically
Scincoid. The tail is long and fragile. A lateral fold is
usually present. The limbs are sometimes reduced to useless
stumps. The few genera and species of this family are strictly
confined to the African sub-region, being found in the whole of
Africa south of the Sahara, and in Madagascar.

Gerrhosatirus, with a strongly developed lateral fold and
complete limbs, occurs in Africa. G, Jlavigularis, of South Africa,
has a total length about one foot.

TetradactyluSy of South Africa, has also a strong lateral fold,
but the limbs are either very short and pentadactyle {T, seps),
or tetradactyle, or they are minute pointed stumps, as in T.
africanus.

Fam. 13. Scincidae. — Pleurodont lizards with strongly de-
veloped osteoderms on head and body, with very feebly nicked,
scaly tongue, with complete cranial arches, and with separated
premaxillaries.

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S60 LACERTAE chap.

The temporal region is covered over, as in the Lacertidae,
with strongly developed, bony, dermal ossifications. Similar
osteoderms imderlie the scales which cover the body and tail.
The tongue is relatively short, not forked behind, and but very
feebly nicked at the tip ; it is covered with scale-like papillae.
Femoral pores are absent.

All the Skinks prefer dry, sandy ground, in which they not
only burrow, but move quickly about, either for protection or in
search of their animal food. In connexion with this sand-
loving and at least temporary subterranean life stands the
frequent reduction of the limbs. Every stage from the fully
developed and functional pentadactyle limb to complete absence
of limbs is represented. There are species within the same
genus with five, four, three, or two fingers or toes. There are
Skinks without fore-limbs, but with vestigial hind-limbs, and
vice versa. The interesting point is that these reductions do
not indicate relationship within the family, but have happened
independently. They are impressive illustrations of convergent
retrogressive evolution.

AhlephariLS, widely distributed in the Old World, has the
lower eyelid transformed into a transparent cover, which is fused
with the rim of the reduced upper lid, exactly as in the Lacertine
genus Ophivps.

All the Scincidae seem to be viviparous, some of them, e.g.
2'rachysaurus, in the strict sense of the word, since the hard or
parchment-like egg-shell has been dispensed with.

The family contains about foiu: hundred species, which have
been arranged in nearly thirty genera, many of them on fanciful
grounds. The family is cosmopolitan, but reaches its greatest
diversity in numbers and forms in the tropical parts of the Old
World, especially in the Australian region, inclusive of the
islands of the Pacific, America, notably South America, has
the smallest number.

Trachysaurv.s, with one species, T, rv(/()sus, inhabits the whole
of the Australian continent. It is easily recognised by the large
and rough scales, and the short and broad stump-like tail. It
is dark brown above with yellowish irregular markings ; the
under parts are yellowish, marked with brown. Embryos of
this species have yellow transverse bands on the back, but these
often fade away before birth. The creature is .strictly vivi-

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SCINCLDAE 56 1

parous, the egg-membrane being very thin, and the two or three
embryos are ripened in uterus-like dilatations of the oviducts.
The period of gestation is about three months, and the birth
takes place, in South Australia, about April. According to
Fischer ^ this species, which is often in the market, is easily kept.
It requires warmth, sand and stones for basking, and water, in
which it soaks itself preparatory to the shedding of the
skin, which takes place half-a-dozen times in the year, and is a
slow process, requiring eight to ten days. The food consists
chiefly of worms, lizards, and snakes, but meat, cabbage, and
lettuce are also taken. The total length is about one foot.

.^^^

Fig. 145. — Trachysaurus rugosus, x^.

Cyclodus a Tiliqua, of Australia, Tasmania, and the Malay
Islands, has stout lateral teeth with spherical crowns. The
imbricating, cycloid scales of the body and the rather short but
pointed tail are quite smooth and shiny. C, gigas, of New
Guinea and the Moluccas, reaches a length of nearly 2 feet.
The general colour is brownish yellow, with broad, dark bands
across the body and tail.

Scincus, of North Africa, Arabia, Persia, and Sindh, has penta-
dactyle limbs, with laterally serrated digits. The eyelids are
well developed, but the ear is hidden under scaly flaps. S.
officinalis, of the Sahara and of Egypt, grows to about 8
inches in length. The snout is peculiarly shaped, cuneiform.
Tlie eyes are very small The scales of the body are perfectly
smooth ; the sides of the belly are somewhat angular. The

1 Zool. Gari, 1882, p. 206.
VOL. VIII 2 0

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562 LACERTAE

whole shape of the creature, the scales, and the digits are
adapted to burrowing and moving quickly through the louse
sand. The general colour is yellowish or brownish above, each
scale with small brown and whitish spots ; the under parts are
uniform whitish. The young are quite beautiful, being uniform
pale salmon-coloured above, silvery white below. When a little
older, yellow spots appear on the flanks and grey bands across
the back. These Skinks live in the absolutely dry reddish-
yellow sand of the desert, in which they may almost be said to
swim about, so swift and easy are their movements. They live

Fia. 146. — Cychidus gigas. x ^.

on insects, while in their turn they are eaten by snakes, and above
all by the Varanus lizards.

Of Mahuia with about forty species, in the whole of Africa,
Southern Asia, and in Tropical America, we mention only M.
(Fiq^repcs) vittata, on account of its partly semi-aquatic life, a
very rare condition among Scincidae. This creature, about
7 inches long when full grown, frequents damp localities in
Tunis and Algeria, where the French call it " Poisson de sable."
It often sits on the floating leaves of Nymphaea alba, and dives
into the water in order to esciipe. Its proper element is, how-
ever, the sand, and for the night it retires under stones. The
general colour is olive brown with a lighter vertebral band and
two narrow whitish lines on each side, sometimes edged with
black. The under parts are yellowish or greenish white.

Ckalvidcs s. Scps s. Gongylus, of the Mediterranean countries

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XII SCINCIDAE 563

also occurs in South-Western Asia. The lower eyelid has a
transparent disc. The body is much elongated, and is covered
with smooth shiny scales. The limbs are very short, or reduced
to mere vestiges.

C%. ocellatus, of the Southern Mediterranean countries, oc-
curring also in Malta and Sardinia, reaches about 10 inches in
length. The snout is conical, the ear-opening a small slit or
hole. The limbs have five fingers and toes. The tmder parts
are uniform silvery white, but the colour of the upper parts is
very variable, mostly olive brown with black spots and irregular
cross-bars, or with dark and light spots; sometimes uniform
bronzy brown with a light upper and a black lateral band.
This Skink seems to have no fixed abode, but digs itself into the
sand wherever it wants to hide. The skin is not shed in flakes,
but, as in most Skinks, it peels off by a process of gradual
desquamation. Fischer's specimens paired towards the end of
December. The gestation lasted 56 days, when nine young
were born, which measured about 75 mm. or 3 inches; when
three weeks old they had increased to nearly double this length.

Ch. lineaius, of Spain and Portugal, and of the South of
France, like Ch. tridactylus of Italy and Xorth-West Africa, has
only three fingers and toes. The fore-limbs are only about one
quarter of an inch in length in large specimens of 10 inches
total length ; the hind-limbs are a little longer. The general
colour is bronzy olive or brown above, in the former species with
nine or eleven darker longitudinal streaks ; uniform, and with an
even number of streaks in the latter species. Ch. hedriagaCy of
Spain and Portugal, has mostly five fingers and toes, and the
limbs are relatively longer in this smaller species ; but it is a
question if these and other species of this genus are not to a
great extent simply individual variations, since the reduction of
the limbs and toes seems to be a very recent feature. Ch,
guentheri, of Palestine, otherwise in every respect like Ch.
tridactyluSy but reaching a length of more than 14 inches,
has the limbs reduced to tiny conical stumps without a trace of
separate digits.

I have caught Seps accidentally under stones or pieces of
bark in sandy districts. On the western coast of Galicia and
Portugal, close to the sea, they frequent the gorse-bushes, on
which they can be seen basking, provided they are approached

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564 LACERTAE

stealthily. They disappear on the slightest alarm, almost
swimming, as it were, with great agility through the prickly
cover, and then hiding and wriggling through the loose sand
between the roots.

The following iSve " families " are composed of degraded
forms of various descent. Most of them lead a burrowing,
subterranean life, in adaptation to which the body has become
snake-shaped or worm-like. The fore-limbs are entirely absent,
except in Cliirotes ; the hind-limbs are absent, or reduced to
small flaps ; the girdles are reduced correspondingly. The skull
is devoid of postorbital, postfronto-squamosal, supratemporal, and
jugal arches. The quadrate bone is mostly immovable. The
eyes and ears are concealed, except in the Pygopodidae.

Fam. 14. Anel3rtropidae. — An artificial assembly of a few
degraded Scincoids. The worm-shaped, limbless body is devoid
of osteoderms. The tongue is short, slightly nicked anteriorly,
and covered with imbricating papillae. Columellae cranii are
present. Anelytroj^sis pajyillosus in Mexico. TypMosaurus and
Feyllnia in South and West Africa.

Fam. 16. Dibamidae, consisting of the genus Dibamus, with
D. novae-guiyieae, in New Guinea, the Moluccas, Celebes, and the
Nicobar Islands. The tongue is arrow-shaped, undivided in
front, covered with curved papillae. Columellae cranii are
absent. The vermiform body is covered with cycloid imbricating
scales without osteoderms. The limbs and even their arches
are absent, but in the males the hind-limbs are represented by a
pair of flaps. Total length of the animal about 6 inches.

Fam. 16. Aniellidae. — The genus AnieUa comprises a few
small worm- or snake-shaped species in California, which seem to
be degraded forms of Anguidae. The eyes and ears are con-
cealed, limbs are entirely absent, the body and tail are covered
with soft, imbricating, more or less hexagonal scales. The tongue
is villose, smooth, and bifid anteriorly. The teeth are relatively
large, few in numbers, recurved, with short swollen bases. The
skull, by reduction, approaches the Ophidian type ; there is no
columella cranii, the postorbital |irch is ligamentous, the pre-
maxillary is single, the nasiils and frontals remain separate, the
pre- and post-orbitals are in contact with each other, excluding
the frontal from the orbit.

A, 2mIcJira. — Silvery, the scales edged with brown ; back and

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XII ANELYTROPIDAE AMPHISBAENIDAE 565

tail with a narrow, brown, median line. Total length, 7 to
8 inches.

Fam. 17. Amphisbaenidae. — Worm-shaped lizards with the
soft skin forming numerous rings, each of which is divided into
many little squares, the vestiges of scales which are otherwise
restricted to the head. The eyes and ears are concealed. Limbs
are absent except in Chirotes, which has short four-clawed fore-
limbs. The pectoral arch, and still more so the pelvic arch, are
reduced to minute vestiges. The tail is very short. The skull
is small, compact, and strongly ossified, in adaptation to the
burrowing life, and is devoid of postorbital and postfronto-
squamosal arches and of columellae. The teeth are either acrodont

"3.
■i AMPHISBAENIDAE.

Fig. 147. — Map showing the distribution of Ami»his])aeni«lae.

or pleurodont. The tongue is slightly elongated, covered with
scale-like papillae, and bifurcates into two long and narrow
smooth points.

The Ampliisbaenas lead an entirely subterranean, burrowing
life, like earth-worms. They are frequently found in ants* nests
or in manure-heaps. Their progression is very worm-like, their
annulated soft skin enabling them to make almost peristaltic
motions and to move backwards as well as forwards. Tliey
crawl in a straight line, with slight vertical waves, not, like other
limbless lizards or snakes, by lateral undulations. The food
consists of worms and small insects. About one dozen genera
with more than sixty species are known, most of which inhabit
the warmer parts of America, the West Indies, and Africa. Four
inhabit Mediterranean countries.

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566 LACERTAE

If the tongue and the dentition be taken as indications of
relationship, the Amphisbaenidae may perhaps be considered as
degraded descendants of Iguanidae, a family which contains
various limbless, burrowing, worm-shaped forms. But it is also
possible that the Amphisbaenidae are not a natural group.
This consideration applies with most force to the genera Amphis-
haena and Ayioj^s, the various species of which occur in America
and in Africa.

Chirotes canaliculatus, the only species of the genus, is the
only Amphisbaenid which still possesses fore-limbs. These are
short, stout, placed close behind the head, and are provided with
four-clawed digits. This species occurs in Mexico and California,
is brownish or flesh-coloured, and reaches a length of about 8 inches.

Amphishaena, with nearly thirty species, in Tropical America
and Africa. On account of the short rounded-off head and the
almost equally blunt tail these creatures are called by the natives
** cobras de dous cabezas," i.e. snakes with two heads, or they are
known as " maes das formigas," i.e. mothers of ants, because of
their predilection for taking up their quarters in the nests of
ants or termites. The scientific name refers of course to their
capability of moving forwards and backwards (a/x<^t9, at both ends,
and ^alvco, walk).

A. fuliginosa, one of the commonest species in South America
and in the West Indies, is chequered black and white. The skin
of the body has about two hundred rings, the tail about thirty.
Total length between one and two feet. A more or less distinct
fold extends along each side of the body from the neck to the
tail, at the level where the dorsal scales originally joined the
ventral scales.

Blanus is the only genus of the Mediterranean province.
B. cinereus, of Portugal, Spain south of the Cantabrian range,
Morocco, and Algeria, reaches a length of 10 inches, but such large
specimens are rather rare. The general colour of the living
animal is pink with a brownish tinge and with minute grey
specks. The lateral lines or folds are well marked, and a stronger
transverse fold is placed behind the head. The body shows
from one hundred and ten to one hundred and twenty-five
rings, the tail from twenty to twenty-two ; each body-ring con-
tains about thirty little squares or remnants of scales. There are
a few pre-anal pores.

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AMPHISBAENIDAE — PYGOPODIDAE 567

I have sometimes found this species in Portugal whilst digging
for earth-worms in manure-heaps and similar moist places, where
they lead the same life as the worms except that they live uj)on
them and upon insects. When kept dry they become very thin
and shrunken, but when put back into moist soil they again become
turgid and supple within a short time. Those which I have
kept in glass jars filled with rich mould throve veiy well,
living upon the tiny insects and worms which infest such com-
post soil ; they dug long tortuous channels, in which they moved
forwards and sometimes backwards, but they never came to the
surface.

Fam. 18. Pjgopodidae. — Pleurodont, snake-shaped lizards,
without fore-limbs, but with the hind-limbs appearing as a pair
of scaly flaps.

The shoulder-girdle is much reduced. The liind-limbs,
although very small and hidden within the scaly, almost fin-
like flaps, still possess five toes. The ischium appears externally
as a small spur on either side of the anal cleft. The eyes are
devoid of movable lids, remaining open and unprotected ; the
pupil is vertical. The ear is either concealed or exposed. The
tongue is fleshy, slightly forked and extensible. The body is
covered with roundish imbricating scales. The tail is very long
and brittle. The few genera of this undoubtedly natural family
of unknown relationship contain in all about ten species,
restricted entirely to Australia, Tasmania, and perhaps New
Guinea. Next to nothing is known about their habits, except
that some of them eat other lizards.

Pygopus lejyidopus is distributed over the whole of Australia.
It reaches a total length of about 2 feet, 16 inches of
which belong to the tail. General colour coppery grey above,
sometimes with several longitudinal series of dark spots.

Lialis hurtoni of nearly the same size and equally wide dis-
tribution has the hind-limbs reduced to extremely small, scarcely
visible, narrow appendages.

Snb-Order 3. Chamaeleontes. — Acrodont Old- World Saurians
v:ith a laterally compressed body, j^^'^l^^f^sile tail , and urll-develojyed
limbs until the digits arranged in ojyposingj grasping y bundles of
two and three resptectively.

The Chameleons are an essentially African family. About
half of the fifty species known inhabit Madagascar, the others

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568 LACERTILIA CHAMAELEONTES

the African continent. One, the common Chameleon, ia North
African, extending into Andalucia; two others occur in South
Arabia and Socotra, and only one in Southern India and Ceylon.
This sub-order is well distinguished from all other Saurians
by several, mostly unique, characters. The tongue is club-shaped
and extremely projectile, to a length equal to that of the body.
L ^^ The head is usually described as

CO^rki^ forming a casque, with prominent

crests and tuberclea There is no
tympanum and no tympanic cavity.
The parietal bones, united into one,
extend backwards far beyond the
occiput, and the tip of this projec-
tion is met by a much -elongated
... supratemporal bone, which, partly

^H CHAMAELEONTIDAE. /. i -fv ^v i u i *

FIG. 148.-Map showing the (Ustri- f^scd With the Squamosal, helps to
bution of Chameleons. enclosc a hugc supratcmporal fossa.

The latter is widely open behind. The postfronto- squamosal
arch and the postorbital arch are strong. The jugal is widely
separated from the quadrate ; the latter stands vertically and is
not reached by the pterygoid. There is no columella cranii.
The pre- and post-frontals often join to form a supra -orbital
roof The nasals are verj^ small and are excluded from the nares,
which are bordered entirely by the enlarged prefrontals and by
the maxillaries. The premaxillaries are small and carry no
teeth. The latter are acrodont, compressed and tricuspid, and
are restricted to the maxillaries and mandibles.

Tlie limbs are peculiar. Not only are they relatively long and
very slender, but two digits are permanently opposed to the other
three. On the hand the first three fingers form an inner bundle
opposed to the outer, or fourth and fifth fingers. On the foot the
inner bundle is formed by the first and second, the outer by the
other toes. The shoulder-girdle is of the ordinary Saurian tyi)e,
but there are no clavicles and no interclavicle. The costal stei;pum
is well developed ; the ribs posterior to those which meet the
sternum are very thin and elongated: they meet and fuse with their
fellows in the medio-ventral line. These hoops are not connected
with their neighbours in front or behind. The tail is prehensile
by being rolled downwards ; it is not brittle and is incapable of
being renewed. The skin is not covered with scales, but with

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ANATOMICAL STRUCTURE

569

granules. The eyes are very remarkable. The eyeballs them-
selves are large, but the eyelids are united into one fold with
a small central opening. However, when the Chameleon is asleep
the margins of this opening sometimes become more slit-like.
The right and left eye can be, and are incessantly, moved
separately from each
other, and the creature
squints terribly. Each
eyeball, together with
the pin-hole eyelid, is
rolled up and down,
backwards and forwards,
independently of the
other eye. This is a
unique fciiture, but it
also occurs in people
who squint badly. The
question " What, and
how, do these creatures
see ? " is therefore quite
idle, especially since in
reptiles binocular vision
does not exist at all and,
consequently, cannot be
disturbed by squinting.

The tongue has
attained an extraordin-
ary development. The

tongue proper (Fig. 152) fig. 149.— a. Dorsal, B, ventral, and C, lateral view
is club-shaped and is of the skuW of Chanweletm vulgaris, x 1. Co^id,

, -.t- ' .1 occipital condyle ; J5^/', ectopterygoid ; y?^, jugal ;

covered with a sticky i,ic, lacrymal ; P(a, palatine ; Par, parietal ; Prf,

secretion. The base or prefrontal ; Pt./, postfrontal ; Pt(j. pterygoid ;

Q, qua«irate ; Sq, squamosal ; To, vomer.

root of the tongue is very

narrow, composed of extremely elastic fibres, and is supported by
a much-elongated copular piece of the hyoid. The elastic part
of the tongue is, so to speak, telescoped over the style-shaped
copula, and the whole apparatus is kept in a contracted state like
a spring in a tube.

A pair of wide, very elastic blood-vessels and special elastic
bands extend from the base into the thick end of the tongue.

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570 CHAMAELEONTES chap.

By rapidly filling the apparatus with blood, and by the action of
certain hyoid muscles, the spring is, so to speak, released, and the
momentum gained by the thick and heavy club-shaped tongueproper
projects it far out of the mouth. The sticky end of the club shapes
itself into an upper and a lower flap, which partly envelop the
prey, and the elastic bands of the far-stretched stalk withdraw the
whole. The detailed working of this ingenious shooting apparatus
is not easy to follow. An ordinary full-grown Chameleon can
shoot a fly at the distance of 7 or 8 inches. The whole
performance is very quick, lasting less than one second. "When
the desired object is very near, only 2 or 3 inches off, the
Chameleon has a certain difficulty in shooting its prey. The
tongue is at first put out slowly, tentatively, the following jerk
is feeble, and it seems as if the apparatus refuses to w^ork unless
it is allowed to shoot out with full force.

Another remarkable and quite proverbial feature of Chameleons
is their changing of colour. This is by no means restricted to
Chameleons, which indeed are rivalled in this respect by various
other lizards, for instance by the Indian Agamoid Calotes and by
the American Ameiva.

The microscopical structure and mechanism of the colour-
changing apparatus is, in Chamaeleon vvlgaris, as follows : —

The epidermis is colourless, and the Malpighian layer is not
particularly modified except that in it are imbedded some iri-
descent cells, with very minute wavy striation on their surface?.
The cutis contains in its leathery tissue a great number of small
and closely packed cells, filled with strongly refractive granules,
chiefly guanine-crystals. These cause the white colour by diffuse
reflection of direct light. The cells nearer the surface are charged
with oil-drops and appear yellow. Large chromatophores are
imbedded in the white granular mass, most of them with blackish-
brown, others with reddish pigment, the granules of which are
shifted up and down, towards and away from the surface of the
cutis, in ramified branches of the chromatophores. When these
branches are contracted the pigment is conveyed back into the
bulbous basal portion of the chromatophores and the skin api)ears
yellow or white. When all the pigment is shifted towards the
surface of the cutis, the animal looks dark, sometimes black. In
intermediate conditions the light is changed into green by
diffraction through the yellowish upper strata and by the finely

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CHANGES OF COLOUR

S7I

Fig. 150. — Diagrammatic section through
the skin of a Chameleon. Highly
magnified. C, deeper portion of the
cutis ; CK three chromatophores, in
various stages of contraction, filled
with black, brown, or reddish pig-
ment ; Ey epidermis ; IF, white layer
of granules ; 1', yellow layer of cells.

striated iridescent cells of the Malpighiau layer. Those parts into
which the chromatophores do not send pigment appear as yellow-
spots. The chromatophores are to a great extent under control
of the will of the Chameleon,
but external stimuli, as heat and
cold and other reflex actions,
also play a great part in their
movements.

For further information on
this subject see Briicke/ P. Bert,^
Pouchet,^ Thilenius,* and lastly
Keller,^ who has written a very
long but rather confused account.

The process of moulting is
curious. When the Chameleon
is in good health the whole
process is accomplished within a
few hours. The skin to be cast
off becomes loose and assumes a
blistered appearance. Sometimes the creature looks as if it were
wrapped up in white, semi-transparent tissue paper. By rubbing
against stones, or between the twigs of trees, the skin comes off
in large flakes, first on the lips, then on the contorted body, and
last on the under surface of the hands and feet. During a rapid
and successful moult the changes of colour go on as usual in the
new skin. Sometimes large flakes of the old skin remain
adherent for days, especially on the top of the head. The
moulting takes place several times in one year. One of my
Cli. vulgaris moulted in January and September, and then not
until June of the next following year. A Ch. pumiius moulted
in the months of May, October, and March.

When they know themselves to be discovered. Chameleons
make themselves as thin as possible by compressing the body or
rather the belly. This is done by means of the peculiarly
elongated abdominal ribs described above. The whole body is
then put into such a position that, by presenting only its
narrow edge to the enemy, it has become as little visible as

» Denk. Ak. Wien. iv. 1852. ^ c*. R, Ass. Franc, Ixxx. 1876, No. 21.

^ J. dc Vanat. physio!, viii. 1872, p. 401.

* Morphol. Arbeit, vii. 1897, p. 515. ^ Arch. Physiol. Ixi. 1895, p. 123.

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572 CHAMAELEONTES

possible. At the same time the Chameleon turns round upou
its twig, so that the latter comes to stand between the observer
and its own body, which may thereby be completely hidden. When
angry, the creature either presents its broadest surfa^, swaying
.to the right and left, or it blows itself up and hisses. The lungs
are very capacious, and, instead of being bag-shaped, end in
several narrow blind sacs which extend far down into the body-
cavity, so that not only the chest but the whole body can be
blown up.

The usual mode of propagation is by means of eggs, but a
few species allied to Ch. pumilns are viviparous. The time of
incubation and of gestation is long. For instance, the pairing of
Ch. vulgaris takes place in the month of August. The eggs are
laid in the last week of October, about fifty to sixty days later.
Sometimes, however, the eggs are retained much longer, since I
have received specimens with ripening eggs in July which did
not lay until the end of October. The eggs are deposited in the
ground and are not hatched until the following February or
March, i.e. about 130 days later. The new-born little creatures
are snowy white, and cannot change or rather assume colour
until after the second week.

All Chameleons are insectivorous and require enormous
quantities of food, which must be alive to be taken. Most of
them prefer Orthoptera, e.g. Locusts and Grasshoppers, and
Lepidoptera. They also eat flies, meal-worms, and cockroaches,
but their tastes differ not only individually but also temporarily.
They require change of diet. One individual will take cock-
roaches greedily, whilst another of the same kind will rather
starve itself than touch one. The same applies to meal-worms.
It is a great but common mistake to suppose that Chameleons
do not require water. On the contrary they drink regularly and
often, generally by licking up drops of water or by scooping them
up with their lips, shoving the snout along the edges of wet
leaves. It is not too much to say that most Chameleons are
short-lived in captivity on account of the want of water. Those
which are sold by the dealers are generally in a parched
condition. Sprinkling the twigs or leaves of their cage with
water works a wonderful change in them ; the dull, apathetic-
looking creatures drink and drink, revive, assume brighter colours,
and will soon take food, which they have until then refused

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CHAMAELEONTIDAE 573

obstinately. Once I have even seen a Chameleon, when put
into the gi^enhouse, make straight for a tank and actually
drink in gulps.

After they have fattened themselves in the autumn, Chame-
leons, at least thoSe of Xorth Africa, withdraw to hibernate in
the ground. But nothing is known about how, when, and
where they do this, nor is it known if tropical species aestivate
during the dry season.

Chameleons are notoriously difficult to keep successfully,
whereby we do not mean the keeping for three to six months.
This is easy enough, since it takes them several months *
to die of starvation. The difficulty is to keep them through
the winter. To enable them to do this, it is absolutely necessary
to fatten them up during the summer and autumn. Otherwise,
although kept in a warm place, they are liable to lose their
appetite in the autumn, when they become restless, probably with
the desire to hibernate. Those few individuals which get over
this critical period, say during the month of October, and do not
refuse food, are probably safe. But those are doomed which
refuse to eat meal-worms or cockroaches or such food as can be
procured easily during the winter.

The origin of the Chameleons is unknown. They form only
one family, Chamaeleontidae, with between fifty and sixty species,
which, with a few exceptions, belong to the genus Chainaeleon.

Ch, vulgaris is the Common Chameleon of North Africa, Syria,
and Asia Minor. It occurs also in a few parts of Southern Andalucia,
for instance near Jerez, and near Nerja, to the east of Malaga, where
it has possibly been introduced. A series of conical, slightly
enlarged granules forms a little crest on the median line of the
throat. A whitish line, which does not change colour, extends
from the chin to the vent. The rest of the skin, with the
exception of a median dorsal series of slightly enlarged tubercles
on part of the back, is composed of small granules. A small
but distinct lobe of leathery skin extends along either side of the
occiput towards the posterior end of the median parietal crest.
Dead or spirit-specimens are usually pale yellow ; living ones are
greenish, usually with differently coloured patches on the sides.
Exceptionally large males reach a total length of about 9 inches,
females reach the length of perhaps a foot, but about half
of the total length belongs to the tail.

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574

CHAMAELEOXTES

CHAP. XII

It is impossible to Siiy what is the cmIoiit of this Chameleon,
since the same specimen may within a few days appear in half-a-
dozen different garlis, not counting minor combinations of colour.
After it has Ijeen watched for several months, when all its
possibilities seem to \*e exhausted, it will pro}»tibly sur|)rise us
by a totally new combination. Xot every S|»ecimen changes
alike : 8<^)nie keep the same apj)earance for a long time, others
change often ; some are partial to sj>ecks, others to large patches.

Fl';, I'll. — f'h'iiiuU'ti'H r\il'ntri<. x S.

In the group of Chameleons shown in Fig. 152 several of
the more usual arrangements of colour have been indicated by
stij^pling and various kinds uf cmss-hatching.

A represents the usual coloration at night. The whole
animal, which has just been stirred up from its sleep in the
dark, is cream-coloured, with iiTt-gular pitches of yellow on the
head, the back, the sides of the l>ody, the legs, and the tail

B has the usual coloration: grey-green, with inniunerable small
darker specks, with two series of pale brown patches on the sides
<if the body, and with one patch on the region of the ear.

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/ Oi fJX>^ rt^on;. ^| . reJ^-htwn m-hlue . -^fefi^iij^ green r^ytHfitr

Fig. 152. — Showing changes of colour in Chameleons. A to D, Chamaeleon vulgaris
(see p. 574). Ckamaeleoii immilm in the right upper comer.

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5 7^ CHAMAELEONTES

C is the same specimen in an excited frame of mind;
it is represented in the act of shooting a fly. The light brown
patches have changed to maroon brown ; and many round golden
yellow spots have appeared on the green parts.

D shows a specimen, coloured like C, within a few
seconds after it has been put into an angry mood, in the present
case by having its tail squeezed. The whole body is blown out,
the thick tongue causes the throat to bulge out, and all the
yellow spots have become blackish green.

Many small spots scattered over the body are usually a sign
of anger. One of the specimens described above was, when fast
asleep in a dark room, dirty white, with about two dozen large
and small round spots of a rich yellow on each side of its body.
Then a lighted lamp was brought into the room without in any
way disturbing the animal. Within sixteen minutes the yellow
spots had vanished completely; the whole body and tail had
become suffused with greenish yellow, which gradually turned to
pale yellowish green, and those parts which in Fig. B are pale
brown, were just distinguishable as pale yellowish-white regions.
The Chameleon was found to be fast asleep, and it kept this
coloration during the rest of the evening. Other specimens
behaved on similar occasions in the same way, but the greatest
interest is attached to the fact that frequently only that side of
the body " greened up " which happened to be exposed to the
light, whilst the opposite side remained whitish. These changes
are not absolutely unconscious ; they are, after all, under the
control of the creature. In order to test the possibility of direct
action of the light, I have taken the precaution of throw^ing the
light of a candle only upon the body, whilst the head was kept
in darkness. No changes of colour took place whilst the animal
was asleep, but when a little light was allowed to sweep across
the closed eye, this soon began to twitch, and although the
creature did not open the eye, the usual changes of colour began
to take place. When the light was removed, the animal soon
re-assumed its whitish appearance. Artificially coloured light, for
instance green, red, or blue glass or paper, has apparently no
influence upon the changes of colour. The Chameleons behave
as they would behave under ordinary conditions. Direct and
hot sunshine however causes them to darken, sometimes to
turn uniform dull black, except for the white median ventral

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xn CHAMAELEONTIDAE 577

line. Occasionally I found one of the specimens described above
deep maroon brown, with dozens of round orange spots. Blue
and red do not seem to be within the range of Ch. vulgaris^ but
the combinations of green, yellow, brown, black, and white, with
their various shades, are almost endless. Sometimes the Chame-
leons do not turn pale during the night, but remain more or less
dull green, with or without brownish patches. Adaptation to
their immediate surroundings takes place to a very moderate degree
only, but as a rule they are brightest, especially in their green
tints, when they are allowed to sit amongst green foliage. The intro-
duction of a branch with fresh leaves generally has a brightening
effect upon those which have previously been confined in a cage
with dry twigs only. Cold does not necessarily make them pale,
but they appear duller, and the changes take place more slowly.
After all, Linnaeus has summed up the little we really know
about the causes of these changes, in the following terse sentence :
" Vivus varios colores assumit secundum animi passiones, calorem
et frigus."

Chameleons are not very amiable. When taken up they
blow themselves out or they bite painfully, and it is a long
time before they are tame enough not to go through various
antics of anger when one approaches them. When taken in
the hand they produce a peculiar faint grunting noise, which,
however, can be better felt than heard. They quarrel much
amongst each other ; and the males, during the pairing season,
are particularly ill-tempered. Each individual selects its own
particular branch to sleep on, if possible a horizontal one,
upon which it crouches down lengthwise, with the head and
belly resting upon the branch. The tail generally makes a turn
round another branch, and the four legs, grasping some support-
ing branch, are put into any, sometimes into an almost incredibly,
awkward position. Although they climb about a good deal
during the daytime, they generally resort to their accustomed
sleeping branch, and they defend this vigorously against would-
be intruders.

Chameleons are most deliberate in their movements, some-
times provokingly slow. Each arm and foot leaves the firmly
grasj)ed ])rancli with great hesitation, and makes with equal
deliberation for some other foothold. It does not matter if the
thigh appears almost twisted out of its joint. The creature will
VOL. VIII 2 p

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578 Cll AM AELEONTES chap.

remain in the most uncomfortable position, forgetting, one might
think, to put one or morQ of its limbs down, but keeping them
instead in the air.

It is most interesting to watch them stalking their prey.
Suppose we have introduced some butterflies into their
roomy cage, which is furnished with living plants and with
plenty of twigs.* The Chameleons, hitherto quite motionless,
perhaps basking with flattened-out bodies so as to catch as many
of the sun's rays as possible, become at once lively. One of
them makes for a butterfly which has settled in the farther
upper corner of the cage. With unusually fast motions the
Chameleon stilts along and across the branches and all seems to
go well, until he discovers that the end of the branch is still 8
inches from the prey, and he knows perfectly well that 7 inches
are the utmost limit to a shot with his tongue. He pauses
to think, perhaps with two limbs in the air, but stability is
secured by a judicious turn of the tail. After he has solved the
puzzle, he retraces his steps to the base of the branch, climbs up
the main stem, creeps along the next branch above, and when
arrived at the 7 inch distance, he shoots the butterfly with
unerring aim. The capacity of the mouth and throat is
astonishing. A full-grown Chameleon will catch, chew, and
swallow the largest moth, for instance a Sphinx ligusiri. When
large, the prey is chewed, but the wings and legs are swallowed
with the rest. Occasionally these parts are bitten ofiF, especially
the prickly long legs of large locusts.

In water Chameleons are quite helplesa Sometimes they
inflate themselves, but they always topple over on to the side,
and the movements of their limbs are absolutely without any
definite purpose.

When the eggs are ripe, and this happens with the Common
Chameleon about the end of October, the female refuses to take
food, and becomes restless. One of my specimens searched
about probing the ground for about a week before she dug
a hole in some more solid soil. This took two days. In the
evening I found her sitting in the hole to the middle of her
body. On the following morning she was still there, but busy
filling the hole with soil and covering it with dry leaves. A
few eggs were lying about outside, two of which at least I
saw her taking up by the hand and putting them on the

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XII CHAMAELEONTIDAE 579

nest, which was found to contain some thirty soft-shelled eggs
closely packed upon each other. During the whole process she
was very snappy, and hissed much when approached. After that
she crept into the twigs as usu«d, but recused to eat, vomited at
once the artificially introduced food, became restless on the sixth
day, crawling about at the bottom of the cage, and died on the
following day. This is the usual fate, almost without exception,
of females after they have deposited their eggs in captivity. The
great number of eggs and their deposition naturally exhausts
them, and they probably want to hibernate at once. The eggs,
which are yellowish, long -oval, about half an inch long and
covered with a parchment-like shell, are very difficult to rear,
chiefly on account of the difficulty of regulating the moisture. They
shrink up when too dry, and they are very liable to become mouldy.
According to Fischer ^ the eggs can be hatched in a large flower-
pot with a layer of horse-droppings at the bottom, then a layer
of 6 inches of slightly moist soil, then the eggs, then another
6 inches of loose soil, with a glass plate covering the top,
securing at the same time ventilation. In this way he succeeded
in hatching several sets of eggs after 125 and 133 days
respectively.

Ch, ccdcaratus, the Indian Chameleon, is found in the southern
half of the Peninsula and in Ceylon, but it is far from common.
It much resembles Ch, vulgaris, but the male is distinguished by a
tarsal process or " spur," covered with skin, on the inner side of
the foot.

Ch. pumilus, the Dwarf Chameleon of South Africa, reaches a
total length of 5 to 6 inches. It has a well-marked, serrated
gular crest, which extends from the chin to the end of the neck.
The chest and belly are without a toothed line, but a strongly
serrated series extends from the occiput over the back and tail
(see the right upper corner of Fig. 152 on p. 575). A row of
enlarged tubercles or scales extends along the sides of the body.
The general colour is green, with a large and long patch of brick-
red on the sides ; small dots and spots of intense red are scattered
over various parts of the body. The changes of colour are rather
limited. At night the Dwarf Chameleon does not turn pale, but
generally keeps its colour. When they are very well the green
is quite saturated, and the large red patch on the side is

1 Fischer, ZooL Gart. 1882, p. 4.

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580 CHAMAELEONTES chap, xii

interrupted by several blue spots. When they are angry or
unhappy the red turns into dirty brown, and the green beconies
quite dull. Sometimes the whole animal turns dull black.

This pretty little species is relatively hardy, being, as a native
of South Africa, accustomed to cold nights. It does well in
an ordinary temperate greenhouse, where it will live for several
years, provided it has an ample supply of flies and meal-worms.
It is viviparous, the young being probably bom in the month
of March or April.

Ch, bijidus, of Madagascar, shows an extraordinary difference
between the sexes. The male reaches the great length of
16 inches, and develops two long rostral processes, which
extend forwards beyond the snout ; these processes are formed of
dense connective tissue, which ossifies in the adult, and they are
covered with scaly skin.

Ch. parsoni, likewise of Madagascar, is the giant amongst
Chameleons, reaching a total length of 2 feet. The male has
two large rostral processes which diverge upwards and outwarda

Brookesia, with several species in Madagascar, may be
mentioned on account of its stunted appearance. The tail is
much shorter than the body and scarcely prehensile ; the scales on
the soles are spinous. Total length only about 3 inches.

Rhampholeon, of tropical continental Africa, with several
species, is likewise remarkable for the stunted and dwarfed appear-
ance, and for the peculiar claws, each of which is furnished with
a second cusp which is directed downwards. The tail is much
shorter than the body. The total length of Rh, spectrum of the
Camaroons is about 3 inches.

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CHAPTER XIII

8A.URIA, continued OPHIDIA — SNAKES

Order H. OVEIDIA— SNAKES,

Saurians which have the right and left halves of the lower jaw
connected h/ an elastic hand.

The Snakes are the most highly specialised branch of the Sauria,
from which they do not differ in any fundamental characters.
The chief modifications consist in the absence of the limbs and
limb -girdles (a feature intimately correlated with the much-
elongated body), and in the swallowing apparatus. The reduc-
tion of the limbs and the elongation of the body also occurs in
many Lacertilia ; in several of the older families of Snakes {e.g,
Typhlopidae and Boidae) vestiges of the hind -limbs and even
of the pelvis are still in existence. Even the peculiar suspensorial
apparatus of the lower jaw approaches that of the Lacertilia
in the burrowing Ilysiidae and in Xenopeltis,

In the majority of the Snakes the quadrate is very loosely
suspended from the squamosal (by some authorities homologised
with the supratemporal bone of other reptiles), and this again
is loosely attached to the lateral parietal region of the skull,
placed horizontally, and elongated so far backwards that the
vertically placed quadrate lies in a plane behind the skull. In
most Snakes the elongated pterygoids are loosely attached to the
inner side of the distal end of the quadrates, and they also often
touch the mandibles. The whole palatal apparatus is movably
attached to the skull, except in some burrowing families. The
right and left pterygoids and palatines are widely separated from
each other. The pterygoids and maxillaries, connected by the
ectopterygoids, are absent, owing to reduction, in the Typhlo-

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S82 OPHIDIA CHAP.

pidae and Glauconiidae only. The premazilla is unpaired and
small, and is rarely furnished with teeth. The latter are always
sharp and recurved, and are lodged in sockets upon the edge of the
supporting bone, with which they become firmly ankylosed.
There is a perpetual succession of teeth. In the majority of
Snakes teeth are carried by the maxillaries, palatines, pterygoids,
and dentaries, rarely by the premaxillaries. The palatal teeth are
restricted to the palatines in Oligodon, Dasj/peltis, and Atractaspis
only.

Peculiar modifications prevail in the poisonous Snakes.
Those maxillary teeth which are at their base in connexion
with the openings of poison -glands (modified upper labial
glands), either have a furrow on the anterior side (Proteroglypha
if the anterior teeth are grooved, e,g, the Cobras ; Opistho-
glypha if some of the posterior teeth are grooved), or the groove
is converted into a canal, as in the Solenoglypha or Viperidae.
The special modification of the maxillaries of the vipers with
their long poison-fangs is described on pp. 587 and 637.

The orbit is generally closed behind by the postfrontal.
Quadrate -jugal, postfronto- squamosal, and other arches are
absent, so that the temporal fossa is quite open (see Fig. 156, .p.
597, and Fig. 155, p. 596). The occipital condyle is distinctly
triple. The mandibles are composed of several bones, but the
coronoid is absent in the Xenopeltidae, Colubridae, Ambly-
cephalidae, and Viperidae ; it is large in the Boidae, reduced to
a nodule in the Ilysiidae.

The parietals are always fused into a large unpaired bone,
which generally forms a sharp crest and partly overlaps the
occipitals ; there is no interparietal or pineal foramen.

The vertebral column consists of many, often nearly three
hundred vertebrae, and these skeletal segments correspond in
number with those of the ventral and transverse scales of the skin.
The vertebrae are procoelous; in addition to the anterior and
posterior zygapophyses they have a pair of accessory articulations
on the neural arches, dorsally to the zygapophyses ; — the " zygan-
trum" carried by the posterior end of the neural arches, its
articular surfaces looking upwards ; and the " zygosphene " carried
by the anterior end and looking downwards. Such accessory
articulations occur also in a few Lizards, e.g. Iguanidae. The
vertebrae of many Snakes have unpaired vertical, blade -like

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XIII ANATOMICAL STRUCTURE 583

haemapophyses on their centra for the more efifective attachment
of the muscles. All the vertebrae, except the atlas, carry ribs.
These articulate by their capitular portions only, and are very
movable in a head- and tail-ward direction. The ribs being long,
and fitting with their ventral ends into the connective tissue of
the sides of the ventral transverse scales, are the principal agents
in pushing the body forwards, the posterior edges of these scales
being sharp and imbricating.

The skin is covered with scales, absolutely devoid of osteo-
derms. When the scales are enlarged they are called shields.
The keel, a common feature, is caused by a slight ridge of
the cutaneous part of the scale. The whole skin is covered
with a thin layer of homy epidermis, which is shed frequently,
at least several times in one year ; the shedding begins at the
lips, and the whole outer skin is turned inside out from head to
tail, retaining every minute detail of the cutaneous scales ; even
the watch-glass-like covering of the eyes is preserved.

The eyes are peculiar in so far as they possess no lids. The
latter are still present in a vestigial condition in the embryo, but
their place is taken by what is probably a modification of the
nictitating membrane, which is drawn over the eye and
covered with a single transparelht scale of the horny skin, like a
watch-glass. The eyes themselves are not movable. The
'• tears," which of course cannot appear on the outside, are
drained off into the nasal cavities by the naso-lacrymal ducts.

The ear is likewise peculiar. There is a long columellar rod
with a fibrous or cartilaginous pad at the outer end, which plays
against the middle of the shaft of the quadrate, an arrangement
which, we must assume, produces a thundering noise in the
internal ear, since every motion of the quadrate during the act
of swallowing conveys the vibrations directly to the fenestra
ovalis. The tympanic cavity, the Eustachian tubes, and the
tympanum are abolished, and no external traces of the ear are
visible. However, in spite of all this. Snakes can hear very well.

The nose is well develoi)ed, and many Snakes, for instance
the Grass-Snake, are guided to their prey as much by the sense
of smell as by the eyes and ears. The tongue is slender, very
protractile and bifid, always moist, and furnished with many
sensory corpuscles. It acts entirely as an additional sense-organ,
hence the incessant play of the tongue of a snake which wants

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5 84 OPHIDIA

to investigate anything. In spite of the protractility of the
tongue, the hyoid apparatus is very small; the hyoid arches
themselves are reduced to mere vestiges near the base of the
first and only branchial arches, which are thread-like and extend
backwards down the throat.

The trachea is very long, and opens far forwards in the
mouth; it can be slightly protruded between the two
halves of the lower jaw so as not to be blocked during the act
of swallowing. This is a laborious process. The snake, having
got hold of its prey with its teeth, generally shifts it into the
most convenient position, in order to swallow the head first. One
half of the mandible is then pushed forwards, then the other
half; the recurved teeth afiford the necessary hold, and the
snake, little by little, draws its mouth-cavity, and later on itself,
over the prey. In fact, it literally gets outside it. Sometimes
with a large victim this process may last for hours ; the whole
mouth and head become painfully distended and the veins swollen
almost to bursting. The snake pushes the prey against a stone or
other obstacle, rests awhile quite exhausted, and begins afresh.
At last the bulk of the prey has passed the mouth, the skin of
the neck is stretched to the utmost, the scales being separated
by wide interstices, the ribs work spasmodically, the victim is
pressed into the shape of a sausage, and the deed is done. In
order to assist deglutition there is a great amount of salivation,
but the often-heard story that Snakes cover their prey with
saliva before they swallow it, is a fable, or based upon faulty
observation, snakes sometimes being forced to disgorge the
half-swallowed prey, which, in such a case, is covered with slime.
One of my tame snakes had swallowed a frog on my table when
a friend entered the room. The snake was frightened, jumped
on to the ground, striking it with its full belly, and thereby
hurting the frog, which squeaked loudly, whereupon the snake
reversed its mechanism and the frog hopped away, none the
worse for its terrible experience.

In correlation with the elongated narrow space of the body-
cavity the lungs are not equally developed, the left being much
smaller than the right. The latter is a very thin-walled, hollow
bag, and the posterior half or third scarcely contains any of the
honey-comb-like respiratory " cells," but acts merely as a reservoir
of air.

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ANATOMICAL STRUCTURE — DISTRIBUTION

58s

The cloacal arrangement is essentially the same as that of
the Lacertilia, but Snakes possess no urinary bladder. The
copulatory organs are stowed away beneath the skin in recesses
of the posterior lateral corners of the shallow cloacal vestibu-
lum. Each organ is generally bifurcated at the free end,
and furnished with little spike-shaped, but scarcely horny, ex-
crescences. On each side of the outer cloacal chamber, in botli
sexes, lies a roimdish gland with an offensive, vStrongly-scented
secretion; that of various Boas smells disagreeably sweet and
musky. The majority of Snakes lay eggs, but most of the
Viperidae and the thoroughly aquatic kinds, besides a few ter-

limittviPERINE. ^CR(n>U.INE. ^^ELAPINE.

Fig. 153. — Map showing the distribution of dangerously poisonous snakes.

restrial forms, are viviparous. The egg-shells are like parch-
ment, with very little or no calcareous deposit, so that they
are always soft ; many embryos are, however, provided with a
little " egg-tooth " on the tip of the snout.

Snakes are intelligent creatures ; some become quite affec-
tionate in captivity, but most of them are of a morose disposition,
and do not care for company.

The geographical distribution of Snakes has been dealt
with in detail in connexion with the various families. Un-
fortunately very few fossils are known. One of the oldest is
Falaeo2?his, of the • London clay (I^wer Eocene). Kemains of
Elapine and of innocuous Colubrine snakes have been found in
the Lower Miocene of Germany ; Crotaline forms are known from
the Miocene of Turkey and North America. All the Plistocene

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586 OPHIDIA

remains belong to recent genera. There are indications that the
Ophidia are a relatively young branch of Beptilia, essentially of
Tertiary date, but the foundations of the distribution of most
of the older families were laid in Miocene timea The older
families, notably those which still possess vestiges of hind-limbs
or of the pelvis, are circumtropical, e.g. Typhlopidae, Boidae.
The few survivors of the Glauconiidae are likewise circum-
tropical, with the exception of Australia. The Ilysiidae occur
in South-Eastern Asia and in tropical South America; their
offshoot the Uropeltidae are restricted to India and Ceylon.
The Colubridae and even many of their sub-families are cosmo-
politan. It is quite possible that the Opisthoglypha and Pro-
teroglypha are not natural groups, but that their respective
conditions have been developed on various occasions and in
different countries. The same applies more strongly to the
Viperidae, a further development of the Opisthoglyphous type.
To judge from their distribution, the Crotaline snakes were
possibly developed in the Palaearctic sub-region ; they spread all
over America, but they were debarred from entering either
Australia or Africa. The Viperidae, on the other hand, are
restricted entirely to the Palaeotropical region and to the
Palaearctic sub-region. The fact that no separating belt of
water existed for them between Europe and Africa, indicates
their being the most recently developed of poisonous snakes.
Madagascar is the only large country which, besides snakeless
New Zealand, enjoys a total absence of poisonous snakes of any
kind, while the Oriental is the only sub-region which suffers
from the presence of numerous species of every sub-family of
poisonous Elapine, Crotaline, and Viperine snakes.

Snake-Poison, — Many Snakes, belonging to diflferent families,
are poisonous, and unfortunately there is no external character,
easily ascertained, by which every poisonous snake can be
distinguished from a harmless kind. If the head is very broad,
this is probably due to the pair of poison-glands on the sides
of the head ; but many harmless snakes can flatten and broaden
their heads in a suspicious way, and, what is much worse, many
of the most poisonous snakes, for instance the Cobras, have a
head as smooth and as sleek -looking as the Grass- or Ring-
Snake, the most harmless of species. It so happens that, with
a few exceptions, for instance among the Crotalines and Vipers, no

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SNAKE-POISON 587

badly poisonous snake has loreal shields, i,e, a pair of shields
intercalated between the nasals and the preoculars, but this
character is obviously no good for any practical purposes. There-
fore, unless you know a snake well enough when you see it, leave
it alone, because a mistake may be fatal.

The poison is secreted in modified upper labial glands, or in a
pair of large glands which are the homologues of the parotid
salivary glands of other animals.^ A duct passes from the
gland forwards along the side of the upper jaw. Just in front
of the fang it doubles on itself, so as to open by a small papilla
on the anterior wall of the sheath of mucous membrane which
embraces the base of the tooth like a pocket. As mentioned
before (p. 582), the poison is conveyed either along a furrow on
the anterior side of the tooth, or the growing substance of the
tooth partly converts the furrow into a canal which opens only
near the end of the tooth. This is a perfectly devilish contriv-
ance, ensuring the conveyance of the poison into the very
deepest part of the wound. The Elapinae have relatively short
fangs, while those of the Vipers, and especially those of the Crota-
line snakes, are much longer, sometimes measuring nearly an inch
in length. The most formidable apparatus is that of the Viperidae,
since in them the maxillaries, each provided with only one
acting fang, and without any other teeth behind, can be erected.
The mechanism is explained in Fig. 154 and Fig. 179 (p. 647).
The apparatus of the upper jaw is so constructed that the
pushing forwards of the horizontal pterygoid bar will, by
acting on the ectopterygoid, rotate and erect the short majcillary.
The pulling forwards is effected by contraction of the spheno-
pterygoid muscle, which arises far forwards from the basal
orbi to-sphenoid region, and is inserted on to the inner dorsal
surface of the pterygoid. The principal closing muscles of the
mouth are the temporo- masseteric muscles (Fig. 179, T,a, and
7\p.) and the inner and outer pterygoid muscles, which latter
arise from the outer surface of the pterygoid bone, or from the
maxillary, and are inserted on to the articular region of the
mandible.

A strong ligament arises from the squamoso- quadrate
junction, and spreads fan -shaped upon the connective tissue

^ For a detailed anatomical account, see West, J. Linn. Soc, xxv. 1895, p.
419 ; xxvi. 1898, p. 517 ; and xxviii. 1900.

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588

OPHIDIA

wall of the poison-gland ; the anterior and posterior ends of the
gland are held by another strong band, which stretches from
the maxilla to the mandibular joint. The whole is so arranged
that the acts of opening the jaws (by the digastric muscles) and
the erection of the fang -bearing maxillaries are enough to
mechanically squeeze the contents out of the poison-gland. A

Fig. 154. — Explanation of the biting mechanism of a rattlesnake. la and 76,
position of the apparatus when the mouth is shut. Ila and lib, position of the
apparatus when the mouth is opened widely ; the spheno-pterygoid muscle {P,e) is
contracted, the pterygoid {Pt) is pulled forwards, the transverse bone or ectopterygoid
{Tr) pushes the maxillary (i/), rotates it and thereby causes the poison-fang {J) to
assume an erect position. Li^ Digastric muscle, contraction of which lowers, or
opens, the lower jaw ; G^ the groove or pit characteristic of the Crotaliue snakes ;
J, poison-fang ; 3/, maxillary ; i^ palatine ; P.e, spheno-pterygoid muscle ; Pm^
premaxillary ; Pt, pterygoid ; Q, quadrate ; <Sy, squamosal ; T,a, insertion of the
anterior temporal muscle, by contraction of which the mouth ia shut ; cf. Fig. 179
(p. 647) ; Tr, transversum or ectopterygoid ; X, origin and insertion of a muscle
and a strong ligament, contraction of which draws the maxillary and its tooth back
into the position of rest and assists in shutting the mouth.

portion of the anterior temporal muscle is attached to the
capsule of the poison-gland.

An excellent account of the nature and of the effect of the
venom of Snakes has been written by Charles J. Martin.^ The
following condensed account has been abstracted from it : —

" The poison is a clear, pale yellow, or straw-coloured fluid,
' Clifford Allbutt's System of Medicine^ vol. ii. London, 1896, p. 809.

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XIII SNAKE-POISON 589

which reacts acid, and contains about 30 per cent of solids, but
this varies much according to the state of concentration. Most
venoms are tasteless, but Cobra poison is said^to be disagreeably
bitter. Dried venom keeps indefinitely, and dissolves readily in
wat^r. It keeps also in glycerine. It contains albuminous
bodies in solution. The venom is, in fact, a pure solution of
two or more poisonous proteids, which are the active agents,
with a small quantity of an organic acid or colouring matter.
The venom is destroyed by reagents which precipitate proteids
in an insoluble form, or which destroy them, e.g. silver nitrate or
permanganate of potash. Hypochlorites have the same effect.
Carbolic acid and caustic potash destroy it only after a day
or two.

"The venom is generally introduced into the subcutaneous
tissue, whence it reaches the general circulation by absorption
through the lymph and blood-vessels. When introduced directly
into a vein, the effects are instantaneous. It is absorbed by the
conjunctiva, but, excepting Cobra poison, not by the mouth or
alimentary canal, provided there be no hollow teeth or no
abrasions. The venom of the various kinds of Snakes acts
differently.

"The symptoms of Cobra poison. Burning pain, followed
by sleepiness, and weakness in the legs after half an hour.
Then profuse salivation, paralysis of the tongue and larynx, and
inability to speak. Vomiting. Incapability of movement. The
patient seems to be conscious, but is unable to express himself.
The breathing becomes difficult. The heart's action is quickened.
The pupil remains contracted and reacts to light. At length
breathing ceases, with or without convulsions, and the heart
slowly stops. Should the patient survive, he returns rapidly to
complete health.

" The symptoms of Eattle-snake poison. The painful wound
is speedily discoloured and swollen. Constitutional symptoms
appear as a rule in less than fifteen minutes: prostration,
staggering, cold sweats, vomiting, feeble and quick pulse, dila-
tation of the pupil, and slight mental disturbance. In this
state the patient may die in about twelve hours. If he recovers
from the depression, the local symptoms begin to play a much
more important part than in Cobra poisoning: great swelling
and discoloration extending up the limb and trunk, rise of

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590 OPHIDIA

temperature and repeated syncope, and laboured respiration.
Death may occur in this stage. The local haemorrhagic ex-
travasation frequently suppurates, or becomes gangrenous, and
from this the patient may die even weeks afterwards. Recoveir
is sudden, and within a few hours the patient becomes bright
and intelligent

" Symptoms of bite from the European Viper. Local burning
pain ; the bitten limb soon swells and is discoloured. Great
prostration,' vomiting, and cold, clammy perspiration follow
within one to three hours. The pulse is very feeble, with slight
difficulty in breathing, and restlessness. In severe cases the
pulse may become imperceptible, the extremities may become
cold, and the patient may pass into coma. In from twelve to
twenty-four hours these severe constitutional symptoms usually
pass off, but in the meantime the swelling and discoloration
have spread enormously. Within a few days recovery usually
occurs somewhat suddenly, but death may occur from the severe
depression, or from the secondary effects of suppuration.

" Symptoms of bite from the Daboia or Vijpera russellL
These resemble the effects of Rattle-snake poison, but sanious
discharges from the rectum, etc., are an additional and prominent
feature. The recovering patient suffers from haemorrhagic
extravasations in various organs, besides from the lungs, nose,
mouth, and bowek. Kidney haemorrhage and albuminuria is a
constant symptom. The pupil is always dilated and insensitive
to light.

" Symptoms of bite of Australian Elapine snakes. Pain and
local swelling The first constitutional symptoms appear in
fifteen minutes to two hours. First faintness, and an irresistible
desire to sleep. Then alarming prostration and vomiting. The
pulse is extremely feeble and thread-like, and uncountable. The
limbs are cold, and the skin is blanched. Respiration becomes
shallow with the increasing coma. Sensation is blunted. The
pupil is widely dilated, and insensible to light. There is some-
times passing of blood. If the patient survives the coma,
recovery is complete and as a rule rapid, without secondary
symptoms. The Australian venom and that of all viperine
snakes, perhaps also that of the Cobra, if introduced rapidly into
the circulation, occasions extensive intravascular clotting. If
the venom is slowly absorbed, the blood loses its coagulability,

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XIII SNAKE-POISON 5 9 1

owing to the breaking down of the red blood-corpuscles, most so
with vipers, less with Australian snjtkes, least so with the Cobra.
The Cobra venom is supposed to extinguish the functions of the
various nerve-centres of the cerebro-spinal system, the paralysa-
tion extending from below upwards, and it has a special affinity
for the respiratory centre. The toxicity or relative strength of
the Cobra venom has been calculated to be sixteen times that
of the European Viper. Snakes can poison each other, even
those of the same kind.

" Treatment — Apply a ligature above, not on the top of the
situation of the bite; twist the string tightly with a stick.
Then make a free incision into the wound. Sucking out is
dangerous! Then bandage the limb downwards, progressing
towards the wound ; repeat this several timea Direct applica-
tion into the widened wound of calcium hypochlorite, i.e.
bleaching powder, is very good, or of a 1 per cent solution of
permanganate of potash, or Condy's fluid Amputation of the
finger is the best remedy of all if a large snake has bitten it.
Do not keep the ligature longer than half an hour. Then let
the circulation return, and apply the ligature again. In any
case, do not keep the ligature on for more than one hour for fear
of gangrene.

" Internal remedies, — The administration of enormous doses of
alcohol is to be condemned strongly ; small stimulating doses are
good, but stimulation can be more effectively produced by
ammonia or strychnia. Hypodermic injection of strychnine, in
some cases as much as one to two grains (but not into a vein !)
has in some cases had good results ; but injection of ammonia,
instead of doing any good, has disastrous sloughing results.
There is only one fairly reliable treatment, that by serum thera-
peutics, the injection of considerable quantities of serum of
animals which have been partially immunised by repeated doses
of snake-venom. Unfortunately this treatment will not often be
available."

Several well-known Mammals and Birds are immune by
nature against snake-venom, but most of them avoid being bitten.
Some birds induce the snake to strike and bite frequently into
their spread-out wings. Such more or less common creatures are
the Mongoose, the Hedgehog, and the Pig, the Secretary bird, the
Honey Buzzard, the Stork and probably other snake-eaters.

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592 OPHIDIA CHAP.

Olassification of Ophidia. — Bumeril and Bibron^ divided
Snakes according to their teeth into Opoterodonts, Aglypho-
donts, Solenoglypha, Proteroglypha, and Opisthoglypha.

J. E. Gray 2 divided Snakes into two sub-orders: Viperina
and Colubrinia. Giinther * distinguished between Ophidii colu-
brifomies, O. colubriformes venenosi (Elapidae and Hydrophidae)
and O. viperiformes. Cope * laid stress upon the modifications
of the squamosal, ectopterygoid, and endopterygoid bones, and
also upon the condition of the vestigial limbs. He divided the
snakes into Scolecophidia (Typhlopidae), Catodonta, Tortricina,
Asinea (the harmless snakes without limb-vestiges), Protero-
glypha, and Solenoglypha.

Boulenger^ has accepted Cope's principles, and, mainly by
combining the Asinea with the Proteroglypha as Colubridae, has
produced a logically conceived system, by far the best hitherto
proposed. It has been followed in the present work. Boulenger's
phylogenetic system stands as follows : —

9 Viperidae
5 Uropeltidae | 8 Amblycephalidae

7a C. Opisthoglypha 76 C. Proteroglypha I

4 Ilysiidae 7 Colubridae Aglypha

I 6 Xenopeltidae

1 Typhlopidae 3 Boldae 2 Glauconiidae

I. No ectopterygoid ; pterygoid not extending to quadrate or to mandible ;
no Bupratemporal (squamosal) ; prefrontal forming a suture with
nasal ; coronoid present ; vestiges of pelvis.
Maxillary vertical, loosely attached, toothed ; mandible edentulous ; a

single pelvic bone. Tijphlopidaej p. 693.
Maxillary bordering mouth, forming a suture with premaxillary, pre-
frontal, and frontal, toothless ; lower jaw toothed ; pubis and
ischium present, latter forming a symphysis. Glauconiidae, p. 594.
II. Ectopterygoid present ; both jaws toothed.

A. Coronoid present ; prefrontal in contact with nasaL

1. Vestiges of hind-limbs ; supratemporal (squamosal) present.
Squamosal large, suspending quadrate. Boidae, p. 596.
Squamosal small, intercalated in the cranial walL Ilysiidae^ p. 594.

2. No vestiges of limljs ; squamosal absent Uropeltidae^ p. 595.

^ Erp4tologic g^n&ale^ Suites d Buffon, vol. vii. Paris, 1852.

2 Catalogue of Stiakes^ British Museum^ London, 1849.

^ Reptiles of British Iiidia^ Ray 6ociety, 1864.

* P, Ac, Philad. 1864, p. 230.

" Caialogue of SnakcSf British Muscuviy London, 1893-1896.

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XIII CLASSIFICATION — TYPHLOPIDAE 593

• B. Coronoid absent ; sqiiamoBal present

1. Maxillary horizontal ; pterygoid reaching quadrate or mandible.
Prefrontal. bone in contact with nasaL Xenopeltidae, p. 605.
Prefrontal not in contact with nasal. Colubridae^ p. 606.

2. Maxillary horizontal ; pterygoid not reaching quadrate or man-
dible. Amhlycephalidaef p. 637.

3. Maxillary vertically erectile, perpendicularly to ectopterygoid ;
pterygoid reaching quadrate or mandible. Viperidae^ p. 637.

For ordinary practical purposes this synopsis is useless, being
based entirely upon anatomical characters, not all easily ascer-
tained. The following characterisation of families may therefore
be preferred : —

Eyes vestigial ; no teeth in the lower jaw ; without enlarged ventral scales.

Typldopidae,
Eyes vestigial ; teeth restricted to the lower jaw ; without enlarged ventral

scales. GlaxLconiidae.
Eyes very small ; head not distinct ; ventral scales scarcely enlarged ; tail

extremely short, ending obtusely and covered with peculiar scales.

Uropeltidae,
With vestiges of the hind-limbs appearing as claw-like spurs on each side of

the vent ; ventral scales transversely enlarged ; eyes functional, free.
Ventral scales scarcely enlarged. Ilysiidae.
Ventral scales transversely enlarged. Boidae.
With a pair of poison-fangs in the front part of the mouth, carried by the

otherwise toothless, much shortened, and vertically erectile maxillaries ;

ventral scales transversely enlarged ; eyes free. Viperidae,
All the remaining Snakes combine the following characters : the maxillaries

are typical, not separately movable, horizontal, with a series of

teeth.^ Tlie mandible is toothed but has no coronoid bone.

There are no vestiges of limbs or of their girdles. The eyes are free.
Dentary movably attached to the tip of the articular bone of
the mandible ; skin beautifully iridescent. Xenopeltidae.
Without a mental groove ; the ends of the pterygoids are free, not reaching

the quadrates. Amblycephalidae,
With a median longitudinal groove l^etween the shields of the chin ; the

squamosal is horizontally elongated, movable ; the pterygoid reaches

the quadrate. Coluhridae.

Fam. 1. Typhlopidae. — Burrowing snakes which have the
whole body covered with uniform cycloid scales, and with the
teeth restricted to the small and transversely placed maxillary
bones. The pterygoids do not extend backwards to the
quadrates, and there are no endopterygoids. The quadrates
slant obliquely forwards, and are attached directly to the pro-

^ Except Oligodon, Dasypeltis and Atractaspis (see p. 582), in which palatal
teeth are restricted to the palatines.

VOL. VIII 2 Q

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594 OPHIDIA

otics, owing to the absence of squamosal bones. The prefix)ntak
are in lateral contact with the nasals. There are vestiges of the
pelvis, reduced to a single bone on each side. The eyes are
hidden by shields of the skin.

The Typhlopidae, mainly composed of the genus Typhlops, with
about one hundred species, are undoubtedly the last living descend-
ants of formerly cosmopolitan, rather archaic, snakes, which in
adaptation to their burrowing life and insectivorous diet have
undergone degradation. They are still widely distributed in all
tropical and sub-tropical countries, some on the solitary Christmas
Island, but not in ifew Zealand. One species, T, vermicularis^
inhabits the Balkan Peninsula and South- West Asia. It is
brown above, yellowish below, and reaches a length of
about 10 inches. The tail is extremely short and ends in a
horny spine. T. hramimcs is widely distributed in Southern
Asia, the Malay Islands, the islands in the Indian Ocean and in
Southern Africa.

Fam. 2. Glauconiidae. — In most respects resembling the
Typhlopidae, but the maxillaries retain their normal position and
are toothless, teeth being restricted to the lower jaw, which is
stout and short. The pelvic girdle and the hind-limbs show the
least reduction found in any recent Snakes ; in the pelvis the
ilia, pubes, and ischia can still be distinguished, the last even
retaining their symphysis ; there are also vestiges of femurs.
About thirty species, nearly all belonging to the genus Glaiiconin,
are found in South-Western Asia, Africa, and the warmer parts
of America, including the West Indies.

Fam. 3. Ilysiidae. — The scales of the cylindrical body are
smooth and small, those on the ventral side are scarcely hirger.
The tail is extremely short and blunt. The head is very small,
not distinct from the neck. The gape of the mouth is very
narrow. Teeth are carried by the mandibles, the pterygoiils,
palatines, maxillaries, and one or two or more by the premaxillae.
The endopterygoids are short. An important cranial feature is
the short quadrates, which stand rather vertically and are con-
nected with the cranium by the squamosals; these are very
small and are firmly wedged in between the upper ends of the
([uadrates and the pro-otic, lateral, and supra-occipital fK)nes ; still
forming part of the cranial wall. Vestiges of the pelvis and
hind-limbs are very incomplete, and terminate in claw-like spm^

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XIII GLAUCONIIDAE — ILYSIIDAE UROPELTIDAE 595

protruding between the Scales on either side of the vent. The
eyes are very small, and are either free or covered by trans-
parent shields. The few, scarcely half-a-dozen, species are found
in South America {Eysia) and in Ceylon, the Malay Islands,
and Indo-China.

Ilysia {Tortrix) sci/tale, the Coral-Snake of Tropical South
America, is a beautiful coral-red with black rings. On account
of its beauty, perfectly harmless nature, and for "cooling
purposes," this snake, which grows to nearly a yard in length, is
sometimes worn as a necklace by native ladies. All the Ily-
siidae lead a partly burrowing life, live chiefly upon worms,
insects, and little Typhlopidae, and are viviparous.

Fam. 4. Uropeltidae. — Burrowing snakes of Ceylon and
Southern India, with a short and rigid cylindrical body and a
very short tail, which ends in a large peculiar shield, often
obliquely truncated. The scales of the body are smooth, and are
little larger on the belly ; the coloration is mostly very beautiful.
The eyes are very small

The Uropeltidae are somewhat intermediate between the
Ilysiidae, Glauconiidae, and Boidae. The pterygoids do not reach
the quadrates ; but ectopterygoids are present ; the quadrates are
veiy small and directly attached to the skull, squamosals being
absent. Teeth are carried by the mandibles and by the maxil-
laries, which are normal in their position. There are no vestiges
of hind-limbs or of the pelvis. The Uropeltidae, of which about
forty species are known, are viviparous, burrow in the ground,
and frequent damp localities, preferring mountain-forests. The
use of the characteristic tail -shield is not clear ; perhaps it
assists these rather rigid creatures in digging, by being pressed
against the ground.

Urojpeltis. — The tail is obliquely truncated, ending in a
roundish, flat shield.

U. grandis s. philippinus. — The latter name seems to have
misled W. Marshall ^ into including the Philippine Islands in the
range of the family, a mistake which is sure to be propagated.
The species, the only one of the genus, is confined to Ceylon ; it
is blackish above, yellow below, frequently with small yellow
spots above and brown spots on the under surface. It grows to
about 18 inches in length.

^ Atlas der Thierverhreitumj^ pt. v. Gotha, 1887.

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596

OPHIDIA

Rldnophis. — The tail -shield is convex and the snout is
pointed. Eh. sanguineus of Southern India is black above with
a bluish gloss, sometimes with small pale specks ; the belly and
several of the lateral series of scales are bright red, spotted with
black. The tail-shield is black and red.

Fam. 6. Boidae. — Typical Snakes, usually large, and with
vestiges of pelvis and hind-limbs, appearing externally as daw-

^>i

Fia. 155. — Skull of Eunectes muritius. x 1. The teeth on the maxillary, palatiue, ami
pterygoid have been omitted. Cdf Columella auris ; Cond, occipital condyle ;
E.P. and KPtg, ectopterygoid or transverse bone ; /', frontal ; Mandy mandible;
M(ij:y maxillary ; Xa, nasal ; Pal, palatine ; Par, parietal ; Pmx, premaxillary ;
Pr/f prefrontal ; Pt./, postfrontal ; Ptfj, pterygoid ; Q, quadrate ; Sq, squamosal ;
Tby turbinal.

like spurs on each side of the vent. The scales of the uppr
surface are usually small and smooth, while those of the ventral
surface form one broad series on the belly, and one or two rows
on the tail The quadrate is carried by the horizontally elongated
squamosal, which rests loosely upon the lateral occipital i-egion.
The prefrontal is in contact with the nasal. Teeth are carried
by the mandibles, the pter}'goids, palatines, maxillaries, and, in
the Pythoninae, by the premaxillaries also. For further details
see Figs. 155, 156.

The Boidae comprise between sixty and seventy species, which

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BOIDAE

597

have been grouped into many genera, on unimportant characters,
referring to the scales and shields of the head. It is doubtful if
they are natural groups, a consideration which detracts much
from their value in the study of geographical distribution.
Even the two sub -families are not free from this reproach.
The range of the family is world-wide, Boidae occurring in all
tropical and sub -tropical countries, including islands, except
New Zealand. A few species live in South -Eastern Europe

ptf.-

Fia. 156. — A, Ventral, B, dorsal, view of the skull of Eunectea murinua.
Lettering as in Fig. 155. x 1.

{Eryx) and in North-Western America. They mostly prefer
wooded districts, especially forests ; climbing trees, assisted by
the short and partly prehensile tail. Others are semi-aquatic,
and a few live in sandy localities. They are all rapacious, and
by preference feed on warm-blooded creatures, which they con-
strict by coils of the body in order to hold, kill, and crush the
victim before swallowing it. Exaggerated notions are enter-
tained about their swallowing capacity. It is obvious that a
large snake, 20 feet long, half a foot thick, and weighing
several hundred pounds, can crush a tiger, a stag, or even a

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598 OPHIDIA

cow ; but common sense tells us where to draw the line when it
comes to the swallowing of the prey. Small game, although of
a bulk apparently far too big for the snake, is so crushed and
mangled that it is tiu-ned into the shape of a sausage preparatory
to the long process of swallowing. The Boidae lay eggs, and
some species incubate them, or rather the female coils herself
round them for the sake of protection. No appreciable amount
of extra warmth is developed. Unfortunately the observations
of one of the best cases on record^ were conducted so imperfectly
that they are of little value.

Sub-Fam. 1. Pythoninae. — With a pair of supra-orbital
bones, intercalated between the prefrontal, frontal, and postfrontal
bones. The sub-caudal scales are mostly in two rowa The pre-
maxilla often carries a few small teeth.

The Pythoninae, comprising about twenty species, are
restricted to the Palaeotropical and Australian regions, with the
sole exception of Loxocemus hicolor in Southern Mexico.

Python, the principal genus, has teeth on the premaxilla.
The rostral, each of the anterior upper labials, and some of the
lower labial shields, contain a deep, probably sensory, pit. The
maxillary and mandibular teeth are long, but decrease from
before backwards. The head is distinct from the neck, and is
covered with symmetrical shields or with small scales. The
scales of the body are small and smooth. The tail is short
and prehensile ; below with two rows of scales. The pupil of
the eye is vertical. The range of the genus extends over the
whole of the Palaeotropical and Australian regions, excepting
Madagascar and New Zealand.

P. spilotes, the " Carpet Snake " of Australia and New Guinea,
is mostly beautifully marked, but is subject to much variation in
colour. The more typical specimens are black above, each scale
with a yellowish dot, with yellow spots or combinations of dots,
more or less arranged in rows. The under parts are yellow. It
reaches a length of about two yards, and spends a great part of
its time in trees.

P. reticulatus is the commonest species in Indo-China and in
the Malay Islands. Four upper labial shields of each side are
pitted. The specific name refers to the bold, dark, lozenge-shaped
markings upon the lighter yellowish or brown ground. A black

^ AV. A. Forbes, P.Z.S. 1881, p. 960.

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XIII BOIDAE PYTIIONINAE 599

line extends over the head from the nose to the neck, and
another on each side from the eye to the angle of the mouth.
The under parts are mostly yellowish, with small brown spots on
the sides.

This is one of the largest species of Python, some specimens
being known which measured about 30 feet in length.

As a sample of folk-lore connected wath this monstrous snake
the following Burmese fable has been recorded by Mason : — ^

" According to a Karen legend all the poisonous serpents derive
their virulence from the Python, which, though innocuous now.
was originally the only one that was venomous. In those days

Fig. 157. — Python spilotcs (the Carpet Snake), x i.

he was perfectly white, but having seduced away a man's wife,
Aunt Eu (Eve), he made her, while she was in his den, w^eave
figures on his skin in the forms which are now seen. At that
time, if he bit the footstep of a man in the road, such was tlie
virulence of his poison that the man died, how far soever that
man might have passed from the bitten track. The Python had
not, however, an ocular demonstration of the fact, so he said to
the Crow : ' Crow, go and see whether people die or not w^hen I
bite the foot-track.' The Crow went to the neighbourhood of a
Karen cabin, and found the people, as is their custom at funerals,
laughing, singing, dancing, jumping, and beating drums. He
therefore returned to the Python, and told him that so far from

^ Burma, its People and Prodvction$, London, 1SS2.

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6oo

OPHIDIA

his efforts producing death, on the contrary they produced joy.
The Python was so angry when he heard this that he ascended
a tree and spit up all his venom, but other creeping things caine
and swallowed it, and people die of their malignancy to this day.
The tree, therefore, from which the Python spat up his venom
became deadly, and its juice is used to this day for the purpose
of poisoning arrows. The Python made the other creatures
promise not to bite without provocation. The Cobra said : * If

iaillJi/rU'ni';.

A'fr^'iM/*

Fio. 158. — Python molurus. x yV-

there be transgression so as to dazzle my eyes, to make my tears
fall seven times in one day, I will bite.' So said the Tiger (whose
bite the Karens esteem as virulent as a serpent's) and others, and
they were allowed to retain their poison. But the Water Snake
and Frog said they would bite with or without cause as they
liked ; so the Python drove them into the water, where their
poison melted away and their bite became harmless."

P. molurus is the species of India and Ceylon, ranging,
however, also into Indo-China. Boulenger quotes W. Elliot ^

^ I^cp. Brit. Ass. 1870. Trans, p. 115.

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XIII BOIDAE — PYTHONINAE — BOINAE 6oi

as the authority for the statement that this species grows to
the length of 30 feet. Only two pairs of upper labials are
pitted The general colour above is greyish or yellowish brown
with a dorsal series of large reddish-brown, black-edged patches,
and on the sides of the body with a series of smaller spots with
light centres. On the head is a lance-shaped marking ; a brown
stripe passes from the eye backwards. The under parts are
yellowish.

P. sehae and P. regius are African species. The former has
two pairs of upper labials pitted, the latter four pairs. P. sehae
is generally pale brown above with dark brown, black -edged
cross-bars, which are usually coniiected by a sinuous dark stripe
along each side of the back. The upper surface of the tail has a
light stripe between two black stripes. The belly is spotted and
dotted with dark brown. .P. sehae ranges over the whole of
Tropical and Southern Africa, perhaps with the exception of
Eastern Africa. P. refjius of West Africa is beautifully marked,
and may be recognised by the dark brown, black-edged band along
the back, sending down triangular or Y-shaped processes on the
sides, which are pale brown. This dorsal band encloses a light
streak on the neck and another on the tail The belly is
yellowish.

These African Pythons grow to a length of about 15 feet, but
specimens so large as this are not often met with. The negi'oes
of certain parts of the coast of Guinea are said to worship them
and to keep them in special temples, where they are regularly
attended to. Their food consists chiefly of small Mammals,
notably rats, and of Birds. A couple of these snakes paired in
the Zoological Gardens of London in the month of June. The
female laid nearly one hundred eggs in the following January,
and incubated them until April, when the embryos were found
to be still unripe.

Sub-Fam. 2. Boinae. — Without supra-orbital bones. The
premaxilla is toothless. The subcaudal scales form mostly a
single row.

The Boinae comprise between forty and fifty species. Most
of them are American, but the genus Eryx inhabits North Africa,
Greece, and South-Western Asia; the genus Enygrus inhabits
New Guinea and many of the Pacific Islands, for instance New
Britain CNeu Poramern), the Solomon, Loyalty and Fiji Islands,

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602 OPHIDIA

and the New Hebrides. Casarea dussumieri is found on Kound
Island near Mauritius ; and two species of Boa and one of Corallvs
represent the Boidae in Madagascar, while all the others live in
Central and South America.

Boa, — The maxillary and mandibular teeth gradually decrease
in size. The scales of the upper parts of the body and tail are
smooth and very small. The rostral shield is enlarged. The
nostrils are placed between two or three nasals, and these are
separated from those of the other side by small scales. The tail
is short and prehensile. The pupil is vertical.

B. constrictor, of South America, has the head covered with
small scales, one of the pre-oculars being enlarged. The eye is
separated from the labials by several series of tiny scales. The
general colour is a delicate " pale brown above, with fifteen to
twenty dark brown cross-bars widening on each side, and, if con-
nected by a dark dorso-lateral streak, enclosing large elongate
oval spots. ... On each side is a series of large dark brown spots
with light centres, most of which alternate with the cross-bars.
On the tail the markings become much larger, brick-red, edged
with black, and separated by narrow, yellowish interspaces."
Under parts yellowish with black dots. Boa constrictor, a name
applied in popular parlance to many species, reaches a length of
more than 10 feet; the largest specimen in the British Museum
measures exactly 11 feet. A few other species inhabit Central
America and the West Indies. B. dumerili and B. madaffas-
cariensiSy both of Madagascar, cannot be separated from the genus
Boa.

A. D. Bartlett ^ has described the following incident : —

" In the evening of 5th October 1892 two pigeons were put
into the cage in which two fine specimens of Boa cort-strictor had
been living on friendly terms since the beginning of the year.
The larger snake seized one of tlie pigeons and the keeper left
the house. The next morning only one of the snakes, the larger
specimen, was visible, and from its enormously extended body
it was evident that it had swallowed its companion, which was
about 9 feet in length. It had no longer the power of curling
itself round, but remained extended nearly to its full length in a
straight line, and appeared to be at least three times its normal
circumference. It was almost painful to see the distended skin,

1 P.Z.S. 1S94, ]). 669.

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XIII BOINAE 603

which had separated the scales all over the middle of the body.
By 2nd November, twenty-eight days later, the snake had not
only digested its companion but had regained its appetite as well
as its normal size, and it immediately swallowed a pigeon put
into its den."

This peculiar case is not one of ordinary cannibalism. It is
rather an unintentional accident. When two snakes happen to get
hold of the same animal (in the present case a pigeon) and begin
to swallow it, the action of swallowing becomes almost mechanical,
the snakes continuing to push their jaws over the prey — which
in the case of a bird or mammal they cannot taste, nor can they
see it — so long as they feel something in the mouth. After the
original prey has been mastered, it is the turn of the opposite

Fig. 1.59. — Head of Eunedes murinits. xl.

snake's head, and if the weaker snake does not give way it is
swallowed by its stronger mate. Grass -Snakes will swallow
several frogs if these are tied together in a string, and other
snakes do the same with mice. There are instances on record
in which a Python swallowed its blanket, which, being absolutely
indigestible, caused its death.

Casarea, the " Eound-Island Snake," differs from Boa chiefly
by the rough and strongly keeled scales, and by the relatively
much longer tail.

Eunectes mu7Hnus, the " Anaconda," is an aquatic Boa. It
differs from this genus mainly by the inner of the three nasal
shields being in contact with that of the other side (see Fig.
159), and by the absence of the little scales between the eye and
the labials ; the snout is, moreover, covered with shields instead
of small scales. The pupil of the eye is normally vertical, but it
had contracted into a round pinhole in the dead but still fresh

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604 OPHIDIA

specimen from which the figure was drawn. The general colour
is dark olive-brown, with large oval black spots arranged in two
more or less alternating rows along the back, and with smaller
black, white-eyed spots along the sides. The under parts are
whitish, spotted with black. The upper parts of this and of
many other dark-coloured species of Boidae are often shiny, with
an iridescent lustre.

The Anaconda combines an arboreal with an aquatic life, a
kind of existence eminently in harmony with the well-watered,
dense forests of Tropical South America, which are the home of
this, the largest of all modern Snakes. It is said to attain a
length of as much as 33 feet. There is no inherent impossi-
bility in such statements, but the giant specimens seem to have
a knack of keeping out of the naturalist's way.

The Anaconda feeds chiefly upon Birds and Mammals, which
it catches either on land, mostly during the night-time, or in the
water. For the latter purpose it lies submerged in the rivers or
floats about leisurely, only the head being above the surface, and
anything suitable is attacked. In other localities the snake, if
so inclined, establishes itself upon the branches of a tree which
overhangs the water, or the track of the game. These aquatic
Snakes seem to be viviparous.

Eryx has the head not distinct from the neck and covered
entirely with small scales. Those of the body are likewise small,
and are either smooth or keeled. The tail is very short. The
anterior maxillary and mandibular teeth are longer than the
posterior teeth. These snakes, most of which are less than
3 feet in length, inhabit the sandy districts of North Africa,
Arabia, and South-Western Asia, extending into Central Asia.
One species, E. jaculus, extends into Greece and the Ionian
Islands. Like the other species it is an ugly creature, pale grey
or yellowish above, witli darker patches and spots. The under
parts are whitish. The scales are smooth on the front half of
the body, becoming keeled further back and on the tail. Total
length under 2 feet. The pupil is vertical.

According to Zander ^ and Werner ^ this snake lives in sandy
localities, digging itself into the sand, or covering the body lightly
with sand and leaving only the eyes and nostrils free. The whole
body is very flabby, and presses itself into any irregularity of the

» Zool. Oart. 1895, p. 330. * Ibid. 1896, p. 85.

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XIII BOINAE — XENOPELTIDAE 605

ground over which the snake creeps. Some specimens live on
lizards, others prefer mice. The prey is caught by the head, and
further secured by several turns of the body of the captor, whose
tail is then turned forwards, round the head of the victim, so
as to form a kind of knot.

Not less striking than their agility is their jealousy, which
is so strong that a snake will occasionally leave the mouse
which it has just strangled in order to seize another snake's
mouse. Sometimes several snakes fight for the same mouse,
coiled together into one inextricable lump so that the mouse
itself is quite invisible. The snakes poke their heads about
in search of the hidden prey, and every attempt of one of
the snakes to free itself, causes the others to squeeze it firmer and
firmer, thinking apparently that the motion was caused by the
lost prey.

Occasionally one of Werner's captives caught several mice
in succession. With these it crawled into a corner, dropped the
mice, and then proceeded quietly to swallow one after another.
After a fortnight the whole repast was digested, and the snake
was ready for more.

Fam. 6. Xenopeltidae. — The single species, JCenopcUis uni-
color, of South -Eastern Asia, including the Malay Islands, has
been raised to the dignity of family -rank on account of the
following combination of characters. The prefrontal bones are
still in contact with the nasals as in the previous families, but
the coronoid bones of the mandibles are absent as in the remain-
ing families. The whole suspensorial apparatus and the lower
jaw itself are peculiar. The dentary bone is movably attached
to the end of the much -elongated articular bone, the movability
being enhanced by the absence of the coronoid element.^ The
quadrate is short and thick, and is carried by the short and
broad sc^uamosal, which lies flat against the skull, resembling in
this respect that of some of the Ilysiidae. Boulenger rightly
considers Xenoj)eUis to be in various ways intermediate between
this family, the Boidae and tlie Colubridae. The head is small
and not distinct from the neck. Tlie eyes are small and have
a vertical pupil. The body is cylindrical, covered above with

^ The same arrangement occurs in the Colubrine genus PoIt/odontophiSj with
about ten siK^cies in South -Eastern Asia, Madagascar, the Comoro Islauda, and in
Central America.

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6o6 OrHIDIA — COLUIJRIDAE chap.

smooth black or brown and highly iridescent scales, hence the
generic name. The ventral scales are white and transversely
enlarged as in the majority of snakes. The tail is short, but
not stunted, measuring about 4 inches in full-grown specimens
of a total length of 3 feet.

Fam. 7. Oolubridae. — This family comprises those snakes
(abqut nine -tenths of all recent species) which combine the
following characters : — ectopterygoids are present : the sciuamosals
are loosely attached to the skull, and carry the quadrates, which
are not reached by the pterygoids : the prefrontals are not in
contact with the nasals : the maxillaries are horizontal and form
the greater portion of the upper jaws : the mandibles lack the
coronoid process or element ; both jaws are toothed.

The best arrangement of this enormous cosmopolitan family
with terrestrial, arboreal, and aquatic forms, is that by Boulenger,
who, adopting Dum^riFs terms, has divided them into three
parallel series.

A. Aglypha, — All the teeth are solid and not grooved.

B. Opisthoglypha, — One or more of the posterior maxillary
teeth are grooved.

C. Proteroglypha, — The anterior maxillary teeth are grooved
or " perforated."

The Aglypha are harmless, non-poisonous. Most of the
Opisthoglypha are poisonous, although few of them are danger-
ously so. The Proteroglypha, which comprise the " Cobras " and
their allies, are deadly poisonous.

Series A. AOLYPHA.

Sub-Fam. 1. Acrochordinae. — The postfrontal bones, besides
bordering the orbits posteriorly, are extended forwards so as to
form the upper border of the orbits, separating the latter from
the frontals. The few genera and species of this sub -family
are mostly aquatic, inhabiting rivers, or estuaries with brafekish
water, and they have been known to swim far out into the sea.
The body is covered with small, frequently granular scales ; in the
typically aquatic forms the body is slightly compressed laterally,
and the ventral scales are scarcely larger than the others. Most
of these ugly snakes inhabit the rivers of coasts of South-Eastem
Asia and Papuasia ; one, StoIiczLaia, is found in the Khasia Hills

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xiii AGLYPHA — ACROCHORDINAE — COLUBRINAE 607

of Xorth-Eastern India ; another, Notliopsis, lives far from its
supposed allies, on the Isthmus of Darien, Central America.

Acrochordus javanicus has no ventral shields. The head is
flat, covered with small granules, with the eyes and nostrils on
the upper surface. The general colour is dull olive -brown,
lighter and spotted beneath. The food consists of fishea Total
length up to 4 feet.

Chersydrus graiiidatus ranges from the coast of Madras to
New Guinea. The body and tail are compressed, and form a
ventral fold, covered with tiny scales like the rest of the body.
General colour grey above, yellow below.

Sub-Fam. 2. Oolubrinae. — The postfrontal bones are restricted
to the posterior border of the orbits. The maxillary and
dentary bones carry teeth on their whole length. The scales
are usually imbricating. This sub-family contains the over-
whelming majority of snakes, about 1000 species, all of them
harmless so far as poison is concerned. None of them reach a
great size, species of 6 or 7 feet in length being rare, e.g.
Zayyienis mucosus, but a few species of the Indian genus Zaocys
8. Coryphodon grow to 10 feet. Most of the Colubrine snakes are
oviparous, but some, e.g. Coronella, are viviparous. Some are
aquatic, or semi-aqiiatic, others are absolutely arboreal, others
again prefer dry, sandy, or rocky localities, according to their
food. The distribution of the sub-family is cosmopolitan, finding
its natural limits only in the permanently frozen under-ground,
a condition which makes hibernation impossible. Most of them
love warmth and like to bask, although many are not fond of the
broiling sun. In the temperate regions they hibernate. As a rule
they are intelligent and some of them become even afifectionate.

Tropidonotus, — The teeth form closely set series on the whole
length of the maxillaries, palatines, pterygoids, and the greater
portion of the dentaries. The premaxilla is toothless. The
teeth of the maxillaries gradually increase in length, the posterior
teeth being the longest. The pupil is round. There is a pair
of internasal shields. The scales covering the body have each
an apical, sensory pit, are mostly keeled, and are arranged in
longitudinal series. The ventral shields are broad ; the sub-
caudals form two rows. This genus, with more than seventy
species, has a wide range, practically over the whole world with
the exception of New Zealand and the southern half of Australia.

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6o8 OPHIDIA

T, natrix, the common Grass-Snake, has a divided, or double,
anal shield. The strongly keeled scales of the body form nineteen
rows. There are normally seven upper labials, the third and
fourth of which border the eye. The usual colour of the Grass-
Snake is olive-grey or brown above, with black spots and narrow
cross-bands. The labials are white or yellowish, with black
sutures. The belly is checkered black and white, more or less
suffused with grey. There are several colour-varieties. The
typical or northern form has a white, yellow, or orange collar,
bordered behind by a black collar ; the pale collar is sometimes
faint or absent. The second variety, rather common in Spain
and Portugal, although not the only form in the Peninsula, has
no collar whatever, and these specimens are sometimes almost
uniformly grey -green above. The third variety, common in
South -Eastern Europe and in Asia Minor, has a well-marked
collar and a yellowish streak along each side of the back. But
there are also almost black specimens.

The usual length of an adult female Grass-Snake is about
3 feet, but very exceptional cases of more than 6 feet are on
record; the males are smaller and more slenderly built. The
range extends over the whole of Middle Europe, Algeria, West
and Central Asia. It does not, however, occur in Ireland or
Scotland. Its northern limit is the southern part of Sweden.

The Grass -Snake prefers moist, grassy localities, with the
neighbourhood of water, chiefly on account of the food, which
consists entirely of fishes and Amphibia, notably of frogs ;
tree-frogs are preferred to anything else ; toads are occasionally
eaten, but mice are never taken.

The Grass -Snake can climb trees or rather shrubs and is
an accomplished swimmer, often spending much of its time in
water for fishing purposes. The fish is caught by the belly and
then generally swallowed on land. The Grass-Snakes appear
in the spring and disappear in the autumn to hibernate in the
ground. They pair, in England, in the month of May or June,
usually on warm and sunny mornings. The eggs are laid from
July to the end of August, mostly in rich vegetable soil, in
heaps of weeds or in manure-heaps. Young snakes lay few^er
eggs than old specimens, which sometimes produce more than
three dozen at a time. The eggs are soft, whitish yellow, about
one inch long, and soon stick together, so that the whole clump

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COLUBRINAE 609

can be taken up at once. As a rule the new-laid eggs dp not
contain any visible sign of the embryo, but it often happens that
the snake has to delay oviposition, and then the embryos are
more or less advanced. This is especially the case with recently
caught specimena The young are hatched in the late summer
or in the autumn, and seem to live at first upon soft insects and
worms. Curiously enough they are easily drowned when they
fall into the water, even in a shallow tank. My tame snakes
have often laid eggs between the stones in the greenhouse ;
the young throve well upon unknown food, but most of them
met their fate in the water. When they are a few weeks old
they are strong enough to take baby-frogs.

The Grass-Snake becomes very tame, learns to distinguish
between different people, allows itself to be handled without
hissing or without voiding the obnoxiously smelling contents of its
cloaca and anal glands, will in time take the offered food from
the hand, and will even crawl up the arm or sleeve and coil
itself up contentedly. . One of the finest specimens, quite green,
without a trace of a collar, and with brownish -red eyes, I
caught in the Guadiana, where it had been fishing in mid-
stream. It swam towards the bank, dived, and hid itself at the
' bottom between rocks. This snake, a female, became very tame.
It never hibernated, shed its skin regularly every few months,
and grew within nine years from 35 inches to 42 inches in
length.

The Grass-Snake is perfectly harmless : although hissing, and
striking out furiously with its head, it never bites, not even
when it is severely handled. Its only defence consists of the
awful contents of the cloaca and the anal glands, the secretion
of which smells of concentrated essence of garlic mixed with
other indescribable odours. The wildest specimens I have ever
met with inhabited a swamp with a little stream to the north of
Oporto close to the coast. To my utter surprise some of them
actually made for me, swimming along rapidly with the head
erect, about 6 inches above the water, and darting forwards
with widely opened jaws, but they did not bite. These and
other kinds of allied snakes require to drink much and often.
Occasionally they drink milk when this is offered them, but
that they suck the udders of cows or the breasts of women is
an idle fable.

VOL. VTIT 2 R

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6io

OPHIDIA

CHAP.

T. miHrinus. — The scales are strongly keeled and form twenty-
one to twenty-three longitudinal rows. The third and fourth labials
border the eye. The anal shield is divided. The eyes and nostrils
are directed upwards instead of sidewards, in adaptation to the
essentially aquatic habits of this species, which lives upon fishes
and Amphibia. The general colour is grey to reddish brown,
with a black zigzag band along the back and a lateral series of
black, yellow-eyed spots. The belly is yellow or red, checkered
with black.

The Viperine Snake bears a general resemblance to the
common viper. It inhabits France, Italy, Spain and Portugal,
and Morocco. Very large specimens attain a length of nearly
3 feet, but the ordinary size of adults is 2 feet. This snake
spends most of its time in the water, but it is often found on

Fig. leO.—Tropidonotus sirtaiis.

land, basking on the top of a low wall or on a low shrub. It
is exceedingly common in Spain and Portugal, where it inhabits
almost every ditch, any standing water or slow river. In the
Alemtejo, when during the rainless and hot summer the small

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XIII

COLUBRINAE 6ll

rivers have nearly dried up, these snakes collect in great
quantities in the remaining stagnant and muddy pools, and
as the stock of suitable fish gets exhausted, are often reduced to
a deplorably emaciated condition. By the month of August
they have become so thoroughly aquatic that they cannot be
kept alive in dry surroundings for twenty-four hours. Those
which I collected generally died, apparently from some kind of
cutaneous suffocation, during the night following their capture.
Taken under other conditions they are very easily kept and
tamed.

I once caught a Viperine Snake in a ditch whilst it was
swallowing an eel of nearly its own length. Both were separated,
and then put into a small bag together with other creatures,
and no more attention was paid to them for several hours.
When I opened the bag again, the snake, undisturbed by my
incessant walking about, was again busily engaged in trying to
get outside that same eel !

T, sirtcdis (Fig. 160) is one of the almost endless varieties of
what is now known by the name T, ordinahis, of North and
Central America.

T. tesselatn^ is closely allied to T. viperinus, which it represents
in South Germany, Italy, South-Eastern Europe, and Asia ; but
the scales form only nineteen rows, and the fourth, or fourth and
fifth labials, border the eye. The usual colour is olive-grey with
dark little spots, and with a dark chevron-shaped band behind
the occiput. The lower parts are yellow or red checkered with
black, hence the specific name.

Zajnenis. — The maxillary teeth are not closely packed ; they
increase slightly in size backwards, and the last two are often a
little larger and separated from the rest by a diastema. The
mandibular teeth rather decrease in size from before backwards,
inversely with the upper teeth. The scales are smooth with
apical pits ; the sub-caudals form two rows. The eye is large,
and has a round pupil. The range of this genus, with about
thirty species, extends over the whole of the Periarctic region.

Z. {Ptyas) miicosus (Fig. 161), the Eat Snake of India, extending
from Transcaspia to Java, is a very common species, often seen in
menageries. Its general colour is brown above, often with black
cross-bands on the hinder part of the body and tail. The under
parts are yellowish. The fourth and fifth labials border the eye.

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OPHIDIA

The scales on the body form only seventeen rows. Another
feature of this species is the prominent ridge of the back-bone,
not only in half-starved but in well -conditioned specimens.

Fig. 161. — Zamenu mucosns (Rat Snake). x |.

The Rat Snake grows to a length of more than 7 feet, and is
as ill-tempered as most species of this genus.

Z. gemonensis s. viridijiavus inhabits France, Italy, the
Balkan Peninsula, and Asia Minor. Its coloration is very
variable. In general it is either green above and yellow
below, hence the appropriate name viridiflavus, or the ground-
colour of the back is greyish or olive -yellow with brownish
spots, which form more or less longitudinal rows on the trunk,
but gradually pass into blackish continuous lines on the tail :
the under parts are yellow or greenish white, often with inany
very small, dark specks. The scales form seventeen or nineteen
rows ; the anal shield is divided. There are two sma-ll post-
ocular scales and one subocular ; of the eight labials, the fourth
and fifth border the eye. This species is very lively, attacks

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COLUBRINAE 613

and bites furiously, climbs well, and when suspended from
branches can protrude half of its length in a horizontal direction.
It eats any kind of Reptile, Bird, or Mammal it can master ;
small animals are swallowed directly, rats and moles are first
killed by constrictions. Large specimens reach perhaps 6 feet
in length.

Z. hifjpocrepis is the representative species in the Iberian
Peninsula and in North-Western Africa. It is rarely more than
4 feet long, and is very pretty, the ground-colour being reddish
or olive-yellow with a row of large, dark brown, yellow-edged
spots along the back. Two rows of smaller spots adorn the
sides ; where the dark spots are large, the pale ground-colour is
restricted to forming rings around the spots, producing a pretty
appearance. The under parts are yellow or orange, with black
spots. On the head is a dark, pale-edged patch in the shape of
a horse-shoe, a feature alluded to by the specific name. Struc-
tural characters are the possession of a row of little subocular
scales, which completely separate the eye from the labials, the
double anal shield, and the small and smooth scales on the body,
which form generally as many as twenty-seven rows.

Z, constrictor. — The American Black Snake. The scales are
smooth, and arranged in seventeen rows; the anal sliield is
divided. The general colour above is uniform bluish-black ;
below slaty, tinged with blue ; the chin and throat are silvery
white, sometimes with a black spot. Large specimens attain a
total length of 6 feet.

Holbrook gives the following exhaustive account of this
species, about which many sensational stories are current even
in would-be scientific periodicals : —

The "Black Snake" is one of the commonest of North
American species. It is extremely active, climbing with facility,
and running with great rapidity, whence it is not uncommonly
called the " Eacer." It frequents shady places, covered with
thick shrubs, on the margins of water. It feeds on mice, toads,
or small birds ; and, as it is an excellent climber, is frequently
seen on trees in search of birds' nests. It is a bold and daring
serpent, enters barns and out-houses without fear, and has
been known to destroy young chickens. Its specific name
constrictor would imply that it suffocates or crushes its prey,
but this according to Holbrook is at least doubtful. In the

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6 14 OPHIDIA

breeding season it is extremely irascible, and w^l frequently
attack persons passing at a distance of several steps; the tail
then quivers with rage, making a quick vibrating motion, which
in forests and among dead leaves sounds not unlike the Eattle-
Snake ; it now elevates the head one or two feet from the
ground, and darts upon its adversary ; luckily its bite is harm-
less, and not more painful than the scratch of a pin.

" It will even descend from trees to attack its enemy if
teazed, yet it does not twine itself around the legs, as L«i
commonly supposed.

" The same power of charming its prey has been attributed
to the Black as to the Eattle-Snake, and with still less appear-
ance of reason ; for this is a nimble animal, and can pursue its
prey, while the Eattle-Snake must lie in wait for his. It is
remarkable that the birds most commonly found ' charmed ' are
the Cat-bird {Turdtis carolinensis) or red- winged Black-bird
{Icterus phoeniceus). These birds choose thick and shady places
on the margins of streams for their residence, and generally
build their nests on such shrubs as the alder ; the latter bird not
unfreqiiently takes the precaution to select such bushes as are
on small islands, or such as have their roots surrounded by
water, and thus their home is more secure. Now the Black
Snake chooses precisely the same localities, knowing probably
the haunts of its prey. The serpent begins the war by besieging
the nest ; the old bird, aware of its intention, attacks it with
fluttering and uncertain motions, accompanied by a plaintive cry
of distress, and is then said to be * charmed/ The snake is at
last either driven off, or it captures the young and not unfre-
quently the old bird too.

" Sometimes the old bird, by her cries, calls in the assistance
of her neighbours to drive away the aggressor. I have seen
more than a dozen birds thus engaged with a large Black Snake
that had probably just committed some depredation, but was
now quietly stretched on a rock, basking in the sun ; and it was
not a Uttle singular that birds of very different genera, and
those seldom seen together, all united in this warfare against a
common enemy, and finally compelled him to seek shelter among
some low, thick shrubs, by tlie violence of tlieir assault."

Zaocys, with about half-a-dozen species in South-Eastera
Asia, is closely allied to Zamenis. Z. carinatus, of the Malay

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COLUBRINAE

615

Islands, grows to 10 feet in length; it is consequently one of
the largest harmless Colubrine snakes. The scales form only
sixteen to eighteen rows. The sub-caudals are double. The
general colour above is dark olive, passing into greenish brown

Fig. 162. — Zactci/s carinatus.

farther back. The under parts are yellowish ; black and yellow
posteriorly. The fifth and sixth labials border the eye.

Coluber, — The maxillary teeth are of equal size, but the
anterior mandibular teeth are the longest. The head is distinct
from the neck. The nasals are distinct ; not fused with the
loreals. The eye is rather large, with a round pupil. The
scales, smooth or keeled, have apical pits ; the ventrals are
roimded or angulate laterally ; the sub-caudals are double. They
all lay eggs and constrict their food. Nearly fifty species in the
Periarctic region.

C. (Elajihis) quatuorlineatus s. quaterradiatus inhabits Italy
and South-Eastern Europe. It occurs also in the Southern
Tyrol. The scales of this large snake, which grows to nearly
6 feet in length, are arranged in twenty-five rows, and are feebly

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6l6 OPHIDIA

CHAP.

keeled. The anal is divided. Adult specimens have a yellowish-
brown ground-colour with a pair of black streaks on each side of
the back. A black line extends from the eye to the angle of
the mouth ; the under parts are yellow, mostly closely spotted
with brown. This snake is good-tempered, and keeps well in
captivity. They live on sparrows, mice, lizards, etc., and are very
fond of eggs. Large specimens can swallow several fowls' eggs
in succession ; the crushed remains of the shells are later dis-
gorged. This handsome snake climbs extremely well in search
of birds and their eggs, and it is not afraid of the water. The
prey is caught either with the teeth or by a rapid twist of tlie
tail ; in any case, the prey is always strangulated by the constric-
tion of coils thrown round it. A sparrow thus secured is
literally passed through the moving coils along the snake's body
into a position convenient for swallowing. Hungry snakes
catch and secure several birds or mice before eating them. My
own specimens became almost affectionately tame, never attempted
to bite, and took food from the hand.

C leopardinus is smaller, but is one of the handsomest snakes
of Southern Italy, South-Eastern Europe, and Asia Minor. It is
closely allied to the previously described species. The ground-
colour is pale brown with a dorsal series of dark brown or
reddish, black-edged, transverse spots, and a lateral alternating
series of smaller black spots, or with two dark brown, black-
edged stripes bordering a yellowish vertebral stripe ; usually with
a forked black mark on the occiput and nape. The imder parts
are white, checkered with black, sometimes with the latter colour
prevailing.

C, Jiavesceiis s. aescidapii is the Aesculap-Snake, for which the
almost unknown name of longissivius has now been unearthed in
deference to the fetish of priority. This snake is of an extremely
graceful and slender build, with a very long tail. Its home is
the South of France, Italy, and South-Eastern Europe. It occurs
sporadically in the Tyrol, for instance near Bozen, in Austria, at
Baden near Vienna, in Germany only in the Taunus, especially at
Schlangenbad, which has received its name from the frequent
occurrence of this snake. This sporadic distribution favours
the idea that these snakes w^ere introduced by the Bomans as
inmates of the temples erected to Aesculapius at such watering-
places. Specific characters are the smooth and shiny scales,

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XIII COLUBRINAE 617

which are arranged in twenty-one to twenty-three rows, the
distinctly angulate ventrals and the double anal and sub-caudals ;
the fourth and fifth of the upper labials border the eye, which has
a round pupil. The coloration is very variable, as a rule olive-
brown above with a dark streak behind the eye ; the upper lips
and a triangular patch on the temples are yellow ; the under parts
are uniform pale yellow. Some specimens are pale golden brown ;
others are very dark, almost black ; while some have four darker
stripes along the body, and lastly whitish specks occur on the
upper surface. Large Aesculap snakes grow to a length of 5
feet. Their food consists chiefly of mice. They become ver}^
tame, although many of them at first bite furiously. Their
climbing capacities are astonishing, the snakes being able to scale
high and vertical walls provided there is the slightest " foothold."
Some of my specimens escaped in the room and were at last
found near the ceiling, resting on the rods of the curtains, up
the folds of which they had managed to wriggle. Boulenger
kept one for many years in a glass cage, where the snake
entwined himself round the branches of a stick and allowed us
to take him with the stick out of its socket and to inspect him.
Being kept in an inhabited room, the snake did not exactly
hibernate, creeping into the moss at the bottom of the cage ; but
it refused to feed, and remained in a rather drowsy condition
coiled up on its favourite stand. During the pairing season they
frequently resort to the water, at Schlangenbad at least ; the
few eggs are deposited under dry moss or in dry, decayed wood,
and are hatched in about six weeks.

C. {Rhinechis) sccdaris has the smooth scales disposed in
twenty-seven rows. The snout is strongly projecting, and has a
V-shaped dark mark on the top; a black streak runs through
the eye, and another black spot lies below the eye. Young
specimens are pale brown with a series of dark H -shaped marks
on the back, suggesting a ladder, hence the specific name. In
the adult these marks are replaced by a pair of brown stripes
running along the back ; the under parts are always uniform
yellow. Total length about 3 feet. This snake is restricted
to the Iberian Peninsula and to the South of France. Most
specimens are very ill-tempered. The young live upon locusts
and small lizards, the old eat mice and small birds. In captivity
they also take dead animals — a rare habit with snakes.

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OPHIDIA

Deiidrophis with about ten species inhabits South-Eastern
Asia and Australia. They are typical Tree-Snakes. The scales
are keeled, and form only thirteen or fifteen rows ; those of the
vertebral row are enlarged ; the ventrals have a pair of suture-
like lateral keels and a notch on each side, arrangements which
are of great assistance in climbing, these snakes being able to
slide up the branches of trees in almost straight lines instead of
having to twist and undulate their way up.

Fig. 163. — Lendrophis punctulaiu^. x\.

D. picnctulalus, of Northern and Eastern Australia, is olive-
brown above, uniform or with black edges and yellow outer
borders to the scales. The upper lips, the throat and anterior
ventrals, are yellowish. Total length up to 6 feet.

Leptophis is a Neotropical genus of Tree-Snakes. The lx)dy
and tlie extremely long, whip-like tail are very slender. The
head is very distinct from the neck ; the eye is large, with a
round pupil. The scales form thirteen or fifteen rows; the
ventrals are sometimes angulate laterally ; the sub-caudals are
double. Z. (Ahaetulla) liocerus is a beautiful snake, green above

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COLUBRINAE

619

with a golden lustre, while the under parts are yellow or white.
The total length of this species amounts to 6 feet, the tail then
being nearly 2 feet long. These graceful Tree-Snakes live upon
small reptiles and birds and their eggs. When shaken out of a
tree or frightened off they let themselves fall down from con-
siderable heights, coiling body and tail like a watch-spring,

Fio. 164. — LqHophis liocerus.

and alighting on the ground upon the spiral, which breaks the
fall.

Coronella. — The teeth are nearly all of equal size and form
continuous series. The scales are smooth and have apical pits ;
the sub-caudals are double. The head is scarcely distinct from
the neck. The pupil is round. This genus, with nearly
twenty species, is widely distributed except in the Australian
region, the northern half of Asia, and South America. We can
mention only the two European species, one of which occurs in
England.

C. austriaca s. lacvis, the Smooth Snake. The scales are
arranged in nineteen rows. Mostly the third and fourth

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CHAP. XIII COLUBRINAE 62 1

labials border the eye. The anal shield is divided. The general
colour is brown or reddish above, often with one or two lighter
stripes, with small dark brown or red spots ; two dark brown or
red stripes on the nape, usually confluent with a large dark patch
on the occiput ; a dark streak extends from the nostril through
the eye to the angle of the mouth. The under parts are red,
orange, brown, grey or blackish, either uniform or speckled with
black and white. The coloration is, however, subject to much
variation, and some specimens strikingly resemble some of the
Common Viper, which is also very variable in its coloration. The
resemblance is enhanced when the Smooth Snake broadens its
head by widening the jaws, as it is in the habit of doing.
Two such similarly coloured specimens are represented in Fig.
165. On closer inspection the differences are great enough, the
harmless snake having smooth scales, and the top of the head
being covered with large shields; while the Viper has keeled
scales, the top of the head being covered mostly with scales, a
vertical (not roimd) pupil, and, moreover, when attacked, usually
coils itself into a spiral disc with the head standing out in the
middle, ready to strike. However, these two species are some-
times mistaken for each other. — The Smooth Snake prefers lizards
as food to anything else, but it also takes mice. The prey is
hunted chiefly in the late afternoon and in the evening, and is con-
stricted by the coils of the snake. When caught or even when
handled after months of captivity, the Smooth Snake bites
deliberately and firmly, selecting a suitable spot, for instance a
finger, opens the mouth widely and almost chews the spot. The
bite is of course quite harmless, and scarcely draws blood, few
of these snakes attaining a length of more than 2 feet. They
are viviparous, bringing forth about half-a-dozen young at a time.
The range of the Smooth Snake extends over the greater part
of temperate Europe, from England and the Iberian Peninsula to
Berlin, and south-eastwards to Asia Minor. In England it occurs
in a few counties only, for instance in Hampshire and in
Dorsetshire.

C. gironclica, of the South of France, Italy, the Iberian Penin-
sula and North-Western Africa, much resembles the English
Smooth Snake, from which it differs in a few points only. The
scales are arranged in twenty-one, rarely in nineteen, rows ;
usually the fourth and fifth labials border the eye ; and the rostral

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622

OPHIDIA COLUBRTDAE

CHAP.

shield, covering the end of the snout, is much broader than high.
The coloration is variable, but thei-^jfe always a pair of elongated
blackish spots or a U-shaped mark on the nape.

Sub-Fam. 3. Rhachiodontinae. — With only a few teeth on
the posterior part of the maxillaries, on the palatines and
dentaries. Some of the vertebrae in the region of the lower neck
have strongly developed hypapophyses, which are directed forwards
and pierce the oesophagus. They are used for filing through or

FiQ. 166. — Dasypeltis scabra. x j.

breaking the birds' eggs which seem to be the chief food of these
snakes.

Basi/pcltis scahra, the only species, inhabits Tropical and South
Africa ; although it reaches scarcely more than two feet and a
half in length, such a specimen is able to swallow an ordinary
fowl's egg. Pigeons' eggs are swallowed by snakes little more
than one foot in length, which seems at first sight quite im-
possible. The swallowed egg distends the skin to its utmost
capacity ; it then slides down further, the snake makes some
slight contortions and the swelling collapses ; after a while the
broken and sucked-out shell is vomited out as a crumpled up

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xui RHACHIODONTINAE DIPSADOMORPHINAE 623

mass. Miss Durham has illustrated this curious process in a
series of drawings.^

Series B. OPISTHOGLTPHA.

One, or a few, of the posterior maxillary teeth have a groove
or furrow in front, which conducts the secretion of the enlarged
upper labial glands. Apparently all these snakes are more or
less poisonous, paralysing their prey before or during the act of
deglutition. So far as man is concerned they are rather harm-
less, since the poison is not very strong, not available in large
quantities, and above all because the small poison-teeth stand
so far back that the snakes cannot easily inflict wounds with
them.

The Opisthoglypha are of considerable morphological interest,
since they connect the Colubridae with the Viperidae, the char-
acteristic poisonous apparatus of which seems to have been derived
from that of the Opisthoglypha by the reduction or shortening
of the anterior portion of the maxillaries and the harmless teeth,
so that the posterior or poison-fangs come to the front.

The Opisthoglypha comprise about three himdred species and
are cosmopolitan, including Madagascar but excepting New Zea-
land. They contain truly terrestrial, arboreal, and thoroughly
aquatic forms.

Sub-Fam. 1. Dipsadomorphinae. — The nostrils are lateral
and the dentition is well developed. Long-tailed, terrestrial, and
arboreal forms. Most of the arboreal species are green above,
often with white or yellow longitudinal bands, while the under
parts are white or yellow. They feed chiefly upon lizards, birds
and Iheir eggs.

Dipsadomorphus s. Dipsas (part). — Typical, very long-bodied
and long-tailed Tree-Snakes, with a vertical pupil. The median
or vertebral row of smooth scales is enlarged ; the broad ventral
scales are bent at an obtuse angle on the sides, the resulting
ridge assisting in climbing. The sub-caudals are arranged in
two rows. Ten to fourteen maxillary teeth are followed by two
or three enlarged, grooved fangs.

D. trigonatus, of India, grows to one yard in length.
Yellowish olive or pale grey above, with a white, black-edged
1 P.Z.S. 1896, p. 715.

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624 OPHIDIA CHAP.

zigzag band along the l)ack, or with a series of white, black-edged
spots.

D, cyanevs, of Northern India, Assam, etc., is a beautiful Tree-
Snake, green above, with the skin between the scales black,
uniform greenish yellow below. Total length up to 4 or 5 feet.

Dipsds, e.g. D, bmephala, — Maxillaries with eleven or more
teeth. Pterygoids toothless. Body strongly compressed, with
thirteen rows of smooth scales ; the vertebral row enlarged :
sub-caudals double ; tail very long. Tropical South America.

Lejptognathus with many species in Central and South
America, like DipsaSy but with teeth on the pterygoids.

Coelopeltis. — Terrestrial and diurnal, with a round pupil. The
row of small maxillary teeth is followed by one or two much
larger, grooved fangs situated at a level below the posterior
border of the eye. The first half-a-dozen mandibular teeth are
much larger tlian the rest. The scales of the adult are more or
less distinctly grooved longitudinally, hence the generic name,
and are arranged in seventeen or nineteen rows. The sub-caudals
form two rows ; the ventrals are rounded ofif laterally. Two
species in the Mediterranean countries and in South-Western
Asia.

C. monspessidaim s. lacertina is one of the largest snakes in
Europe, reaching a length of 6 feet, of which the tail takes
up 18 inches. Olive -brown or yellowish or reddish abovo,
frequently with small, dark, light-edged spots. The sides are
often blackish, with whitish specks. The xmder parts are
yellowish white, with or without brownish markings. Some
specimens are very green, with a dull blackish neck. One of the
specific names of this terrestrial snake is the latinised form of
Moutpellier ; the other refers to the shape of the head, which is
not unlike that of a lizard, partly owing to the concave forehead.
This species inhabits rather dry localities studded with shrubs,
where it hunts for lizards, birds, and mice. It is sure to attract
notice by its loud hissing when it is disturbed. When driven
into a corner it strikes out furiously, but does not, as a rule, bite.
I have caught some which after a few days became quite gentle.
Small animals become torpid a few minutes after they have been
bitten.

Maeroprotodon cucuUatns occurs in Andalucia, the Balearic
Islands, and in Xorth Africa. The dentitioh is peculiar. The

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XIII ELACHISTODONTINAE HOMALOPSINAE 625

fourth and fifth maxillary teeth are enlarged, followed by an
interspace, then follow several small teeth, and lastly the two
enlarged, grooved teeth. The sixth mandibular tooth is very
long, separated by a space from the much smaller posterior teeth.
The general colour of this sand-loving snake is pale brown or
grey above with small spots or streaks on the tnmk, and with a
large black patch behind the head extending over the sides of the
neck, hence the specific name. The under parts are bright red or
yellowish, sometimes spotted with black. Total length under 2 feet.

Snb-Fam. 2. Elachistodontinae. — With only a few teeth on
the posterior part of the maxillary and dentary bones, and on
the palatines and pterygoids. Some of the vertebrae in the
thoracic region have much -developed unpaired hypapophyses,
which are directed forwards and pierce the dorsal wall of the
gullet. In this respect Elachistodon luestermaniii, of Bengal, the
only species, bears a striking resemblance to the South African
Ai^\y^\\(Aon\i Dasypcltis (see p. 622), and it is probable that this
apparently very rare Indian snake also swallows eggs. It is
brown above, with a yellowish vertebml stripe ; yellowish below.

Sub-Fam. 3. Homalopsinae. — The nostrils of these absolutely
aquatic and viviparous snakes are valvular, and are situated on the
upper surface of the snout. The eyes are small with vertical
pupils. The two dozen species, mostly very ugly, inhabit the
rivers and estuaries of the East Indies from Bengal to North
Australia. Some species have very small and narrow ventral
scales, reqalling the Hydrophinae, or the burrowing snakes, none
of which use tlieir ventral scales for locomotory purposes.

Homalopsis huccata, Cerberus rhynchoj^s and Hypsirhina, e.g.
H. plumbed, have well-developed ventral scales ; the other scales
of the first two genera are keeled, those of the third are smooth.
In Hipistes the whole head is covered with very small scales ; all
the scales of the body are smooth except the very narrow
ventrals, which have double keels. If. hydrinns, of Siam and the
Malay Peninsula, has a compressed body, and in its general
appearance much resembles the Hydrophinae. It lives, like its
allies, upon fishes, and it swims far out into the sea.

Series 0. PROTEROGLYPHA.

The anterior maxillary teeth are deeply grooved, or so folded
VOL. VIII 2 s

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626 OPHIDIA CHAP.

as to appear hollow or perforated. Behind these enlarged
poison-fiAngs the maxilla carries a series of smaller, solid teeth ;
hence the term " proteroglyphous," which means that the anterior
teeth are grooved, in opposition to " opiftthoglyphous." Both
series have been developed independently.

The Proteroglypha are all extremely poisonous, mostly
viviparous, and widely distributed over the whole of the
Australian, Palaeotropical and Neotropical regions, with the
exception of Madagascar and New Zealand ; they extend north-
wards into the warmer parts of North America, and they also
range over a great portion of the Palaearctic sub-region, being found
in North Africa and South-Westem Asia. They form two
natural sub-families : Elapinae, with cylindrical tails, and Hydro-
phinae or Sea-Snakes, with laterally compressed tails.

Sub-Fam. 1. Elapinae. — The tail is cylindrical. The Elapinae
comprise nearly 150 species, which have been grouped into a

^ ELAPINAE. fflilMAJA. ^ <y '

Fia. 167. — Map showing the distribution of the Elapine Snakes.

great number of, mostly somewhat imaginary, genera. In
Australia they constitute the great majority of Snakes, there
being besides the deadly Elapinae only a few Pythons and
Typhlopidae, and very few Colubrinae.

Naja. — The pair of large and grooved poison-fangs are
separated by an interspace from one to three small, faintly
grooved teeth near the posterior end of the maxillaries. The
scales are smooth and without pits, and are arranged in fifteen
to twenty-five oblique rows on the trunk, although more occur in
the region of the neck ; the vertebral row is not enlarged. The
head is but slightly distinct from the neck. Each nostril lies
between two nasals and the internasal. The sub-caudals form
two rows. The pupil is round. The neck-region can be expanded

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XIII

ELAPINAE

627

into a hood by the spreading and moving headwards of the ribs.
Several species in Southern Asia and in Africa.

JV. tripvdians (the "Cobra"). — The coloration varies much.
The typical form is yellowish to dark brown with a black and
white spectacle-mark on the dorsal side of the hood, and with a
large black and white spot on each side of the corresponding
under surface. Other specimens are uniform pale brown to
blackish grey, without any markings on the hood. The Cobra is

^rT'j?^^*f

Fig, 168. — iVq/a tripvdians (the Cobra), x \,

widely distributed, from Transcaspia to China and to the Malay
Islands ; in the Himalayas it ascends to about 3000 feet above
the level of the sea. Very large specimens are said to attain
more than 6 feet in length, but a cobra of 5 feet, inclusive of
the tail of 9 inches, is considered large. The Cobra prefers places
which afford it a convenient hole to retire into ; for instance,
deserted hills of termites, ruins, heaps of stones and stacks
of wood, and it has the disagreeable habit, like the harmless
Rat-snake, Zamenis raucosus, of making itself at home in inhabited
houses, probably attracted by the rats. Its chief food consists
of small Vertebrates ; — frogs, lizards, rats, occasionally fishes and

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628 OPHIDIA

small birds. It drinks much, and hunts chiefly in the late
afternoon and in the evening, although it possesses a round
pupil. It avoids hot sunshine. Many observations show that
the cobras live in pairs, otherwise they do not take much notice
of each other or of other kinds of snakes. The female lays about
a dozen soft-shelled eggs as large as those of pigeons.

This cobra is used by Indian oonjurera The " dance " is
the habit of these snakes of erecting themselves, when agitated,
upon the hinder third or quarter of their length, whilst they
spread out the hood and sway the head and neck to the right
and left, always in an attitude ready for striking. They are
docile and by nature not vicious. Most of the performing
cobras have their teeth drawn, and they then know well that
they cannot bite. They only strike at the hand, just as
uninjured specimens soon avoid biting into the iron rod with
which they are lifted up in menageries. The drawing of the teeih
is an operation which has to be repeated, since reserve -teeth
soon take the place of the lost pair.

I cannot refrain from relating an abstract of a ridiculous
episode which happened in the Munich Aquarium in the
year 1882. One of six specimens of the African si>ecies JN'rT/a
haje was missing. The police closed the establishment, which
during the following eight days was turned inside out without
any other effect than that two other, harmless, snakes w^ere dis-
covered. Twice the building was fumigated with sulphur, until
the Cobra was at last found suffocated, fifteen days after the
beginning of the search. This snake caused the owner of the
Aquarium a loss of nearly £1500. But . the cruel joke was,
that during the commotion the man who had collected and sold
the six snakes declared upon oath that their teeth had been so
well drawn and the germs of possible reserve-teeth had been so
thoroughly destroyed that the snakes were rendered absolutely
harmless. But he was not believed, in spite of a commission of
professors and doctors appointed, who experimented upon the
remaining five Cobras with sulphur and did not find any poison-
fangs, " although the mouth was probed and poked into as far
down as the larynx."

Cobras have quite a number of enemies. Peafowl and
Jungle-cocks are said to be partial to young snakes; pigs eat
them greedily, and are to a certain extent immune against

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XIII ELAPINAE 629

their bite. The same applies, according to the most recent
observations, to the famous Mongoos. Sir E» Tennent, in his
Natural History of Ceylon, quoted several times in the present
book, makes the following remarks about the immunity of this
little creature : —

" I have found universally that the natives of Ceylon attach
no credit to the European story of the Mongoos {Herpestes griseus)
resorting to some plant, which no one has yet succeeded in identi-
fying, as an antidote against the bite of the venomous serpents
on which it preys. There is no doubt that, in its conflicts with
the cobra and other poisonous snakes, which it attacks with as
little hesitation as the harmless ones, it may be seen occasionally
to retreat, and even to retire into the jungle, and, it is added, to
eat some vegetable. ... A number of plants, such as the Ojphi^
oxylon serpentinum and Ophiorhiza mungos, the Aristolochia
indica, the Mimosa octandria, and others, have each been asserted
to be the Ichneumon's specific. ... If the Ichneumon were in-
spired by that courage which would result from the consciousness
of security, it would be so indifferent to the bite of the serpent,
that we might conclude that, both in its approaches and its
assault, it would be utterly careless as to the precise mode of
attack. Such, however, is far from being the case ; and next to
its audacity, nothing can be more surprising than the adroitness
with which it escapes the spring of the snake under a due sense
of danger, and the cunning with which it makes its arrangements
to leap upon the back and fasten its teeth in the head of the
cobra. It is this display of instinctive ingenuity that Lucan
celebrates where he paints the Ichneumon diverting the attention
of the Asp by the motion of his bushy tail, and then seizing it
in the midst of its confusion. See Pharsalia, lib. iv. verses
729-734."

There is a widespread belief in the eflBcacy of " Snake-
stones," which are generally pieces of charred bone, well polished,
occasionally pieces of chalk or some similar porous substance,
which, if pressed upon the bleeding wound, are supposed to
absorb the poison. Snake-charmers profess to prepare such
"stones," and to preserve the composition as a secret. The
manufacture is a lucrative trade. The Boers bought them,
imported from India, at high prices. Mr. Selous saw one, or
heard of one, that was kept as an heirloom. Snake-stones are

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630 OPHIDIA CHAP.

also made, and used, in Mexico, of charred hartshorn ; they are
called " piedras ponsonas."

The use of the Snake-stone, called " Pamboo-Kaloo," has prob-
ably been communicated to the Singhalese by the itinerant
snake-charmers who resort to the island from the coast of
Coromandel.

Although Sir E. Tennent describes several instances of
the successful application of snake-stones as well authenticated,
he has never himself been an eye-witness. Although two cases
have been fully described, they do not at all exclude the possi-
bility, nay the probability, that the Tamils imposed upon the
Europeans in order to sell the snake-stones.

" No doubt the snake-stones, owing to their porous nature,
adhered to the bleeding wound, became saturated with blood,
and then fell off. Very likely, in case of a poisonous bite, some
of the venom would be sucked up too, but we do not know if
those snakes were still in the possession of their poison-fangs.
Properly conducted experiments with snake-stones have proved
as little efficacious as the application of dry cup.

" Theoretically snake-stones as quick absorbent agents of the
blood with the poison are good; they will certainly prevent
some of the poison from entering the system, but that would, at
best, be a partial cure only.

"In March 1854 a friend of mine was riding, with some
other civil officers of the Government, along a jungle path in
the vicinity of Bintenne, when he saw one of two Tamils, who
were approaching the party, suddenly dart into the forest and
return, holding in both hands a Col)ra de capello, which he had
seized by the head and tail. He called to his companion for
assistance to place it in their covered basket, but in doing this,
he handled it so inexpertly that it seized him by the finger, and
retained its hold for a few seconds, as if unable to retract its
fangs. The blood flowed, and intense pain appeared to follow
almost immediately ; but with all expedition the friend of the
sufferer undid his waist-cloth, and took from it two snake-stones,
eacli of the size of a small almond, intensely black and highly
polished, though of an extremely light substance. These he
applied, one to each wound inflicted by the teeth of the serpent,
to which they attached themselves closely ; the blood that oozed
from the bites being rapidly imbibed by the porous texture of

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xiii ELAPINAE 63 1

the article ajifplied. The stones adhered tenaciouslj for three or
four minutes, the wounded man's companion in the meanwhile
rubbing his arm downwards from the shoulders towards the
fingers. At length the snake-stones dropped off of their own
accord ; the suffering of the man appeared to subside ; he twisted
his fingers till the joints cracked, and went on his way without
concern. Whilst this had been going on, another Indian of the
party, who had come up, took from his bag a small piece of
white wood, which resembled a root, and passed it gently near
the head of the cobra, which the latter immediately inclined
close to the ground ; he then lifted the snake without hesitation,
and coiled it into a circle at the bottom of his basket. The
root by which he professed to be enabled to perform this opera-
tion with safety he called the " Naya-thalic kalanga " (the root of
the snake-plant), protected by which he professed his ability to
approach any reptile with impunity."

The following narrative, communicated to Sir E. Tennent by
H. E. Reyne, of the Department of Public Works, Colombo, seems
to exclude the possibility of deception : —

"A snake-charmer came to my bungalow in 18 54, 'requesting
me to allow him to show me his snakes dancing. As I had
frequently seen them, I told him I would give him a rupee if he
would accompany me to the jungle and catch a cobra that I
knew frequented the place. He was willing, and as I was
anxious to test the truth of the charm, I counted his tame
snakes, and put a watch over them until I returned with him.
Before going I examined the man, and satisfied myself he had
no snake about his person. When we arrived at the spot, he
played on a small pipe, and after persevering for some time, out
came a large cobra from an ant-hill, which I knew it occupied.
Oil seeing the man it tried to escape, but he caught it by the
tail and kept swinging it round until we reached the bungalow.
He then made it dance, but before long it bit him above the
knee. He immediately bandaged the leg above the bite, and
applied a snake-stone to the wound to extract the poison. He
was in great pain for a few minutes, but after that it gradually
went away, the stone falling off just before he was relieved.
When he recovered he held a cloth up, which the snake flew at,
and caught its fangs in it ; while in that position, the man
passed his hand up its back, and having seized it by the throat,

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632 OPHIDIA

he extracted the fangs in my presence and gave them to me.
He then squeezed out the poison on to a lea£ It was a clear
oily substance, and when rubbed on the hand produced a fine
lather. I carefully watched the whole operation, which was
also witnessed by my clerk and two or three other persons."

N, haje is the common hooded cobra of Africa, the " Aspis,"
so called on account of its shield or hood — the " Spy-Slange " of
the Boers. As a rule the spectacle-marks on the neck are absent
or indistinct, the general colour varies much, either brown above,
yellowish beneath, with or without brown spots ; or dark brown
above with yellowish spots, dark brown beneath ; or blackish
above and beneath. The name Spy-Slange, meaning Spitting
Snake, refers to the habit which this and other African Cobras
have of letting the poison drop from the mouth like saliva when
they are excited. This is not a particularly economical habit,
nor is it of the slightest use to the snake.

N, (Ophiophagvs s. Hamadryas) hungarus s. daps is the
*' Hamadryad " or " Snake-eating Cobra" or " King Cobra." It
has a well dilatable hood ; the very variable coloration is
yellowish to black, with or without an olive gloss. Many
specimens have more or less distinct dark cross-bands or rings
around the body, while others are olive above with black-edged
scales, and others again are very dark above and beneath. The
distinctive, specific character is the small number of scales, these
forming only fifteen rows on the middle of the body, nine-
teen or twenty-one on the dilatable neck. There is a pair of
large occipitJil shields behind the parietals.

This snake reaches the length, enormous for a poisonous
snake, of 12 feet or more. Its size and very poisonous nature
make it the curse of the jungla It ranges from India to South
China, and to the Philippines. The food seems to consist entirely
of other snakes.

Sepedon haemachates is another hooded snake in South Africa,
where it is known as the "Einghals," i.e. banded neck. It
differs from Naja by the absence of small teeth on the maxil-
laries behind the fangs, and by the strongly keeled scales, which
form nineteen rows. The general colour is black above variegated
with yellow or pale brown ; the under parts are also black, often
with one or two whitish bands across the lower portion of the
neck.

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The Rev. G. Fisk ^ mentions the case of two young " Ringhals,"
of 10 and 9 inches in length, having been attacked and partly
devoured by a mouse, supposed to be Dendromys melanotis, which

Fig. 169. — Sepedon haemachates (the "Ringhals"). x^.

was put with the snakes in a band-box. On the habits of the
Einghals see Symonds.^

Bungarus. — The scales are smooth, and form thirteen to
seventeen rows. The spine is very prominent, and the median
row of scales which covers the ridge is much enlarged. There is
no dilatable hood. In other respects Bungariis is closely allied
to Naja ; about half-a-dozen species, in South-Eastern Asia.

B.fasciatus reaches a length of 5 feet. The general colour
is bright yellow, alternating with blackish rings.

B. coeruleiis s. candidus is the dreaded " Krait," occurring in
the whole of the Indian sub-region. It is dark brown or bluish
black with narrow cross-bars or white specks, or it is alternately
barred brown and yellow ; the imder parts are uniform white.
Total length rarely 4 feet.

The " Krait " seems to cause more deaths in India than any
other snake, since it is veiy common, especially in Bengal and in
» P.Z.S. 1887, p. 340. 2 P.Z.S. 1887, p. 489.

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634 OPHIDIA

CHAP.

Southern India, and often creeps into the houses. It lives
chiefly on rats, lizards, and snakes.

CaJlophis. — With only thirteen rows of smooth scales. The
head is small, not distinct from the neck. The small eye has a
round pupil. The short tail has two ventral rows of scales.
The whole body is cylindricaL Several small species, one or two
feet in length, in South-Eastern Asia. C, macclellandi in India
and Indo-China is reddish brown above, yellow below, with
regular, equidistant, black, light-edged cross-bands or rings. Total
length up to 2 feet.

BoHophis differs from Callophis mainly by the enormously
developed poison-glands which, instead of being restricted to the
head, extend along the anterior third of the body, gradually
thickening, and terminating in front of the heart with club-
shaped ends. Owing to the extension of these glands, which
can be felt through the skin as thickenings at the end of the
first third of the body, the heart has been shifted farther
back than in any other snake. Several species in Indo-China
and in the Malay Islands, D. intestinalis with many colour-
variations.

Australia suffers from an abundance of Elapine snakes, of
which we will mention only the three commonest.

Fseudechis e.g. Fs. porphyriaceus, the " Black Snake " of
Australia, has seventeen rows of smooth scales on the body, a few
more on the neck, which however is not, or is only slightly,
dilated. A few of the sub -caudal scales are undivided, the rest
are paired. The head is distinct from the neck; the pupil is
round. Total length up to 5 or 6 feet. The general colour
above is black, with the outer row of scales red at the base :
the ventral scales are red with black edges. The females are
generally more brown than black, and are therefore sometimes
known as " Brown Adders." They live on small mammals, birds,
lizards and other snakes.

Notechis scutatus s. Hoplocepkalus curtus, the " Tiger Snake,"
has rather small eyes with round pupils. The head is distinct
from the cylindrical body, which is covered with fifteen to nine-
teen rows of smooth scales. The sub-caudals are single. The
head of this variably coloured snake is mostly black, the body
olive brown with dark cross-bands ; towards the tail the color-
ation becomes more uniformly blackish. The under parts are pale

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XIII ELAPINAE — HYDROPHINAE 63 5

yellow. The range of this very common snake extends over
Tasmania and Australia.

Acanthophis antarcticris, the " Death Adder," is easily recog-
nised by the peculiar tail, the end of which is laterally compressed,
beset with a few rows of enlarged imbricating scales, and terminates
in a thin horny spine. The head is distinct from the neck, and
flat ; the eye has a vertical pupil. The short and thick body is
covered with twenty-one or twenty-three rows of keeled scales.
The anterior caudals are single, the posterior double. The
colours of the upper parts are a mixture of brown, reddish and
yellow, with dark cross -bands. The belly is pale yellow, often
spotted with brown or black. The end of the tail is yellow,
reddish brown or black. The total length of this stout and
ugly viviparous creature remains under 3 feet. It is widely
distributed from South Australia to *the Moluccas. The use of
the peculiar tail very probably consists in attracting or fixing the
attention of small animals ; the snake, lying coiled up on a dry
and sandy spot, slightly raising and vibrating the tip of the tail.

Maps is an entirely American genus, with many species, most
of which are extremely prettily coloured, red and black in alter-
nate rings being a favourite pattern. The maxillaries carry no
teeth behind the poison-fangs. The scales of the body are smooth
and form fifteen rows. The tail is short. The small eye has
mostly a vertical pupil. The head is very small, not distinct
from the neck. The squamosal and quadrate bones are short, and
the gape of the mouth is so limited that these beautiful snakes,
although possessing strong poison, are practically harmless to
man. One of the prettiest is U. corallinus of the forests of
Tropical South America and the Lesser Antilles. The whole
body, above and below, is adorned with about twenty deep black
rings, which are edged with yellow and again separated by red
rings equalling in width the black ones. Sometimes the red
rings are dotted with black, and the black dots may form addi-
tional rings between the red and the yellow. Total length
under 3 feet.

Sub-Fam. 2, Hydrophinae (Sea-Snakes). — The tail is strongly
compressed, sometimes the body also. All the scales are
small, and there are often no enlarged ventrals. The eyes are
small, with round pupils. All these snakes are very poisonous
and live in the sea, often at considerable distances from the land,

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636

OPHIDIA

with the exception of one species of IHstira, D, semperi, which
is confined to the land-locked freshwater Lake Taal at Luzon in
the Philippines. They live on fish, and range from the Persian
Gulf to Central America. In conformity with their absolutely
aquatic life they are viviparous, and they die when kept out of
the water for any length Of time. About fifty species are known.
Enhj/drina valakadien s. hengalensis has scales with a small
tubercle or keel, which is stronger in the males ; the ventrals are
very small, forming a scarcely enlarged series. The maxillaries

Fig. 170. — Enhydrina tHtlakadien (left upper tigiire) and Hydrophia obscura
(right lower ligure). x \.

carry two or more small grooved teeth in addition to the poison-
fangs. The back is olive or dark grey, with black transverse
bands, which are most distinct in the young. The under parts
are white. This species ranges from Persia to the Malay
Islands.

Hydrophis e.g. H. ohscura. — The body is long ; the head and
neck are very slender, the body becoming much thicker farther
biick. The small teeth behind the poison-fangs are not grooved.
The ventral scales are very small, the others are keeled, strongly so

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XIII AMBLYCEPHALIDAE ^\MPERIDAE 637

in the males. The general coloration of this Sea-Snake, which
reaches about one yard in length, is dark olive-green above with
yellowish cross-bars, which form complete rings round the slender
part of the body. Other specimens are pale olive, with dark
cross-bands. This species occurs in the Bay of Bengal and the
Malay Archipelago.

As a rule Sea-Snakes are not found in mid-ocean. After
leaving Ceylon, the steamer meets them again in the Straits of
Malacca. Those which occur near the south coast of Japan, e.g.
Distira cyanocincta, are found there only in the summer, and
are probably carried there by the south-west monsoon.

According to Semper the gravid female visits the shores of
low islands, there to give birth to its young between the rocks,
and she remains with her offspring for some time. Semper
once found a large female, probably Flaiurus fasciatus s. coin-
hrinus, coiled up amongst rocks, and between the folds were at
least twenty young, each already about 2 feet long.

Boulenger ^ has written an interesting popular account of
Sea-Snakes.

Fajn. 8. Amblycephalidae. — Some thirty species of Neotropical
and Oriental Snakes have been separated from the Colubridae on
account of the pterygoids, which are widely separated from the quad-
rates, the posterior ends of the pterygoids not reaching beyond the
level of the occipital condyle. This condition can be ascertained
when the mouth is opened widely. The prefrontals are not in
contact with the nasals. The squamosals are reduced to pad-
like vestiges. Externally the Amblycephalidae are easily dis-
tinguished from the Colubridae by the absence of a longitudinal
median mental groove. The head is thick, very distinct from
the neck, and gives these harmless snakes a " poisonous " appear-
ance. The pupil is vertical.

Ainhlycephalus, e.g. A^ monticola. — Maxillaries short, with
only five or six teeth. Sub-caudals in two rows. Body com-
pressed, covered with fifteen rows of scales. South-Eastern Asia.

Fam. 9. Viperidae. — The maxillaries are very short, movably
attached to the prefrontals and ectopterygoids, so that they can
be erected together with the large poison-fangs, which (besides
reserve- teeth) are the only maxillary teeth. The prefrontals are
not in contact with the nasals. The squamosals are very loosely
^ Natural Science^ i. 1892, p. 44.

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638 OPHIDIA CHAP.

attached. For further details see Fig. 180. The poison-fangs
are perforated, having a wide hole on the anterior side at the base,
in connexion with the large poison-gland ; the hole leads into a
canal, which opens gradually as a semi-canal on the anterior
surface of the distal third or quarter of the tooth. As usual in
poisonous snakes, several reserve-teeth are stowed away behind
the acting fang. . When the latter is broken off or has ser\'ed its
time it is cast off at the base, and the next reserve tooth takes
its place. The supply of resei*ve-teeth is indefinite, half-finished
teeth down to mere germs constantly growing.

All the Viperidae are very poisonous, and all, except the
African Atractaspis, are viviparous. They include terrestrial,
arboreal, semi -aquatic, and burrowing types. The family is
cosmopolitan, excepting Madagascar and the whole of the
Australian region ; it is divided into Vipers and Pit- Vipers.

Sub-Fam. 1. Viperinae (Vipers). — There is no sensory

■iViPERINAE.

Fig. 171. — Map showing the distribution of the Sub-Family Viperinae. Corsica and
Sardinia shouhl be black in the map.

external pit between the eye and the nose, and the maxillary is
not hollowed out above. The Vipers are absolutely restricted to
the Old World, ranging over the whole of Europe, Africa, and
Asia, with the exception of Madagascar ; their northern extension
is limited only by the permanently frozen condition of the under-
ground. Nine genera with about forty species are known.

Causus with a few species in Africa and Azemiops feae in
Upper Burmah are the only vipers which have the head covered

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XIII

VIPERINAE

639

with large symmetrical shields, while in the other genera the
head-shields are broken up into scales or small shields. Causus
rhomheatus is very common in Africa, from the Gambia to the
Cape. It reaches a length of a little more than 2 feet. Pale
olive-brown above, usually "with a dorsal series of large rhombic
or V-shaped dark brown, sometimes white-edged spots, and with
a dark arrow-shaped mark on the occiput ; under parts yellowish
white or grey.

Bitia s. Echidna, — Very much like Vipera, but the nasal

Fio. n%—BUU arietans (Puff Adder), x J.

shields are separated from the rostral by small scales, and the
postfix)ntal bone is very large. Several species in Africa.

The head is very distinct from the neck, chiefly owing to the
large poison-glands and to its being, like the body, much depressed.
The small eye has a vertical pupil, and is separated from the
labials by a series of small scales. The scales are keeled, and
form many, from twenty-nine to forty-one, rows ; the tail is very
short, with two rows of scales below.

In B, arietans, the " Puff Adder," the nostrils are directed
upwards. This ugly brute is yellowish to orange brown above
with regular, chevron-shaped dark bars or other markings, helping

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640 OPHIDIA

CHAP

to conceal the creature when it is lying on sandy and stony
ground ; the under parts are yellowish white. The PuflF Adder
reaches a length of 4, or very rarely 5 feet, ranging all over
Africa, except the north coast, and extending into Southern
Arabia. It is very slow, and trusts to not being discovered when
lying in the dry grass; when approached it inflates the body
and hisses loudly with a puffing sound, watches the enemy with
raised and characteristically bent head and neck; but it bite«
only when actually touched or attacked. The efifect of the bite
is very dangerous. Its prey consists chiefly of small mammals,
which are hunted during the night.

B. {Echidna) naskorriis, of Tropical West Africa,, has two or
three enlarged scales above the supranasals ; they stand upon
erectile tissue so as to form horn-like elevations. This " Xose-
horned Viper " grows to a length of 4 feet, and is rather prettily
marked ; the ground-colour is pui-plish or reddish brown, with a
vertebral series of large, pale, dark-edged spots and oblique
crosses. The young are at birth as much as one foot in length,
and are very tastefully coloured.

Cerastes and Echis prefer to burrow in sand. The lateral
scales are smaller than the dorsals, and arranged obliquely with
serrated keels, so that the snakes can cover themselves with sand
by lateral shovelling motions of the sides of the body.

Cerastes cornut us, the " Horned Viper" of North-Eastern Africa,
from Algeria to Arabia, extending also into Palestine, has the
sides of the ventral scales bent angularly, with an obtuse keel on
either side. Above each eye stands a large horny, spiky scale.
The upper parts are pale yellowish brown, mostly with dark
spots arranged in several longitudinal rows. The under parts are
white. This, or perhaps C. vipera, which has no horns, is
supposed to be the species which has become famous through the
suicide of Cleopatra.

About twenty years ago a number of "Horned Vipers" were
brought to the Zoological Gardens of London, and attracted
attention by their unusually long horns. It was found that
some wily Egyptian snake-catcher had tried to manufacture a
new species by taking specimens of the hornless C. tnpera and
inserting a pair of hedgehog's spines, pushing them upwards
through the mouth.

The " Horned Viper " attains a length of two feet and a half.

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VIPERINAE 641

S.c-
k,

IT-

In the daytime it is invisible, being buried in the sand with
only the eyes, nostrils, and the " horns " appearing above the
surface.

Vipera, — -The head is distinct from the neck, and is covered
with small scales and a few larger shields. The eye is separated
from the labials by scales ; the nasals are in contact with the
rostral shield or separated by one naso-rostral shield. The scales
on the body are strongly keeled ; they are in two rows on the
short tail. This genus with about ten species ranges over
Europe, Asia, and the greater part of Africa.

F". herus, the Common European Viper (see Fig. 165, p. 620).
The snout is not turned up at the end ; between the small head-

FiQ. 173. — Cerastes comuttts^ the "Homed Vijier" (right), and Vipera amnwdytesy the
" European Nose-horned Viper " (left), x 1.

scales there is generally a pair of well -developed parietal and
frontal shields. The scales of the trunk form twenty-one rows.
The coloration is very variable, there being grey, brown, red, or
black specimens in the same country, and the much -spoken- of
black zigzag line along the back is so often indistinct that it is a
character not to be relied upon. Usually the grey, yellowish,
olive, brown or red ground-colour is set oft' by a dark zigzag band
along the spine, and by a series of lateral spots ; an oblique or
St. Andrew's cross or two diverging bold streaks of dark brown
or black are usually present on the back of the head, and there is
a dark streak behind the eye. The under parts are grey, brown,
or black, uniform or speckled ; the end of the tail is usually
yellow or red. According to Boulenger, who is making a special
study of the individual variations of Vipers (concerning colour,
scaling, number of vertebrae, etc.), some specimens are entirely

VOL. VIII 2 T

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642

OPHIDIA

CHAP.

black in the males through extension of the black markings,
in the females through darkening of the ground-colour. Males
are usually distinguishable from females by darker, deep black
markings and lighter ground-colour. The females are mostly
larger than the males. The largest specimen in the British
Museum measures 700 mm. = 28 inches, but a viper 2 feet
long may be considered a very large specimen. The Common
Viper has a wide range, from Wales to Saghalien Island, and from
Caithness to the north of Spain. It agcends the Alps to a con-
siderable altitude, up to 6000 feet. J. Blum^ has published an
elaborate statistical account of the Viper in Germany, un-
fortunately confining himseK strictly to the political frontiers.

According to the map
attached to his work, the
Viper is common all over
Germany with the excep-
tion of South - Western
and parts of Middle Ger-
many. It is absent in
Alsace, the Bavarian Pala-
tinate, Ehenish Prussia,
Hesse, the northern half
of Baden, Wiirtemberff.
and Franconia, countries
which, speaking broadly,
have a warm subsoil, composed of Red Sandstone and Basaltic
formation. As a rule the Viper prefers heaths, moors, and mixed
woods with sunny slopes. Brambles, clumps of nettles, hedges,
the edges of little copses, heaps of stones, are favourite places of
retreat, affording shelter, holes, and the vicinity of mice, which
form its chief sustenance. At harvest-time it is often found in
cornfields, and it frequently hides in the sheaves. Vipers are
fond of basking on certain spots, on the top of a stone, the
stump of a tree, or a patch of sand : a shower of rain or even
passing clouds drive them back into their holes. They are
eminently nocturnal, when they regularly " beat " their district,
biting and paralysing their prey before swallowing it. A fire
kindled at night is sure to draw vipers near ; the same applies
to other vipers, for instance Cerastes, which appears in perplexing
* Verhreitung dcr KreuzotUr in Deutschland, Frankfurt a. M. 1888.

Fio. 174.— Skin of Viper, x 1. (From White's
History of Selbome. )

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xin VIPERINAE 643

numbers at the camp-fire. They cannot climb, and they avoid
going into water. The pairing takes place as a rule from March
to May, a number of individuals, mostly males, collecting around
the females, and forming entangled lumps of snakes ; parturition
takes place in the following July and August. In exceptionally
warm winters they have been known to pair in December, having
left their winter-quarters. They hibernate for about six months,
more or less according to the climate, congregating in great
numbers, sometimes in dozens. With very rare exceptions
Vipers do not take food in captivity, but prefer starving them-
selves to death. The bite is as a rule not fatal The seriousness
of the case depends of course upon many circumstances, as for
instance the state of concentration of the venom, the position and
depth of the bite, and last but not least upon the general condi-
tion of health of the victim. General depression aggravated
by nervousness, weakness of the bitten limb, occasional breaking
out of the wound, are of frequent and protracted occurrence.
(See also p. 590.)

V. aspis is a more southern and western European Viper,
occurring from France to the Tyrol, and in Italy. The snout is
slightly turned up at the end, and still more so in V, latastei
of Spain and Portugal. In V. amiiwdytes, of South-Eastern
Europe, the raised portion is produced into a soft, scaly
appendage (see the lower figure on p. 641). Vipers are some-
times unpleasantly common in certain localities. This was for
instance the case at the drill-ground near Metz,and the military
authorities paid a price for each viper delivered to them. The
supply of the latter increased to an alarming extent until the
German authorities discovered that a regular trade had been
established across the frontier, and that the French Lorrainers
were importing vipers briskly.

V, russelli, the " Daboia " or Russeirs Viper, is one of the
scourges of India, Ceylon, Burma, and Siam. The scales form
about thirty rows on the body. The upper surface of the head
is covered with small, imbricating, usually keeled scales. The
general colour is pale brown above with three longitudinal series
of black, light-edged rings, which sometimes encircle reddish spots.
The under parts are yellowish white, uniform, or with small
crescentic black spots. Total length up to about 5 feet. The
poisoning symptoms are described on p. 590.

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644

OPHIDIA

CHAP.

Sub-Fam. 2. Orotalinae (" Pit-Vipers "). — With a deep cavity
or pit between the eye and the nose, lodged in the hoUowed-
out maxillary bone. This pit is lined with a modified continua-
tion of the epidermis, and is amply supplied
with branches from the trigeminal nerve. It
is undoubtedly sensory, but we do not know its
function. A good anatomical account of this
organ has been given by West.^ Some of the
Pit- Vipers have a rattle at the end of the tail ;
these are the Eattle-Snakes. The rattle is
composed of a number of homy bells which fit
into each other. The oldest or terminal bell is
in reality the horny covering of the tip of the
tail, and with each moult or shedding of the
skin the youngest bell becomes loose, but is held
by the new covering which has been developed
in the meantime. There is thus produced an
ever-increasing number of loosely-jointed bells,
^'''o^l^.'r^"^ °^ but now and then most or all the bells break

Rattle-Snake.

(Prom White's jyw- ofif, probably when they are worn out, and a

ton/ 0/ sdbame.) ^^^^ g^^ -g gradually developed. Rattles with

a dozen bells are, for instance, very rare. They naturally

increase in bulk with the age of the snake, but the number of

joints is no indication of the snake's age.

-Jj 9'^' --^

I OROTALINAE,

Fig. 176. — Map showing the distribution of the Sub-Family Crotalinac.

Pit- Vipers have a very wide distribution. They are divided
into four genera with about sixty species. Eattle-Snakes are

^ J. Linn. Soc. xxviii.

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XIII

VIPERIDAE CROTALINAE

645

restricted to America, but other Pit- Vipers occur in North and
South America and in the southern half of Asia.

Ancistrodon. — Without a rattle. The upper surface of the
head is covered with nine large shields, but the internasals and
prefrontals are sometimes broken up into scales. The scales of
the body have apical, sensory pits. About ten species, some in
Central and North America, others in the Caspian district {A,
hcUys), in the Himalayas (A, hiincdayanus), in Ceylon, Java, etc.

hi^

«^|3

Fro. 177.— ^«cw^rorfon/>WCTran« (Water- Viper), x ^.

A, piscivorus s. Trigonocephalus cenchris (part), the " Water-
Viper," inhabits North America from Carolina and Indiana to
Florida and Texas. The general colour is reddish to dark brown,
with darker cross-bands or with Q-shaped markings ; a dark,
light-edged band extends from the eye to the angle of the mouth.
The under parts are yellowish, spotted with black, or the latter
is the prevailing colour. Total length up to 5 feet. The
Water- Viper is semi-aquatic and lives chiefly on fishes, but it
also eats other snakes and various Amphibia, Birds, and Mammals.
This snake is very good-humoured in captivity, and becomes

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646

OPHIDIA

CHAP.

easily tame. A gentleman in Berlin, rather too much addicted
to niaking pets of poisonous snakes, had a pair which propagated
regularly. When I was a boy he invited me to feed the young
Water -Vipers with fishes cut into strips, and I enjoyed this
immensely until he warned me not to touch the mother, which
might bite strangers.

A, contortrix s. IVigonocephaliis cenchris (part), the " Moccasin-
Snake " or " Copper-head," is one of the few poisonous snakes

Fig. 178. — Ancistrodon contortrix {MoccasinSnAke or Copper-hetid). x\.

which possess a loreal shield, i.e. a shield intercalated between
the pre-oculars and the nasals ; below it lies the pit. The general
colour is yellowish to pink or pale brown, with dark brown or
red cross - bars or triangular marks. The under surface is
yellowish or reddish, speckled with grey or brown, and with a
lateral series of large blackish spots. Total length of full-grown
specimens about one yard. The Moccasin-Snake ranges from
Massachusetts and Kansas to Northern Florida and Texas. It
prefers swampy localities or meadows with high grass, where it
hunts for small Mammals and Birds.

Lachesis. — Without a rattle. The upper surface of the head
is covered with very small shields or with scales. About forty

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CROTALINAE 647

species in South - Eastern Asia and in Central and South
America.

L, (Bothrops s. Craspedocephaliis) lanceolatus inhabits nearly
the whole of South America, extending into Mexico and the
Lower Antilles, e,g, Martinique, Guadaloupe, and Santa Lucia,
where it is known as the " Fer-de-Lance," and is the curse of
the sugar-plantations on account of its being so very common
amd so deadly poisonous. The Mongoose was introduced as
a possible antagonist, but the little Indian Mammal wisely left
the dangerous reptile alone, and has in some places established
himself as another pest — as a destroyer of poultry. The Fer-de-

Fig, 179. — Head of Lachesis lanceolatus after removal of the skin, x 1. />, Duct, bent
upon itself, from the poison-gland into the tooth ; Dig, digastric muscle or opener
of the jaw; Nj nostril; P.G^ poison-gland; S.Gr^ sensory groove or pit; *S.Q,
point of junction of the squamosal and quadrate ; T.a, Temp.Uj anterior, and T.p^
posterior, temporal muscle.

Lance grows to a length of 6 feet, establishes itself everywhere —
in swamps, plantations, forests, in the plains and in the hills —
and is very prolific, producing, according to its size, dozens of
young which are 10 inches long, very active and snappy.

L. (Trimeresurus) gramineus s. viridis, to mention one
Asiatic species, grows to less than 3 feet in length, is bright
green above, sometimes with faint blackish bars ; green, yellow,
or whitish below, and with a light streak along the outer row of
scales. The end of the tail is usually bright red. This beautiful
snake has a prehensile tail and is arboreal. Its range extends
ovtm the whole of India, to Hong-Kong and to Timor, and even
into the Andaman and Nicobar Islands.

Sistrurus. — With a rattle. The upper surface of the head
is covered with nine large shields. A few species in North
America east of the Eocky Mountains, e.g. S. miliarius.

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648

OPHIDIA

CHAP.

CroUdus, — With a rattle. The upper surface of the head is
covered with small scales. Range from Southern Canada and

Prax

fru^f;

PwixfC>

Fio.

180. — SkuU of a Rattle-Snake, Crotalus durissus, x 1. ▲, Lateral view, jaws
slightly opened ; B, ventral view ; C, lateral view, the jaws opened folly in the
position of striking ; D, dorsal view. Compare this with the diagrammatic figures
on p. 588, where the mechanism has been explained. Col, Columella auris ; Cond,
condyle ; Or (in B), sphenoidal crest for the attachment of the powerfully developed
ventral cranio-cervical muscles ; KP, ectopterygoid or transverse bone ; F, frontal ;
Max, maxillary ; P, parietal ; P^, post-orbital process ; Palf palatine ; Pmx, pre-
maxillary ; Pr/, prefrontal ; Ptg^ endopterygoid ; Q, quadrate ; Sq, squamosiiL

British Columbia to Northern Argentina, but not in the West
Indian Islands. About ten, mostly closely-allied species.

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XIII CROTALINAE 649

The efiTect of the poison of Eattle-Snakes has been discussed
on p. 589.

C. horridus is the common Eattle-Snake of the United States ;
C. confluentvs is the • species in Western and C. durissus the
common species in South -Eastern North America. Very large
Kattle-Snakes, C. durissus, attain a length of 8 feet, others not
often more than five. They prey chiefly upon small Mammals,
hunting for them at night. In the daytime they are also
about, mainly in order to bask. Although they occasionally
take to the water in pursuit of their prey, they dislike being
wetted by rain, withdrawing then into their holes, appropri-
ating as a rule those of ground-squirrels, rats, and Prairie-
dogs. The often-repeated story about Eattle-Snake^ living in
neighbourly friendship in the holes of Prairie-dogs, together
with the little Prairie-owls, is an exaggeration. We do not
know how many of the original inmates are eaten. Pairing
takes place in the spring. During the cold months they
hibernate under ground, often in considerable numbers.

Rattle-Snakes have few enemies besides man and pigs. The
latter kill and eat them wherever they can. The rattle is
decidedly useful to the snake as an instrument of warning off
any approaching possible enemy, since no snake likes to bite
unless in self-defence or in order to kill its prey. The noise of
the rattle is very loud in dry weather, much duller on clammy
days ; it is a shrill sound like that of a rattling alarm-clock,
and a well-conditioned snake in a room can make conversation
well-nigh impossible, and can keep on rattling for half an hour
or longer. The rattle is kept in such rapid lateral vibra-
tions that it shows only a blurred image, the rattle standing
with its broader sides vertically, not horizontally. They endure
captivity for many years, and become tame enough not to hiss
and to rattle whenever they are approached.

C, horridus is grey - brown above, usually with a rusty
vertebral stripe and with V- or M-shaped blackish cross-bands ;
the under surface is yellowish ; the end of the tail is blackish.
The supra-ocular shields are smooth and much narrower than the
scaly space between them, and there is only one pair of inter-
nasals.

C, durissus s. adama7iteus differs from the previous species
chiefly by possessing two pairs of internasals; and the dark

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6so

OPHIDIA

CHAP. XIII

markings on the body form a handsome pattern of rhombs with
lighter centres and yellowish edges. This is the largest species
of Rattle-Snake, reaching a length of 8 feet.

C, confluentus has broader, transversely striated, supra-ocular

m.ep.

Pio. 181. — CrotcUus durissus s. adamantevs (Rattle-Snake), x J.

shields. The specific name refers to the continuous series of
large brown or red rhomboidal spots on the back.

C terrificiis ranges from Arizona to Argentina, and is the
only species of Rattle -Snake in South America. It differs
from the others by having a pair of prefrontal shields behind
the pair of internasals.

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INDEX

Every reference is to the page : words in italics are names of genera or species ; figures
in italics indicate that the reference relates to systematic position ; figures in thick
type refer to an illustration ; f. = and in following page or pages ; n. = note.

A^xlominal armour, of Cricotus^ 287 ; of

Microsauri, 289 ; of Prosauri, 290
Abdominal ribs, of Rhynchocephali, 292,

298; of Dinosauria, 414; of Megalo-

S(turu8j 421
Aberrant scaling of Lacertilia, 495
Ablepharus, 660 ; eyelids, 494
A canthoilaciylus vulgaris, 659
Acanthopkis atUarcticus, GS6
Acentrous vertebrae, i.e, those without a

centrum or body, 4
.'1cm, 186, 189; A, gryllua, 207 f.
Acrochordinae, 606
Acrochfyrdiis javanicuSf 607
Acieosaurus^ 489
Actitiodon, 83 ; 287, 288
Adams, visit to the Mugger-peer, 455 f.
Adaptive characters of Anura, 142
Adhesive apparatus, of tadpoles, 57, 57 ;

of Tree-frogs, 187; of Thoropn, 209;

of finger -discs of Raninae, 239 ; of

Geckos, 505, 606
Adurosaurus, 807

Aestivation, of Crocodiles, 457 ; of Tor-
toises, 357, 365, 404
Aetosaurus^ 432 ; Ae^ferratus^ 4^8, 448
AgcUychnis, 189, 206
Agama, 520; A. mnguinolenta, 620 ; A.

stellio, 521, 631
Agamidae, 513, 615 f.
Agaviurcu, tail, 506
Agassiz, on habits of Alligator Turtle, 341 ;

of Trionyx, 407
Age of Chelonia, how to estimate, 326 ;

great age attained by Tortoises, 369, 376,

377 ; see also Growth, rate of
Aglossa, 139, I4O, I48 f. ; dUtribution,

143
Aglypha, 592, 606 f.
Ahaetulla s. Leptophis, 618, 619
Aistopodes, 81

Aldabra, gigantic tortoises of, 373 f., 375
Algae, destructive to shell of tortoises, 357
Allantois, an embryonic outgroiK'tli from

the ]>08terior part of the gut, acting as

a respiratory organ, 278
Alligator, 4^0, 4^^ ^« » -^' mississip-

piensis, ^67 f. ; skull, 468 ; nesting,

469 ; A. sinensis, 471
Alligator Turtle, 340
Allopleuron hofmanni, 380 *
AUosaurus, 4^^

Alpine, Newt, 126 ; Salamander, 119
Altitude, high, in which Anura have been

found, 181
Alytes, 157 f. ; A. cistemasi, 160; A,

obstetricans, 158; urino-genital organs,

49
Amblycephalidae, 592, 593, 637
AmblycephtUus monticola, 637
AmUyrhynchus, 528 ; A. cristatus, 633
Amblystoma, skull, 17, 94, 96, 109, 110 f.,

112; A. jeffersonianuvu 111; A, ina-

vortium, 116 ; A. opacum, 110; A.per-

simile, 111; A. punetatum, 110; A.

talpoidewnu, 110 ; A. tigrinum, 111 t. ;

metamorphosis of, 112 f.
Amblystomatinae, 102, 109
Ameiva, 549
Amnion, a membrane round the embryo,

278
Amphibia, 3 f. ; definition, 5 ; systematic

position, 5 ; numbers of species, 4
Aniphicondylous, i.e. the occipital part of

the skull articulates with the neck by a

right and a left knob, 4
AinphigwUhodon, 185 ; A. gtientheri, 188
Amphignathodontinae, 139, 188
Amphisbaena, 666 ; A./nliginosa, 566
Amphisbaenidae, 614^ 666 f.
Amphiuma, 88, 96 ; ^. meaiis s. tridactyla,

100, 101

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652

INDEX

Amphiumidae, 94i 97

AmphoduSj 210; A. itmcherif 211

Anaconda, 603, 608

Anaidet = Autodax (q.v.), 107

Anal sacs of Chelonia, used as additional
respiratory organs, 330

Anarosaurtis pumiliOf 477

Anchiaaunu, 416, 417, 421; skull of A,
coeluruSf 421

Ancistrodon^ 64S; A. contortrtx, 646, 646 ;
A. ha/j/s, 645 ; A. himalayantis, 646;
A, piscivoruSf 646, 646

Anderson, on nest of Oavialis^ 452

Andrews, on Amblystonui, 110

A ndnas sdieuchztri, 84

Anelytropidae, 514^ 564

A iveJytropsis papilloaus, 564

Anguidae, 513, 537 f. ; distribution, 501,
629

Anguis fragilis, 639, 639

Aniella piUchra, 564

Aniellidae, 514, 564

Annandale, on habits of Calotes, 518 ; of
Liolepis, 627 ; of Rhacophorus, 247 ; of
Varanus salvator, 544

A nodonta, as food of Trionyx, 407

A nodojitohyla, 236

Anolis, 528 ; A. carolinejisis, 629

Aiiomodoutia, 309

Anura, 7 ; characters, 138 ; classification,
139 f., 14f ; phylogenetic tree of, 142

Anus, asymmetrical position of, 60

A]>oda, 84 f. ; affinities, 88 ; distribution,
89 ; eyes, 86 ; skin, 87 ; skull, 84, 86 ;
spermatozoa, 87 ; tentacular apparatus,
88 ; vertebrae, 86 ; visceral arches, 86

Archaeopteryx, 417

Archegosaurus, vertebrae, 13, 82, 287

Arcifera, of Cope, 140 ; of Boulenger, 140

Arciferous, type of shoulder-girdle, 24, 26

Avion, slug, eaten by tortoises, 363

Arrau-turtle [Podocnemia), 391 f.

Arteria cutanea magna, 144 ; A. sacralis
of Anura, 144

ArthrolepiU, 241, 242; A. seychellensis,
243, 243

Ascaphiis, 153

Asterophrys^ 161

Athecae, 333, definition of name, 337

AUnntosavrus, 415, 4IO \ -^1- immanis,
419, 420

Atlas and Axis, i.e. first and second
cervical vertebrae ; of Cryptohranckus,
13 ; of Crocodilia, 283 ; of Chelonia,
283, 316 ; of Sphcnodon, 288, 294 ;
atlas fused with axis, 307

Atoposauridae, 453

Atractanpis, 638 ; dentition, 593 n.

Atria, the thin-walled receptive parts (au-
ricles) of tlie heart

Auditory columellar apjuiratus, of Am-
phibia, 24 ; of Anura, 29

Australian, Anura, spawning time and
habiU of, 201 ; Lacertilia, 502

Autodax, 96, IO4, lOI ; A. luffubris, 107;
A. iecanus, 1&7

Autosanri, 491 f.

Axis ; see Atlas

Axolotl, 65, 112 t, 112 ; Keoteny of, 65,
112

Azemiops feae, 638

Balancers of Amphibia, 45

Bapiajiodan, 483, 484

Barfurth, on absorption of Tadpole's tail,

61
Bartlett, on Boa constrictor, 602 ; on Pipet,

162
Basiliseus, 528, 630 ; B. amerieanus, 530,

680
Bates, on habits of Podocnemis, 392 f.
Batrachomyia, fly infesting Bufonidae, 177
Batrachophrynus, 224; B. macrostomus,

225 ; B. brachydaclylus, 224
Batrachopsia, 161
Bcdrachotepa, 96, IO4
Bairachylodes, 24I
Batrachyperus, 96 ; B. sinefuia, 109
Baur, on Sphargis, 336
BdeUopkis, 90
Bedriaga, on Axolotl, 114 ; synopsis of

Urodelous Larvae, 59 n.
Bell, J., on classification, 8
Bell, Napier, on habits of Iguana, 531
Belodon, 305, 434, 448
Bemmelen, on Sphargis, 336
Berg, on Spelerpes fuscus, 106
Bert, quoted, 571 n.
Bidder's organ, 49, 52
Biedermann, on change of colour in ffyla,

35
Binls not related to Dinosaurs, 416 f.
Biti-s arietans, 639, 689 ; B, nasicomis,

640
Black Snake, of Australia, 634 ; of North

America, 613
Blainville, de, on classification, 7
Blanns cinereus, 666

Blood, shape of red corpuscles, 4 ; tem-
perature, 67 f.
Blood-sucker = (Motes opkiomachtu, 519
Blum, quoted, 642 n.
Boa, 602 ; B. constrictor, 602 ; B. dumertliy

602 ; B. madagascarifnsis, 602
Boettger, on influence of dimateand country

upon reptiles, 492 f.
Boidae, 692, 696 f. ; skull, 696, 69T
Boinae, 601 f.
Bovibinator, 164 f, IBB ; habits, 156 f. ;

tadpoles, 167 ; abnormal vertebrae, 22 ;

slioulder-girdle, 26 ; urino-genital organs,

49 ; B. igneus, 164, IBB ; B. padiypus,

155
Bothrops, 647

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INDEX

653

Boulenger, claadflcation of Amphibia Cau-
data, 9 ; on vertebrae of Pelobates, 20 ;
on vertebrae of Bombinatarj 22 ; number
of phalanges in Anara, 27 ; on poison of
Amphibia 86 ; on vocal sacs, 48 ; on
modes of fecundation and nursing habits,
54, 56 ; synopsis of Tadpole8,59 n. ; on
tadpoles of Hana opUlhodon^ 260 ; on
clas8itication of Anura, 140, 141 ; on
Pt/xi,152 ; on Scaphiapus aolitantiSyl66 ;
on Alligator sinensiSy 471 ; on Lanth-
aiwtus^ 542 ; on aberrant scaling, 495 ;
on Helodei-may 540 n. ; on classification
of Snakes, 592 ; on Sea-Snakes, 687 ; on
SphargUy 336

BoulengenUoj 90

Box-Tortoises, S62, 364, 865

Brachial plexus, of Aunra, 39

BrachycephaluSy 226, 227 ; B, ephippiuniy
231

Brachyl&phus, distribution, 601, 528

Brain, of ScaphvgiuUhuSy 485 ; small size
of, in Dinosaurs, 425

Branchial arches, of Urodela, 16 ; of Anura,
42

Branchiosauri, 80

Branchiosau-niSy skull, 80 ; B. aalamand-
roides^ 80

Brauer, on development of Apoda, 92 ; on
nursing habits of ArthrdeptiSy 243

Breeding of AxolotI, 113

Breviceps, shoulder-girdle, 25, 225, 226,
'2J7, 232 ; B. vwsmmbicua, iS2

Brithcpusy 308

Brongniart, on classification, 7

BrontosauruSy 415, 4^8 ; B. excdsus, 418

Brontozauniy 415, 417; B. gigantcum, 420

Brood -pouches, of Anura, 151, 248 ; of
Hyla goeldii, 198 ; of Notoiremay 202 ;
of Rhinodemia, 228

Brookesiay 680

Brol^^l Adder, GS4

Brown Frog, Common, 251 f., 256

Briicke, quoted, 571

Buchholz, on ChiromaniiSy 244 f«

Budgett, on breeding habits of PhylJo-
mediutay 204 ; on Paludicolay 220 ; on
LejridobatraehuSy 218 ; quick develop-
ment of Phryniacus, 231 ; on Bv/o
marinusy 179

Bvfo, sacral vertebra, 22 ; shoulder-girdle,
26 ; nrino-genital organs, 49 ; develop-
ment of adhesive apparatus, 67 ; B.agva^
178; B.americavvSy 178 ; B. calamitay
181 f. ; B. ceratophn/Sy 179 ; B. empusus
and B. peltocephalusy dermal ossifica-
tions, 179 ; B. jerboa, 166 ; B. lenti-
ginosusy 17 8 y 179 ; map of distribution,
167, 168y 169 f. ; B. warinvSy 178 ; B.
mauritanica s. jHintherimiy 184; -S*
viehnwstictuSy 177 y 179 ; B. quercinus^
178; B. variabilis = viridisy 180 ; B.

viridisy 180 f., 493; B. vulgarUy 170 f.,
172 ; large-sized specimens, 171 ; im-
mured in buildings, 174 ; diseases, 176 ;
distribution, 177

Bnfonidae, lS9y 166 f. ; distribution, 167 ;
affinities, 166

Bufoniformes, 139

Bullfrog, of America, Bona catesbianoy
261 ; of India, Callula ptUchrOy 234 ;
Jtana tigrinay 261

Bungarus coeruleus s. candiduSy 633 ; B.
faaciatvsy 633

Butler, on fat-bodies, 500

CabritOy 551

CacapuSy shoulder - girdle, 25, 225, 226,

228
Cacosteniumy 225, 227
Gaimaiiy 460y 4'^^ : vomer, 435 ; C. nigtTy

47 ly 472; C. paipebrosusy 471; C.

sclerops, 471 ; C. trigonal us y 47 It 4"^ 2
Calcareous deposits in the skin of Am-
phibia, 81, 34
Catliphora sUvaticay fly infesting Bvfoy

176
Callopkis macdellaiidi, 634
CallueUay 285, 236
Callulay 226, 228y 234; CpulckrOy habiis

of, 234 f.
Callulopsy 225, 228
Calophrynusy 225, 227
Calotes, 517 ; C. nmnUy 518 ; C mystacevSy

619; C. ophioniachuSy 519; C. versi-

COl&Ty 518

CalyptocephalxiSy 179, 212y 215

CamptomuruSy 420

Capitosaurxis^ 83

Carapace, 321 f., 819, 320, 822. 323;
posterior portion movable in CtJii/xiSy
364i 866 ; carapace of tortoises, evolu-
tion of, 387 ; composition of, 324 f. ;
reduction of component elements, 325 ;
reduction in thickness, 373 ; correlative
changes, 328 ; of Sphargij*y 336 f. ; of
ChelonCy 379 ; of TestndOy 822 ; of Pleu-
rodira, 389 ; reduction in Trionychidae,
325 ; fenestration, 325 ; with hinge in
CinyxiSy 364, 865

Cardioglossay 274

Carettochelydidae, 313, 314

CareltochelySy 337, 389, 390 ; C. insculpttty
404 ; absence of homy shields, 325

Carpet Snake. 698, 699

Carpus {see also Linibs), of EryopSy 286 :
of Sphenodon, 294 ; of Chelonia, 320,
820 ; of Eusnchia. 440

Casarea, 003

Case, on Sphargis, 336

Cassina, 240

CausHs^ 03S ; C. rhombefftuSy 639

Centrolenc geckoideum, 211

Cerastes cornutus, 040, 641

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654

INDEX

Ceratobatracliidae, 141

Ceratobatrachinae, 139, ^87 f.

CercUobcUrachua giientheri, 237

Ceratohyla, 211

Ceralojiiora, 617; C. stoddarti, 617 ; C.
Unnentif 617

Ceratopkrys, 212, 216 f. ; C. comuta, 216 ;
C. dorsata, 215 ; C. anuUa, 216, 217

Ceratopsia, 430

CeratosaurtiSf 418,. 416, 417 ; C. nasiconiis,
422, 422

Cerberus rhynchops, 626

CetioaauniA, $19

Chalarodon, 628; geographical distribu-
tion of, 501

Ohalcides, 662 ; Ch. bedriagae, 663 ; Cfh.
guentheriy 563 ; Ch, lineatus, 563 ; Ch.
* ocellatus, 563; Ch, tridactylm, 563

Chamaeleon, 673 ; Ch. biJidiiSy 580 ; Ch.
calcaratiiSf 579 ; Ch. paraoniy 580 ; Ch.
pumihis, 579, 575 ; Ch, vulgaris, 573,
674, 676

Chamaeleontes, 667 f. ; distribution, 568 ;
skull, 568, 669 ; tongue, 569 f.; colour-
changing mechanism, 570, 671, 573 f.;
eggs, 572

Chamaeleontidae, 673 f.

Chaniaeropa ?iumilis, dates of, eaten by
Testitdo, 367

Chameleon, misnamed Calotes, 618 ;
misnamed Polychrus, 529

Chauvin, Marie von, on Axolotl, 113 ; on
Salamandra cUra, 120

Chelodina, suppression of neural plates,
324 ; iutergular shields, 389, 815 ;
skull, 399 ; Ch. longicollis, 402 f., 403

CheloM, skull, 817 ; skeleton, 820 ;
plastron, 821 ; shields, 827 ; intergular
shields, 325 ; Ch. viydas, 381 f. ; various
modes of fishing, etc., 382, 883 ; Ch,
imbricataf 384, 386

Chelouemydidae, 380

Chelonia, 312; number of species, 312;
affinities of, 312 ; classification, 313 ;
key to living families, 314 ; plastron,
names of the horny shields, 815, 821,
325; vertebrae, 314 f., 816; skull,
280, 817, 856, 864, 8T9, 400, 405;
skeleton of Tfjitiulo, 819; of Chehme, 320;
IKctoral arch, 318 ; pelvis, 319 ; plas-
tron, bones of, 821 ; limbs, 320 ; lx)ny
shell, 321 f., 822, 823; evolution of,
337 ; evolution of the homy shields,
326 f., 827 ; regeneration, 329 ; sense-
organs, 329 ; digestive apparatus, 330 ;
respiration, 331 ; growth of Chrysemys,
349

Chclonidae, SIS, 314, 378 f.; affinities of,
380

Chelydidae, 313, 314, S90 ; distribution,
332, 383

Chehjdosaurus, 82, 287

Chelydra, 328 ; Ch, serpentina, 33S
Chelydridae, 313, 314,338; distribution

of, 882
ChdydropsU, nuchal plates, 324
Chelys finibriata, 4OO ; skull, 400, 401;

intergular shields, 325
Chersydrus gramdalus, 607
Chioglossa, 96, 116; Ch, lusUanica, 131
Chirixalus, 24I
Chirdeptes, 209, 213, 221 ; Ch. platyeeph-

alut, 221
ChiroinaiUiSy 238, 24I, 244 / Ch, petersi^

244 *' Oh, xerampelina, 244
ChiroteSf 564 ; Ch, canal iculalus, 600
Chirotherium, 83
Chlamydosaurus kingi, 522, 628
Choanae, or inner nasal openings, 47
Chorda dorsalis, the axial rod between the

gut and the spinal cord, around which

the vertebrae are formed, 12
<^i&rophUus, 186, 189; Ch. omatits, 208
Chromatophores, 35
CJirysemys, costal plates of; 325 ; green

colour of, 328, 846 f.; colour of iris,

329 ; Ch, wncinna, 346, 849, 860 ; (Ju

elegans, 346 ; Ch. picta, 346, S47, 348 ;

Ch. rubriventn's, 846
Chthonerpeton, 87, 90
CimoliasauruSy -^78 ; C. australist ^75 / ('.

cantabrigiensis, 478 ; C. chilensix, ^75 ;

C. haasti, 478
Cinosternidae, 313, 314, 342 ; distribution,

882
Ciiwstemum, 342 f. ; arrangement of

neural plates, 324 ; C. leueostoMum, S42,

344 ; C. odoraium, 342, 848 ; C. pcnn-

sylvanicum, 342, 344
Cinyxis belliatia, 366 ; C. erosa, 364, 366 ;

C. humeana, 364
CisfecejjfuUus, 310
Ci^tiuii), arrangement of neural plates, 824;

C. Carolina, 361 f., 364 ; colour of iris,

329
Claosaurus, 4^9
Clarke, on habits and development of

AUigaiiyr, 467
Classification of Amphibia, historical

account, 7 f.
Clawed Toad {Xenopus), I46 f.
Claws or nails of Amphibia, 32
Cleithra=the pair of additional clavicles :

of Stegocephali, 79 ; of Parciasatint»,

304, 305
Clanmys, 356 f.; C. caspiea, 358; C.

inscidpia, 369 ; C, leprosa, 356 f., 363 ;

skull, 866
Clepsydrops, 308
ClidaMes tortor, 4^0
Cloaca, of Chelonia, 380 ; of Crocodiles,

445 ; of Lacertilia, 498
Cnemidophwus, 649 ; C. sexlineatus, 549
Cobra, 627, 627

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INDEX

655

Coccyx. 8, Os coccygeum, of Anura, 20, 21,

22
CoecUia, S9

Coeciliidae, S9 f. ; distribution of, 89
CoelopeltiSt 6i4 >' (^- monspesmUana s.

lacertina, 624
CoeluruBt 4^5 ; C, ffracUis, 4^3
Colombo, gigantic .tortoise of, 377
Coloration, warning colours of Amphibia,

38, 156 ; protectivfl, of Amphibia, 191,

238, 252 ; of deserticolous reptiles, 494
Colosth^us, 238, ^4S
Colour, changes of, in Anura, 35 ; in

Calotes, 518, 519, 520 ; in Geckos, 509 ;

in Lacertilia, 498 ; mechanism of chang-
ing, in Chameleons, 570, 671
Coluber^ 615 f.; C. aescvlapii=Jlavescen8=^

longiuimus, 616 f.; C. leopardinits, 616 ;

V. [Rhinechis) scaUiris, 617
Colubridae, 59S, 606 f.
Colubrinae, 607 f.
Columella cranii, 496, 650, 551
Columellar auditory chain, of Amphibia,

4 ; of Anura, 29 ; of Crocodiles, 446 ;

of Lizards, 496
Comoro Islands, Tortoises of, 373
CompsognaihtLSy 4^5, 416, 417; C longipes^

423, 425
Condyle, occipital, of Theromorpha, 802 ;

exaggerated importance of its character,

285
C(molophua suhcristatua, 633
Conus arteriosus, continuation of the heart

beyond the ventricles so far as it con-
tains valves, 6
Cope, on classification of Amphibia, 9 ; of

Anura, 140, 141 ; on Siren, 136 ; on

hand - skeleton of Eryops^ 286 ; on

Sphargis, 336 ; clarification of Lacertae,

513 ; classification of Snakes, 592
Cophopknjnej 167, 168
Cophyla, 236
Copper-head, 646, 646
Copulatory organs, of Lacertilia, 499 ;

absent in SvhemKion, 294 ; of Clielonia,

330 ; of Snakes, 585
Coqni, 214
Coronella, 619 ; C. austriaca s. laerijt, 619,

620 ; C. girondica^ 621
Cornufer, 24U 243 ; C. comigiUus, 244 !

C. joknstoni, 2^3 ; C. solonwnis, 244*

C. unicolor^ 244
Con/thonianiUf 189, 207 ; C. greeningi,

207
Costal plates of Chelonia, 324 f., 322, 32S
Craapedoc^luilua^ 647
Crested Newt, 125, 126
Cricotus, 285, 285 ; C. heferodihis, 287
OrinioL, 213 ; spawning, 223
Crocodilia, 431 f. ; skeleton, 434 f. ;

skull, 280, 434 f.; atlas and axis, 283,

481, 439 ; affinities, 432 ; teeth, 437 ;

skin, 442 ; dermal armour, 442 ; skin
glands, 443 ; tongue, 443 ; respiratory
organ8,444 ; "diaphragm," 444 ; digestive
organs, 444 ; cloaca, 445 ; heart, 445 ;
ear, 445 ; eye, 446 ; geographical dis-
tribution, 446, 446 ; voice, 447 ; habits,
447 ; propagation, 447 ; classification,
448

Crocodilidae, 454

Orocodilw, 450, 454 f. ; teeth, 437 ; skin
glands, 443 ; C. actUus, 446, 449 ; C.
americamis, 4^^ *' skull, 466 ; C.
biporcaius —porosus, 458; rate of growth,
459 ; C. cataphractii8, 465 ; C. iiUer-
medius, 4^6; C. johiistmii, 4^^ i O.
niloticu8=^ vulgaris, 46O f., 449, 461 ;
habits, 462 f. ; C. palustris, 449, 454 ;
skull, 466 ; C. porosity, 458; skull, 468 ;
C. vulgaris, 449, 460 f.

Crotalinae, 644

Crotalus, 648; rattle of, 644; O. ada-
mantetis, 649, 660 ; C. cmiJiv^rUus, 649,
650; C. durissus, 648, 649, 660; C.
horridus, 649 ; C. ierri/icus, 650

Cryptobranckus, 84, 96, 99 ; fossil, 84 ;

C. alleghaniensis, 97 ; C, japonkaa, 98,
99

CryptoclidiLs, shoulder-girdle, 474, 476,

478
Cryptodira, SIS, 338
Cryptopaophis, 89; C. mtUtiplicattia, 92
Cryptotia, 213
Cutis, of Amphibia, 33 f.
Cya^iodua, 311

Cyclanorbia, 4II ; nuchal plate, 324
Cyclodentia, 4II
Cyclodua s. Tiliqmi., 561; C. gigas, 561,

662
Cyclorhamphus, 212
Cynognathva, 301, 302, 303; C. berryi,

307 ; C. craieronotvs, 306 ; Cpiatycepa,

307
Cystignathidae, 139, 209 f. ; distribution,

161
Cystigiiathinae, 139, 211 f.
Oystiynatkus — Leptodactylua, 210, 218

Dab= Uroniaatix, 526, 626

Dabnia, 643

iJactylethra ; see Xenoptia, I46 f.

Darwin, on dmoiophua, 532 n. ; on

tortoises of Galapagos Islands, 377
DaaypelUs acahra, 622, 622 ; dentition,

593 n.
Davison, on breeding of Amphiuma, 101
Dawsonia, 289
Death Adder, 635
Denburgh, van, on Autodax, 107
Dendrobalea, 272 ; D, braccatua, 273 ; D.

tinctoriua, 272, 27S ; T). irivittatua, 273 :

D. typographua, 273 ; various uses of
its poison, 38

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6s6

INDEX

Dendrobatinae, 1S9, 237, 272 f. ; dUtribu-

tion, 239
Dendrqphisy 618; /). pundtdattis, 618 ,

618

Deudrophrynisciuae, 139, 224

Dendrophryniscus brevipoUicatiis, 224,

Dentition, of snakes, 582, 592, 593 ; see
also Teeth

Dermal armour, of Oricotus, 287 ; of Micro-
sauri, 289 ; of Prosauri, 290 ; of Thero-
morpha, 302 ; of Ohelonia, 321 f., 337 ;
of Diuosauna, 415 ; of Pseudosuchia,
483 ; of Parasuchia, 434 ; of Crocodiles,
442

Dermal ossification in Annra, 179, 190,
210

Darmatemydidae, 313, 314, ^41 : distribu-
tion of, 332

lJn'}natemy8 inawi, 34 L 342

JJennatochdj/s coriacea, 333 f., 834

Demiophis, 89, 93 ; D. thouiensis, 93

Deserticolous reptiles, 493 f.

Desmognathinae, 102

Desniogiiatkus, 96, 102 ; D. fuscus, 102,
103

DeiUerosaiirus, 308

Developmeut, of Anura, 56 f., 57 ; of
horny teeth, 68 ; of Apoda, 92 ; of
CrocodUiis, 465 ; of AlligcUor, 467

Diadectes, SOS

Diadevwdon, 309

Diarjlemi, 185, IS9 ; D. jordani, 207 ; D.
petasata, 207

Diaphragm, of Anura, 144 ; of crocodiles,
444

Diapophyses (the lateral or "transverse"
processes of the neural arches) of Anura,
138, 141

Dibamidae, J/^, 564

Dihamus nov(U-guliie(ie, 564

Dicamptaion, 96 ; D. ensatus, 109

Dicloiiius = Iladrosanru^, 429

Dicywjdon, 301, 302, 303, 310; skull,
280; D. leonicejjs, 310; D. orietUalis,
310; D, fMjriceps, 310

Digestive apparatus, of Chelonia, 330 ; of
crocoiiiles, 444 ; of Lacertilia, 498

Digits = Fingers and Toes. Number of
digits in Urodela, 15, 16 ; in Anura,
26 ; terminal plialanges, 26 ; number
of joints, 27 ; adliesive discs, 27 ; vari-
ability in numbers, 563 ; digits of
Eryops, 286 ; of Crocodiles. 441 ; of
Plesiosauri, 475 ; of Geckos, 505

Dinietrmlimy SOS

Dimorphoilon maci'onyx, 486

Dinosauria, .^/i ; affinities of, 415 ; anal-
ogies with Birds, 416

hiplocynodon^ 448 ; />. hastinrjsiae, 4^4

DipliMiocus long us, 419 f. ; skull, 419

Diphnrrtehronl 2S7, 288

Dipsadomorphinae, 623 f.

Dipaadomorphus, 623 ; D. cyaneus, 6t4 ;
D. trigonatus, 623

Dipsas biicephala, 624

Discoglossidae, 139, 152 f.

Diacogloasua, urino-genital organs, 49, ISS;-
D,pictu8, 153 f.

Dissorcphus mvUidnctus, 82

Distira q^anocincttif 637,; D, aemperi, 636

Distribution, geographical ; see Maps

Dolichosauri, 4^9

DdUkaaaurus longicoUis, 4S9

Dolichosoma longissimum, 81

Ddiophis intestinalis, 634

Dollo, on Sphargis, 886

Draaunct, 547 ; D. guiuneyisis, 649

Dracoy 516; D, diissumieri, 516; D.
vdans, 516, 616

Dumeril, 7, 139 ; and Bibron, on classi-
fication of Suake8, 592

Dwarf Chameleon, 579

Dyscophinae, 139, 235 f.

Dyscophtcs, 236 ; Z>. afUongUi, 236

EcheneU rtnwra^ naed for turtle fishing,
382

Echidna s. Bilis, 639, 689

Ediis, 640 ; E, arenicola, deserticolous,
493

EdalorhiHO, 212

Ear, of Chelouia, 330 ; of Crocodiles, 445 f. ;
of Snakes, 583

Ear-opening of deserticolous reptiles, 494

Eggs of Amphibia, 53 ; mode of deposi-
tion in Amphibia, 54-56 ; of leklhy-
ophis, 91 ; and spermatophore of Triton
viridescens, 128 ; nursing and taking
care of, 55 ; by Pipa, 151 ; by Alytfs^
159 ; by Rhacophorus reticuUitus, 248 ;
by Notoirenui, 188, 202; by Amphi-
gnaUudon, 188 ; by Uyla goelilii, 198,
198 ; by Leptodactyht^ mystacinus^ 219 ;
by Rhinodeniuif 228 ; by Rhacophorus,
248; by Desmognaihua fuscus, 103,
103; number of: m Rnfovulgaris^llh ',
in Bufo viridis, 181 ; in Hyla arborea,
193 ; in Rana escnlenta, 270

Eggs of Reptilia : SphenwioH, 299 ; Cliel-
onia, 331 ; Testudo graeca, 869 ; T.
ihera, 369 ; T. elegans, 871 ; T. poly-
phcmrcs, 372 ; Emys orbicularis, 355 ;
Cleinmys leprosa^ 358 ; Ckelone mydas,
382 ; Tfudassochelys caretta, 387 ; Podo-
cnemis expansa, 393 f., 398 ; Trionyx^
408 ; mode of laying by Emys, 355 ;
by Podociiewis, 393 ; used commercially,
394 f. ; enormous destruction of, 395,
399; CrocodUtts, 463, 464 f. ; AiH-
gator, 470; eggs and nest of GarialiA,
452 ; Lacertilia, 499 ; increasing in size
after deposition, 499 ; Geckos, 506, 508,
509, 511; Tarentolxi, 509; Laceria
viridis, 555 ; Chameleons, 672

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INDEX

657

Egg-sac, of Salamandrella, 110

Egg-tooth, of Lacertilia, 499

Eimer, on habits of Lacerta, 552 ; on Z.
coendeaf 558

Elachistodon teeatermanni, 625

Elachistodontinae, 626

E/apkis 8. Coluber^ 615 t.

Elapinae, 626

Maps corallinus, 6S5

Elasmosauridae, 478

Elasmosaurus, Jpl8

Elginia, 301, 304; E. mirabUis, skull,
280, S05

Elosia, 212

Eiseya, 389, 399

Emerald Llzanl, 555

Emery, on hand-skeleton of Eryops, 286

Empedias malaria, SOS

Emyda, 4II

Emydura, 389, 399

Emys, 350 f . ; E, blanditigi, 355; E,
eura2xiea = orbicularis, 351 f., 868

Enaliosauri, 4^6

Endothiodotif 307

Engyatoma, 227 , 231; £. carolinetue,
232

Engystomatidae, 139, 225 f.

Engystomatinae, 139, 225 t

Engystmnops, 166, 168

ErUhydrina valakadien s. beTigalennSy 636,
686

Enygrus, 601

Kospkargis, 336, 337

Epichordal type of vertebrae, 20, 145

Epidermis, of Amphibia, 31 f. ; sense-
organs in, 33

Equisetum, eaten by Urmnastix, 525

Ereynias, 551 ; deserticolous, 493

ErpetosiLchus, 4^3

Eryops, 285, £86; trunk-vertebrae, 286,
288, 804 ; E. megacephaliis, 286

Eryx, 604; JS. jacidus, 604; deserti-
colous, 493

Escuerzo = Cera^qpAry*, 216

Espada, on Rhinodenna, 228

Enblepharinae, 512

Euchirosaurus, 83, 287

Eunectes viurinus, 603, 608

Euprgpes vittata, 562

Euproctus= Triton, 130

Ewrystemum, 380

Eustachian tubes, of Anura, 29 ; 01 Pelo-
batidae, 161 ; of Aglossa, 143

Eusuchia, 4^4

Eye, of Apoda, 86 ; of Chelonia, 329 ; of
deserticolous reptiles, 494 ; of Chame-
leons, 569 ; of Snakes, 583

Eyed Lizard, 556, 666

Eyelid, of Geckos, 504, 512 ; transparent
in Chdodina, 329 ; lower, transparent
in Lacertidae, 551 ; in Scincidae, 560

VOL. VIII

Fasting, of Chrysemya, 847

Fat-bodies, of Amphibia, 49, 52 ; of Lacer-
taia,500

Fecundation, various modes of, in
Amphibia, 54 ; in AjMKia, 87

Fer-de-Lance, 647

Perreiro = ffyla /aber, 196 f.

Feylinia, 564

Fingers, number of, in Urodela, 15
number of joints in Anura, 26, 27
terminal modifications of, in Anura, 26
mechanism of adhesive discs in Hylidae^
187

Fire Salamander, 115

Firmistemal, type of shoulder-girdle, 24,
26

Finuisternia, of Cope, 140 ; of Bonlenger,
140

Fischer-Sigwart, on growth of Alytes, 159
f. ; on growth of £u/o, 175 ; on gesta-
tion of Chalcides, 563

Fletcher, on spawning of Australian frogs,
201, 223

Flower, S. S., on habits of Bhacophorus,
249; Phrynella pdlicaris, 233; Callula
pidchra, 234

Flying Dragon, 516

Flying Frog, Rhacopfiorus, 245 f., 246

Foot, tridactyle, in ffallopus, 423 ; bird-
like in Compsognathus, 423

Fore-limb, of Urodela, 15 ; of Anura, 26 ;
of Proreptilia, 286 ; of Microsauri, 289 ;
of Prosauri, 290, 298 ; of Theromorpha,
802 ; of Chelonia, 320 ; of Dinosauria,
414, 423, 425, 427 ; of Crocodilia, 440 ;
of Pl&siosauria, 475 ; of Ichthyosauria,
481 ; of Pterosauria, 485 ; of Pythono-
morpha, 489 ; of Lacertilia, 497

Gage, on Triton viridescens, 129

Galapagos Islands, tortoises of, 372, 877 f.

Galesaurus, 307

Gampsosteonyx, 271 ; O. batesi, 238, 240

Gasco, on spawning of newts, 124

Gastrechmia, 140, 141, 232

Gastrocentrous vertebrae, defined, 282

Gaupp, on frogs' respiration, 47 n.

Gavialidae, 451 f.

Gavialis, 435, 436, 451; G, gangetiais,

452; skull, 449, 462
Gavialositchus, 453
Gecko, 511; G. stentor, 511 ; G. verus=

guttatus=verticillatu8, 511
Geckolepis, deserticolous, 493
Geckones, 502 f. ; distribution, 500, 608 ;

adhesive apparatus, 505, 606 ; voice,

506 ; reproduction of tail, 506 ; eyelids,

504, 512
Geckouidae, 507 f.
Geckoninae, 507 f.
Gegenbaur, on classification, 9
Gegenophis, 87, 90

2 u

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658

INDEX

Geihia, 310

Genital organs, of Amphibia, 48 f., 49

GenyophryMf 2S6 ; G, thoinsoni, 236

Crenyophrynidae, 141

Genyophryninae, 139, 236

Geographical distribution, principles of,
69 f. ; regions and sub-regions, 74 f.
(for details see also Maps) ; of Apoda,
89 ; of Urodela, 96, 96 ; of Anura, 143,
161, 167, 185, 239 ; of Chelonia, 331 f.,
832, 333; of Crocodilia, 446; of
Lacertilia, 500 f., 516, 629, 648, 662,
666, 568; of Snakes, 686

Geomolgey 96

GeosaiiruSf 4^1

GeotritoUy 97

GeotrypeteSj 89

Gerrhonotus, 638 ; G. coeruleiiSt 538

Gerrhosauridae, 514, 559

Gerrhosau rus flavigularU, 559

Gharial, 4^3 ; see also Gavialis

Gigantic Tortoises, 372 f.

Gila Monster, 541

Gills, definition, 40 ; development of, 41,
43 ; retention of, 40 ; exteiiial and
internal, 43 f. ; operculum of, 44 ; of
Nototrenm, 203

Gill-clefts, 42 ; of Urodela, 42 ; of Anura,
42

Girtanner, on musical appreciation of tor-
toises, 368

Glass-Snake, 538

Glaucoma, 594

Glauconiidae, 592, 594

Glyphoglossus, 225, 226, 228, 233; G,
molossm, 233

Goeldi, on Uyla faJbcr, 197 ; on habits of
Podoenemis expansa, 397 f.

GomphogjuUhns, 308, 309

Gondtoanosaunis, 83

Gongylus, 562

Goniopholidae, 4^3

Goniopholis, 448, ^55 ; G, crassidens, 453 ;
G. siimis, 453

Gordonia, 301, 303, 310; skull, 280

Grass- Frog, 251 f., 266

Grass-Snake, 608 f.

Greek Tortoise, 365 f.

Green Lizard, 555

Green Toad, 180

Green, or Edible, Turtle, 381 f.

Groenberg, on Pipa, 149

Growth, rate of, in Testudo ihera, 370 ;
Chry.temt/s picta, 349 ; JSmys orbiadaris,
351, 355

Gular shields of Chelonia, 316

Guudlach, on Leptod<i4:tyliu% J219

Gunther, 140 ; on gigantic Tortoises, 374 ;
on classification of Snakes, 592

Gutzeit, on horny teeth of Tadpoles, 68

Gymrnxiartylus, tail, 506, 512, 612 ; de-
serticolous, 493

Gymuophiona, 84 f*

GymnophxA, 90

GyymiophthalMus, aberrant scaling, 495

Haast, on habits of Sphefvodon, 299

Hadroaaurus mirabUis, 429

Haeckel, on classification, 9

Hallopus victor, 423

Hamadryad, 632

Hand-skeleton, escalation of second finger

in Eryops, 286
Haptoglossa, 96
Hardun = i4^afna stdlio, 581
Hatteria — see Sphenodon, 298 f.
Hawksbill-Turtle, 384 f.
Hay, on Sphargis, 337
Hearing of Chelonia, 330
Heart, modification of, in lungless Am*

phibia, 47
Hedonic glands (^dom^, lust), 443
ffeleioponts, 213, 222 : H, albopttnaatns.

222; H. pictm,222
Helix virgata, eaten by ffyla coendea,

200
Hdwlerma horriduf/i, 540; H. sugpectum,

540, 641
Helodermatidae, 513, 540 f.
Hemidactyhis iurcicus, 508, 608
Hemiphractiuae, 139, 210 f.
Hemiphradus, 210
Ilemisus, 225, 226, 228, 232; shouhler-

girdle, 25; H. guttntum, 232; H.

sudanense, 232
Hensel, on Bnfo niarinns^ 179 ; on tadpoles

of Thoropa, 209 ; on nest-building of

LeptiKlactylus^ 219
Herodotus, on Crocodiles, 462
Herpele, 90

Htrpestes griseiis (Mongoos), 629
Hil>eruation, temperature of blood during,

68 ; of Tortoises, 347, 349, 354, 358,

360, 363, 365, 369, 376 ; of Crocodiles,

447
Hinckley, on tadpoles of Hyla versicolor^

195
Hind-limbs, of Urodela, 15 ; of Anura,

27 ; of Prosauria, 289 ; of Theromorpha,

302, 305 ; of Chelonia, 321 ; of Dino-

sauria, 414, 423, 425, 427, 429: of

Crocodilia, 440 ; of Plesiosauria, 476 : of

Ichthyosauria, 480 ; of Pterosauria, 4S6 ;

of Lacertilia, 497 ; of Ophidia, 593, 594.

596
Uipistes hydrimis, 625
Holbrook, on the Black Snake, 613: on

habits of Afligator, 470 f.
Holoblastic eggs ; the whole mass of the

egg undergoes the process of cleavage,

53
Homalopsinae, 625
Homalopsis bttccata^ 625
Homing of turtles, instances of, 386

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INDEX

6S9

Homoeosaunis pnlchdhiSt 292

Hinnopholis^ deserticolous, 493

Homothermoas, defined, 68

Hoplocephaltu curtus, 634

Hoplurus, 538 ; geographical distribution,
501

Horned Toad = C<wa<opAry*, 215 f., 216,
217

Horne<l Viper, 64O, 641

Horny nail, on tail of Chelonia, 328

Horny scales, of Chelonia, 328

Homy shields, of Chelonia, 314, 816, 822,
328, 326 f., 827 ; their growth, 326

Homy teeth, of Auura, 68

Hose, on reproduction of tortoise-shell, 386

Howes, on development of Sphenodan^ 298

Humerus of Sphetwdmi^ 294

Hutton, on Starred Tortoise, 370 f.

Huxley, on classification, 9

Hydraspis, 389 ; skull, 399

Hydromedusa, 589, 404 ; H, Uctifera, 4O4

Hydrophinae, 635

Uydrophis obsctira^ 636 , 686

HydrosaumSy 543

Hyla, 189 f. ; H. aHwrea, 190 f., 190
var. meridiotudiA, 191 ; var. 8avignyi=
Japonica, 191 ; ff, aurea, 201 f. ; spawn
ing, 201 ; H. carolineiisis a. lateralis,
194 ; H. coeridea, 198 f., 199 ; spawn
ing, 223 ; H. dusynotus and H. nigro-
^rutculataf dermal ossifications of, 190
H. eioitu/i, 201 ; spawning, 228 ; H.
Jaher^ peculiar nursing habits, 196 f.
H,fa)Kfrali8, 194; H- goeldii, 198, 198
female with eggs, 198; H, itiaxima,
196 ; H. nebidosa s. luteola^ 197 ; nest
building, 198 ; B. poly taenia, 198; H,
squirelhi, 194; H, vasta, 195; II.
versicolor, 194 f.

Hylaeformes, 139

Ilylaenbatrachtis croyiy 83

Hylaevsaurus, 4^5

Hylumbates, 238, 240

Hyldla, 186, 189, 203

Hylitlae, 139, 185 f.; distribution, 186,
186 ; mechanism of climbing, 187 ;
map of distribution, 186 ; distribution,
186

Hylinae, 139, 189 f.

Hylixfdus, 2;J8, 242

Ilylodes, 2li ; //. viartinicensis, 214 f-,
214 ; H. abbreviatif3 = Thoropa miliaris,
209

llylojumiis, 288, 289

Uyloplesion long icostat urn, 289

Hylopsis, 212 : U. platycepfuilus, 234

Hyhrrhim^ 212

Hymenochirus, 143, 144, 149

liynobiua, 96, 109

Hyoid apparatus, of Urodela, 16 ; of
Anura, 31 ; of Chelonia, 318 ; of
Chelys, 400 ; of Lacertilia, 496

Ilyperodapedon gordoni, 29S
Hyperoliu, 213 ; spawning, 223
Hyperphalangeal limbs, of Eusuchia, 441 ;

of Ichthyosauri, 480
HypogeophU, 87, 89, 92; H. aUemans,

92; H, rostratus, 92
ffypopachus, 226, 227, 235
HypsHophodofi foxi, 4^
Hypsirhina plundyea^ 625

Iberian Water-tortoise, 357 f.
Ichthyodea, distribution of, 96
Ichthycpkts, skull, 86, 88, 89 f., 91 ; /.

glutinosa, 90, 91 ; /. inonochrous, 90
Ichthyopsida, 5, 277
Ichthyopterygia, 476
Ichthyosauri, 433 f.
Ichthyosauria, 478 f. ; skull, 281, 479 ;

vertebrae, 480 ; limbs, 481 ; shoulder-
girdle, 481
Ichtfiyosaunts, 4^3; I, coinmunis, 4^3;

L campylodon^ 4^3; I. quadriscissus,

488 ; /. tenuirostria, 4^^ » I' trigonodon,

483
Idiochtlys, 380

Iguana, 306, 528, 531 ; I. tuberculata, 531
Iguanidae, 513, 528 f. , distribution, 501,

629
Iguanodon, 416, 417, ^^7; Lberniasar-

tensis, 428, 428 ; I. mantdli, 427
Iheriug, on breeding habits of PhyUo-

mediisa, 205 f., 206
Ikeda, on nursing habits of Rfiacqphorus,

248
Ilysia, 595
llysiidae, 592, 594
Inframarginal shields, 326, 316
lutergular shields of Chelonia, 325, 816
Iris, colour of, in Chelonia, 329
Ixalus, 238, 24I

Jaw, lower, of Salamandra, 17 ; of Uro-
dela, 18 ; of Anura, 30

Keller, quoted, 571 u.
Keraterpeton, 81, 288 ; K. crassum, 81
Kidneys of Amphibia, 48 f., 49
Kliuckowstroem, on Pi^ta, 149
KoIImann, on Neoteny, 64
Krait, 633

Labial glands of Helodenna, 498

LahyrinthHlon, 83

Labyrinthodonta, 8S

iMcerta, 553 ; L. axjdis, 554 *' ^' muralis,
557 ; L. ocellata, 556 t., 666 ; L. jmter,
556 ; L. srhreibcri, 555 ; L. tangitana.,
556 ; L. viridis, 555 ; skull, 660 ; L.
vivipara, 553

Lacertae, 513 f.

Lacertidae, 514^ 549 f. ; skull, 660 ; distri-
bution, 662

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66o

INDEX

Lacertilia, 491 f. ; skeleton, 494 f. ; skin,
497 ; change of colour, 498

Lachesis fframineiu, 647 ; L. lanceolatuSt
64$, Wl

Land-tortoises, S64 f.

Lauthanotidae, 514', 541 1

Lanthanotus b&meetisis, 541

LaosauruSy 4^7

Lariosaunu, 473, 474 ; L, balsami, 477

Larvae, oi Icklhyophia^ 91 ; of Hypogeophis,
92 ; of Amblyatotna, 112 ; of Triton
waiai, 181

Latreille, on classification, 7, 8

Laurenti, on classification, 7

Leathery Turtle = ^^Aa/v/w, 333 f., 3S4

Lechriodonta, distrihution of, 95 ; defined,
102

Leptdobairachus, glS^ 218

Lepidophymay 547

Lepospondylous, defined, 79

Leptobrachium^ 161; L. caruiensey 166

Leptodactylusy 21iy 218 f.; L. albilaJbria^
219 ; L. myatacinusy 219 ; L. ocellatus,
219 ; L. typhoniwy 219 f.

LeptogncUhuSy 624

LeptophUy 618 ; L. iiocerus, 6l8y 619

Leslie, on XeTwpiiSy 146

Leuckart, on classification, 8

Leydig's dnct, defined, 48, 49

Lialis burioniy 567

Limbs, of Amphibia, 26, 27 ; Stegocephali,
79, 83 ; ErynpSy 286 ; Microsauri,
289 ; Prosauri, 291 ; SphetvodoHy 298 ;
Theromorpha, 302 ; Pareiasauri, 305 ;
Chelonia, 320, 319, 320; Sphargis,
335 ; Chelonidae, 379 ; Dinosauria,
414 f., 418, 420 ; Compsognathu^, 423 ;
Stegosaiiri, 426 f. ; IguanodoUy 428,
428 ; Eusnchia, 440 ; Plesiosauria,
475, 475 ; Lariosaurujf, 477 ; Ichthyo-
s.iiiria, 480 ; Pterosauria, 485, 486 ;
Lacertilia, 495 ; Geckones, 505 ; Cham-
eleons, 568 ; reduction of, in Lacertilia,
497 ; in Ophidia, 593, 594, 596

Limjimiynastes, 2l3y 222 ; spa waning, 223

LiuutoniediisOy S12

Linnaeus, on classification, 7

Liodo7i haumurUnsUy 4^)0

LioUpis beliiy 527

Liopelmay 153 ; L. hochatetteriy 160

Lmphrynty 225, 227

LiosaKntSy 529

Lissamphibia, 84 f.

Lizard, Common English. 55S ; Emerald,
555 ; Eyed, 556', 656 ; Green, 555 ;
Sand, 554 ; Wall, 557

Lizards, 4(fl f.

Locality, sense of, in Tortoises, 368,
387

Loggerhead Turtle, SS7 ; individual
varieties of shields, 327, 388

Longevity, of Test ado daudiniy 376 ; T,

graecii, 369 ; T. t&em, 369 ; T wmeiret,

877
Loxooemus bioolor, 598
LoxommOy 88

LucUia bt^fonivora, fiy infesting Biifo, 176
Lungs, definition, 40 ; suppression of, 46 ;

of Aglossa, 144 ; of Lacertilia, 499
Luth, or Leathery Turtle, 333 f., 384
LycosauruSy S07
Lygoaomoy distribution, 501
Lymph-spaces, in the cutis of Anuia, 34
Lyriocephalua acutatuSy 517, 618
Lytdomoy 836, 880

MabuiOy 562 ; distribution, 501 ; eyelids,
494 ; M. viltoJttay 562

Macrodemmysy 326 ; J/, tanmincki, $40 f»
840

Macroprotodon cucullcUuSy 624

Macrorhyuchidae, 4^1

Madagascar, Lacertilia of, 502

Malacoclemmys terrapiiiy 359 f. ; com-
mercial breeding-farms, 360

Malpighian, body, 49 ; stratum, 32

Mammalian afiSnities of Theromorpha, 303,
309

Manculusy 96, lOSy 106; JI. qvadridiffi-
tatiuy 106

Mandible, composition of, in Crocodiles,
437; very Mammalian in GomphofftM-
thusy 309

Mantelloy 274

Mantophrynty 225, 227

Maps showing geographical distribution, of
Coeciliidae, 89 ; Urodela, 95 ; Aglossa,
148 ; Cystignathidae, Discoglossidae,
Pelobatidae, 161 ; Bofonidae, 167 ;
Hylidae, 186; Ranidae, 289; Chely-
didae, 882, 888 ; Chelydridae, 832 ;
Cinostemidae, 882 ; Dermatemydldae.
882 ; Pelomedusidae, 882 ; Platyster-
nidae, 882 ; Trionychidae, 388 ; Croco-
dilia, 446 ; Geckonidae, 608 ; Agamidae,
616 ; Anguidae, 529 ; Iguanidae, 529 ;
Zonuridae, 529 ; Varanidae, 643 :
Lacertidae, 552 ; Amphisbaenidae, 565 ;
Chaniaeleontes, 568 ; Snakes, danger-
ously poisonous, 686 ; Elapinae, 626 :
Crotalinae, 644 ; Viperinae, 688

Marbled Newt, 126

Marginal plates of Chelonia, 325, 332, 33S

Marginal shields, 326

Marsh, on Axolotl, 115

Marsh Crocodile, ^55

Marshall, on distribution of Uropeltidae,
595

Mascarene Islands, tortoises of, 373 f.

Mason, on habits of CcUotesy 519 ; on
Python legends, 599 ; on Varanusy 544

MastodansauruSy 83

'^a.t&nmtvi= Ch^iys ^rnbriatay 400, 401

Mauritius, gigantic tortoises, 873 f., 376

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INDEX

66 1

Mecodontaf distribution of, 96; defltied,

102
Megalixalusy 288, 240
Megalophrya, 161 ; tadpole, 60
MegohloaaurvSi 416 ; M, buMindi, 4^1
Megalotritony S3
Afelanerpetan, 81, 289
Melanobatmchvs, 226, 228
Meiosaurus, 287
Menobranchus lateralis, 1S2
Menojwma, S7

Mento-Meckelian cartilages, 30
Meroblastic eggs ; part of the egg only

undergoes the process of cleavage, 53
Merrem, on classification, 8
Mesosauridae, 476

Mesosaurus, ^7tf ; M. tetiuidens. Iff 6
Metamorphosis of Tadpoles, 56 f.
Metasternum of Anura, 26, 26 ; taxonomic

value, 141 ; definition, 26
Metatarsalia of Theropoda, 420
MetojnaSy 83

Jfetoi^oceros cornutus, 632
Metriorhynchidae, 4^1
Metriorhynchus, atlas and axis, 283, 431,

451
Metzdorff, on Axolotl, 113
Micrixalus, 241
Microgomphodim, 308, 309
Microhyla, 228
MicTosauri, 288
Midwife-toad, 158
Mimosa (plant), 629
MuAania, 390

Mirophyes^ 213 ; spawning, 218
Mixosaurus, limbs, 480, 481, 4^^
Molfie — see Triton, JJ2
Moloch harridusy 527, 627
Mongoos and Cobra, 629
Monitor, 543

Morosaurus grandisy 4^9 ; pelvis, 419
Mosasauri, 48U f.

Mosamnrns^ 489 ; M. camj^rL, 4UO
Moult of Geckos, 510 ; of Chameleons,

571 ; of Snakes, 583
Mud-diver, 165
Mud-turtle, 342
Mugger, 454

Miiller, J., ou classification, 8
Miillerian duct, 49, 51
Musical a])preciation of Tortoises, 368
Mifof)airachus, 166, 167, 168; M. goiddi,

284. 227, 236
Mystriosaurus, 432, ^5i

Nails or claws of Amphibia, 32

iV(7;W, 626; JV". bungarns s. elaj)s, 633 ;

y. Juije, 628, 6S2 '; S. tripudiaru% 627,

627
JVannobatrackus, 238, 24O
Naosaurtis clat-iger, 308
Natterjack, 181

NavUinus elegans, 506

Neck, mode of withdrawing in Chelonia,

328 f.
Nectes, 166, 168; N. aubasper, 169
Xectophryne, 166, 168; N. afro, 169; N.

tuberculosa, 169; N. guentheri, 169;

N. hosei, 169 ; N. misera, 169
Necturus, pelvis, 15, 96, 132 ; N. 7n€icu-

latus, 132
Neoteny, 63 f. ; defined, 64
Nephrostomes, 48, 49
Nephrurus asper, tail, 606
Nerves, spinal, of Amphibia, 88 ; cranial,

39
Nest, of Orocodihis, 463 ; of Gavialis, 452
Neural plates, of Chelonia, 323 f., 822, 328 ;

suppression of plates, 324 ; in Pleuro-

dira, 389 ; of Dermatetnys, 842
Neusticosaurus, ^77
Newt, Common, 127 ; Crested, 125, 126 ;

Marbled, 126; Spotted, 127
Newton, K T., on fossil Keptiles, 803 n.
Nile Crocodile, 461
Nodosaunis, 430
Nose-horned Viper, 64O
Notaden, 166, 169 ; iV. bennetti, 167
Notechis scutatus, 634
Nothosauri, 476 f.
Nothosauridae, ^77
Noihosaurus, 474 ; N. mirabilis, 477
Notoceutrous vertebrae, defined, 19
Notochord = Chorda dorsalLs, q.v.
NotoiremiL, 189; S. eoniuium, 203; N. .

viarsupiatum, 202; iV. oviferwn, 202;

peculiar gills of embryos, 203 : IC.

pygmaewn, 202 ; y. testudineuni, 20 J
Nuchal plate of Clielonia, 323 f. ; of

Pleurodira, 389
Nuchal shield of Chelonia, 326, 827 ; of

Pleurodira, 389, 399
Nuptial excrescences of Anura, 33
Nursing, habits, ot'Arthroleptis seychelle7isis,

243 ; of Chiromantis rufesrens, 244 ; of

liha/^opfumis, 248 ; of RJiinoticrina, 228

f. ; of Pipa, 151 ; oi Hyla falter, 196 f. ;

of H. nebidosa, 198 ; of ii. goeldii, 198 ;

of Nototrema, 203 ; of Phyllomedusa,

204 f. ; of Leptodactylus, 219 f . ; of

eggs by Desmognatkus, 103, 103 ; by

Autoddx, 108
XyctUHdrachiis, 24O
Nyctinmntis nigiceps, 189, 206
Ni/diaxilus, 238

Occipital condyle, of Reptilia, 278 ; ex-
aggerated importance of, 285 ; of Thero-
niorpha, 302 ; of Pareiasauri, 305 ;
of Cynognathus, 307 ; of Orateronohcs,
307 ; of Dicynodon, 310 ; of Eusuchia,
437 ; of Amphisbaenidae, 496

Odontaglossa, 140

Oiigodan, dentition, 593 n.

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662

INDEX

0mo9aurus= StegosauruSf 4^6
Omosternum of Anura, 35 ; tazonomic

valae, 141
Onychodactylw, 96 ; 0. Japonictis, 109
' Operculam of gills, 44
(Iphidtrpetont 81
Ophidia, 491, 581 f.
OphiophoffuSf G3'2
OphiopSf 551
Ophioxylon (plant), 629
OphiaauruSy 638 ; 0. apus^ 538 ; 0.

gracilis^ 538
Ophthal}nomuruSf limbs, 481, 481, 484
Opisthocoelous, definition, 12
Opisthoglossa, 140
Opisthoglypha, 59£, 606 f., 623 f.
Oppel, on cla.ssiticatiou, 7
OreobatrachiiSy 24t
Oreophripiella, 227
Omithocheirus, 486
Omithomimus, 417 ; 0. grandis^ 4^9
Ornithopoda, 425, 426
Ornitfwpsisy ^79
Ornithoscelida, 416
Ornithoatichiuf, 4-iS
Orthocosta, 288, 289
Orthopoda, 4'^ 4
Ossifications, dermal, in Annra, 31, 34 179,

190, 210, 211
0.steoderm.s = ossifications in the skin, of

Spkar(jis, 837 ; of Caiman, 337 ; of

Lizanis, 504, 513, 514
Osteolaeniuji, 450 ; 0. tefmspis, 460
Oiiarau, 54J

Oiul4;n(HloHy 301 ; 0. rugosiia, 310
Ovary, 49
Oviduct, 49, 51

Owen, on fossil Reptiles, 303 u.
Oxydactyla, 140
Oxyyhssus, 239, 241

Pachytriton, 96, 115 ; P. hreripes, 133
Painted Terrapin, 347, 348 ; rate of

growth, 349
Pahtrobatrachus, vertebral column, 22,

145
PcdaeMJuitleria lonyimud^ita, 201 ; skull,

280, 304
P(i!uduu>/(i, resembles Knyystoinnps, 166,

2 12, 221 i; P./u,scomaculaia, 220] P.blli-

f/onigera, 221
Pa/iuihirt, as food of Trianyx, 407 I

ParasteiTiuni = the sum total of the AMb-

minal ribs, q.i\ ; of iSphenodan, 298 ; of

Crocodilia, 440 ; of Ichthyosauria, 480
Parasuchia, 4'^ J
Pareiasauri, 301, 302. 304
Pareiasaun's baiiit\ 30^
Parrots, feathers dyed with poison of Den-

drobiites, 272
Pt'/(tb(Uf,s^ variation of vertebrae, 19 ;

sacral vertebra, 22, 161 f. : /'. fuscus.

162; P.cullri^es, 163,1^ ; /*. syriaeus,

164
Pelobatidae, 139, 160 f. ; distribution. 161
Pdodytea, 161, 165; P, puncUUns, 165;

P. caucasieus, 166
Peioniedusa galeata, 391
Pelomedusidae, 313, 314; distribution,

832, 390 f.
Pelosaurus, 81
Pelvic, plexas of Annra, 89
Pelvis, of Urodela, 15 ; of Anura, 22, 27 ;

of Kryops, 286 ; of Microsauri, 289 ; of

Sphewidon, 298 ; of Tlieromorpha, 302 ;

of Pareiasauri, 305 ; of Cynognathug^

307 ; of Dicynodon, 810 ; of Chelonia,

319, 819, 820 ; of Pleunxlira, 388, 389 ;

ofDinosauria, 414; of Eusuchia, 441 ;

of Plesiosauria, 476 ; of Icbthj'osauria,

480 ; of Pterosauria, 485 ; of Pytbono-

morpha, 489 ; of Lacertilia, 496'
Perennlbranchiata, 8, 9 ; not a natural

group, 65
Petrels living with Spfienodon, 299
Petrobales, 288, 289
Phalanges, number of, in Urodela, 15 ; in

Anura, 26, 27, 238 ; in Stegocephali,

79 ; in Palaeohatteria, 291 ; in Chelonia,

820, 321 ; in CMone, 879 ; in ^'kelido-

murui, 425 ; in CampUtsaunts^ 427 ;

in Laosaurus, 427 ; in Iguaiwdon, 428 ;

in Eusuchia, 441 ; in Plesiosauria 475 ;

in Lariosaunis, 477 ; in Ichthyosauria,

481, 481 ; in Pterosauria, 485, 485 ;

shape in Anura, 138 ; peculiar in Pijfa,

151
Phanerc^lossa, 152
Phaueroglosses, 139
Phanei'otis, 213

Phisalix, on poison of Amphibia, 37
Pholidosaurits, 451
Phractamphibia, 78 f.
Phrynaglosses. 139
Phrynella, 227 ; Ph. jwllicaris, 233
Phn/niscNS, 226, 227, JJO ; Ph. ingncans,

220
Phrynvbatrachvs, 241
Phrynocara^ 235, i36
Phrynocephodm, 521 ; deserticolous, 493 ;

coloration, 494 ; Ph. helwscopus, 523 ;

Ph, intei'svapularis, 622; Ph. mygtaccu^

532
Ph ryi\ (>denna^ 24 1
Phrynoviantis, 226, 228
Phrynopsis, 241
Phrynosoma, 305, 533; Ph. coronaium^

524, 636 ; Ph. cornuttnn, 533, 634
PhyUi^iiPs, 242; Ph. bkdor, 242; PK

trinitatis, 242
Phyllodactylm, 507 ; Ph. europaeus, 607
Phyflod ramus, 238, 242
Phylloniedusa, 189, 203 f. ; PA. bicvlor^
S03 ; Ph. dacnicolor, 203; PK hyiwchon-

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INDEX

663

drialiSf breeding habits and develop-
ment, 204 ; Ph. iheringif £05 ; breeding
habits, 206

Phytogeny, of Amphibia, 66 ; of Annra,
142 f. ; of Reptilia, 282 ; of Lacertilia,
515 ; of Ophidia, 592

Physignathtis l&meuri, 523^ 624

Pigment in the skin, 34

Pipa. 148, 144, I49 f., 160

Pit- Vipers, 6U

Placodoutia, Sll

Placodm, 301 ; P, gigas, Sll

Plastron, of Chelonia, 316, 321, 321;
provided with hinges, 323 ; sexual
characters of, 331 ; movable in Emys,
350 ; of Chelonidae, 321, 821, 822,
880 ; of Pelomedusidae, 390 ; of Chely-
didae, 399 ; of Trionychoidea, 406

PlatecarpuSf 490

Platemysy suppression of neural plates, 824 ;
skull, 399

Plathander= Xennpus^ I46 f.

Platurus fcLsciatus s. colubrinust 6S7

Platydactyla, 140

Platydacfi/hts facetanus, 509, 608

Phityhylay ^36

Plaiypelis, 235, ^236

Platystemidae, SI 4, 326, 345

PI aty sternum mefjacephoduviy 345

PlectromantUy tli

Plesiochdya, 380, 389

Plesiosauri, -^77' f.

Plesiosauria, -^75 f. ; vertebrae, 474

Plesiosauridae, 478

Plesiosawnts, 476, 478 ; P. cwiybeari, 478 ;
P. dolicfiodirus, 478

Plethoclon, 94, 96, lOj, 106; P, eryihro-
notifs, 107 ; P. glntinosus^ lOG

Plethodontinae, 102, lOS

Plethodontohyla, 235, ^36

Pleurodira, 313, 388 f.

PUiiromnrus, ii04

Pleurosternum, 3J)0

Hioplatecarpus^ 489

Pliosauridae, ^77

Pliosaurus grandis, 477

Plover, Egyptian, and Crocodile, 462

Podocnemis, 390, 391 ; P. expanm, 391 f. ;
Bates, on habits of, 392 f.

Poikilothermous, defined, 67

Poison, of Amphibia, 37, 38 ; peculiar use
of, 272

Poison -aj)paratus, of HelodennOy 540 ; of
Snakes, 586 f.

PolacanUius, 4^5

Pulychrus viarmoratus, 529

PdyodontophiSy 605 n.

Portschinsky, on parasitic flies, 177 n.

Postpubis, of Dinosaurs, 414, 424, 426

Pouchet, quoted, 571

Predentary bone, of Dinosaur, 424

Prehallux, of Anura, 28

Prepubia, of Diuosauis, 414, 424, 426

ProganochelySf 389

Proganosauria, 476

Proreptilia, S85

Prosauri, S90

Prosauria, 288

Prostherapis, 2^

Protection of Amphibia by poison, 38

Proteidae, 94, 96, 132 f.

Proteroglossa, 140

Proteroglypha, 625

Proteus, 96 ; P. anguinus, 133, 184

Protorosauri, 290, 304

Protorosaunis lincki, S91

Protoaphargis, 336

Protostega, 336

ProtrUon, 80, 81

Psamvwdromua hiapaniats, 558; P. algirus^

558
Psamjnoaaurns, 643
Pseph()den/ia, 337
Psephophonis, 336, 337
Psetuiechis porphyriaceus, 634
Pseudis, 212, 213; P. paradoxa, 213 f.
Pseudobranchvs, 96 ; P. atriatus, 137
Pseudocentrous, defined, 79
PseMdophrynt, 166, 167, 168 ; spawning,

223 ; P. auatralia, 168 ; P. bibrotii, 168
Paeudopua, aberrant scaling, 495 ; P.

pallaai, 538
Pseudosphargia, 336
Pseudosuchia, 432
Ptenopua, 507 ; desertlcolous, 494
Pteraiwdon longtcepa, 487
Pt«ranodonte8, 487

Ptemohyla, 179, 189; P.fodiena, 207
Pterodactyli, 486
Pterodactylua longiroatria, 487 ; P. aped-

abilia, 487
Pterosauri, 486
Pterosauria, 484 f«
Ptyaa=Zamenia, 611
Ptychozoon, tail, 506 ; P. hoinalocephaluin,

512, 612 ; adhesive apparatus, 606
Pubis, of Dinosaurs, 414, 424, 426
Puff Adder, 639, 639
Pygopodidae, 514, 567
Pygopics lepidopiis, 567
Pythmi, 598 ; P. molurtia, 600, 600 ; P.

regitis=P. sebae, 601; P. reticutaius,

598; P. spilotea, 598, 699
Pythoninae, 598 f.
Pythononiorpha, 487 f.
Pyxis arachnoulea, 365

Radde's "law of the steppe," 493
Pajia, 241, 249 f. ; sacral vertebrae, 22 ;
shoulder-girdle, 26; urino-geuitnl organs,
49 ; Tadpoles' homy teetli, 68 ; vocal
sacs, 250 ; nuptial excrescences, 250 ;
large glandular complexes, 250; distribu-
tion, 251 ; species with finger • discs,

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664

INDEX

250 ; R, qfghanot 250 ; R, agiiis, S67 ;
K albolabris, 250 ; R. altkola, 250 ; R,
arvaliSf 257 ; R, eatabianay S61; R,
chaJconota, 250 ; R. chloronota, 250 ; R
cla^nataj 26iy 263 ; R corrugaia, 250 ;
R curtipeSf 250 ; R. cya^wphlyctU^ 250 ;
R, eUgans, 250 ; R ^rythraea, £50 ; R.
esciUetUa, 263 ; mechanuim of tongue,
268 ; vocal sacs, 269 ; var. chinensU,
267 ; var. Ussoiiae, 265 ; var. ridtbundcL,
264; var. typica^ 265 ; R. foiitinoUl%
262 ; R glandulosa, 250 ; R gracilis^
261 ; R graeca, 259 ; R. guppyU 261 ;
72. halecina, 26S ; R hexadadyla, 250 ;
R, iberica, 258; R, latastei, 259; R.
liebigi, 250 ; R. mascariejisis, 250 ; R.
nwntezumae^ 250 ; R. mttgiens^ 261 ; R.
opiatkodoTif 260 ; R. oxyrhynchus^ 250 ;
R. rugosOj 250 ; R, silvaii^Mj 259 ; R.
temporalis, 250 ; R. teniporarmy 251 f.,
265 ; R, tigriim, 261

Ranidae, 1S9, 2S7 f.

Ranidens, 96 ; R. sihiricus, 109

Raniformes 189, 140

Raninae, 139, 237, 238 f. ; distribution,
239

Rappia, 241

Rat Snake, 611, 612

Rattle of Rattle-Snake, 64«

Rattle-Snake, 648 f., 648, 660

Reduction of limbs, in Urodela, 16 ; ia
Lacertilia, 497

Regeneration, in Amphibia, 66 f. ; of tail in
Spkenodon, 298 ; of shell in Chelonia,
329 ; of horny shields in Chelonia, 329,
386 : of Uil in Lacertilia, 495 ; of tail
in Geckos, 506

Regions, geographical, 74 f.

Reproduction of Tortoise-shell, 386

Reptilia, defined, 277 ; principal characters
of, 278 ; classification of, 279 ; diagram
of affinities of principal groups, 282 ;
affinities to Mammalia, 303, 309

Respiration, mode of, in Chelonia, 331 ;
assisted by anal sacs, 330

Respiratory organs, of Amphibia, 40

Rhachiodontinae, 6;22

RMcopiwrus, 151, 186, 238. 241, 244,
246 ; Rh. letvcomyatax, 247 ; tadpoles,
249 ; Rh, maculatita^ nesting, 248 ; Rh.
vmdagasmrietisis, 245; Rh. nuunmiis,
245 ; Rh. pardalis, 246, 246 ; Rh.
reinwardti, 247 ; Rh. retkuliitiis, 248 ;
Rh. schlegdi, nesting, 248

Rhamplwleon spectrum, 580

Rhaviphorhyiich^LS longicavdatiis, 486 ;
RJi. phylhtrxLs, 486 ; Rh. inuensteri,
487

Rhamphosuchns crcusidais, 4*^3

Rhinatrema, 89

Rhinenuja, 389, 399

RhiiLOchelys, 390

Rhinoderma, 226, 237, 228; Rk, darwini,
228 i.

Rhinophis, 91 ; Rh. sanguineus, 596

Rhxnophrynus, 166, 168-; Rh. dorsaiis^
185, 227

Rhinophrys, 167

Rhombophryfu, 225, 227

Rhynchocephali, 292

Rhynchosaurus, 292

Rhytidosiefis, 83

Ribs, of Urodela, 14 ; of Anura, 21 ; of
Microsanri, 288 ; of Sphenodon, 297 ;
of Theromorpha, 302 ; of Qynogtuithu.%
807 ; of Microgomphodon, 309 ; of
Chelonia, 815, 320, 824 ; of Dinosauria,
413 ; of Crocodilia, 438 ; of Parasuchia,
434 ; of Eusuchia, 439 ; of Lacertilia,
495 ; of Geckones, 504 ; much elongated
in certain Iguanidae, 529 ; meeting
ventrally in Chameleons, 568

Ridewood, on hyoid apparatus of Anura.
31

Ringhals, 632, 683

Ritter and Miller, on AiUodax, 107

Robinson, on peculiar use of Varanus, 546

Rodriguez, gigantic tortoises, 374

Rostral bone of Ceratopsia, 480

Round Island snake, 603

Sacral vertebrae of Anura, 21 , 22
ScdaviandrcL, 115 f. ; trunk -vertebra, 14 :

skull, 17 ; lower jaw, 17 ; distribution,

96, 115 f. ; S. aim, 119 f. ; S. eaucasica,

121 ; S. maeviosa, 115 f.
Salamandrella, 96, 109; & keyserlingi,

109 ; S. achrenki, egg-sac, 110
Salamandridae, 94, 103
SalatiMndrina, 96, 115; S. perspiciUafa,

122 ; skull, 17
Salamandrinae, 102, 115
Sarasin, P. and F., 10 ; on Coeciliae, 88 :

on IcJithyophis, 90
Sauria, 491 f.

Saurichnites salamandroides, 83
Sauropoda, 4I8
Sauropsida, 5, 277
Sauroptei^gia, 476
Saurosternwn, 291
Scales of Apoda, 87
Scaling, aberrant, 495
Scaphwphryne, 225, 226, 227
Scaphiopiis, 161, 164 ; S. sdltarius, 165
Scapteira, deserticolous, 494
Scapula, attached to thoracic vertebrae,

487
Scdidosaurns, 416 : S. harrisotii, 4^5
Scheuchzer, on Homo diiuvii testis, 84
Schlegel, on Oryptohranchus, 100
Schuberg, on mechanism of finger-discs of

Hylidae, 187
Schwalbe, on Salamandra atra, 120
Scincidae, 514, 559 f.

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INDEX

66 s

Seinctu officinalis^ 661

Sciurus bicolor^ sqairrel, 248

Scolecomorphusj 90

Sea Snakes, 63o

Seeley, on fossil Reptiles, 303 n.

JSeeleya, SI, 289

Segmental duct, 49

Sense-organs, ofChelonia, 329 f. ; of Croco-
diles, 445 f.

Sepedon haetHOcfuUeSy 6S£, 683

Seps, 562

Seychelles, gigantic tortoises of, 373

Shell of Chelonia, 321 f., 319, 320, 821,
322, 323, 327 ; partial regeneration of,
329 ; correlated changes, 828

Shields, homy, of Chelonia, 322, 328,
325 f., 827 ; evolution of, 326 f. ;
individual variation in, 326, 327 ;
periodical peeling of, 328

Shoulder-girdle, of Urodela, 14 ; of Annra,
24, 26, 138 f. ; arciferous, 24, 26,
138 ; lirmistemal, 24, 26, l38 ; of
Aglossa, 144 ; of Microsauri, 289 ; of
Protorosauri, 290 ; of Theroniorpha, 802;
of Pareiasauri, 305 ; of Dicynodonf 310 ;
of Chelonia, 818, 819, 320 ; of Dino-
sauria, 414 ; of Eusiichia, 440 ; of
Plesiosauria, 474 ; of CryptocJidns, 476 ;
of Ichthyosauria, 480, 481 ; of Ptero-
sauria, 485 ; of Pteranodon, 487 ; of
Pvthonomorpha, 488 ; of Lacertilia,
496

Shufeldt, on Axolotl, 114 ; on Heloderma,
540 n.

Simosaurus, 477

Sijihonops, 86, 87, 90

Siredon (Axolotl), 112

Sireih 96 ; »S. lacertimt, 136, 136

Sirenidae, 94, 96, 136

SUtnirus miliarivs^ 647

Skeleton, figured, of Ttslndo, 319 ; oiChdone,
820 ; of Brontmaurusj 418 ; of Cerato-
saunis, 422 ; of Stegnmurus, 426 ; of
Iffuanodmi, 428 ; of Tricemtops, 430 ;
of Pterodactyl us, 486

Skin, of larval Amphibia, 31 ; shedding of,
32 ; glands, 32 ; pigment, 34 ; change
of colour, 35 ; poison, 36 ; of Apoda,
87 ; of Pi pa, 149 ; forms receptacles for
eggs, 151, 248 ; of Eusuchia, 441 f.

Skiu-glands,of Crocodiles,443 ; of Lacertilia,
497 : of Geckon(^s, 504 ; of Snakes, 583

Skull, of Urodela, 16 f., 17; of Ambly-
stmna, 17 ; of Salamandrina, 17 ; of
Salamandra, 17 ; of Anura, 28 f. ; of
Apoda, 84, 86

Skull, of Reptilia, 280, 281:— of Prorep-
tilia : Cricotus, 287 ; Eryops, 286 :— of
Microsauri, 289 : — of Protorosauri, 280 ;
PaUieohaiieria, 280, 291 :— of Rhyncho-
cephali {Sphenodon), 280, 296, 295 :—
of Theroniorpha, 280, 301, 303 ;

Elginia, 280, 305 f. ; Cynognathus, 280,

306 ; GoTd(m%€L, 280, 310 ; Dicynodon,
280, 310 ; Theriodontia, 306 ; Mam-
malian resemblances, 308 f. ; Lycosauras

307 ; ICndotkiodon, 807 ; Gompho-
gncUhus, 308 ; Anomodontia, 809, 280 ;
Oudenodon, 310 ; Placodus, 311 :— of
Mammalia, generalised, 281 : — of Che-
lonia, 316 f., 280 ; Sphargis, 335 ;
Chelydridae, 280, 338 ; Chelydra, 280,
838 ; Chelonidae, 817, 378, 879 ; Che-
lone, 817, 378 ; Thalassochelys, 879 ;
Clenimys, 366 ; Test-udo, 864 ; ChelySf
400,344 ; Trionyx, 406, 404 ; Chiysemys,
280, 346 ; Cisludo, 280, 361 : Pleurodira,
388, 400 ; Pelomedusidae, 390 ; Cliely-
didae, 399, 400; Trionychoidea, 404,
406:— of Dinosauria, 412 f., 422;
Anchisaunis, 421 ; Ceralosawtis, 422 ;
Diplodocns, 410 : — of Crocodilia, 280 ;
Pseudosuchia, 432 ; Parasuchia, 433 ;
Eusuchia, 434 f. ; Gavialis, 462 ; Croco-
dilus americanvs, 466 ; C. niloticus,
460 ; C. palustris, 466 ; C. porosus^
468; Alligator, 468: — of Plesiosauria,
473 ; Nothosaurus, ^Til : — of Ichthyo-
sauria, 479 ; Ickthyosaiirus, 281 : — of
Pterosauria, 484 ; iJimorphodon, 281 : — *
of Pythonomorpha, 488 ; Clidastes,
281 : — of Lacertilia, 281 ; Geckones,
504 ; Agamidae, 281, 516 ; Uromastix,
281 ; Iguanidae, 528 ; Anguidae, 537 ;
Helodermatidae, 540; Varanidae, 281,
642, 543 ; Varamis, 281 ; Tejidae, 547 ;
Lacertidae, 281, 550, 660 ; Lacerta,
281 ; Sciucidae, 559 ; Amphisbaenidae,
565 ; Chamaeleontes, 568, 669 :— of
Ophidia, 281, 696, 697, 688 ; Eunectes,
696, 697 ; Orotahts, 688

Slow-worm, 539, 639

Slugs eaten by tortoises, 363

Smell, sense of, of Chelonia, 830

Smith, the, - Hyla faber, peculiar nursiug
habits, 196 f.

Smooth Snake, 619, 620

Snakes, 5S1 f. ; skull. 581 f. ; 281, 688,
696, 697 f. ; vertebrae, 582 ; general
anatomical structure, 583 f. ; geographical
distribution, 585 f., 686 ; classification,
592 f.

Snake-charming, 631 .

Snake-poison, 586 f.

Snake-stones, 629 f.

Snapping Turtle, 33<S f.

Soft-shelled Turtle, 408

Sound produced by rubbing of scales of
TeratoscincHs, 507

Spade-foot, 162

Spelcrpcs, 94, 96, 97, 103, 104, 106 ; S.
altamazanicus 104 : S, hilinealus, 104 /
S. fuscus, 104, 105 ; tongue, 106 ; ^S*.
inf meatus, 104; S. lineolus, 104; S.

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666

INDEX

parvipeSf 104; S. porphyriticus^ 105; S,

aalnwnetUf 105; S. subpcUmattu, 104;

S. unifonnia, IO4
Spencer, on habits of Chirol^tea, 221 f.
Spermatophores, 5S, 128
Spermatozoa of Amphibia, 52 £
Sphargidae, SIS, SI4, SS3 f. ; affinities,

336 ; morphology of shell, 837
SphargU coriacea, 3S3 f., 834 ; absence of

horny shields, 325
Sphenodon, 288, 290, 305, 306, 432 ; S,

punctatutju, 293, 294 ! skull, 296 ;

cervical vertebrae, 297 ; habits, 298 f.
Sphenopkryii€y 225, 227
Sphemsauras, 82, 287
Spiny-tailed Lizard, 524 f*
Spiracle, development, 45
Spotted Newt, 127
Spy-Slange, 6S2
St. Helena, gigantic tortoises introduced,

375
Staganolepis, 4S4
Stannius, 8, 139 ; on vertebrae of Pdo-

bates, 20
Staurotypits saJvinij 342
Stegocephali, 78 f.; St. Lepospondyli, 5(7 f.;

St. Temnospondyli, 81 f. ; St. Stereo-

spondyli, 83 f. ; vertebrae, 78 f . ;

shoulder-girdle, 79 ; dermal armour, 79
Stegosauri, ^i5
Stegoftaunis armatus, 4^5 ; S. ungulatus,

426, 426
SletifHiacti/lns, deserticoloas, 494 ; sleeping

attitude, 509
Stereocf/dopSy 227 y 231; S. incrassatuSf

231
StereorhachiSy 308

Stereospondylous vertebrae, defined, 284
Stemot/vaem<s, 324, 389, 390; S. derbianus,

391 ; shields ot 327
Sternum, of Urodela, 15 ; of Anura, 25 ;

taxouoniic value, 141, 142 ; of Sphen-

odon, 297 f. ; Protorosauri, 290 ; Dino-

sauria, 414 ; Eusuchia, 440
Stewart, quoted, on Heloderma, 540 n.
Stinkpot Terrapin, 342
Suboccipital (first spinal nerve) of Anura,

144
Subregions, geographical 74 f.
Syrrhopus; 2\2
Systomata, 139

Tadpoles, horny teeth of, 58 f. ; of
Megaltyphn/Sy 59, 60 ; absorption of tail,
61 f. ; of Xenopus, 147, 148 ; of
BombinatoVy 157 ; of Alytes, 159 ; of
Hyla arborea, 193 ; of H. versicolor,
195 ; of Bh/o viridis, 181 ; of B.
ca/atnUa, 183 ; of B. vvJgaris, 176 ;
of Thoropa miliaris, 209 ; of Pseudis
paradoxa, 213 ; of Hylodes marthiicen^is,
214 ; of Rhinodennn darwini, 229 ; of

Arthroleptis seychdlensis, 243 ; of
Rana temporarta, 256; of R. opiMho-
dmty 260 ; of R. esctdetUa, 270

Tail, of Anura, 21, 24 ; its absorption, 61 ;
of Chelonia, 328 ; of Geckos, varioos
shapes, 506 ; reproduction of, 506

Tarentoiu mauriianic<i, 508, 509 f.

Tarsus (see also Limbs), of Chelonia, 819,
820, 321 ; of Dinosanria, 416, 418, 420,
421, 423, 426 ; of Theropoda, 420 ; of
CompaogncUhus, 423 ; of Iffitanodtyf^
428

Teeth, of Anura, 30, 138, 139 ; substitutes
for, 30, 68, 218, 237 ; of Apoda, 86 ; of
Rhyrukosaurus, 292 ; of Homoeoaanmg,
292 ; of Rhynchocephali, 292 ; of Sphen-
odmiy 296 ; of Theromorpha, 301 ; of
Etginia, 306, 280 ; of Cfynognathns, 306;.
280 ; of Lycosaurua, 307 ; of Galeaattrus,
307 ; of Endothwdoriy 307 ; of Aw-
pedias, 308 ; of StereorhacMs, 308 ; of
Gomphognathus, 308 ; of Tritylodon,

309 ; Mammalian resemblances, 309; of
Anomodontia, 309 ; of Dicynodtm, 280,

310 ; of Gordonia, 280 ; of Placodv*^
311; of Sauropoda, 418 f., 419; of
Tlieropoda, 420 f., 422 ; of Orthopoda,
424 f.; of Eusuchia, 437 ; of Ichthy-
osauri, 479 : of Snakes, 582

Tejidae, 514, H'^ f-

Teju, 548, 548

Teleosauridae. 450

Teleosaurus, 4^1

Teierpeton elffinense, 291

Temnospondylous vertebrae, defined, 284

Temperature of blood, 67 f. ; of water for
Crocodiles, 460

Teunent, on immunity of Cobras, 629 f. ;
on turtles at Ceylon, 384, 386 ; on
habits of Owcodilus palustris, 4^fi f. ;
on habits of C. porosus, 459 ; on peculiar
use of Varanus, 546 ; on habits of
Gecko, 511

Tentacular apparatus of Apoda, 45, 86, SS

Tepkro)iieloj)Oti, 493

Te-raioscinciis, deserticolous, 493 ; eye, 494 ;
T. scincifs, 507

Terrapin, 359 f.

Testis, 49

Testudinidae, 313, 314^ 345 ; distribution,
332

TestmlOy 365 ; skeleton, 322, 828 ; shields
of, 327; T. ahingdmii, 878, 378; T.
atUis, 372, 377 ; T. dmidini, 375, 376 :
T. clegans, 370 f. ; T. el^harUina, 374 ;
T. eleplmntopus, 378; T. ephippiina.
378 ; T. gigatUea, 374; T. gracca, 365
f. ; habits, 367 ; eggs, 369 ; great age,
369 ; T. grandUifen, 373 ; T. korsfiddi^
370 ; T.ihera, 366; age attained, 369 ;
rate of growth, 370 ; T. marginata,
367 ; T. perpigniana, 372 ; T. poly-

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INDEX

667

phemusy 371 f. ; T. sunieireit 376 ; T,
vosmaeri, 873, 877

Tetradadyliu, 5S9 ; T. a/ricanus, 559 ; T,
seps^ 559

Tetrapoda, Credner's name for "four-
footed " creatares in opposition to the
tishes, which have fins, 4, 11

Thalassemydidae, 880

Thalasaochelys earetta, individual variation
of shields, 326, 827, S87; skull, 879

Thecophora, definition of term, 337, 338

Theobald, on Varanusy 544

Theriodontia, 306

Tlieromorpha, 300, 301 ; skull, 280, 801 ;
their affinity to Mammals, 303 f., 309

Theropoda, 420

Thilenius, quoted, 571 n.

ThoracosauruA, 4^1

ThorUiSy 96, 108 ; Th. penruUulMS, 103

Tharopoy 186, 189; Th. miliarUy 209

Tipper Snake, 634

Tiliqua s. Cydodvs^ 561

Toes, number of, in Urodela, 16 ; in Anura,
28 ; of Geckos, structure, 505, '605

Tomistomay 435, 436, 450 ; T. schlegeliy
453

Tongue, of Amphibia, nerve-supply, 39 ;
shape of, in Anura, 47; of Spelerpes,
106 ; absent in Aglossa, 145 ; of Rana
escvlentay 268 ; of Crocodiles, 443 ; of
Lacertilia, 498 ; of Chameleons, 569 f.

Tortoise, Greek, 365 f. ; habits, 367 f. ;
Moroccan, 366 ; habits, 367 f. : Starred,
370 f. ; Gopher, 371 f. ; Gigantic Land-
Tortoises, 372 f.

Tortoises = C'helonia, 31 2 f.

Tortoise-shell of commerce, 386

Trachea, of CYocodiles, 443

TrachysauruSy 560, 560 ; T. ntgosiM, 560y
561

Tree-frogs, 185 f. : change of colour, 35

TrematosauriiSy 80, S3

Truxratops, 413 ; T. prorsusy 430y 430 ; 71
jlaMIatuSy 430

Triclutbatrachm, 240 ; T, rohustns, 271

Tt'igonoctphalus cenchri^y 645, 646, 646y
646

Trimerorhachis, 82

Trionycbidae, 313 ; distribution, 333

Trionvchoidea, 313y 314, 4^4 f- ; habits,
407

Trumj/y, nuchal plate, 324 ; skull, 406 ;
plastron, 406 ; number of costal
plates, 325; T./emx, 4O8, 409; T.
/ortnosa, 411* 411 ; T. gangeticus, 4IO,
410 ; T. huruniy 410 ; T. triungnis, 4IO

Triprion, 179, 185, 189; T. jH'tamtvs,
207

Trirhachiodony 309

TrUony H2 f., 96, 115, 125, 128. 181 ;
fossil, 83 ; sperm atoph ores, 68 ; T. alpes-
tris, 123, 126; T. as2>ery 123, 130; T.

blasiu, 126; T. hoscaiy 123, 127 ; T.
cristcUus, 122, 125, 125; T. hagm-
viueUeriy 123; T. helveiicuSy 127; T,
italicusy 127 ; T. marmoratuSy 122, 126 ;
T. montadoniy 127 ; T, montanusy 123,
130; T. painiatusy 127; T. poii-etiy
123 ; r. punctata =:vulgarisy 127 ; T,
pyreiiaeusy 130; T, pyrrhogaster, 123,
128 ; T. rusconiiy 123, 130 ; T, sinensis,
123, 128 ; T, tatniaiv^-'mdgarisy 1Z7 ;
r. iwosusy 123, 128; T. viridescenSy
123, 128; egg, 128; T, vittatuSy 122,
128; r. mdgaHsy 123, 127 ; T. tcalUi,
123, 130y 181

Tritylodoiiy 301, 309

Tropidonotusy 607 ; T. natrix, 608 f, ; T.
ordinatus, 611; T. sirtaliSy 616, 611;
T. tesadcUuSy 611

Tropidosauray 558

TupinandnSy 548; T. tegxiixin, 648 ; T.
nigropunctcdusy 548y 648

Turtles, 378 f. ; skuU, 817, 379 ; skeleton,
820 ; plastron, 821 ; on Laysan Islands,
883 ; Green or Edible, 381 f. ; Hawks-
bill, 384y 386

Tylototritmy 96, 115 ; T. andeisoniy 130 ;
T. verrucosus, 132

Tympanic cavity, reduction of, in Anura,
30 ; in Ophidia, 583

Tympanum of Aglossa, 143

Typhlomolgey 96 ; T, rathbuni, 135

TyphlonecteSy 87, 90 ; T. compressicavda,
93

Typhlopidae, 592y 593 f.

Typhlopsy 91; T. braminuSy 694; T.
remiiadariSy 594

TyphlosauniSy 564

TypJdotriUm, 94, 96, 102; T. s})elaeusy
103

Cra^otyphlnsy 86, 89

Ureter, 48 f., 49

Urino-geuital organs, 48 f., 49

Uroc(yrdylns, Sly 288

Urodaeum of Chelonia, 330

Urodela, 94 f. ; geograjihical distribution,

96
rmmastij', 524; ^« acanthitiumSy 526y

526 ; i\ hardwichi, 525
ITropeltidae, 592, 695
Vropeltisy 595 ; U, grandis, 595
rrtqAateSy 512
Uroplatinae, 512
Urostyle, of Anura, 23 ; of Chelonia, 328

Varanidae, 514. 542 f. ; skull, 642 ; dis-
tribution, 643

Vara mtSy 543; V, goiddi, 546 ; V.griseu-Sy
skull, 542; V, niloticus, 543; V,
sfdvatoTy 543 f., 646

Vas deferens, 48 f., 49

Vertebrae, procoelous, definition, 19, 138 ;

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668

INDEX

acentrous, ue, without a centre or body,
4 ; amphicoelous, defined, 12 ; of Uro-
dela, 11 ; gastrocentrou.s defined, 2S2 ;
lepospondylons, 5 ; defined, 78 ; noto-
centrous, 4 ; defined, 19 ; opisthocoe-
lous, defined, 12, 188 ; pseudocentrous,
4, 78 ; Htereospondyloas, defined, 79,
284 ; temnospondylous, 18 ; defined, 79,
284 ; development of — in Urodela, 12,
18 ; in Anura, 19 ; of trunk of Sala-
mandra, 14 ; epichordal, 20 ; sacral, of
Anura, 22 ; shifting forwards of sacral
attachment of ilium, 23 ; of Reptilia,
composition of, 283, 288 ; trunk-
vertebrae of Eryops^ 283, 286, 286 ; of
Cricotus, 287 ; of Microsauri, 289 ; of
Sph4inodoii, 294, 296, 297 ; atlas and axis
of Sphenodon, 283 ; of Theromorpha,
302 ; of Pareiasauri, 305 ; atlas fused
with axis in Cynognaih\iSf 307 ; of
Dimetrodon, with peculiar processes,
308; of Chelonia, 314 f . ; atlas of
TrionyXj 288 ; of ChelySf 283 ; of Dino-
sauria, 413 ; hollow in Dinosaurs, 415,
420 ; of Eusuchia, 438 f. ; atlas and
axis of CrocodUuSf 288 ; of Metrio-
rhynchusy 283 ; of Pterosauria, 485 ; of
Ichthyosauria, 480 ; of Pythonomorpha,
488; of Lacertilia, 494; of Geckones,
503 ; of Snakes, 582
Vertebral column, instance of greatest
shortening, 144 ; of Urodela, 11, 13 ; of
Stegocephali, 78 ; of Anura, 18 f., 21, 22 ;
Paiaeobatrachus, 22; Pipa, 22, 143;
Hy}nenochirtiSy 22, 143 ; Boinhinator^
22 ; Xenopvs, 21, 143 ; of Apoda, 86 ;
number of vertebrae of ProtorosauruSj
291 ; of Palacohatteria, 291 ; of Homoeo-
sauru^f 292 ; of SpJienodon, 297 ; of
Cynogimthus^ 306 ; of Ig^utnodon, 428 ;
of Eusuchia, 440 ; of Plesiosauria, 474 ;
of Elasmosauridae, 478
Vesiculae seminnles, 49, 51
Viper, Common, 641 f., 620, 642
Vipera, 64 1 ; V. (wnnodyfes, 641, 643;
V. aspis, 64$ ; V. hems, 6 41, 642, 620 ;
V. latastd, 643 ; V. russdli, 643
Viperidae, 592, 593, 637

Viperinae, 6S8

Viperine Snake, 610

Vis, de, on Chtamydogaurut^ 523

Viviparous, Chameleon, 572; Lacertilia,

499 ; Geckos, 506
Vocal sacs, 47 f. : of Paludicola, 220 ; of

Rhinodermat used as brood-pouches, 228
Voeltzkow, on nesting of Crocodiles, 462 f. ;

on Testudo' daudini, 375
Voice, 47

Wagler, 8

Wallace, on RhacopJiortigf 246 f.

Wall-Lizard, 557

Warning, attitudes, of Bcmbinator, 157 ;

colours, 88, 116, 156 ; of Helod^rma^

541
Water- Viper, 645, 646
Weismann, on Azolotl, 64, 114
Werner, on Eryx, 604
White's aged Tortoise, 369
Wilder, on DesmogncUhvSf 103

Xantusia, 547

Xantusiidae, 514, ^47

XenobcUrachtiSj 225 ; X ophiodon, 228

Xenopeltidae, 59Sy 605

Xenopeltis unicolor, 605

Xm<ipu8, 143 ; distribution, 148, 144, 146

f. ; X calcaratus, 146 ; X laevis, 140 f.,

147 ; X mueHeriy 146
Xenorhinu, 228
Xenosauridae, 513, 636
Xenosatinis grandiSf 536

Zachamns, 212

Zarrvenis constrictor, 613 ; Z.gemonnisis s.

viridiflaviis, 612; Z, hippocrepis, 613;

Z. muc4mu, 611, 612
Zanclodmi, 417, 421
Zander, on habits of Agama, 520 ; of

Phrymcephalus, 522 ; of Eryx, 604
Zaocya carinatus, 614, 816
Zatachys, 82
Zeller, on spermatophores, 53 ; on Protats,

134
Zonuridae, 513, 536
Zonunis derbianus s. giganUus, 536, 637

END OF VOL. VIII

Printed by R. & R. Clask, Limited, Edmburgh,

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THE CAMBRIDGE NATURAL HISTORY

Edited by S. F. Harmer, ScD., F.R.S., Fellow of King's College,
Cambridge, Superintendent of the University Museum of Zoology;
and A. E, Shipley, M.A., Fellow of Christ's College, Cambridge,
University Lecturer on the Morphology of Invertebrates.

To be completed in Ten Volumes. Svo. Price 17 s, net each

Intended in all respects to be a Standard Natural History accurate
enough to be of use to the Student, and at the same time popular enough for
the general reader who desires trustworthy information as to the structure
and habits of all members of the Animal Kingdom, from the Protozoa to the
Mammals. The Volumes are fully illustrated by original figures drawn
where possible from nature. When complete the Series is one which should
be indispensable in all Libraries, whether public or private.

WORMS, LEECHES, ETC.

VOLUME II

Flat Worms. By F. W. Gamble, M.Sc. Vict., Owens College.— Nemeftines.
By Miss L. Sheldon, Newnham College, Cambridge. — Thread-worms, etc.
By A. £. Shipley, M.A., Fellow of Christ's College, Cambridge. — Rotifers.
By Marcus Hartog, M.A. Trinity College, Cambridge, D. So. Ix)nd., Professor
of Natural History in the Queen's College, Cork. — Polychaet Worms. By
W. Blaxland Bekham, D.Sc. Lond.,Hon. M.A. Oxon., Professor of Biology in
the University of Otago.— -Earth-worms and Leeches. By F. £. Beddard,
M.A. Ozon., F.R.S., Prosector to the Zoological Society, London. — Gephyrea,
etc. By A. £. Shipley, M.A., Fellow of Christ's College, Cambridge.—
Polyzoa. By S. F. Harmer, M. A., F.R.S., Fellow of King's College, Cambridge.

CAMBRIDGE REVIEW,—** Several of the groups treated of in this volume are
unknown by sight even, to the general reader, and possess no popular name what-
soever ; and as only a few insignificant details are known of the habits of the
animals composing them, their treatment in the volume before us has necessarily
been to a large extent anatomical. This circumstance renders the book of especial
value to students, more particularly as in some cases the articles on the groups in
question are the first comprehensive ones dealing with their respective subjects. . . .
Most of the articles are of a very high order of merit — taken as a whole, it may be
said that they are by far the best which have as yet been published. . . . We may
say with confidence that the same amount of information, within the same compass,
is CO be had in no other zoological work."

NA TURAL SCIENCE,— **T]nB second volume of the Cambridge Natural History
is certain to prove a most welcome addition to English Zoological literature. It
deals with a series of animal groups, all deeply interesting to the specisJist in
morphology ; some important from their economic relations to other living things,
others in their life-histories rivalling the marvels of fairy-tales. And the style in
which they are here treated is also interesting ; history and the early observations
of the older writers lend their charm ; accounts of habits and mode of occurrence,
of life, in a word, from the cradle to the grave, are given in ample detail,
accompanied by full references to modern and current literature. The whole is
admirably illustrated."

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SHELLS

VOLUME III

Molluscs and Brachiopods.

By the Rev. A. H. Cooks, M.A., A. £. Shiplxy, M.A., and F. R. C. Bsed, M.A.

TIMES, — " There are very many, not only among educated people who take
an interest in science, bat even among speciaJists, wno will welcome a work of
reasonable compass and handy form containing a trustworthy treatment of the
Tarious departments of Natoral History by men who are familiar with, and
competent to deal with, the latest results of scientific research. Altogether, to
judge from this first volume, the Cambridge Natural History promises to Ailfil all
the ezpectationa that its prospectus holds out."

FIELD. — "We know of no book available to the general reader which affords
such a vast fund of information on the structure and habits of molluscs."

KNOWLEDGE,— ** It succeeding volumes are like this one, the Cambridge
Natural History will rank as one of the finest works on natural history ever
published."

ATHEN^UM, — "The series certainlv ought not to be restricted in its circula-
tion to lecturers and students only ; and, if the forthcoming volumes reach the
standard of the one here under notice, the success of the enterprise should be
assured."

INSECTS AND CENTIPEDES

VOLUME V

Peripatus. By Adam Sedowick, M. A. , F. E. S. — -Myriapods. By F. G. Sikoulik,
M.A.— Insects. Part I. By David Sharp, M.A. CanUb., M.B. Edin.,
F.R.S.

FIELD. — "Although written for the student and the specialist, the book is not
the less adapted to all intelligent readers who wish to make themselves thoroughly
acquainted with the habits, structure, and the modern classification of the animals
of which it treats. To such it cannot be recommended too strongly."

SCIENCE GOSSIP. — * * Every library, school, and college in 3ie country should
possess this work, which is of the highest educational value."

Frof. RAPHAEL MELDOLA, F.It.S., F.C.S., in kU PresiderUtal Address to ths
Entomological Society of London, said : — "The authors of this volume are certainly
to be congratulated upon having furnished such a valuable contribution to our litera-
ture. When its successor appears, and I will venture to express the hope that thia
will be at no very distant period, we shall be in possession of a treatise on the
natural history of insects which, from the point of view of the general reader, will
compare most favourably with any similar work that has been published in the
English language."

ENTOMOLOOISrS MONTHLY MAGAZINE.— '*^^ venture to think the
work will be found indispensable to all who seek to extend their general knowledge
beyond the narrowing influence of exclusive attention to certain orders or groups,
and that it will take a high position in 'The Cambridge Natural History' series."

INSECTS— Part II

VOLUME VI

Hymenoptera continued (Tubnlifera and Aculeata), Coleoptera, Strepsipteim.
Lepidoptera, Diptera, Aphaniptera, Thysanoptera, Hemiptera, Anoplora.
By David Sharp, F.R.S.

SATURDAY REVIEW.— '''Di. Sharp's treatment is altogether worthy of the
series and of his own high scientific reputation. But in a work of this sort it is
not only necessary that information should be accurate, but also that it shall be
presented to the eye, so far as illustrations and printing are concerned, in such a
way as to render its matter as easily intelligible as possible, and readily usable
for purposes of reference. Under both these heads we nave nothing but commenda-
tion for Mr. Sharp's treatise. The illustrations are indeed beautiful, and the use
of the heavy type for tlie headings of the various sections and leading paragraphs
materially helps the reader in the progress of his study. Certainly tnis is a book
that should be in every entomologist's library. "

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DAILY NEWS, — *'It would be hard to say too much in praise of this most
admirable volume. It is too often the case that scientific books are written in a
dull and uninteresting style. The reader will find nothing of that kind to complain
of here. The descriptions are clear, the illustrations are excellent ; while, as in the
previous volumes of the series, printing and P&per sre all that oould be desired."

SPEAKER, — '* Amateur naturalists will nnd the volume of the greatest possible
assistance, while serious workers will welcome it as an extremely convenient hand-
book, in which the latest results of original research at home and abroad are clearly
and succinctly summarised. No book so comprehensive and of such value has
appeared since Westwood's CUuiifictUum marked an epoch in the literature of
entomology sixty years aso."

LITERATURE.^" ^Q may confidently expect it will take a similar position
to that which Westwood's Introduction has so long occupied. ... An immense
amount of well-selected matter, much of which is by no means easy of access, has
been brought together and made available for instant reference by an index extend-
ing over twenty-four pages, in double columns."

AMPHIBIA AND REPTILES

VOLUME VIII
By H. Gadow, M.A., r.R.S. [Ready,

BIRDS

VOLUME IX

By A. H. Evans, M.A., Clare College, Cambridge. With numerous Illustrations
by G. E. LoDOB.

IBIS, — '* Mr. Evans has produced a book full of concentrated essence of informa-
tion on birds, especially as regards their outer structure and habits, and one that
we can cordially recommend as a work of reference to all students of ornithology."

NATURE NOTES.—** We venture to predict that, of the ten volumes of which
this excellent series is planned to consist, none will secure a wider popularity than
Mr. Evans's treatise on birds. Strange as it may appear, among the manv books
on birds- that have appeared of late years, we do not recall anv that covers the same
ground. . . . We are grateful to the author for the mine of valuable information
which he has crowded between his two covers."

SCIENCE O0SSIP.—**GQnetdX readers will find this work most useful in
obtaining a proper understanding of birds, and will be assisted by the effective
diaeram of a hawk in the introduction, showing the recognised names of every part
of the exterior appearance. The expressions used in naming the various portions
are fully explained on the adjoining page. As we have already said, the illustra-
tions are admirable. The book is a useful addition to any library, as it treats of
nearly every known kind of bird throughout the world."

NATURALISTS' JOURNAL.—** As a descriptive handbook the student can
have no better. . . . We have no hesitation in saying every ornithologist ought to
possess the book."

SATURDAY REVIEW.— '** The expert and the novice alike must be at once
delighted by the accuracy and the beauty of the illustrations. ... It is astonish-
ing to note the mass of information the author has been able to bring together. . . .
With a little practice any observant person would soon learn bv the help of this
volume to track down any bird very nearly to its ultimate place in classification."

LITERATURE.—** The classification— in the main that of Dr. Hans Gadow— is
admirable. . . . The general account prefixed to the Orders and Sub-Orders is
excellent ; the illustrations, drawn for the greater part by Mr. Lodge, are admirablv
characteristic. In a serviceable introduction the author has shortly dealt with
feathers, colour, moult, structure, and the leading facts of migration. The book,
as a whole, is a reflection of conscientious and accurate labour. . . . The accuracy
of the work is beyond all challenge."

SPEAKER. — "It is safe to conclude that all workiufi^ ornithologists will place
this volume on their shelves within easy reach, ana not far from Newton's
* Dictionary of Birds ' ; and we have great pleasure in recommending it to all who
take an intelligent interest in birds and bird-life as an admirable introduction and
guide to the study of the subject. . . . The illustrations by Mr. G. E. Lodge are
admirable. All reach a very high level of excellence, and for a few no praise can
be too high."

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COMPLETE LIST OF SERIES

To be completed in Ten Volumes. 8vo, 1 7«. net eadk
VOLUME I

Protozoa, Marous Haetoo, M.A., Trinity College (Professor of Natural History in
the Queeu's College, Cork) ; Spong:e8, W. J. Sollab, Sc.D., F.B.S., St John's
College (Professor of Geology in the University of Oxford) ; Jellj-fish, Sea-
Anemones, etc, S. J. HiCKsoN, M.A., Downing College (Beyer Professor of
Zoology in The Owens College, Manchester) ; Star-fish, Sea- Urchins, etc,
£. wTMaoBbide, M.A., St. John's College (Professor of Zoology, McGill
University, Montreal).

VOLUME II

Flat Worms, etc., F. W. Gamble, M.Sc. Vict. (Demonstrator and Assistant-
Lecturer in Zoology in The Owens College, Manchester) ; Nemertines, Miss L.
Sheldon, Newnham College; Thread-worms, etc, A. £. Shipley, M.A.,
Christ's College ; Rotifers, etc., Marcus Habtgq, M.A., Trinity College,
D.Sc. Lond., (Professor of Natural History in the Queen's College, Cork);
Polychaet Worms, W. B. Benham, D.Sc. Lond., F.KS., Hon. M.A. Oxon.,
(Professor of Biolosy in the University of Otago) ; Earth-worms and Leeches,
F. £. Brddard, M.A. Oxon., F.R.S. (Prosector to the Zoological Society);
Gephyrea, A. E. Shiplbt, M.A., Christ's College ; Polyzoa, S. F. Harmeb,
M.A., F.R.S., King's College. [lUady.

VOLUME III

MoUuscs, A. H. Cooke, M.A., Kine^ College; Brachiopods (Recent), A. E.
Shipley, M.A., Christ's College ; Brachiopods (Fossil), F. R. C. Reed, M. A.,
Trinity College. llleady.

VOLUME IV

Spiders, Mites, etc., C. Warburton, M.A., Christ's College (Zoologist to the Royal
Agricultural Society); Scorpions, Trilobites, etc., M. Laurie, B.A., King's
College, D.Sc Edinb., (Professor of Zoology in St. Mungo's College, Glasgow);
P7cnog:otiids, etc, D'Arcy W. Thompson, C.B., M.A., Trinity College
(Professor of Zoolo^ in University College, Dundee) ; Crustacea, W. F. R.
Weldon, M.A., F.R.S., St John's College (Linacre Professor of ComparatiTs
Anatomy in the University of Oxford).

VOLUME V

Peripatns, A. Sedgwick, M.A., F.R.S., Trinity Collt>ge; Centipedes, etc, F. G.
Sinclair, M.A., Trinity College; Insects, Part I., D. Sharp, M.A., F.R.S.

[Beady.
VOLUME VI

Insects, Part IL, D. Sharp, M.A., F.R.S. [JReady.

VOLUME VII

Balanoffiossus, etc, 8. F. Haruer, Sc.D., F.R.S., Eing^s College; Asddiana
andAmphiozus, W. A. Herdman, D.Sc. Lond., F.R.S. (Professor of Natural
History in University College, Liverpool) ; Fishes, T. W, Bridge, So.D.,
Trinity College (Professor of Zoology in Birmingham University). [In the Press,

VOLUME VIII
Anqihibia and Reptiles, H. Oadow, M. A„ F.R.S., Ring's College. [JReady.

VOLUME IX
Birds, A. H. Evans, M.A., Clare College. [Beady.

VOLUME X

Mammals, F. E. Beddard, M.A. Oxon., F.R.S. (Prosector to the Zoological
Society). [In the Preee,

MACMILLAN AND CO., Ltd., LONDON.

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