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

Harvard College 



CUm of 1839 


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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 


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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. 


I 90 I 

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S 7670.ll^(x)j 

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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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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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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 

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 

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 

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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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. 


Cambridge, December 19, 1900. 

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Preface v 

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



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


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 

Nkoteny— Regeneration— Temperature — Geographical Distribution . 63 

Stegocephali or Labyrinthodonts— Lissamphibia— Apoda ... 78 

Lissamphibia {continvbd) — Urodela 94 

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

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Definition and CHARArrERs— Position of the Class Reptilia in the 

Phylum Vertebrata— Classification— Skull and Vertebrae . 277 

Proreptilia — Prosauria— Theromorpha 285 

Chelonia— Atheoae— Thecophora 312 



Plesiosauria — Ichthtosauria— Pterosauria— Pythonomorpha . . 473 



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

INDEX 651 

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PHALI (p. 78) 


C Branch 10- 



phali Lepo- 1 sauki (p. 80). 

(p. 80) I 

phali Temso- 
(p. 81) 

' (p. 84) 


(p. 94) 


(p. 81). 



(p. 143). 


(p. 138) 



(p. 152) 


[ Amphiumidae 
(p. 97). 

Salamandridae I 
(p. 102) 1 

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


(p. 152). 


(p. 160). 

(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). 


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



(p. 288) 


(p. 300) 


(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) 



(p. 431) 


(p. 473) 


(p. 478) 


(p. 484) 


(p. 487) 


(p. 838) 


(p. 412) 


1 Ornit 
I (p. 4 


(p. 418). 

(p. 420). 


(p. 424) 


(p. 430). 

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

Ea8nchia(p. 434) 

' Nothosauri 

(p. 476) 


(p. 477) 


(p. 483). 


(p. 486) 

' Dolichosauri 

(p. 489). 

, (p. 489). 

f Cryptodiua 
(p. 388) 

Pleurodira I 

(p. 388) ] 

(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). 
(p. 404). 

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

. (p. 487). 

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(p. 502) 


fp. 491) 


(p. 491) 

•i Lacertae 
(p. 513) 

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). 




(p. 606) 

(p. 623) 

(p. 625) 


(p. 637). 


(p. 6.37) 

(p. 606). 
Colli hriuae (p. 607). 
, (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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' '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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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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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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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 

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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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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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 ! 


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 

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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DiPNOA. [The Amphibia] 

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


'Mutabilia [with metaniorphoBiB, e.g. 

Amphipneusta [Perennibranchiate Uro- 

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

Twith temporary gills: Eiaudata + Caudata 

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. 


Wagler, 1830, "Systema Amphibionun." 

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


Ranae I. AGLOSSA. 

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

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- 
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 


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

•' 5^/y»7, neck ; yui'w, close. 

Digitized by 



2. Batrachia. A0L08SA. 

PHANER0GL08SA : Systomata = 
Buibninae. Without manubrium 

' stemi. 
Raninae. With manubrium. 
Hyloidea. With adhesive finger- 
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- 
second edition of the "Grundzuge" (1870) they are divided into 
Huxley, 1864, "The Elements of Comparative Anatomy." 

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

Haeckel, 1866, "Generelle Morphologie." 

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

Steqocephali, Gymnophidia, Urodela, Proteidea, Trachy«tomata, 
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- 
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. 

Digitized by 



Siil)-Cla68 II. Urodela. 

Order 1. Proteidae : Proteus. 

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


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. 

Digitized by 







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. 



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. 

Digitized by 



(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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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. 

Digitized by 



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 

Digitized by 



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 

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 

Digitized by 




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. 

Digitized by 




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 

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 

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 

Digitized by 



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 

Digitized by 



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, 

Digitized by 



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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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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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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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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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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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- 

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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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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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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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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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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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} 


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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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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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- 


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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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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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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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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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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. 


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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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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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 

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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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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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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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 

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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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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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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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. 

Digitized by VjOOQIC 



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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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 

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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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 ; segmental duct ; T, testis ; (/V, ureter ; 
V.d, vas deferens ; F.«, vesicnla seminalis. 


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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 

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 

Digitized by 



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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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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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 

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. 

Digitized by 



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. 

Digitized by VjOOQIC 


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 

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 

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 

Nursing habits are verj' common amongst the Anura. 

Digitized by 



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 

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 

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. 

Digitized by 



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. 

Digitized by 



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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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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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, 

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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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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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 

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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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 

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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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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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- 


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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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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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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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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 

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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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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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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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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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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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 

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. 

Digitized by 



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. 

Digitized by 



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 

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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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. 










^ 1 

K! 1 










5t ' 









Proteidae . 




















+ i 

Pelobatidae . 





Bufonidae . 

'' -H 









1 -4- 








' Hemiphractinae . 


! 1 

' Cystignathinae . 

1 + 



1 i 

Genyopiiryninae . 


Engystoioatinae . 



























+ ! 

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

Digitized by 





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. 

Digitized by 



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, 

Digitized by 




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. 


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 

Digitized by 



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. 


Mostly with rather long ribs and with chiefly ventral 


Digitized by VjOOQIC 


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 

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. 

Digitized by 



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. 


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 

Digitized by 



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. 

Amj>hibia loithout dermal armour. 


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 

Digitized by 





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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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 

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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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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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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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 


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

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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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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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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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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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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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 


Fig. 16. 


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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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 


Westeni PaUearctic. 



6 S 



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 


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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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 


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

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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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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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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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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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 


the e<i<^ 

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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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 

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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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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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 

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

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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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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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 
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 

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

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 


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 

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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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: 



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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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 

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. 


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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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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- 

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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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 

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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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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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 

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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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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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- 

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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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 

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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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 

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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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 

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

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, 

Digitized by 



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 

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 

Digitized by 


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. 

Digitized by 



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. 

Digitized by 



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 

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. 

Digitized by 




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 


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. 

Digitized by 


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 

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. 

Digitized by 



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 

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 

Digitized by 




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, 

Digitized by 



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. 

Digitized by 





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. 

Digitized by 



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. 

Digitized by VjOOQ IC 




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 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 :- — 

Digitized by 



^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 — 


B. Finnistemia. 

a. SiAcral diapophyses dilated — 


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

p. 227. 

^^^^^ Dyacophinae, p. 235. 

SiEd. p. 236. 

- Engystomatviiaef p. 225. 

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

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


' - 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 


Digitized by 



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). 
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- 

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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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 

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 

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 

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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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 

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 


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 


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

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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 

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 

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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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. 


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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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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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h krW"^ 

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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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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 

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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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 

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 

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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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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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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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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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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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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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 

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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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- 


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 

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. 
New Guinea 
j3. Sacral vertebra with two 
India and Malaya 

b. Vertebrae opisthocoelous. 

{Ceylon and Malayan Islands . 
New Guineii . 

PelobateSj p. 162. 

PelodyteSy p. 165. 

Leptobrachium, p. 166. 

Mi'f/alophrySy p. 60 (Fig. 11). 

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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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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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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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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 

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 

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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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 

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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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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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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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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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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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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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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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 

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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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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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 

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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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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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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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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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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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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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 


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 


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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Most, if not all, Hylidae are climbers, and manjr lead an 
arboreal life, but it does not follow that all the " Tree-frogs " are 

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 

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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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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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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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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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, 



Fig. 37.— //y« arborea, var. meridioncdis. South European Tree-frog, x 1. 

although this colour is perhaps the most common in the arboreal 

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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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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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 

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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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 

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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 

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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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 

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 

" 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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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 

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 

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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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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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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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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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 

Digitized by VjOOQ IC 

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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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- 

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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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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" 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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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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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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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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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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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 


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. 


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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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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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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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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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 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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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 soil at the bottom, but 
only one was visible on arrival The other wns in si tit- tlie 

Digitized by VjOOQ IC 

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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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\ ^^^ 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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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 

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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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 

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 

^ Proe. Linn, Soc. N,S. W. (2), iv. 1898, p. 357. 

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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 

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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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 


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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 

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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absent, and this loss is compensated for by exceptionally strong 

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. 

Palate i\ith dermal papillae. Africa. 

Breviceps, p. 232. 
With palatal dermal folds. Madaga.scar. 


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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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. 


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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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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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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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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and the hind-limbs are short. The male has a subgular vocal 

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$. 

Digitized by 



(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, 

Digitized by 



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 

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. 

Digitized by 



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 

Digitized by 



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. 

Digitized by 



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- 

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 

Digitized by 



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 

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. 

Digitized by 




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 


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 

Digitized by 


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* 

Digitized by 



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 

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* 


Digitized by 



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. 

Digitized by 



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 marbled, whitish below. The 

' Zool. Jahrh. Syst. xii. 1898, p. 89. 

Digitized by 



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 

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. 

Digitized by 



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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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- 

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 

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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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 

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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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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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 

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 

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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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 

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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quite transparent and thin stratum, in this case the turgid 

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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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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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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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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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 


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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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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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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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Rocky Mountains, southwards into Mexico, northwards into 

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 

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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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 

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 

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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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 


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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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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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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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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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 


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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 


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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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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. 


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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 

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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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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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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 


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 


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 

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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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, 

„ 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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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 : — 


Ichthyosauria Plesiosauria 







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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the centra or bodies of the vertebrae are formed by the pairs 
of interventralia, while the basiventralia are reduced, persisting 



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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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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proreptiua prosauria theromorpha 


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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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 

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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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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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.) 


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 

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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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. 


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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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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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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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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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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 



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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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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». 60.— The 

first three 

cervical vertebrae of 


1, 2, 3, 4, 


; c, . C„ 

centra; Ny 

iVj, neural 



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 

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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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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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 

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. 


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 

The various genera exhibit such a diversity of structure, 
shape, and size, and many are still so imperfectly known, that 

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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 

Digitized by 


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 

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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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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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. 


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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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 


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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. 


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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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 

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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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. 


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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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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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 


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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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 

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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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 — 
ty # y//n/u, n I* -. pij^^jyg^^^ernidae 

[Testudinidae — Chelonidae 
, Trionychoidea Trionychidae 

* Cat, Chelonians, Brit. Mus. 1889. 

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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- 

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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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- 

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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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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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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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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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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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. 


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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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, 
for the toes. In Felomedusa all the fingers possess two phalanges 
only, owing to fusion of the first and second phalanges with each 

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." 


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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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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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 





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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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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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 

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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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 

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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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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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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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. 





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 


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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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 

^ /> 


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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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 


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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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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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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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. 

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of a Chelonian, its age would speak greatly in favour of the 
Athecae being a very primitive and independent group. 


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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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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(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 

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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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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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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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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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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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 

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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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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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 


Flo. 76. — Chryscmys jji'dOy " Painted Terrapin. 


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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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 

Growth of Ch, picta, after Agassiz. 



Length of 

Breadth of 

Height of 

Length of 










Second . . . , 26-5 









Fourth . 















Seventh . 





Eighth ( 6 ) 





Ninth ( 6 ) 





Tenth ( 6 ) 





Eleventh ( 6 ) 





Fourteenth ( 6 ) 





TAventy-fifth ( 9 ) 





Old 9 . 





Very old 9 






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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which stand so closely together that they form almost median 

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. 


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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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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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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I^HCTl, ' 



^^Pl i^''it^' 

^|b il'' 


9H||k^ t 






Ito^Hbl ^f 





'»r mUm ^^ 










UjQp 1 1 


HPt jf^'^' 



1'' wllif^^ 



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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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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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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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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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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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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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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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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 

(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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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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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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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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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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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 

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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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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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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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. 


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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 

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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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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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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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 

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. 

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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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 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 

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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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 

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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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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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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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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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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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- 

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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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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:2 ^ 



eS ^ 


o bo 

^ s 

3 o 

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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 

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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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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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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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 

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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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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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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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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in Madagascar. S. derbianus in West Africa, from the Gambia 
to Angola, is the largest species, with a shell nearly one foot in 

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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to the prowling jaguar, which turns them over on to their backs 
and neatly cleans out the flesh with its sharp and powerful 

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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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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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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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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opinion of the settlers on the Upper Amazon is, that the turtle 
has very greatly decreased in numbers, and is still annuaUy 

"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 

" 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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he Strikes it with a second arrow. With the increased hold 
given by the two cords he has then no difficulty in landing his 

" 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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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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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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by the frontals. The fifth and eighth cervical vertebrae are 

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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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 



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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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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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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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 

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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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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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 

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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Islands, and to Africa from the Nile to the Senegal and to the 

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 

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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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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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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Hudson river ; but it is not found in any other Atlantic stream 
between that and the Savannah river, a distance of nearly 800 

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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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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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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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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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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for the insertion of the long caudi- femoral or long adductor 

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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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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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 

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. 


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Fuhes simjyle, with symphysis. Premaxillae with teeth. 

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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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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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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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 

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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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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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. 


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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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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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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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 

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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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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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 

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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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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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 

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. 


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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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. 


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 

The vertebrae are amphicoelous. The first and second 


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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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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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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 

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 

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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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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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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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, 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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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 

Digitized by 



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