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SNAK 



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G.A.BOULENGER 



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THE SNAKES OF EUROPE 



UNIFORM WITH THIS VOLUME 

THE LIFE OF CRUSTACEA 
BRITISH FRESHWATER FISHES 
THE OX AND ITS KINDRED 
THE LIFE OF THE MOLLUSCA 



THE 
SNAKES OF EUROPE 

BY 

G. A. BOULENGER 

LL.D., D.SC, PH.D.. F.R.S., F.Z.S. 



WITH FOURTEEN PLATES AND FORTY-TWO FIGURES IN THE TEXT 



METHUEN & GO. LTD. 
36 ESSEX STREET W.G. 
" LONDOl ' 



First Published in igi3 



PREFACE 

THERE is no work in the English language 
dealing with the Reptiles of Europe. I have 
therefore endeavoured to supply this desideratum, so 
far as the Snakes are concerned, by drawing up in a 
concise form an account of what is known of their 
characters, their distribution, and their life-histories. 
Professor Sordelli, of Milan, having kindly acceded 
to my request to reproduce some of the beautiful 
figures drawn by him for the work published in 
collaboration with the late Professor Jan under the 
title of" Iconographie Generale des Ophidiens," I have 
been able to supplement my descriptions with illus- 
trations which leave nothing to be desired from the 
point of view of accuracy. A few drawings have 
been made specially for this book by Mr. J. Green. 
I have further to acknowledge the permission given 
by the Trustees of the British Museum, the India 
Office, and the Zoological Society, to reproduce a few 
figures from previous publications of which I am the 
author. 

In order to render this little book more useful, the 
account of the Snakes of Europe has been preceded 



vi THE SNAKES OF EUROPE 

by an Introduction summarizing what is known of 
Snakes generally. 

I have purposely avoided overburdening a work of 
this kind, which aims at concision, with bibliographi- 
cal references and synonymic lists. I am sure my 
readers will be thankful for being spared this display 
of erudition. Whenever I have had to compile, and 
to trespass on ground that is not my own, I have 
been careful to draw only from the writings of the 
most trustworthy authorities. The descriptions of 
the species are based on the collection in the British 
Museum, which has been considerably increased 
since the publication of the Catalogue of Snakes 
(1893-1896). I have also had access to Monsieur F. 
Lataste's rich private collection, now under my care, 
and Dr. R. Gestro has kindly entrusted to me for 
study the collection of Italian Snakes in the Genoa 
Museum. I am indebted to Dr. L. W. Sambon for 
the chapter on Parasites, which he has written at 
my request. 

To all who have helped me I beg to tender my 

hearty thanks. 

G. A. B. 



CONTENTS 



INTRODUCTION 

CHAPTER I'AGE 

I. Definition and Classification - - i 

II. External Characters — Integument - - 8 

III. Coloration - - - - 29 

IV. Skeleton - - - - - - 40 

V. Dentition - - - - - -53 

VI. Poison Apparatus — Different Kinds of 

Poisons - - - - - 62 

VII. Nervous System — Sense Organs - - 73 

VIII. Viscera - - - - - -77 

IX. Organs of Reproduction ; Pairing; Oviposi- 

tion ; Development - - - 82 

X. Habits - - - - - 91 

XI. Parasites - - - - - - 107 

XII. Distribution - - - - - 118 

XIII. Snakes in Relation to Man - - - 133 

SYSTEMATIC ACCOUNT OF THE SNAKES 

OF EUROPE 

First Family: TYPHLOPIDjE 

Genus TYPHLOPS, Schneider - - - 144 

1. Typhlops vermicularis, Merrem — The Greek 

Blind-Snake - 144 

vii 



viii THE SNAKES OF EUROPE 



Second Family : BOID^E 

PAGE 

Genus ERYX, Daudin - - - - - 147 

2. Eryx jaculus, Linnaeus— The Javelin Sand-Boa - 147 



Third Family: COLUBRID^E 

Genus TROPIDONOTUS, Kuhl - - - 152 

3. Tropidonotus natrix, Linnaeus— The Grass- 

Snake, or Ring-Snake - - - - 152 

4. Tropidonotus tessellatus, Lauren ti — The 

Tessellated Water-Snake - 160 

5. Tropidonotus viperinus, Latreille — The Vi- 

perine Water-Snake .... 165 

Genus ZAMENIS, Wagler - - - - 170 

6. Zamenis gemonensis, Laurenti — The European 

Whip-Snake - - - - - 170 

7. Zamenis dahlii, Fitzinger — Dahl's Whip-Snake - 177 

8. Zamenis hippocrepis, Linnaeus — The Horseshoe' 

Whip-Snake \- - - - 179 

Genus COLUBER, Linn^us - - - - 181 

9. Coluber quatuorlineatus, Lacepede— Aldro- 

vandi's Snake - 182 

10. Coluber dione, Pallas — The Dione Snake - 185 

11. Coluber longissimus, Laurenti — The ^Escula- 

pian Snake - - - 187 

12. Coluber leopardinus, Bonaparte — The Leop- 

ard Snake ------ 191 

13. Coluber scalaris, Schinz — The Ladder Snake - 194 

Genus CORONELLA, Laurenti - - - 196 

14. Coronella austriaca, Laurenti — The Smooth 

Snake - - - - - - 197 

15. Coronella girondica, Daudin — The Southern 

Smooth Snake - 202 



CONTENTS ix 

PAGE 

Genus CONTIA, Baird and Girard - - - 205 

16. Contia modesta, Martin— The Dwarf Snake - 205 

Genus CGELOPELTIS, Wagler - - - 207 

17. Ccelopeltis monspessulana, Hermann — The 

Montpellier Snake ----- 208 

Genus MACROPROTODON, Guichenot - - 212 

18. MACROPROTODON CUCULLATUS, I. Geoffroy — The 

False Smooth Snake - - - - 213 

Genus TARBOPHIS, Fleischmann - - - 216 

19. Tarbophis fallax, Fleischmann — The Cat-Snake 217 
2 . Tarbophis iberus, Eichwald — The Caucasian Cat- 
Snake - - 219 

Fourth Family: VIPERID^E 

Genus VIPERA, Laurenti - - - - 221 

21. Vipera ursinii, Bonaparte— Orsini's Viper - 221 

22. Vipera renardi, Christoph — Renard's Viper - 227 

23. Vipera berus, Linnaeus — The Northern Viper, or 

Adder ------ 230 

24. Vipera aspis, Linnaeus— The Asp Viper - - 239 

25. Vipera latastii, Bosca — Lataste's Viper - - 247 

26. Vipera ammodytes, Linnaeus — The Sand-Viper, 

or Long-Nosed Viper ... - 249 

27. Vipera lebetina, Linnaeus — The Blunt-Nosed 

Viper, or Kufi - - - - - 257 

Genus ANCISTRODON, Palisot de Beauvois - 261 

28. Ancistrodon halys, Pallas— Pallas's Pit-Viper - 262 

Index - - - - - - - 265 



LIST OF PLATES 

PLATE FACING PAGE 

I. Typhlops vermicularis, Eryx jaculus - 144 
II. Tropidonotus natrix and vars, cettii and 

PERSA - - - - - -152 

III. Tropidonotus tessellatus, T. viperinus and 

VAR. AUROLINEATUS - - - l6o 

IV. Zamenis gemonensis and vars. persica and 

VIRIDIFLAVUS ----- 170 

V. Zamenis gemonensis, var. caspius, Z. dahlii, 

Z. HIPPOCREPIS - - - - 176 

VI. Coluber quatuorlineatus and var. sauro- 

MATES, C. DIONE - - - - 1 82 

VII. Coluber longissimus, C. leopardinus and 

VAR. QUADRILINEATUS - - - 1 88 

VIII. Coluber scalaris- - - ■ - 194 

IX. CORONELLA AUSTRIACA - - - - 196 

X. CORONELLA GIRONDICA, CONTIA MODESTA - 202 

XI. COZLOPELTIS MONSPESSULANA, M ACROPROTODON 

CUCULLATUS, TARBOPHIS IBERUS, T. FALLAX 208 

XII. VlPERA URSINII, V. RENARDI, V. BERUS - 220 

XIII. VlPERA ASPIS, V. LATASTII - - - 240 

XIV. VlPERA LEBETINA, V. AMMODYTES, ANCISTRODON 

HALYS ------ 250 



XI 



THE SNAKES OE EUROPE 



INTRODUCTION 
CHAPTER I 

DEFINITION AND CLASSIFICATION 

SNAKES, Ophidia— regarded by some authorities 
as an order of the class Reptilia, by the author 
as a sub-order of the order Squamata, which includes 
besides the Lizards, Lacertilia, the Chameleons, 
Rhiptoglossa, and the extinct Dolichosauria and 
Mosasauria — may be defined as greatly elongate 
scaly Reptiles without limbs, or with mere vestiges 
of the hind pair, without movable eyelids, without 
ear-opening, with elongate, deeply forked tongue 
retractile into a basal sheath, with transverse vent 
and paired copulatory organs, and with the two 
halves of the lower jaw independently movable, 
connected at the symphysis by an elastic ligament. 

The latter character alone distinguishes them 
from all Lizards, but no single Lizard possesses all 
the others in combination. 

In their most highly developed form these Reptiles 
are adapted for rapid reptation and for swallowing 



2 INTRODUCTION 

prey much exceeding their own calibre ; hence the 
bones of the skull, on which a prehensile function 
devolves, are loosely attached to the cranium by 
ligamentous elastic tissue, or articulated in such a 
manner as to permit a wide buccal expansion ; whilst 
the absence of a sternum and the mobile attach- 
ment of the ribs allow a corresponding dilatation of 
the body as the prey descends into the digestive canal. 

The fatal venom which many of these Reptiles 
possess has so impressed the mind of men, even the 
scientific, that for a long time snakes were primarily 
divided into poisonous and non-poisonous, a classi- 
fication in which the more important characters, 
derived from the general structure, and especially 
from the skull, were subordinated to the physio- 
logical. Such a system was far from reflecting 
natural relationships. Besides, as our knowledge 
progressed, drawing a distinction between poisonous 
and harmless snakes became more and more diffi- 
cult, so many snakes previously regarded as harmless 
proving to be poisonous in various degrees — at least 
enough to paralyze the small prey on which they 
subsist, if not to be of serious danger to man. 

In the division into families, as followed in this 
work, the presence or absence of a poison organ is 
left out of consideration. Further, in this as in 
many other groups of the animal kingdom, external 
characters do not furnish trustworthy indications 
for higher divisions, and the definitions of the 



DEFINITION AND CLASSIFICATION 3 

families are therefore based exclusively on osteo- 
logical characters. For those who wish to name 
snakes with facility, the key which concludes the 
chapter on External Characters will, however, remedy 
this defect, and suffice for the identification of all 
the European species without any reference to their 
anatomy. Many attempts have been made to 
furnish an easy criterion for the distinction of harm- 
less from poisonous snakes, but the characters 
hitherto suggested with this object can only be ap- 
plied successfully to the small number of representa- 
tives in a limited area. Thus, in Southern Australia 
it might be stated that all snakes showing the regular 
nine large shields on the upper surface of the head 
are dangerous to man, whilst those with small 
shields or scales are harmless ; but in most parts of 
Europe this criterion would have to be reversed. In 
some countries the shape of the pupil might be used 
for the purpose, in others the size of the ventral 
shields, or the presence or absence of a loreal shield, 
between the nasal and the preocular, and so on. 
But when we have to deal with the snakes of the 
whole world, about 2,000 species, of which nearly 
one-third are poisonous to a greater or less degree, 
every attempt at a definition of the two categories 
without regard to the dentition breaks down. Only 
those who have made a study of the snakes of the 
world can make a guess from the general appearance 
as to an unknown form being poisonous or not, and 



4 INTRODUCTION 

even they may sometimes feel embarrassed, unless 
the dentition be examined ; the mistakes which 
have occasionally been made by some experienced 
herpetologists are proof sufficient of the fallacy of 
external characters for this purpose. 

The Ophidia are divided into nine families, the 
first, third, seventh, and ninth of which have repre- 
sentatives in Europe : 

I. No transverse (ectopterygoid) bone ; pterygoid 
not extending to quadrate or mandible ; no supra- 
temporal ; nasals in contact with prefrontals ; 
coronoid present ; vestiges of pelvis. 

Maxillary loosely attached to lower surface of 
cranium, toothed; lower jaw edentulous; a single 
pelvic bone i. Typhlopid^. 

Maxillary bordering mouth, forming a suture with 
premaxillary, prefrontal, and frontal, toothless ; 
pubis and ischium present, latter forming a sym- 
physis 2. Glauconiid^e. 

II. Transverse bone present ; both jaws toothed. 

A . Coronoid present ; nasals in contact with pre- 
frontals, 
i. Vestiges of pelvis ; supratemporal present. 
Supratemporal large, suspending quadrate 

3. BoiDiE. 

(Subfamilies : Pythonincz, Boince.) 
Supratemporal small, intercalated in the cranial wall 

4. Ilysiidjs. 



DEFINITION AND CLASSIFICATION 5 

2. No vestiges of pelvis ; supratemporal absent 

5. Uropeltid,e. 

B. Coronoid absent ; supratemporal present. 

1. Maxillary horizontal ; pterygoids reaching 

quadrate or mandible. 

Nasals in contact with prefrontals 

6. Xenopeltid.e. 
N asals not in contact with prefrontals 

7. COLUBRID.E. 

Three series: A. Aglypha (subfamilies: Acrochor- 
dince, Colubrince, Dasypeltince) ; B. Opisthoglypha 
(Homalopsincz, Dipsadomorphincz, Elachistodontince) ; 

C. Proteroglypha {Hydrophiince, Elapincz). 

2. Maxillary horizontal, converging posteriorly 

towards palatine ; pterygoid not reaching 
quadrate or mandible 8. Amblycephalid.e. 

3. Maxillary vertically erectile perpendicularly to 

transverse bone; pterygoid reaching quad- 
rate or mandible 9. Viperid^e. 

(Subfamilies : Viperince, Crotalince.) 

The technical terms employed in the above 
synopsis will be found explained and illustrated by 
figures in the chapter on the Skeleton. 

No serial arrangement can express the affinities 
of the various groups as conceived by the classifi- 
cator ; a diagram therefore follows to show the 
author's views as to their interrelationships, and 
possibly their phylogeny. Leaving aside the 
Typhlopidae and Glauconiidae, which should be re- 



6 INTRODUCTION 

garded as burrowing types independently derived 
from some Ophidian form less specialized than any 
with which we are at present acquainted, and prob- 
ably without direct relationship to the Lizards, the 
family Boidae, and more especially the Pythons, 
claim the position of ancestral group, from which 
all other snakes may have been derived. 

Viperidas Amblycephalidae 

I 
Colubridae opisthoglyphae Colubridae proteroglyphae 



Uropeltidas 

I 
Ilysiidas Xenopeltidae Colubridce aglyphae 

I ! l 

I 

Boidae 

Further remarks on this subject in the chapter 
on Dentition. 

It is to be regretted that paleontology cannot 
help us at present as concerns the lines of evolution, 
the comparatively few fossil Ophidians known, from 
the Lower Eocene upwards, the remains of which 
can be identified with some measure of certainty, 
being either non-poisonous types (Boidce, Ilysiidce, 
Palceophiidce, Colubvidce) or Viperidce (Viperines from 
the Miocene of France and Germany, Crotalines from 
the Miocene of North America). The vertebrae from 
the Puerco Eocene of America, on the limit between 
the Cretaceous and Eocene periods, described as the 



DEFINITION AND CLASSIFICATION 7 

oldest snake remains, Helagras, Cope, are stated to 
approach the Lacertilian type. 

Whether the vertebrae named Symoliophis, 
Sauvage, from the chalk of France, and Coniophis, 
Marsh, from the Laramie Cretaceous of North 
America, are Ophidian, as claimed by their descri- 
bes, or Dolichosaurian, cannot be decided without 
further material. 



CHAPTER II 

EXTERNAL CHARACTERS— INTEGUMENT 

THE form varies enormously, worm-like in some, 
comparatively short and heavy, elongate and 
more or less slender, or extremely gracile and almost 
filiform, in others. In this respect our common Grass- 
snake occupies a central position, and for this reason 
is termed a moderately slender form, anything above 
or below this standard being described as com- 
paratively short or elongate. Our shortest and 
stoutest European Snakes are the Vipers, especially 
Vipera ursinii ; our longest and slenderest, the Coluber 
and Zamenis, especially Zamenis dahlii. These 
extremes in both directions are, however, far sur- 
passed by many exotic snakes, as we find on 
comparing, for instance, one of the African Puff- 
adders (Bitis), with certain Oxybelis and Leptognathus 
from Tropical America. The body may be some- 
what rigid, as in some burrowing and ground snakes, 
not unlike in appearance to our Slow- worm and other 
limbless Lizards; or extremely flexible, as in many 
Pythons and Boas and in the Tree-snakes generally. 
This flexibility may be accompanied by a vertical 
compression of the body in relation with an arboreal 

existence, whilst sluggish snakes, such as most of the 

8 



EXTERNAL CHARACTERS 9 

Viperidse, may be remarkable for the flattening of 
the body, which they may further increase when 
basking in the sun or in order to assume a more 
formidable appearance on the approach of an 
enemy. This power of flattening out the whole or 
the anterior part of the body is possessed by many 
snakes, poisonous as well as harmless, and reaches 
its highest degree in the Cobras of India and Africa, 
the expanded anterior part being known as the 
" hood," from the Portuguese name " Cobra di 
capello." 

Thoroughly aquatic snakes are often short and 
heavy, but some of the marine forms, or Hydrophids, 
may be extremely slender, with the posterior part of 
the body compressed. In some of these Sea-snakes 
the gracility of the anterior part, or "neck," as it 
has been called, contrasts very strikingly with the 
great girth of the body towards the tail, and sug- 
gests a limbless Plesiosaur. 

The tail, the part of the body behind the trans- 
versely cleft vent, is most frequently about one- 
fourth or one-fifth of the total length ; but it may be 
much shorter, even reduced to a mere stump, as in 
the Typhlops, or, at the opposite extreme, enter for 
one half in the length of the snake, as in the 
African Xenurophis. This organ may taper gradually 
to a fine point ; or end abruptly, as if mutilated ; or 
terminate in a horny spine, such as we see in some 
of the Typhlops or in the Australian Death-adder, 



io INTRODUCTION 

Acanthophis, or in a series of horny segments which 
are vibrated like a rattle, as in the well-known 
Crotalus of America, to which we shall refer again at 
the end of this chapter. In some of the burrowing 
Uropeltidae, the very short tail is obliquely truncated, 
with indurated shields above, and acts as a trowel. 
And, finally, the marine snakes of the subfamily 
Hydrophiinae are distinguished by a strongly com- 
pressed, oar - shaped tail, with rounded vertical 
outline. In a few forms, arboreal or aquatic, the 
tail is more or less prehensile. 

Males generally have a longer tail than females, 
and the genital organs, which are lodged in its base, 
cause a swelling of that region which contrasts with 
the more gradually tapering extremity of the female, 
thus affording a means of distinguishing the sexes 
externally in the majority of snakes. 

The rudimentary hind limbs of Boid snakes, to be 
mentioned further on in the description of the 
skeleton, terminate in a claw-like horny spur, which 
appears on each side of the vent in the male, and 
sometimes also, though less distinctly, in the 
female. These spurs are probably of use in facilita- 
ting the pairing, an explanation which appears the 
more plausible from the fact that the snakes pro- 
vided with them have the copulatory intromittent 
organs destitute of the erectile spines which are 
present in most others. 

The head varies in shape as much as the body. 



EXTERNAL CHARACTERS n 

Although never actually compressed, except in the 
rostral region, it may be very narrow and elongate, 
whilst in the opposite extreme it may be strongly 
depressed, and so broad behind as to be abruptly 
defined from the anterior part of the body, or 
" neck." This feature is very marked in some of 
the Viperidae, and this has given rise to the incorrect 
generalization that poisonous snakes are distin- 
guished from the harmless by a broad and flat 
head, notwithstanding the fact that some of the 
most dangerous, such as the Mambas, Cobras, and 
Kraits, have a comparatively narrow or small head, 
not or but slightly defined behind, whilst, on the 
other hand, the very opposite condition obtains in 
not a few of the harmless Colubrids. 

Leaving the Typhlopidae and Glauconiidse aside 
for the present, snakes have a wide gape, cleft far 
beyond the vertical of the eyes, with, when closed, 
one or two notches in front for the passage of the 
protrusible, bifid tongue. In most snakes this chink 
is in the lower border of the rostral shield, capping 
the tip of the snout, and allows free passage to the 
whole tongue; in the Hydrophids, or Sea-snakes, 
there are two notches in the lower border of the 
rostral shield, through which only the bifid end 
of the tongue can be protruded. The eyes, varying 
from minute to enormous, are usually free from the 
surrounding shields, and may move under a trans- 
parent cap like a watch-glass, which appears to 



12 INTRODUCTION 

represent the lower eyelid of Lizards. The view as 
to this homology is derived from our knowledge of 
various conditions in certain series of Lizards of the 
families Lacertidae and Scincidae, where we find a 
transparent disc appearing like a small window in 
the movable lower eyelid, gradually increasing in 
size so as to occupy the whole of the lower eyelid, 
which finally becomes fused with the rudimentary 
upper lid and loses its mobility. In Ilysia and in 
most of the Uropeltidas, the transparent disc over 
the eye is confluent with a thick horny shield of 
which it occupies the middle. 

The pupil is usually circular or vertical, rarely 
horizontal. In some forms it is difficult to decide 
whether it is round or vertically elliptic ; in others, 
like the Boas and Vipers, for instance, it is decidedly 
vertical, and contracts to the same extent as a cat's. 
In some Water-snakes, and in Sea-snakes generally, 
the round pupil may contract to a mere dot. The con- 
traction of the pupil is independent on the two sides. 

The snout, or the part of the head anterior to the 
eyes, may be short or long, rounded or pointed, de- 
pressed or compressed, sometimes projecting strongly 
beyond the mouth, turned up at the end, or ter- 
minating in one (Langaha) or two (Herpcton) long 
scaly dermal appendages. In some burrowing forms 
it is provided with a more or less trenchant hori- 
zontal or vertical edge. When the sides of the snout 
(loreal region) form an angle with the upper surface, 



EXTERNAL CHARACTERS 13 

the angle is termed the "canthus rostralis," which 
may be intensified by the loreal region being concave. 

The deep pits which are sometimes present on 
the lips or between the nostril and the eye (loreal 
pit) will be alluded to further on under Sensory 
Organs. 

The nostrils are either lateral, or, in the aquatic 
forms, directed upwards, sometimes entirely on the 
upper surface of the snout. 

Most snakes have a longitudinal groove on the chin 
(mental groove) to allow for the distension caused by 
the lateral movements of the rami of the lower jaw. 

In the Typhlopidse, the head passes gradually into 
the vermiform body, and the small mouth is situated 
on the under surface of the projecting snout ; the 
head so resembles the extremely short tail, and the 
mouth is so similar in shape and position to the vent, 
which is close to the posterior extremity of the snake, 
that such creatures are often believed by non-critical 
observers to have a head at each end. The eyes are 
very small, and covered over by the semi-transparent 
head-shields, or they may be completely concealed. 
There is no mental groove. It is much the same 
with the Glauconiidae, which have, however, a some- 
what less abbreviated tail. In both, the nostrils 
often open on the lower side of the snout, which may 
be excavated so as to appear hooked in profile, or 
may be provided with a sharp cutting horizontal 
edge. 



14 INTRODUCTION 

Snakes are covered with epidermal folds in the form 
of scales and shields, the shape and arrangement of 
which affords important characters for their classifica- 
tion. Dermal ossifications are absent. 

The scales on the body are usually elliptic or 
lanceolate and imbricate, forming straight longitudinal 
and oblique transverse series, and they are replaced 
on the belly and under the tail by transverse shields 
mostly corresponding in number with the series of 
scales, and also with the vertebrae. The body of the 
Typhlopidae and Glauconiidse is uniformly covered 
with polished, closely adherent, rounded, overlapping, 
sub-equal scales, without even an indication of ven- 
tral shields. Jn some of the Acrochordinae, aberrant 
aquatic Colubrids, the scaling consists, above and 
beneath, of small juxtaposed, sometimes spinose 
granules, the skin being suggestive of the shagreen of 
sharks. In the marine snakes of the subfamily 
Hydrophiinae, the ventral shields are often absent or 
merely indicated, and the scales are mostly juxtaposed 
or feebly imbricate, sometimes tetragonal or hexag- 
onal, and occasionally studded with spinose tubercles. 
In the more typical Ophidia the imbricate scales may 
be long and narrow or short and broad, with every 
intermediate step between the two extremes; smooth 
or furnished with a longitudinal ridge or keel, or even 
several keels ; nearly equal in size or with the median 
or outer series more or less enlarged, the longitudinal 
series in odd, rarely in even number ; instead of run- 



INTEGUMENT 15 

ning in longitudinal series parallel with the axis of 
the body, as is the rule, they are sometimes disposed 
obliquely, and among those in which we meet with 
this peculiarity several genera are further remarkable 
in having some of the oblique lateral scales furnished 
with a serrated keel, to which we shall again allude 
in the chapter on Habits, when dealing with the 
rustling sounds produced by certain snakes. The 
number of longitudinal series of scales on the body 
varies from 10 (Herpetodryas) to nearly 100 {Python, 
Boa) ; in the European species from 17 (Contia 
modesta) to 50 {Eryx jacuhis). The scales are some- 
times furnished near the end with one or two shallow 
impressions, termed "apical pits," which afford indica- 
tions for the distinction of genera and species ; unless 
of a lighter or darker colour, as is often the case, 
these pits are not always easy to see, except in a 
strong light and with the aid of a powerful magni- 
fying glass. 

The ventral shields, also called " gastrosteges," 
usually occupy the whole width of the belly ; but 
they may be much narrower — in Eryx, for instance. 
They are sometimes bent at an angle on the sides, and 
this angle may even form a sharp keel, accompanied 
by a notch in the posterior border, corresponding to 
the keel, as in several of the more arboreal genera of 
Colubrids. The shields under the tail, termed sub- 
caudals or "urosteges," are sometimes similar to the 
ventrals, but more often disposed in pairs ; in certain 



16 INTRODUCTION 

species or individuals some of the subcaudals are 
single, and the others paired. When the number of 
subcaudals is given in the descriptions, each pair is 
reckoned as one, and the conical or spine-like shield 
which caps the end of the tail is not included. These 
numbers afford important characters for the definition 
of species, and sometimes also for the distinction of 
sexes. The subcaudals are nearly always much fewer 
than the ventrals, but the difference is often not so 
great in the males as in the females, the tail of 
which is usually shorter in proportion to the body. It 
is noteworthy that in many species, if the number of 
subcaudals (C.) be added to that of the ventrals (V.), 
the total is nearly the same in the male as in the 
female, however much the respective numbers may 
differ when taken separately. The following figures 
may be given by way of example, taken from British 
specimens : 

Coronella austriaca : S V. 154; C. 58=212 

? V. 165; C. 48 = 213 

Vipera herns: $ V. 138; C. 35 = 173 

? V. 144; C. 29=173 



>» f> 



Although this rule is by no means universal, and 
does not apply at all to some species, it will be found 
to hold good in many cases, and is of interest in 
showing that the changes that have taken place in 
the vertebral column (the vertebrae corresponding in 
number to the shields), according to the sexes, have 



INTEGUMENT 17 

been by a modification of the character of the seg- 
ments about the anal region, a conversion of trunk 
vertebrae into caudals, or vice versa. In dealing with 
certain species — of Vipers, for instance — it is impor- 
tant, for systematic purposes, to keep the counts of 
shields distinct for the two sexes. 

The shield which covers the vent, the anal shield, 
is either single or divided into two. 

Some snakes have the head covered with scales or 
small tubercles similar to those on the body, but in 
the great majority the lepidosis is in the form of 
large symmetrical juxtaposed shields, the shape, pro- 
portions, and number of which furnish some of the 
most important characters for the distinction of 
genera and species. These head-shields belong to 
two primarily different types, from each of which all 
further modifications may be regarded as derived by 
alteration in shape or by disintegration. The first 
type is that shown by the Typhlopidae and Glau- 
coniidas, which is explained by the figure on the 
next page. 

The rostral, which is usually the largest of the head- 
shields, extends to the upper surface of the head, of 
which it may occupy the greater part. In the 
Glauconiidae, the ocular usually borders the mouth. 

As may be seen by a comparison of the first figure 

with the second, the arrangement of the head-shields 

is essentially different from that which prevails in the 

Colubrids and the majority of other snakes. 
2 



i8 



INTRODUCTION 



The second type is exemplified by the head of a 
member of the genus Zamenis. 

In the descriptions, temporals 2 + 3 means two 
superposed temporals in the first row, three in the 
second. The internasals and the temporals, and the 
loreal and the preocular, are sometimes absent, and 
the prefrontal or the internasal may be single. One 
or two large shields are in rare cases present behind 
the parietals, and are called occipital. 

A breaking up into smaller shields takes place in 
many snakes. In the Pythons, for instance, the 








Fig. 1 — Head of Typhlops braminus. (From "Fauna of 

British India") 

/, Frontal ; ip, interparietal ; I, labial ; n, nasal ; 0, ocular ; 
p, parietal ; po, preocular ; prf, prefrontal ; r, rostral ; so, supra- 
ocular. 

frontal may be divided into two by a longitudinal 
cleft, and separated from the prefrontals by small 
shields. In some Vipers, such as V. berus and 
V. ursinii, in which the frontal and parietals, though 
reduced in size, usually preserve their primitive 
condition, the former is normally separated from the 
supraocular by a series of small shields, and the 
internasals and prefrontals are broken up ; in these 



INTEGUMENT 



19 



snakes the small shield or shields behind the 
rostral are termed " apical," and those on the upper 
edge of the snout are termed " canthals." The shield 
which, in Vipers, separates the rostral from the nasal is 




Fig. 2— Head of Zamenis ventrimaculatus. (From " Fauna of 

British India") 

cs, Chin-shields (anterior) ; cs', chin-shields (posterior) ; /, frontal ; 
in, internasal ; I, loreal ; la, labial (upper) ; la', labial (lower) ; 
m, mental ; n, nasal ; p, parietal ; pf, prefrontal ; pro, preocular ; 
pto, postocular ; v, rostral; sbo, subocular ; so, supraocular; 
t, temporals (first row) ; t' , temporals (second row) ; v> first 
ventral. 

called " naso-rostral." Allusion has been made above 
to the scaly dermal appendages which terminate the 
snout in certain genera. Some Viperidae are furnished 



20 INTRODUCTION 

with horn-like erect spines above the eyes or at the 
end of the snout, which add greatly to their sinistral 
appearance. 

The periodical shedding of the outer layer of the 
epidermis in a single piece, including even the 
covering of the eye, is one of the most striking 
peculiarities of snakes, although paralleled in the 
Lizards of the family Anguidae, to which our 
British Slow-worm belongs. The skin becomes 
detached at the lips, and is turned inside out 
from head to tail, without any sort of laceration 
when the snake is in good health. These exuviae are 
transparent, but often carry a certain amount of 
pigment, especially those of the Vipers, in which the 
characteristic dark markings are perfectly visible ; 
they usually exceed the length of the reptile, owing 
to stretching. In Sea-snakes the epidermis is cast 
piecemeal, and sloughing is a longer operation than 
in ordinary snakes. 

In Rattlesnakes each piece of the rattle, or 
" crotalon," in which the tail terminates, represents 
a retained portion of the sloughed epidermis. This 
remarkable appendage looks like a number of horny 
rings, but it consists in reality of hollow, bell-like 
pieces, similar to the terminal one, or "button," each 
with a circular constriction, in which the incurved 
free edge of the following piece fits, thus keeping the 
pieces together without impairing the mobility 
necessary to produce the rattling sound for which 



INTEGUMENT 21 

the apparatus is intended. At each exuviation one 
bell-shaped horny piece is added. The number of 
segments in the rattle is, therefore, not an index to 
age, as formerly believed ; nor is it to the number of 
exuviations, for whilst segments are being added at 
the base of the apparatus the terminal ones break 
off and are lost. A Crotalus sixteen months old may 
have six pieces to the rattle if there have been six 
exuviations and no loss. No rattle appears ever to 
comprise more than about twenty pieces, even in old 
specimens. The size of the terminal button shows 
whether it was formed at birth or at any later period, 
no growth taking place in the horny tissue. 

So far as trustworthy records are concerned, the 
largest snakes known, the Malay Python reticulatus 
and the South American Anaconda, Eunectes murium, 
reach a length of 25 to 30 feet. Measurements of 
skins must be accepted with caution, as a skin may 
easily be stretched to once and a half its real length ; 
in estimating the exact length from such a stretched 
skin, it is necessary to deduct the interstitial spaces 
showing between the scales, and about one-fourth of 
the scale to allow for the overlap. The smallest 
snake known is 4 inches long (Glauconia dissimilis) . 
The largest European snake (Coluber quatuorlineatus) 
is reported to reach a length of 8 feet ; the smallest 
(Typhlops vermicularis) does not exceed 14 inches. 



22 INTRODUCTION 

Key to the Identification of the European 
Snakes from External Characters only 

I. Eyes minute, under the head-shields; mouth 
small, inferior ; body vermiform, covered with 
uniform scales above and beneath ; vent close to 
the end of the body, ■ the extremely short tail 
ending in a small spine Typhlops vermicular is. 

II. Eyes very small, with vertical pupil; upper 
surface of head covered with small scales ; 
ventral shields much narrower than the body ; 
tail short, ending obtusely ; subcaudals single, or 
mostly single ; scales smooth or feebly keeled, in 
40 to 50 rows Eryx jaculus. 

III. Eyes small, moderate, or large; ventral shields 
at least nearly as broad as the body ; tail taper- 
ing to a point ; subcaudals paired. 
A. Pupil round; upper surface of head with nine 
large shields ; no upper labial in contact with 
the parietal ; anal shield usually divided. 
1. Dorsal scales strongly keeled, with paired 
apical pits ; a single anterior temporal. 
a. Nostrils lateral ; internasals broadly trun- 
cate in front. 
Scales in 19 rows ; normally 1 pre- and 3 post- 
oculars ; usually 7 upper labials, third and fourth 
entering the eye; ventrals 157-181 ; subcaudals 
50-88 Tropidonotus natrix. 



EXTERNAL CHARACTERS 23 

b. Nostrils directed upwards ; internasals 
much narrowed in front. 
Scales in ig rows ; normally 2 pre- and 3 or 4 post- 
oculars ; suboculars sometimes present ; usually 
8 upper labials, fourth or fourth and fifth entering 

the eye; ventrals 160-187 ; subcaudals 48-79 

Tropidonotus tessellatus. 

Scales in 21 (rarely 19 or 23) rows ; normally 1 or 2 

pre- and 2 postoculars ; usually 7 upper labials, 

third and fourth entering the eye ; ventrals 

147-164; subcaudals 46-72. Tropidonotus viperinus. 

2. Dorsal scales smooth or feebly keeled; 

normally a single loreal. 

a. Two or three superposed anterior tem- 
porals (very rarely one) ; nostril usually 
between two nasals. 
a. A subocular below the preocular. 
* Scales smooth, in 17 or 19 rows. 
Two upper labials entering the eye ; preocular not 
in contact with the frontal ; scales with two apical 
pits ; ventrals more or less distinctly angulate 

laterally, 160-230; subcaudals 87-131 

Zamenis gemonensis. 
Two upper labials entering the eye ; preocular usually 
in contact with the frontal ; scales with a single 
apical pit ; ventrals very distinctly angulate later- 
ally, 205-218 ; subcaudals 98-132. Zamenis dahlii. 
** Scales in 23 to 29 rows (usually 25 or 
27), with two apical pits. 



24 INTRODUCTIOxN 

Upper labials usually separated from the eye by a 
series of suboculars; preocular in contact with 
the frontal ; scales smooth ; ventrals very dis- 
tinctly angulate laterally, 222-258 ; subcaudals 

77-107 Zamenis hippocvepis. 

Two upper labials entering the eye; preocular not 
in contact with the frontal ; scales feebly but dis- 
tinctly keeled; ventrals not angulate laterally, 
195-234; subcaudals 56-90. Coluber quatuorlineatus. 
Two upper labials entering the eye ; preocular not 
in contact with the frontal ; scales smooth or 
faintly keeled ; ventrals not or but very obtusely 

angulate laterally, 172-214; subcaudals 50-80 

Coluber dione. 
ft. No subocular ; scales smooth, or faintly 
keeled on the posterior part of the body. 
* Ventrals more than 200 ; scales with 
two apical pits. 
Snout obtuse; rostral broader than deep; scales in 
21 or 23 rows; ventrals distinctly angulate later- 
ally, 212-248 ; subcaudals 60-91 

Coluber longissimus. 
Snout obtuse ; rostral broader than deep ; scales in 
25 or 27 rows ; ventrals not angulate laterally, 
222-260; subcaudals 68-90 ... Coluber leopardinus. 
Snout pointed, strongly projecting ; rostral deeper 
than broad, wedged in between the internasals ; 
scales in 25 to 29 rows ; ventrals not angulate later- 
ally, 201-220 ; subcaudals 48-68. Coluber scalaris. 



EXTERNAL CHARACTERS 25 

*■* Ventrals not more than 200 ; scales 

mostly with a single apical pit. 

Rostral at least as deep as broad, often wedged in 

between the internasals ; usually 7 upper labials, 

third and fourth entering the eye ; scales in 19 

(rarely 21) rows; ventrals 153-199; subcaudals 

41-70 Coronclla austriaca. 

Rostral broader than deep ; usually 8 upper labials, 
fourth and fifth entering the eye; scales in 21 
(rarely 19 or 23) rows ; ventrals 170-200 ; sub- 
caudals 49-72 Coronclla girondica. 

b. A single anterior temporal ; nostril in a 
single nasal ; scales smooth, with single 
apical pits, in 17 rows; ventrals 150-191 ; 

subcaudals 53-78 Contia modesta. 

3. Scales longitudinally grooved in the adult, in 
17 or 19 rows ; two loreals ; canthus rostralis 
strongly marked; frontal very narrow, in 
contact with the preocular; ventrals 160-189 ; 
subcaudals 68-102 ... Caelopeltismonspessulana. 
B. Pupil vertical or vertically subelliptic (some- 
times appearing round in Macroprotodon). 
1. Scales smooth, mostly with single apical pits ; 
upper surface of head with nine large shields. 
Frontal 1^ to 2 times as long as broad ; loreal sepa- 
rated from the eye by the preocular ; one upper labial 
usually in contact with the parietal ; scales in 19 to 
23 (rarely 25) rows; ventrals 153-192; anal divided; 
subcaudals 40-54 Macroprotodon cucullatus. 



26 INTRODUCTION 

Frontal i\ to ij times as long as broad, much 
shorter than the parietals ; loreal entering the 
eye; scales oblique, in 19 or 21 rows; ventrals 

186-222; anal divided ; subcaudals 48-73 

Tarbophis fallax. 

Frontal ij to i| times as long as broad, nearly 

as long as the parietals ; loreal entering the eye ; 

scales oblique, in 19 or 21 rows ; ventrals 203-235 ; 

anal entire ; subcaudals 54-70 ... Tarbophis iberus. 

2. Scales keeled, with two apical pits ; anal 

shield entire. 

a. No pit between the nostril and the eye ; 
upper head-shields small, if present ; nasal 
separated from the rostral by a naso- 
rostral ; eye separated from the upper 
labials by suboculars. 

a. Snout not turned up at the end ; supra- 
ocular usually extending posteriorly 
beyond the vertical of the posterior border 
of the eye; frontal and parietal shields 
usually well developed ; usually a single 
series of scales between the eye and the 
upper labials. 
Snout obtusely pointed, flat above, or with the 
canthus slightly raised ; rostral usually in contact 
with a single apical shield, rarely with two ; 6 to 9 
upper labials, usually 7 or 8 ; scales in 19 rows, 

rarely 21 ; ventrals: S 120-135, ? 125-142 

Vipcva ursinii. 



EXTERNAL CHARACTERS 27 

Snout pointed, with raised canthus ; rostral in con- 
tact with a single apical shield ; 8 or 9 upper 
labials; scales in 21 rows, rarely 19; ventrals : 

c? 130-148, ? 130-150 Vipera renardi. 

Snout truncate or broadly rounded, flat above or 
with slightly raised canthus ; rostral in contact 
with two apical shields, rarely with one ; 8 or 9 
upper labials; scales in 21 rows, rarely 19 or 23; 

ventrals: c? 132-150, $ 132-158 Vipera bems. 

/3. Snout usually more or less turned up at 
the end or produced into a scaly dermal 
appendage ; supraocular not extending 
posteriorly beyond the vertical of the pos- 
terior border of the eye ; frontal and 
parietals often absent or very small ; 
2 or 3 series of scales between the eye and 
the upper labials ; 9 to 13 upper labials ; 
scales in 21 or 23 rows, rarely 19 or 25. 
Snout simply turned up, the raised portion bearing 
2 or 3 scales ; rostral not more than once and a half 
as deep as broad; ventrals: 0*134-158, $ 141-169. 

Vipera aspis. 
Snout simply turned up or produced into a small 
appendage, the raised portion with 5 or 6 (rarely 3) 
scales ; rostral i\ to 2 times as deep as broad ; 
ventrals: o* I 25-i46, ? 135-147 ... Vipera I atastii. 
Snout produced into an appendage covered with 
10 to 20 scales ; rostral not reaching the summit 
of the rostral appendage; ventrals: S 133-161, 
? 135-163 Vipera ammodytes. 



28 INTRODUCTION 

7. Snout not turned up at the end ; supra- 
ocular narrow or broken up into several 
small shields ; upper surface of head 
with small, usually keeled scales ; two or 
three series of scales between the eye and 
the upper labials ; scales in 23 to 27 rows, 
usually 25 ; ventrals : S 151-177, ? 153- 

180 Vipera lebetina. 

b. A pit between the nostril and the eye ; 
upper surface of head with 9 large shields ; 
nasal in contact with the rostral ; third 
upper labial entering the eye; scales in 
23 rows ; ventrals 149-174 ; subcaudals 
3 1-44 A ncistrodon halys. 



CHAPTER III 
COLORATION 

IN dealing with the coloration, we have first to 
distinguish between the colour and the markings. 
The former is very often highly variable among 
snakes of the same species, to say nothing of the 
changes which may take place with age or with 
the condition of the individuals, whether before or 
after exuviation ; it is not unusual to find among 
specimens from the same locality a great range of 
variation, from greyish-white to brown, or red, or 
black, as, for instance, in our Common Viper. The 
latter afford more important characters, and often 
furnish valuable indications for the distinction of 
species ; but even the disposition of the markings is 
subject to great individual variations, more likely to 
mislead than to help the inexperienced student in 
the discrimination of species. It is therefore always 
advisable to resort in the first instance to structural 
characters for the purpose of specific identification, 
and to fall back on coloration only as a means of 
confirmation. If we were to be guided by colour 
and markings alone, how could we believe that an 

adult four-lined Coluber qaatuorlineatus is of the 

29 



3 o INTRODUCTION 

same species as the handsomely spotted Coluber 
sauromates ; and yet, if we compare the young of 
these two snakes we find them to be absolutely 
identical in their markings, and, in the absence of 
any structural differences, we are forced to conclude 
that they only represent two forms of the same 
species, of which the latter is the more primitive. 

It is nevertheless a fact that, with a few excep- 
tions, the markings, however variable they may be, 
are reducible to certain fundamental patterns to 
which the innumerable variations may be traced 
back, and their derivation followed and scientifically 
explained. Let us consider, for instance, another 
species of Coluber, highly variable in its markings : 
C. leopardinus, of which the typical form, so called 
from having been the first described and named, is 
not by any means to be regarded as the most 
primitive. 

First, we must take for granted that the markings 
of all such snakes, whether consisting of spots, 
stripes, or bars, start from a regular arrangement, 
which may be theoretically represented by four 
paired longitudinal series on the head and body : 
(i) Dorsal series (D) ; (2) Dorso-lateral (DL) ; (3) 
Lateral (L) ; (4) Ventro-lateral (VL). The first 
starts from the middle line of the head, and is con- 
tinued along the spine ; the second occupies the 
space between the first and third, which originates 
at the tip of the snout, passes through the eye, and 



COLORATION 31 

is continued on the temple and along the side of the 
body; the fourth follows the lower lip, and extends 
along each side of the belly. Bearing this in mind, 
we find that the variety of C. leopardinus named 
schwoederi, with a vertebral series of paired spots, is 
to be regarded as the most primitive, from which we 
can derive, on the one hand, the true leopardinus by 
imagining a transverse fusion of the spots of series D 
into a single row, some of the spots often actually 
revealing, in their biscuit shape, their dual origin ; 
whilst, on the other hand, confluence of the paired 
spots of the same series into two longitudinal stripes 
produces the variety named quadrilineatus (see 
Plate VII.). In this particular instance, the paired 
series D has fused into a single streak on the head, 
and the series L appears to have departed from its 
primitive course to extend on the upper surface of 
the head, both in front of and behind the eye. 

Many snakes show an interocular band extending 
from lip to lip, through the eyes, across the inter- 
orbital region. In others the lateral stripe L may 
bifurcate in front of the eye, an upper branch 
extending across the snout, through transverse 
fusion of series D and DL, and it may also bifurcate 
in like manner on the temporal region, fusing with 
the corresponding marking on the other side to form 
a W-shaped figure. The pattern of markings on the 
upper surface of the head is, however, often very 
complicated, and hence difficult of explanation. 



32 INTRODUCTION 

As a second example of the derivation of patterns, 
we may mention Viper a aspis, which varies enor- 
mously as to its mid-dorsal markings, forming, in 
different individuals or even on different parts of the 
body, single or paired spots, a zigzag band, or trans- 
verse bars ; all these are derived from the paired 
spots of series D. Each pair of spots may fuse and 
form transversely oval or elliptical spots or bars, or 
the spots may assume an alternate disposition from 
which, through confluence, the zigzag or sinuous 
band results. Thus, spotted and striped patterns 
may be traced to a common origin, however 
fundamental the difference between them appears at 
first sight. If the elements of the four series, D, 
DL, L, and VL, unite transversely with each other, 
and also with the spots on the ventral surface, we 
obtain ringed forms such as the Coral-snakes. 
That the black nuchal collar of our common Grass- 
snake is actually formed by the fusion of the spots 
of three originally distinct series has been proved by 
tracing the development of the markings in the 
embryo. 

In various species a pair of light streaks extends 
along the back, bordering the D area, without inter- 
fering with the other markings, as we see, among 
European snakes, in some specimens of Tropidonotus 
natrix and viperinus, and Vipera bents. 

Although it sometimes happens that a definite 
system of markings prevails throughout a genus, 



COLORATION 33 

such as the annulate form in the South American 
Elaps, this is far from being universally the case ; 
many closely allied species, or individuals of the 
same species, may be distinguished by very different 
patterns. Even on the same individual we may find 
two opposite types of markings without any transi- 
tion, as in two Central American species of widely 
different genera, Polyodontophis annulatns and Zamenis 
mexicanus, in which the anterior part of the body is 
annulate or barred, and the rest longitudinally 
striped. 

It is also a remarkable fact that very often the 
two sides of the body are not alike in their markings, 
appearing as if formed of the union, on the median 
line, of the right and left halves of two individuals. 
Thus it may happen, in annulate forms, that some of 
the annuli are broken exactly in the mid-dorsal and 
mid-ventral lines, and that the halves do not corre- 
spond in number on the two sides. In the hand- 
some South American Lachesis altematus, which 
derives its specific name from the two series of 
large C-shaped, dark, light-edged markings which 
adorn its back, these markings are not always alter- 
nating, as is the rule ; but some may lie opposite 
to each other and back to back, this being due to 
the fact that the numbers of the markings do not 
correspond on the two sides. In one specimen I 
count twenty-four of these markings on the left side, 
and twenty-seven on the right. This shows tnat 
3 



34 INTRODUCTION 

great importance cannot be attached to the number 
of the markings, for systematic purposes. In fact, 
in some Coral-snakes, Elaps fulvius for instance, the 
number of annuli may vary from twelve to fifty-two, 
with every gradation between the extremes. The 
bilateral asymmetry to which we have alluded pro- 
duces the chess-board arrangement of the ventral 
spots in many snakes. 

Among the markings which call for investigation 
as to their meaning, we must allude to the presence, 
in some Colubrids, of a small, light, dark-edged spot, 
or of a pair of light dots close together, in the 
middle of the parietal shields or on each side of the 
suture between these shields, which correspond in 
their position to the parietal organ of many Lizards. 
May not this marking be in some way correlated 
with sensory organs, like the apical pits on the scales 
of the body ? And what is the explanation of such 
bizarre signs as the spectacle or the eye-spot on the 
hood of the Indian Cobra ? At present it is as 
inexplicable as the lugubrious emblem on the thorax 
of the Death's-head Moth. It cannot be suggested 
that it is a warning mark intended to terrify intruders, 
for when the Cobra is at rest the hood is folded, and 
the characteristic marking is not displayed ; whilst 
as soon as it is aroused, and the hood expanded, it 
faces its enemy in such a way that the spectacle, or 
ocellus, is not to be seen. 

First among the most brilliantly coloured snakes, 



COLORATION 35 

of which there are many, stand the Coral-snakes, 
Elaps, of America, mostly annulate with red, yellow 
or white, and black. This striking coloration obtains 
also in diverse harmless snakes inhabiting the same 
part of the world, and this coincidence has been 
adduced in favour of the theory of mimicry, corre- 
lated with that of natural selection, which accounts 
for the resemblance as being of advantage to a 
harmless species, which is thus mistaken for one 
notorious for its deadly poison, and advertised as 
such by its brilliant colours (warning coloration). 
But other poisonous and much more dangerous 
snakes are not, as a rule, endowed with brilliant 
colours. It is true that these also may have their 
mimics : the Krait, Bungarus cceruleus, and Lycodon 
aulicus, in India, the Pit-viper, Ancistrodon hima- 
layanus, and Psammodynastes pulverulentus, in the 
Himalayas and Assam, are good examples of such 
cases. On the other hand, there are equally striking 
instances of what one would regard as mimics if 
they only occurred together; thus, there is no better 
case of general resemblance between a poisonous 
and a harmless snake than we find in the Indian 
Cobra and the Coluber corah of tropical America, 
where Cobras are absent, or between a Viper and 
the Boid Enygrus asper, from New Guinea, where 
no Vipers exist. 

Without attempting to offer any suggestion to 
account for the similarity of markings which prevails 



36 INTRODUCTION 

in certain parts of the world, attention may be drawn 
to the predominance of longitudinal dark and light 
stripes in the Indo-Malayan representatives of the 
American Elaps, shared by many innocuous snakes 
of similar form inhabiting the same region, and 
to the striped tails common to various Colubrids 
of Madagascar, as if the snakes of a district had 
agreed to conform to certain fashions in dress. 

It is further noteworthy, in relation to the theory of 
warning coloration, that many Uropeltids, innocent 
burrowing creatures living underground or concealed 
under stones or rotting tree-trunks in the forests of 
Southern India and Ceylon, hardly ever showing 
themselves in daylight, are among the most striking 
for their bright yellow or red and black markings. 
We may point out at the same time the very marked 
resemblance in form and coloration between the 
Uropeltid Melanophidium bilineatum, and the Apodal 
Batrachian Ichthyophis glutinosus, both occurring 
together in Southern India. 

The colour of snakes often harmonizes with their 
surroundings. Thus, many Tree-snakes, Boid, Colu- 
brid, or Viperid, are of a bright green, like the 
foliage in which they are concealed. On the other 
hand, other Tree-snakes are not green, or only some 
specimens are green, as in the genera Dendraspis and 
Dispholidus. Desert-snakes are of the yellowish or 
reddish colour of the sand or rock on which they 
live, and in species whose range extends over different 



COLORATION 37 

districts the desert individuals are paler, without or 
with less distinct markings, as compared to their 
fellows among other surroundings. In addition to 
their markings, some snakes are adorned with a 
metallic iridescent gloss, due to a fine striation of 
the scales. 

The iris is often metallic, gold, bronze, or copper- 
red, and the black streaks of the head sometimes 
extend over it. 

Although, unlike many lizards, snakes are unable 
to rapidly alter their colours, some produce a 
semblance of this phenomenon when inflating their 
neck or body ; this is due to the presence of dark 
and light markings or of a bright pigment in the 
interstitial skin, which is not seen when the scales 
overlap. Thus, in the Indian Tree-snake Dryophis 
mycterizans the skin between the green or brown 
scales in the anterior part of the body is black 
and white, producing a striped pattern when the 
neck is inflated; the skin of the same region is 
bright vermilion in the Malay Tropidonotus sub- 
mini atus ; many more examples could be quoted. 
The spectacle marking on the hood of the Indian 
Cobra involves the scales as well as the interstitial 
skin. 

As a rule there are no sexual differences in colour. 
Yet these are so marked in our Common Adder that 
the sex of a specimen can nearly always be recog- 
nized by the coloration. This is, however, the 



38 INTRODUCTION 

exception, even in the genus to which the Adder 
belongs. A nuptial dress is unknown in snakes. 

A special livery for the young is rather exceptional, 
but very often the new-born is more vividly coloured 
than its parents, and in many black varieties the 
young is similar to the typical form. Some green 
Tree-Boids (Chondropython and Cor alius caninus) are 
not green, but yellowish, cream-colour, or pinkish, 
when young, the green appearing around the white 
spots, which are the remains of the ground colour, and 
gradually spreading over the whole body. Conversely, 
the young of a variety of the Pit-viper Lachesis 
wagleri, common in the Malay Peninsula, is green, 
and the adult black and yellow. In the young of 
Gvayia ornata, a West African Water-snake, the 
markings of the young are to those of the adult 
like positive and negative in photography, the white 
bars, forked on the sides, which extend across the 
black back of the former being gradually trans- 
formed into black bars on a light ground in the 
latter ; in such a case it is impossible to decide 
whether the dark or the light parts are to be con- 
sidered as the ground colour. 

That the skin of many snakes contains soluble 
colouring matter of a special kind is well known, 
green snakes, such as Dryophis prasinus and Lachesis 
gramineus staining the spirit in which they are pre- 
served. Chemists have not yet paid attention to 
this question, which requires investigation. 



COLORATION 39 

Melanism is frequent in snakes, and sometimes 
affects all individuals in the same locality. It 
seems undesirable to bestow varietal names on such 
aberrations, as is so frequently done by systematists, 
any more than we should in the case of albinos. 
Melanism may be produced in two ways : by an 
extension of the black markings, which invade the 
whole surface, as in the males of Vipera bents; or by 
a general darkening of the ground colour and of the 
markings, as in the females of the same species. In 
the latter case, the markings reappear under certain 
lights or after a prolonged sojourn in spirits. Some- 
times, as in Zamenis gemonensis, the uniform black 
colour appears only as the snake approaches the 
adult condition, the young having the normal 
livery. 

Partial albinism is rare; perfect albinism, charac- 
terized by absence of black pigment in the eye, rarer 
still. Cases have been observed, among European 
species, in Tropidonotiis natrix and tessellatus, in 
Coluber longissimus, and in Coronella austriaca. 



CHAPTER IV 
SKELETON 

THE typical Ophidian skull is characterized by 
a solidly ossified brain-case, with the distinct 
frontals and the united parietals extending downwards 
to the basisphenoid, which is large and produced 
forward into a rostrum extending to the ethmoidal 
region. The nasal region is less completely ossified, 
and the paired nasals are often attached only at their 
base. The occipital condyle is either trilobate and 
formed by the basioccipital and the exoccipitals, or a 
simple knob formed by the basioccipital ; the supra- 
occipital is excluded from the foramen magnum. 
The basioccipital may bear a strong, curved ventral 
process or hypapophysis (in the Vipers). 

The prefrontal is situated, on each side, between 
the frontal and the maxillary, and may or may not 
be in contact with the nasal ; the postfrontal, usually 
present, borders the orbit behind, rarely also above, 
and in the Pythons a supraorbital is intercalated 
between it and the prefrontal. 

The premaxillary is single and small, and as a 
rule connected with the maxillary only by ligament. 
The paired vomer is narrow. The palatine and 

40 



SKELETON 



4i 



^ ra -prf sor f plf epg p pro ste 



F 



<7lt—. 




PP tw 




q' act, see, 



ste 6 a, £ 



Fig. 3 — Skull of Python amethystinus. (From British Museum 

Catalogue of Snakes) 

an, Angular; ar, articular; bo, basioccipital ; bs, basisphenoid ; 
coy, coronoid ; c.a, columella auris (stapes) ; d, dentary ; eo, ex- 
occipital ; epg, ectopterygoid (transverse) ; /, frontal ; m, maxil- 
lary ; n, nasal ; p, parietal ; //, palatine ; pm, premaxillary ; 
prf, prefrontal ; pro, prootic ; pg, pterygoid ; ///, postfrontal ; 
q, quadrate ; so, supraoccipital ; sor, supraorbital ; s/>, splenial ; 
ste, supratemporal ; tic, turbinal ; v, vomer. 



42 INTRODUCTION 

pterygoid are elongate and parallel to the axis 
of the skull, the latter diverging behind and ex- 
tending to the quadrate or to the articular extremity 
of the mandible ; the pterygoid is connected with 
the maxillary by the ectopterygoid or transverse 
bone, which may be very elongate, and the maxillary 
often emits a process towards the palatine, the latter 
bone being usually produced inwards and upwards 
towards the anterior extremity of the basisphenoid. 
The quadrate is usually large and elongate, and 
attached to the cranium through the supratemporal 
(often regarded as the squamosal). In rare cases 
(Miodon, Polemon) the transverse bone is forked, and 
articulates with two branches of the maxilla. The 
quadrate and the maxillary and palatopterygoid 
arches are more or less movable to allow for the 
distension required by the passage of prey, often 
much exceeding the calibre of the mouth. For the 
same reason, the rami of the lower jaw, which con- 
sist of dentary, splenial, angular, and articular 
elements, with the addition of a coronoid in the 
Boidae and a few other small families, are connected 
at the symphysis by a very extensible elastic ligament. 

The hyoid apparatus is reduced to a pair of cartila- 
ginous filaments situated below the trachea, and 
united in front. 

There are various modifications according to the 
genera. A large vacuity may be present between the 
frontal bones and the basisphenoid (Psanwiophis, 



SKELETON 



43 



Ccelopeltis) ; the maxillary may be much abbreviated 
and movable vertically, as in the Viperidae ; the 
pterygoids may taper and converge posteriorly, 
without any connexion with the quadrate, as in the 
Amblycephalidae ; the supratemporal may be much 




ccr 



an 




V" V 




cor 



Fig. 4 — Skull of Typhlops lumbvicalis. (From British Museum 

Catalogue of Snakes) 

Lettering of the bones as in Fig. 3 

reduced, and wedged in between the adjacent bones 
of the cranium ; the quadrate may be short or ex- 
tremely large ; the prefrontals may join in a median 
suture in front of the frontals ; the dentary may be 
freely movable, and detached from the articular 
posteriorly. 



44 



INTRODUCTION 



The deviation from the normal type is much 
greater still when we consider the degraded, worm- 
like members of the families Typhlopidae (Fig. 4, 
p. 43) and Glauconiidas (Fig. 5), in which the skull is 
very compact and the maxillary much reduced. In 




pro 



60 Ho 

Fig. 5 — Skull of Glauconia macvolepis. (From British Museum 

Catalogue of Snakes) 

Lettering of the bones as in Fig. 3 

the former this bone is loosely attached to the lower 
aspect of the cranium ; in the latter it borders the 
mouth, and is suturally joined to the premaxillary 
and the prefrontal. In both the tranverse bone and 
the supratemporal are absent, but the coronoid ele- 
ment is present in the mandible. 



SKELETON 



45 



The principal modifications of the skull in the 
European genera may be contrasted as in the 
following synopsis : 

I. Quadrate articulating with the cranium, supra- 
temporal absent; mandible much shorter than the 
skull, with coronoid bone ; maxillary small, on 




-an 



Fig. 6 — Skull of Tropidonotus natrix. (From British Museum 

Catalogue of Snakes) 

Lettering of the bones as in Fig. 3 

lower aspect of cranium ; pterygoids not extend- 
ing to quadrate ; nasals forming long sutures with 

the premaxillary, prefrontals, and frontal 

Typhlops. 

II. Quadrate suspended from the supratemporal ; 
mandible at least as long as the skull ; pterygoids 
extending to quadrate or mandible. 



4 6 



INTRODUCTION 



A. Mandible with coronoid bone ; nasals in sutural 
contact with frontals and prefrontals ; trans- 
verse bone short, not projecting much beyond 
cranium ; maxillary not half as long as 






Fig. 7 — Skull of Zamenis gemonensis. (From British Museum 

Catalogue of Snakes) 

mandible, which is not longer than skull (to 

occiput) Eryx. 

B. No coronoid bone ; nasals isolated. 

i. Maxillary elongate, not movable vertically. 
a. Maxillary half as long as mandible. 
Supratemporal half as long as skull, projecting far 



SKELETON 



47 



beyond cranium ; mandible much longer than 

skull Tropidonotus. 

Supratemporal not half as long as skull, projecting 
far beyond cranium ; mandible much longer than 
skull Zamenis. 




Fig. 8 — Skull of Coluber loiigissimus. (From British Museum 

Catalogue of Snakes) 

Supratemporal not half as long as skull, projecting 
but slightly beyond cranium ; mandible much 
longer than skull Coluber. 

Supratemporal not half as long as skull, not pro- 
jecting beyond cranium ; mandible not longer than 
skull Coronella, Contia. 



4 8 



INTRODUCTION . 



b. Maxillary not half as long as mandible, 

which is longer than skull ; supratemporal 

not half as long as skull, projecting beyond 

cranium. 

Quadrate longer than supratemporal ; maxillary 




Fig. 9 — Skull of Coronella austriaca. (From British Museum 

Catalogue of Snakes) 

much longer than quadrate, nearly straight in front 
of prefrontal ; a large vacuity between the frontal 

bones and the basisphenoid Ccelopeltis. 

Quadrate not longer than supratemporal ; maxillary 
little longer than quadrate, strongly curved in front 
of prefrontal Macroprotodon. 



SKELETON 



49 



Quadrate longer than supratemporal ; maxillary 
little longer than quadrate, nearly straight in front 
of prefrontal Tarbophis. 



W f />/-" p rr°.sie 



or. 




Fig. io — Skull of Vipera lebetina. (From British Museum 

Catalogue of Snakes) 

Lettering of the bones as in Fig. 3 

2. Maxillary much abbreviated and erectile ; 
supratemporal not half as long as skull ; 
mandible much longer than skull ; basioc- 
cipital with a strong process. 
4 



50 INTRODUCTION 

Maxillary bone solid Vipera. 

Maxillary bone hollowed out Ancistrodon. 

The vertebrae number 130 to 500 — in the European 
forms 147 (Vipera ursinii) to 330 (Coluber leopardinus). 

The vertebral column consists of an atlas (com- 
posed of two vertebrae) without ribs ; numerous pre- 
caudal vertebrae, all of which, except the first or first 
three, bear long, movable, curved ribs with a small 
posterior tubercle at the base, the last of these ribs 
sometimes forked; two to ten so-called "lumbar 
vertebrae " without ribs, but with bifurcate transverse 
processes (lymphapophyses) enclosing the lymphatic 
vessels ; and a number of ribless caudal vertebrae with 
simple transverse processes. When bifid, the ribs or 
transverse processes have the branches regularly 
superposed. 

The centra have the usual cup-and-ball articulation, 
with the nearly hemispherical or transversely elliptic 
condyle at the back (procoelous vertebrae), whilst the 
neural arch is provided with additional articular sur- 
faces in the form of pre- and post-zygapophyses, broad, 
flattened, and overlapping, and of a pair of anterior 
wedge-shaped processes called zygosphene, fitting 
into a pair of corresponding concavities, zygantrum, 
just below the base of the neural spine. Thus the 
vertebrae of snakes articulate with each other by eight 
joints in addition to the cup-and-ball on the centrum, 
and interlock by parts reciprocally receiving and 
entering one another, like the joints called "tenon- 



SKELETON 



5i 



and-mortice " in carpentry. The precaudal vertebrae 
have a more or less high neural spine which, as a 
rare exception (Xenopholis), may be expanded and 
plate-like above, and short or moderately long trans- 
verse processes to which the ribs are attached by a 
single facet. The centra of the anterior vertebrae 
emit more or less developed descending processes, 






B 



t 







Fig. 11 — Posterior Precaudal Vertebrae of Lioheterodon (A) and 
Heterodon (B). (From British Museum Catalogue of Snakes) 

a, Back view ; b, lower view ; c, side view. 

or haemapophyses, which are sometimes continued 
throughout (Fig. n, A), as in Tropidonoius, Vipera, 
and Ancistrodon, among European genera. 

In the caudal region, elongate transverse processes 
take the place of ribs, and the haemapophyses are 
paired, one on each side of the haemal canal. In the 
Rattlesnakes the seven or eight last vertebrae are 
enlarged and fused into one. 



52 INTRODUCTION 

No snake shows any rudiments of the pectoral 
arch, but remains of the pelvic are found in the 
Typhlopidce, the Glauconiidce, the Boidcz, and the 
I ly slides. In the first these vestiges are reduced to a 
single bone (ilium ?) on each side ; in the second they 
consist of ilium, pubis, and ischium, the latter form- 
ing a ventral symphysis, and a rudimentary femur ; 
whilst in the third there is a long ilium, attached to 
the lower branch of the first bifurcate transverse pro- 
cess of the lumbar vertebrae, bearing three short 
bones, the longest of which, regarded as the femur, 
terminates in a claw-like spur which, in males at 
least, usually appears externally on each side of the 
vent. 



CHAPTER V 
DENTITION 

IN the most generalized snakes — those which 
show the nearest approach to lizards — teeth are 
present not only on the rami of both jaws, but also 
on the premaxillary bone, on the palatines, and on 
the pterygoids. A reduction of the dentition takes 
place in various genera, in which the teeth of either 
the upper or the lower jaw, and of the palatines or 
pterygoids, or both, may be absent, and the pre- 
maxillary is devoid of teeth in the great majority, 
including all European representatives, of the Ophidia. 
In the egg-eating snakes of the genera Dasypeltis 
and Elachistodon the dentition is very much reduced, 
in accordance with the peculiar regime, and this 
deficiency is compensated by the development on 
some of the anterior thoracic vertebras of long, tooth- 
like processes (hypapophyses) directed forwards, and 
capped with a remarkably dense, vitreous tissue 
simulating enamel, the function of these tooth-like 
processes being to break the shell of the egg within 
the gullet, where none of its contents are lost, the 
shell being afterwards rejected through the mouth in 
the form of a pellet. 

53 



54 INTRODUCTION 

With the exception of the worm-like Typhlopidse, 
which are provided with a few teeth in the upper jaw 
only, European snakes have teeth on the maxillary, 
palatine, pterygoid, and dentary bones. Unless the 
maxillary be strongly abbreviated and modified in 
connexion with the poison apparatus, as in the 
Viperidae, the teeth in the jaws as well as on the 
palate form single longitudinal series ; they are 
elongate, conical, with or without a sharp posterior 
edge, more or less recurved, acutely pointed, some- 
times needle-like, and directed backwards, as behoves 
their function, which, in addition to attack and 
defence, is to prevent the retrogression of the prey 
in the act of prehension and deglutition. A notable 
exception occurs in the genus Iguanognathus, from 
Sumatra, all the teeth having spatulate crowns 
ribbed along the outer side. Unfortunately, nothing 
is known as to the food of this remarkable snake. 
The teeth are coated with a thin layer of enamel. It 
was held, for a time, that the glossy outer coating was 
only due to a denser structure of the dentine. As in 
all living Reptiles with the exception of the Crocodiles, 
the teeth are not implanted in true sockets, but 
simply ankylosed to the bone on which, when de- 
tached, their slightly enlarged base, or rather the 
bony tissue on which it rests, leaves a shallow im- 
pression, or pseudo-socket. In the process of biting 
or feeding, some of the teeth are frequently lost, and 
are readily replaced by others lying in reserve in the 



DENTITION 55 

gum at the inner side, and becoming fixed to the bone 
soon after a vacancy occurs. Such replacement teeth, 
of different grades of development, form several series, 
so that in a snake like our common Tropidonotus the 
mouth may contain four times as many teeth as are 
functional, without reckoning different earlier stages 
of tooth germs which escape ordinary observation, 
being placed vertically one above the other. 

Three types of teeth, connected by every inter- 
mediate step, are distinguished : the solid, the 
grooved, and the canaliculated or tubular, so-called 
" perforated " ; the third, as we shall explain, being 
only a further modification of the second. In the 
grooved tooth, a sulcus runs along the anterior or 
outer surface, its object being to convey into the 
wound the secretion of a poison gland. It varies in 
depth according to the species, and may be so slight 
as to escape detection without a very strong magnify- 
ing glass. In some the sulcus may be very deep and 
wide, forming a canal round which the tooth folds to 
the extent of its borders nearly meeting ; from this 
condition the so-called "perforated" fang is derived 
through the complete fusion of the borders of the 
tooth, and the obliteration of the line of union except 
at each extremity. The structure of such a fang may 
be best understood by imagining a tooth, lined all 
round with the same layer of dentine and enamel, 
being flattened out in a vertical plane and then 
folded over, the outer edges coalescing on the front 



56 INTRODUCTION 

median line in such a way that the inner wall of 
the tooth is in reality the anterior surface, and 
the outer wall the posterior surface, of the ordinary 
tooth. 

Grooved teeth, with open canal, are situated either 
at the anterior extremity (Proteroglyphs) or at the 
posterior extremity (Opisthoglyphs) of the maxillary 
bone, usually followed or preceded by a series of solid 
teeth, which in some cases may likewise show a 
more or less distinct groove. Such may also be 
present on the teeth of the lower jaw, as in the Euro- 
pean Ccelopeltis, in some specimens of which a faint 
groove is visible on the outer side with the aid of a 
strong lens. 

The tubular fangs of the Viperidae are inserted on 
the posterior extremity of the much abbreviated and 
erectile maxillary bone, which bears no other teeth. 
The Proteroglyphs (Cobras, Coral-snakes, Sea-snakes) 
and the Solenoglyphs (Vipers, Pit-vipers, Rattlesnakes) 
may be regarded as the diverging extremes in the 
development of the poison apparatus, both culmina- 
ting in forms with tubular fangs, the former as de- 
rived directly from the Aglyphs (harmless snakes), 
the latter from the Opisthoglyphs, likewise evolved out 
of the Aglyphs. That the insertion of the poison 
fangs of the Viperidae is really on the posterior 
extremity of the maxillary bone is evident from the 
condition of the bone in its recumbent position, 
especially in the African Viper, Causus, which in 



DENTITION 57 

several respects departs less markedly from the Colu- 
brid type than our European Vipers. 

The poison fangs of the Viperidse appear to be 
movable, folding in the mouth when at rest, and 
erected, or even thrust forward, when ready to act. 
This, however, is simply due to the mobility of the 
maxillary bone, to which they are ankylosed as in 
all other snakes. There are normally two equally- 
developed fangs, close together and side by side, to 
each maxillary, followed by several replacement fangs 
loosely attached behind them, usually in two series of 
four. When the two fangs are in situ, they of course 
both function in the act of biting, although only one 
is in relation with the single poison duct ; often, how- 
ever, there is only one fang in position, either the 
right or the left, the place of the other being indi- 
cated by a shallow socket which will soon be filled by 
one of the posterior reserve fangs moving forward and 
becoming ankylosed to the bone. Snake-charmers 
who extract the poison fangs of the snakes they use 
for their performances have therefore to renew the 
operation frequently, unless they amputate the bone 
on which the fangs are inserted, an injury which the 
creature does not long survive. 

The dentition of the snakes in which the maxillary 
bone is not movable vertically falls under three 
divisions : the Aglyphs, in which the teeth are all solid ; 
the Opisthoglyphs, in which one or more (usually 
two) of the hindermost teeth are provided with a 



58 INTRODUCTION 

groove ; and the Proteroglyphs, in which grooved or 
canaliculated teeth are situated in front, followed or 
not by solid teeth. Beyond these three principal 
divisions, the dentition furnishes important characters 
for the classification, although that importance has 
sometimes been over-estimated. The maxillary teeth 
may be equal in length (Isodonts), or the anterior the 
longer (Lycodonts), or the posterior the longer, in- 
creasing gradually in size (Coryphodonts) or abruptly, 
without (Syncranterians) or with a diastema, or break, 
in front of them (Diacranterians). These categories 
are, however, so completely connected as to preclude 
their use in taxonomy beyond helping to define 
genera. The number of maxillary teeth and the 
relative proportions and disposition of the mandib- 
ular teeth also afford useful generic characters. 

The European genera may be arranged as follows, 
according to the dentition : 

I. Teeth few, disposed in a transverse series in the 
upper jaw only Typhlops. 

II. Teeth in both jaws and on the palatines and 
pterygoids. 
A . A series of solid teeth along the maxillary ; no 
grooved teeth. 
t. Anterior maxillary and mandibular teeth 

longest; 9 or 10 maxillary teeth Eryx. 

2. Maxillary teeth equal, or increasing in size 
posteriorly. 



DENTITION 59 

a. Mandibular teeth 17 to 30 ; maxillary teeth 
15 to 22. 

Posterior maxillary teeth longest ; mandibular teeth 
subequal, more than 20 Tropidonotus. 

Posterior maxillary teeth longest ; mandibular teeth 
not more than 20, posterior smallest Zamenis. 

Maxillary teeth subequal ; mandibular teeth 20 to 25, 
posterior smallest Coluber. 

b. Mandibular teeth 14 or 15, subequal ; maxil- 
lary teeth 12 to 15. 

Maxillary teeth increasing in size Coronella. 

Maxillary teeth subequal Contia. 

B. One or two enlarged grooved fangs behind the 
series of solid maxillary teeth. 
14 to 17 subequal solid maxillary teeth, forming a 
continuous series ; 21 to 23 mandibular teeth, 

anterior strongly enlarged Ccelopeltis. 

g to 11 solid maxillary teeth, fourth and fifth or fifth 
and sixth enlarged, followed by an interspace ; 
sixth mandibular tooth fang-like, followed by an 

interspace Macroprotodon. 

9 or 10 solid maxillary teeth, forming a continuous 
series, decreasing in length posteriorly ; anterior 

mandibular teeth strongly enlarged Tarbophis. 

C. Maxillary with only two large canaliculated 
fangs side by side, one of which may be miss- 
ing ; anterior mandibular teeth longest 

Viper a, Ancistrodon. 



6o 



INTRODUCTION 



In counting the teeth for the purpose of using this 
key, care must be taken to ascertain the full number, 
as it frequently happens that one or more are miss- 
ing ; but their place is indicated by the shallow 
pits in which their base was implanted, the over- 




a 



*mmm*^^ 




Fig. 12 — Maxillary and Mandible of — (a) Tarbophis fall ax ; 
(b) Ccelopeltis monspessulana ; (c) Macroprotodon cucullatus. (From 
British Museum Catalogue of Snakes) 

looking of which might convey the impression of a 
hiatus such as is characteristic of certain genera — 
Macroprotodon, for instance. Needless to say, the loose 
teeth which are in reserve on the inner side of the 
jaws or behind the tubular fangs are not taken into 
consideration, the numbers given being those of 



DENTITION 6 1 

functional teeth only. Although as a rule the teeth 
can be counted easily, on a specimen preserved in 
spirit, by simply pushing aside the lips and gums 
with the finger, it is sometimes necessary to remove 
and clean the bones of the jaws, an operation which 
does not require much skill. 



CHAPTER VI 

POISON APPARATUS— DIFFERENT KINDS OF 

POISONS 

~^HE gland which secretes the poison is a modifi- 
■■■ cation of the parotid salivary gland of other 
Vertebrates, and is usually situated on each side of 
the head below and behind the eye, invested in a 
muscular sheath. It is provided with large alveoli 
in which the venom is stored before being conveyed 
by a duct to the base of the channelled or tubular 
fang through which it is ejected. 

In the Vipers, which furnish examples of the 
most highly developed poison apparatus, although 
inferior to some in its toxic effects, the poison gland 
is very large and in intimate relation with the 
masseter or temporal muscle, consisting of two bands, 
the superior arising from behind the eye, the inferior 
extending from the gland to the mandible. When 
the snake bites, the jaws close up, causing the gland 
to be powerfully wrung, and the poison pressed out 
into the duct. From the anterior extremity of the 
gland the duct passes, below the eye and above the 
maxillary bone, where it makes a bend, to the basal 
orifice of the poison fang, described above (p. 55), 

62 



POISON APPARATUS 



63 



which is ensheathed in a thick fold of mucous 
membrane, the vagina dentis. By means of the 
movable maxillary bone (supra, p. 49) hinged to the 
prefrontal, and connected with the tranverse bone 
which is pushed forward by muscles set in action by 
the opening of the mouth, the tubular fang is erected 

d g e 6 c a' 




Fig. 13 — Poison Apparatus of Rattlesnake: Venom Gland 
and Muscles (Lateral View). (After Duvernoy) 

a, Venom gland ; a\ venom duct ; b, anterior temporal muscle ; 
b', mandibular portion of same ; c, posterior temporal muscle ; 
d, digastricus muscle ; e, posterior ligament of gland ; /, sheath 
of fang ; g, middle temporal muscle ; h, external pterygoid 
muscle ; i, maxillary salivary gland ; /, mandibulary salivary 
gland. 

and the poison discharged through the distal orifice 
in which it terminates. 

In some of the Proteroglyphous Colubrids, as we 
have seen, the poison fangs are not tubular, but only 
channelled and open along the anterior surface ; and 
as the maxillary bone in these snakes is more or less 
elongate, and not or but slightly movable vertically, 



64 INTRODUCTION 

the poison duct runs above the latter, making a bend 
only at its anterior extremity, and the tranverse bone 
has not the same action on the erection of the fangs. 
Otherwise the mechanism is the same. 

In the Opisthoglyphous Colubrids, with grooved 
teeth situated at the posterior extremity of the 
maxilla, a small posterior portion of the upper 
labial or salivary gland is converted into a poison- 
secreting organ, distinguished by a light yellow 
colour, provided with a duct larger than any of 
those of the labial gland, and proceeding inward 
and downward to the base of the grooved fang ; the 
duct is not in direct connexion with the groove, but 
the two communicate through the mediation of the 
cavity enclosed by the folds of mucous membrane 
surrounding the tooth, and united in front. 

The reserve or successional teeth, which are 
always present just behind or on the side of the 
functional fang of all venomous snakes, are in no 
way connected with the duct until called upon to 
replace a fang that has been lost. It could not be 
otherwise, since the duct would require a new 
terminal portion for each new fang; and as the 
replacement takes place alternately from two parallel 
series, the new poison-conveying tooth does not 
occupy exactly the same position as its predecessor. 

Two genera, Doliophis among the Elapine Colu- 
brids, and Causus among the Viperids, are highly 
remarkable for having the poison gland and its duct 



POISON APPARATUS 65 

of a great length, extending along each side of the 
body and terminating in front of the heart. Instead 
of the muscles of the temporal region serving to 
press out the poison into the duct, this action is 
performed by those of the side of the body. 

When biting, a Viperid snake merely strikes, dis- 
charging the venom the moment the fangs penetrate 
the skin, and then immediately leaves go. A Pro- 
teroglyph or Opisthoglyph, on the contrary, closes 
its jaws like a dog on the part bitten, often holding 
on firmly for a considerable time. 

The poison, which is mostly a clear limpid fluid of 
a pale straw or amber colour, more rarely greenish, 
sometimes with a certain amount of suspended 
matter, is exhausted after several bites, and the 
glands have to recuperate. 

It must be added that the poison can be ejected 
otherwise than by a bite, as in the so-called Spitting 
Snakes of the genera Naia and Sepedon. The fact 
that some of these deadly snakes when irritated are 
in the habit of shooting poison from the mouth, at a 
distance of 4 to 8 feet, even apparently aiming 
at a man's face, has been too often witnessed in 
India and Malaya, and especially in Africa, from the 
days of the ancient Egyptians, for any doubt to 
subsist as to their being endowed with this faculty, 
but the mechanism by which this action is produced 
has not been satisfactorily explained. In all proba- 
bility, the poison escapes from the sheath of 
5 



66 INTRODUCTION 

mucous membrane surrounding the base of the 
fangs, and is mixed with ordinary saliva, the mem- 
branes of the mouth perhaps acting as lips, in which 
case the term " spitting " would not be incorrect. The 
spitting, which may take place three or four times 
in succession, has been observed to be preceded by 
some chewing movements of the jaws. If reaching 
the eye, the poisonous fluid causes severe inflamma- 
tion of the cornea and conjunctiva, but no more 
serious results if washed away at once. 

Snake poisons is a subject which has always 
attracted much attention, and which has made great 
progress within the last quarter of a century, es- 
pecially as regards the defensive reaction by which 
the blood may be rendered proof against their effect 
by processes similar to vaccination — antipoisonous 
serotherapy. The studies to which we allude have 
not only conduced to a method of treatment against 
snake-bites, but have thrown a new light on the 
great problem of immunity. They have shown that 
the antitoxic serums do not act as chemical antidotes 
in destroying the venom, but as physiological anti- 
dotes ; that, in addition to the poison glands, snakes 
possess other glands supplying their blood with 
substances antagonistic to the poison, such as also 
exist in various animals refractory to snake poison, 
the hedgehog and the mungoose for instance. 
Unfortunately, the specificity of the different snake 
poisons is such that, even when the physiological 



POISONS 67 

action appears identical, serum injections or gradu- 
ated direct inoculations confer immunity towards 
one species or a few allied species only. Thus, a 
European in Australia who had become immune 
to the poison of the deadly Notechis scutatus, 
manipulating these snakes with impunity, and 
was under the impression that his immunity ex- 
tended also to other species, when bitten by a 
Denisonia superba, an allied Elapine, died the follow- 
ing day. In India, the serum prepared with the 
venom of Naia tripudians has been found to be 
without effect on the poison of Naia bungarus, the 
two species of Bnngarus, and the Vipers Vipera 
russelli, Echis carinatus, and Lachesis gramineus. 
Vipera russelli serum is without effect on Colubrine 
venoms, and on those of Echis and Lachesis, In 
Brazil, serum prepared with the venom of Lachesis 
lanceolatus has proved to be without action on 
Crotalus poison. These examples, and others which 
could be given, show that the hopes which were 
at first entertained as to the benefits to be conferred 
on mankind by the serum treatment were somewhat 
over-sanguine — at least as regards countries like 
India, where, different kinds of poisonous snakes 
occurring together, it is sometimes impossible to 
know by which the bite has been inflicted. 

Chemistry teaches that snake venoms consist for 
the most part of solutions of modified proteids, and 
all attempts to separate the toxic principles from 



68 INTRODUCTION 

such proteids have hitherto been unsuccessful. 
Accordingly, at the present time we must regard 
such toxic principles as residing in some special 
grouping of a portion of the atoms in the complex 
venom proteid molecule. The analysis of their 
physiological actions has proved them to be made 
up of a great many more constituents than would be 
imagined from their chemical composition. 

The effect of the poison of Proteroglyphous , 
Colubrids (Hydrophids, Cobras, Bungarus, Elaps, 
Pseudechis, Notechis, Acanthophis) is mainly on the 
nervous system, respiratory paralysis being quickly 
produced by bringing the poison into contact with 
the central nervous mechanism which controls 
respiration ; the pain and local swelling which 
follow a bite are not usually severe. 

Viper poison {Viper a, Echis, Lackesis, Crotalus) 
acts more on the vascular system, bringing about 
coagulation of the blood and clotting of the 
pulmonary arteries ; its action on the nervous 
system is not great, no individual group of nerve-cells 
appears to be picked out, and the effect upon respira- 
tion is not so direct ; the influence upon the circulation 
explains the great depression which is a symptom 
of Viperine poisoning. The pain of the wound is 
severe, and is speedily followed by swelling and dis- 
coloration. The symptoms produced by the bite of 
the European Vipers are thus described by the best 
authorities on snake poison (Martin and Lamb) : 



POISONS 69 

The bite is immediately followed by local pain of a 
burning character ; the limb soon swells and becomes 
discoloured, and within one to three hours great 
prostration, accompanied by vomiting, and often 
diarrhoea, sets in. Cold, clammy perspiration is 
usual. The pulse becomes extremely feeble, and 
slight dyspnoea and restlessness may be seen. In 
severe cases, which occur mostly in children, the 
pulse may become imperceptible and the extremities 
cold ; the patient may pass into coma. In from 
twelve to twenty-four hours these severe constitu- 
tional symptoms usually pass off; but in the mean- 
time the swelling and discoloration have spread 
enormously. The limb becomes phlegmonous, and 
occasionally suppurates. Within a few days recovery 
usually occurs somewhat suddenly, but death may 
result from the severe depression or from the 
secondary effects of suppuration. That cases of 
death, in adults as well as in children, are not 
infrequent in some parts of the Continent is 
mentioned in the last chapter of this Introduction. 

The bite of all the Proteroglyphous Colubrids, 
even of the smallest and gentlest, such as the Elaps 
or Coral-snakes, is, so far as known, deadly to man. 
The Viperidse differ much among themselves in the 
toxicity of their venom. Some, such as the Indian 
Vipera russelli and Echis carinatus, the American 
Ancistrodon, Crotalus, Lachcsis mutus and lanccolatus, 
the African Camus, Bitis, and Cerastes, cause fatal 



70 INTRODUCTION 

results unless a remedy be speedily applied. On the 
other hand, the Indian and "Malay Lachesis seldom 
cause the death of man, their bite in some instances 
being no worse than the sting of a hornet. The 
bite of the larger European Vipers may be very 
dangerous, and followed by fatal results, especially 
in children, at least in the hotter parts of the 
Continent ; whilst the small Vipera ursinii, which 
hardly ever bites unless roughly handled, does not 
seem to be possessed of a very virulent poison, and, 
although very common in some parts of Austria- 
Hungary, is not known to have ever caused a serious 
accident. 

It is noteworthy that the size of the poison fangs 
is in no relation to the virulence of the venom. 
The comparatively innocent Indo-Malay Lachesis 
alluded to above have enormous fangs, whilst the 
smallest fangs are found in the most justly dreaded 
of all snakes, the Hydrophids. 

Little is known of the physiology of the poison of 
the Opisthoglyphous Colubrids, except that in most 
cases it approximates to that of the Proteroglyphs. 
Experiments on Ccelopeltis, Psammophis, Trimero- 
rhinus, Dipsadomorphus, Trimorphodon, Dryophis, 
Tarbophis, Hypsirhina, and Cerberus, have shown 
these snakes to be possessed of a specific poison, 
small mammals, lizards, or fish, being rapidly 
paralyzed and succumbing in a very short time, 
whilst others {Eteirodipsas, Ithycyphus) do not seem 



POISONS 71 

to be appreciably venomous. Man, it is true, is not 
easily affected by the bite of these snakes, since, at 
least in most of those which have a long maxillary 
bone, the grooved fangs are placed too far back to 
inflict a wound under ordinary circumstances. There 
are, however, exceptions. A case was reported a 
few years ago of a man in South Africa nearly dying 
as a result of the bite of the Boomslang, Dispholidus 
typus, the symptoms, carefully recorded, being those 
characteristic of Viperine poisoning, an important 
fact to oppose to the conclusions, based on the 
physiological experiments on Ccelopeltis, which 
appeared to disprove the theory that the Viperidae 
may have been derived from Opisthoglyphous 
Colubrids. 

Experiments made with the secretion of the 
parotid gland of Tropidonotus and Zamenis have 
shown that even Aglyphous snakes are not entirely 
devoid of venom, and point to the conclusion that 
the physiological difference between so-called harm- 
less and poisonous snakes is only one of degree, just 
as there are various steps in the transformation of an 
ordinary parotid gland into a poison gland or of a 
solid tooth into a tubular fang. 

The question whether all snakes are immune to 
their own poison is not yet definitely settled. Most 
snakes certainly are, and it is a remarkable fact that 
certain harmless species, such as the North American 
Coronella getula and the Brazilian Rhachidelus brazili, 



72 INTRODUCTION 

are proof against the poison of the Crotalines which 
frequent the same districts, and which they are able 
to overpower and feed upon. The Cribo, Spilotes 
variabilis, is the enemy of the Fer-de-lance in St. 
Lucia, and it is said that in their encounters the 
Cribo is invariably the victor. Repeated experi- 
ments have shown our Common Snake, Tropidonotus 
natrix, not to be affected by the bite of Vipera berus 
and V. aspis, this being due to the presence, in the 
blood of the harmless snake, of toxic principles 
secreted by the parotid and labial glands, and 
analogous to those of the venom of these Vipers. 

The Hedgehog, the Mungoose, the Secretary Bird, 
and a few other birds feeding on snakes, are known 
to be immune to an ordinary dose of snake poison ; 
whether the pig may be considered so is still un- 
certain, although it is well known that, owing to its 
subcutaneous layer of fat, it is often bitten with 
impunity. The Garden Dormouse (Myoxus quercinus) 
has recently been added to the list of animals 
refractory to Viper poison. 



CHAPTER VII 

NERVOUS SYSTEM— SENSE ORGANS 

THE brain is small and of very oblong shape. It 
consists of smooth cerebral hemispheres, small 
optic lobes, a still smaller cerebellum, and long olfac- 
tory lobes ; the pineal body is not accompanied by a 
parietal organ. The spinal accessory cranial nerve 
is absent, and the sympathetic system is but feebly 
developed. 

The eyes have been noticed above (p. 12). When 
normally developed they are susceptible of a slight 
movement under the transparent disc, quite indepen- 
dent from the cornea, which covers them, and from 
which they are separated by the so-called " lacrymal 
chamber." There are two lacrymal glands, one in 
front and one behind ; the lacrymal duct opens into 
the posterior nares. A sclerotic bony ring is absent. 

The olfactory organ proper is little developed, but 
is accompanied by an accessory organ, Jacobson's 
organ, consisting of a pair of pediculate, cup-shaped 
sacs, between the nasal sacs and the roof of the 
mouth, encapsuled by the vomers and the turbinal 
bones, lined by olfactory epithelium, and opening in 
the mouth just in front of the choanae. As this 

73 



74 INTRODUCTION 

organ, richly provided with nerves, communicates 
with the inside of the mouth, its function may be to 
smell the prey as it passes through previous to de- 
glutition. Snakes cannot be credited with a keen 
sense of smell, although undoubtedly guided by it 
during the nuptial period. 

In the more thoroughly aquatic snakes, the nostril 
may be closed, when respiration is suspended, by a 
spongy tissue, which acts as a stopper, and such nos- 
trils are called "valvular," although a valve is not, in 
the strict sense, present ; when the animal breathes, 
the nostril is opened by a compression, through special 
muscles, of the cavernous tissue. In some Sand- 
snakes the narial opening may be reduced to a 
crescentic slit. 

The sense of hearing is not much developed. 
Tympanum, tympanic cavity, and Eustachian tubes 
are absent. In the typical snakes a long columellar 
rod (the stapes), with a fibrous or cartilaginous pad at 
the outer end, extends from the fenestra ovalis in the 
cranium to the quadrate, but in the degraded burrow- 
ing forms the stapes is a small bony plate closing the 
fenestra ovalis. 

With one exception (Eryx jaculus, which is said by 
Schreiber to lap like a lizard), the tongue is not used 
for drinking or for the prehension or gustation of 
food, nor for hissing, but is a tactile organ protruded 
on any object the snake wishes to probe. It is slender 
and deeply bifid at the end, smooth, very protractile, 



SENSE ORGANS 75 

often quite to the length of the head, and furnished 
with many sensory corpuscules. It is darted and 
vibrated on the least excitement, and is usually looked 
upon by the ignorant as a " sting." In most snakes 
it is much pigmented, dark brown or black ; in a few 
it is flesh-coloured or bright red. The tongue is 
entirely retractile into a sheath below the glottis and 
opening in front of it ; it is always withdrawn into 
the sheath when the snake bites or feeds. 

Other organs, which, in the absence of a satisfactory 
explanation of their use, have been termed ''organs of 
a sixth sense," reside in the head-shields and scales of 
many snakes, and in the deep pits on the sides of the 
head which are characteristic of various Boidse and 
a few Colubridae. 

Scales often show, near their posterior extremity, 
one or two small light spots or impressions, caused 
by a thinning of the epidermis, which have been 
called "apical pits"; they appear to coincide with the 
terminations of nerve fibres extending along the 
epidermal folds of the skin. Similar organs some- 
times form series on the borders of some of the head- 
shields, this being particularly noticeable in the 
Typhlopidae. 

The large and deep pit situated between the nostril 
and the eye (loreal pit) in the Crotaline Viperidse 
— whence the name Pit-vipers, or that of " cuatro 
naricas" which is bestowed on them by the Spaniards 
of Mexico — is divided into two chambers: an outer 



76 INTRODUCTION 

with large external orifice, and an inner, rather more 
posterior in position and occupying an excavation on 
the outer face of the maxillary bone. The inner 
walls of these chambers are very thin and membra- 
nous, and form a partition separating the two, except 
for the presence of a minute opening ; this partition is 
stretched across the hollow of the maxillary bone like 
the membrane of a drum, and is supplied with blood- 
vessels and nerves, the latter terminating in cells of 
variable form. The use of the organ, thus situated 
at the base of the poison fang, and therefore in close 
proximity to the sphincter of the poison duct, is still 
unknown. 

Several of the Boidae, such as Python and Cor alius, 
have deep pits in some of the upper and lower labial 
shields, or also on each side of the rostral shield ; 
these problematic organs are in all probability also 
sensory. 



CHAPTER VIII 

VISCERA 

N most snakes there is a very marked asymmetry 
-*- of the viscera and their blood-supply, the organs 
of the right side being anterior to, as well as larger 
than, those of the left. 

The heart in most cases is situated between the 
anterior seventh and the anterior fourth of the body ; 
it may be much farther back, beyond the anterior 
third, in Doliophis, Platurus, and some Viperidae and 
Amblycephalidae, in the middle in Chersydrns. It is of 
rather elongate form, enclosed in a pericardium in 
which it lies freely, and has a sinus venosus, two 
auricles, and a single ventricle divided by a septum. 
Three arteries leave the ventricle, the pulmonary and 
two systemic arches. The right systemic arch gives 
off the carotid artery, which in many snakes, the 
common Grass-snake for instance, may branch into 
two, or in others be double from its origin. The 
anterior abdominal vein is single in most snakes, 
double in some Boidse, and conveys blood from the 
ventral body-wall to the liver. The caudal vein is 
continued as the renal portal. Veins which have 
been regarded as remains of the two posterior cardinal 

77 



78 INTRODUCTION 

of lower Vertebrates have been found in some of the 
Boidae. 

The bifurcate transverse processes of the vertebrae 
at the limit between the body and tail enclose the 
lymph-hearts, which are large and more or less 
elongate, metamerically divided into several cham- 
bers, the right often more developed than the left. 
The thymus gland lies on each side of the trachea, 
near the heart, and the thyroid gland is in the middle 
line, close to the base of the carotid artery. 

The trachea is long, and the tracheal rings may be 
complete in front and incomplete behind, or incom- 
plete throughout. The bronchus opens at once into 
the more or less elongate, usually single lung, with or 
without a rudiment of a second, which seems to be 
constantly the left ; in some snakes the lung extends 
nearly to the cloacal region. In most of the Boidae 
there are two well-developed lungs, the left shorter 
than thq right. The lung has highly cellular walls in 
front, and becomes thin-walled, smooth, or but little 
vascular, behind, where it may receive its blood from 
the systemic and not from the pulmonary circulation. 
In the Typhlopidae and Viperidae, as well as in 
some of the Boidae, Colubridae, and Amblycephalidae, 
the posterior end or the greater part of the trachea 
may have its wall enlarged and provided with air cells, 
resembling the normal lung, with which it is usually 
continuous ; this has been called the " tracheal lung," 
but, although serving as an accessory breathing organ, 



VISCERA 79 

it is not a prolongation of the true lung, nor does it 
represent the missing left lung, as has been believed 
by some authors. 

The glottis has a longitudinal slit, and can be pro- 
jected forwards when the pharynx is obstructed by a 
voluminous prey. An epiglottis is usually absent, or 
represented by a rudiment. It is, however, present in 
some large American species of Coluber (Pityophis), said 
to produce, when hissing, a loud and hoarse sound 
which has been compared to the bellow of the bull — 
hence the popular name of Bull-snakes by which they 
are known. It has also been found in a few allied 
species from Mexico, for which the genus Epiglottophis 
has been proposed. This epiglottis is a narrow, thin 
flap, erect in front of the glottis; it is not hinged, and 
therefore not capable of falling down to cover the 
opening of the windpipe during the process of swallow- 
ing, its function evidently being to increase the sound 
produced by the escape of the air from the windpipe. 

The larynx is represented by two longitudinal 
bands of cartilage, united by transverse bands ; it is 
extremely long in some snakes (Leptognathus). 

The oesophagus, which may be extremely elon- 
gate, sometimes measuring almost one-third of the 
digestive canal, passes into the tubular or sac-like 
stomach, often with thickened walls, which itself 
gradually or abruptly merges into the narrower in- 
testine. The windings of the small intestine are 
connected by ligamentous tissue, and enclosed in 



80 INTRODUCTION 

a common sheath of peritoneum. In several of 
the Glyphodont Water-snakes (Homalopsinse and 
Hydrophiinae), the intestine is much convoluted ; in 
Herpeton it is even longer than the body, although 
when coiled occupying only one-fourth of that length. 
The rectum is sometimes very short, sometimes 
rather long, and its anterior portion may have a short 
caecum ; it may be divided by transverse septa, with 
median or lateral perforation. 

In snakes which swallow hard-shelled snails, the 
anterior part of the intestine has its inner wall fur- 
nished with zigzag muscular folds producing a reticu- 
late appearance, followed farther down by transverse 
and then longitudinal folds. In these snakes the 
intestine is abruptly constricted behind the stomach, 
at which point the shells are broken or crushed after 
their contents have been digested ; whilst in the egg- 
eating snakes, in which the eggshell has to be broken 
previous to its contents reaching the stomach, the 
cesophagus is narrowed in front of the latter, at the 
point where the tooth-like ventral processes of the 
vertebrae project and pierce the wall of the cesophagus 
in order to aid in this function, after which the broken 
shell is rejected through the mouth. 

The more or less elongate, feebly-lobed kidneys are 
placed in the posterior part of the body, often extend- 
ing nearly to the cloaca ; the right is usually a little 
longer than the left, or extends a little farther for- 
ward, or even may commence where the other ends. 



VISCERA 8 1 

The suprarenal bodies are narrow and elongate, placed 
on the renal veins or on the vena cava inferior. 

The ureters leave the hind ends of the kidneys, and 
open through the side-walls of the cloaca on a papilla 
which in the males contains also the opening of the 
vas deferens. There is no urinary bladder. The 
genital organs will be mentioned in the next chapter. 
The liver is usually long and narrow, measuring one- 
fifth to one-fourth the length of the body, on the right 
side of the alimentary canal, commencing just behind 
the heart or farther back. It is exceptionally short 
in Chersydrus. It is sometimes divided by transverse 
furrows. Its posterior extremity is bilobate, and the 
left lobe usually extends beyond the right, although 
the reverse has been observed in some snakes. The 
gall-bladder, which may be absent, is remarkable for 
its distance from the liver. The pancreas, elongate 
but comparatively small, is located near the spleen, on 
the left side of the alimentary canal, at a considerable 
distance from the liver. 

The peritoneal part of the body-cavity is subdivided 
into a number of spaces or ccelomic compartments 
enclosed in serous capsules — viz., a posterior or intes- 
tino-genital, a gastric on the left side, and a pair 
round the liver, corresponding to its two lobes. 

Fat-bodies are much developed, either in the form 
of small separate lobes, or as a continuous, much 
folded band, on each side of the body. 



CHAPTER IX 

ORGANS OF REPRODUCTION; PAIRING; 
OVIPOSITION ; DEVELOPMENT 

THE genital glands are situated anterior to the 
kidneys, the right extending farther forward 
and often larger than the left. The testes are 
elongate. The vas deferens is closely folded proxi- 
mally, and runs along the outer side of the kidney 
into the cloaca close to the ureter. The ovaries are 
elongate, and consist of two lamellae, with a lymph- 
space between them. The oviduct extends from 
near the anterior extremity of the ovary to a common 
chamber, or vagina, which is above the rectum and 
opens into the cloaca ; this vaginal chamber may be 
more or less completely divided into two. 

The males are provided with a pair of intromittent 
organs, or hemipenes, each connected with one of 
the caudal vertebrae by a muscle {retractor penis) which 
often exceeds it in length. These organs are cylin- 
drical or club-shaped and hollow, with the inner 
surface divided into numerous cavities and beset 
with papillae, and usually also with hard spines, of 
which those towards the apex may be greatly de- 
veloped, folded against the walls, and directed 

towards the extremity. Such spines are absent in 

82 



ORGANS OF REPRODUCTION 83 

the snakes provided with claw-like rudiments of 
hind limbs. The cavities of the hemipenis are 
connected by a branch with the dorsal artery, and 
it is by a flow of blood into them that erection of 
the organ is accomplished. Each hemipenis is 
lodged in a cavity on each side of the base of the 
tail; when protruded it turns inside out, and the 
inner surface becomes the outer, the papillae and 
erected spines serving to maintain a firm hold 
in the vagina, from which the organ cannot be 
withdrawn except by invagination. It has been 
observed that the presence of spines on the hemi- 
penis is associated with much tougher vaginal 
walls. The organ is grooved along its entire 
length, the groove being the sulcus spermaticus, 
which, when the edges of the two hemipenes meet, 
forms with its fellow a canal to convey the semen 
into the oviduct ; this sulcus may be bifurcate, as in 
the Viperids and some Colubrids. 

Anal pockets, secretory organs on each side of the 
vent and lodged in the base of the tail, seem, in 
females, to be the homologues of the hemipenes ; 
but this view cannot be held, since the same organs 
are present, though smaller, in males also, situated 
dorsally to the hemipenes. The glands with which 
they are provided produce the strong and offensive 
odour which appears to be a means of defence in 
our Grass-snake and other species, and which also 
serves to bring the sexes together, the glands being 



84 INTRODUCTION 

more active during the breeding season. A Viper- 
catcher in France is said to obtain good results by 
rubbing his boots with these glands, as a means of 
attracting the snakes in the spring. 

In European species pairing takes place in spring, 
sometimes again at the end of summer or in 
autumn. After hibernation the testes of the males 
are rather voluminous, and the sperm-ducts are often 
full of spermatozoa. The male gets alongside the 
female, sometimes seizing her round the neck with 
his jaws, and remains stretched out against her or 
twists the posterior part of his body in a few coils 
around hers. In the Vipers the bodies of the pairing 
individuals are completely entwined. The male 
then endeavours to bring the two anal orifices 
together, and when he has succeeded in getting the 
female to distend her cloacal opening, the intro- 
mittent organs are suddenly everted into the vagina. 
The union of the sexes sometimes lasts only a few 
minutes, but usually an hour or more; it has even 
been observed to last a whole day. Several copula- 
tions may take place at intervals of a few days. 
Many snakes are gregarious during the breeding 
season, and great numbers of males have been seen 
wriggling round the females, forming with their 
coils huge lumps or an entangled mass like a ball. 
The more or less prehensile tail with which 
thoroughly aquatic snakes, such as Hydrophis and 
Acrochordus, are provided, is no doubt of use in 



PAIRING 85 

facilitating the pairing, when it has to take place 
in the water. Our European Water-snakes pair on 
land. 

During the rutting season a slight pressure on the 
base of the male's tail may cause the protrusion of 
the hemipenes, and so may a violent blow on the 
spine of the reptile. Thus, recently killed specimens 
of our Adder, with the organs everted, have more 
than once been taken by the ignorant for snakes 
with hind limbs, a mistake which must be pardoned 
when we remember that male embryos of the slow- 
worm and of snakes, in w r hich the hemipenes are 
normally everted, have been described by zoologists, 
who should have known better, as examples showing 
external vestiges of limbs. 

The spermatozoa soon make their way up the 
oviducts, in which the ripe ova have previously 
descended, or which gradually descend shortly after, 
these ducts becoming dilated in consequence. There 
are usually more eggs in the right than in the left 
oviduct, although the reverse has occasionally been 
observed. 

Some snakes lay eggs shortly after impregnation, 
or a few weeks later; in others the young undergo 
their development within the oviducts, each envel- 
oped in a thin, transparent, membranous capsule, 
which is torn immediately before or immediately 
after parturition, such species being termed " ovovi- 
viparous." Just before oviposition the female curves 



86 INTRODUCTION 

the base of the tail upwards, in order to extend the 
cloacal opening. The eggs are all produced together, 
usually at intervals of a few minutes, and generally 
adhere to one another by means of a sticky fluid 
secreted by the oviducts, thus forming a clump. 
In ovoviviparous snakes the young are born in 
succession, in the course of a few hours or of a few 
days. In many oviparous species it is the rule for 
freshly-laid eggs to contain more or less developed 
embryos, and Coronella punctata is said to produce 
thin-shelled eggs which hatch in less than half the 
time required for the eggs of its American congeners 
under the most favourable circumstances. There is 
thus almost every degree between oviparity and 
ovoviviparity. 

These two modes of parturition bear no relation to 
the natural affinities of snakes. Thus, the European 
Coronella austriaca is ovoviviparous, and its North 
American congeners are oviparous ; whilst, curiously, 
it is the inverse in the genus Tropidonotus. It was long 
believed to be an invariable rule for the Viperidae to 
bring forth live young, the name Viper being derived 
from this well-known peculiarity, but it has now been 
ascertained that the South American Lachesis mutus, 
the Indo-Malay Lachesis monticola, and the African 
Causus and Atractaspis, lay eggs. All exclusively 
aquatic snakes, such as the Hydrophiinse, are ovovi- 
viparous, and thus dispensed from going on land for 
parturition. 



OVIPOSITION 87 

The yolk entirely fills the eggshell ; there is no 
albumen, or, if any exists, it is so much reduced as to 
easily escape observation. The eggshell in oviparous 
species contains a small amount of lime, and is 
not hard, but tough and parchment-like, white or 
yellowish; it is usually smooth, but in Pythons its 
surface is studded with minute pores, and in the 
American Zamenis constrictor it is rough, as if 
sprinkled over with loose grains of salt. The shape 
varies from a short oval to a long ellipse. It has 
been observed in some snakes that the eggs, on 
leaving the cloaca, are of an elongate shape, sugges- 
tive of a short cigar, and immediately after assume 
a more oval form. After they have been laid, the 
eggs absorb moisture and thus increase in size, espe- 
cially in width ; eggs which are at first twice as long 
as broad may be almost globular just before the birth 
of the young. 

The number of eggs or young of one brood varies 
much according to the species, and also according to 
the age of the mother, large females usually producing 
a higher number and of a larger size than smaller 
specimens of the same species. Our European 
Zamenis, Coluber, and Coronella produce only 2 to 15 ; 
our Tropidonotus, 15 to 48; our Vipers, 3 to 22. 
Among exotics we may mention, as the most prolific, 
Bitis nasicornis, up to 47 young; Tropidonotus fasciatus, 
Abastor erythrogr animus, and Farancia abacura, 50; 
Lachesis lanceolatus, 60 ; Vipera russelli, 63 ; Boa con- 



88 INTRODUCTION 

stridor, 64 ; Tropidonotus ordinatus, 78 ; Pseudaspis 
cana, 80 ; Python molurus, nearly 100 eggs. 

The eggs are deposited in holes without any sort 
of nest, under moss or decomposing leaves, in 
accumulations of saw-dust, or in manure-heaps. 
In many cases it has been observed that the female 
remains for some time with her eggs or young, and 
in the large Pythons a sort of incubation takes place, 
the female remaining coiled in a spiral over the mass 
of eggs for six to eight weeks ; an increase of several 
degrees in her temperature at that period has been 
ascertained by experiments conducted with every 
possible care, a remarkable fact in the case of a 
so-called " cold-blooded " animal. 

The numerous reports of young snakes seeking 
refuge in their mother's gullet have not been sub- 
stantiated by satisfactory scientific evidence, and, 
although it is perhaps wise to say that the question 
remains an open one, it may be mentioned that, in 
Europe at least, trained observers who have devoted 
special attention to the habits of Vipers, in districts 
where these reptiles are exceedingly abundant, have 
never come across an instance of the form of maternal 
solicitude with which these snakes in particular have 
been credited. Not a single reported case of a 
female snake swallowing her young for protection 
rests on satisfactory evidence. 

The embryo is closely coiled up in a spiral. Just 
before birth it is distinguished by a large, convex 



DEVELOPMENT 89 

head, with large, prominent eyes, and a compara- 
tively short body, the scales and ventral shields 
being much shorter than later in life. The umbilicus 
is situated in the posterior part of the body, from 
six to ten times as far from the head as from the 
vent. Long after birth the umbilical slit remains 
visible, and affords a means of distinguishing very 
young snakes from older examples of smaller species. 
In oviparous species the embryo is provided with a 
very conspicuous egg-tooth, pointing forwards and 
projecting from the notch in the lower border of the 
rostral shield ; this egg-tooth is much reduced, and 
sometimes very indistinct, in the ovoviviparous 
species. The function of the egg-tooth is to cut 
through the tough eggshell. This, after the young 
has left it, shows one or several slits in its anterior 
extremity, cut as clean as if with a sharp knife. The 
egg-tooth becomes loose soon after birth, and is shed 
within a few hours or a few days, sometimes even 
before birth in ovoviviparous species. 

Frequent cases have been observed of dicephalous 
embryos or young, which may live for a short 
time ; there are even records of a three-headed snake, 
stated to have been seen at Lake Ontario, and of 
snakes with two heads and two tails. 

Unless prematurely born with a considerable mass 
of vitellus attached to the umbilicus, the young 
immediately after birth resent all interference, hiss- 
ing, snapping, or puffing themselves up, after the 



go INTRODUCTION 

manner of their parents. The first shedding of the 
outer coating of the epidermis follows soon after 
birth ; not before then does the young take to food. 

No snake appears to be able to breed before it is 
four years old. 

Well-authenticated instances of different species 
interbreeding are unknown, but specimens inter- 
mediate between Viper a bents and V. aspis, and 
between V. berns and V. ammodytes, have been 
assumed, with much probability, to be hybrids. 



CHAPTER X 
HABITS 

SNAKES may be grouped, according to their 
mode of life, in five principal categories, 
gradually merging into each other, or two of them 
not infrequently found combined in one and the 
same species. These categories are : — Ground-snakes, 
Sand-snakes, Burrovving-snakes, Tree- snakes, and 
Water-snakes. 

Ground-snakes may be defined as living above 
ground, and only occasionally climbing bushes or 
entering the water. Among European genera, 
Coronella and Vipera are perfect examples of this 
type, whilst Coluber and Zamenis approach the 
Tree-snakes in often ascending bushes, or even 
trees. 

Sand-snakes are adapted for living on loose 
sand, in which they seek concealment. Such are 
Lytorhynchus and some Psammophis among the 
Colubridse, Cerastes among the Viperidse. Eryx con- 
nects this category with the next. 

Burrowing-snakes live chiefly underground, and 
often have the visual organ atrophied in consequence, 
as in Typhlops ; all the Typhlopidae, Glauconiidae, 

9i 



9 2 INTRODUCTION 

and Uropeltidae, belong to this category ; the Viperid 
Atractaspis is also a burrowing type. 

Tree-snakes spend the greater part of their life on 
bushes or trees. Cor alius among the Boidae, Den- 
drophis and Dendraspis among the Colubridae, Athens 
and various species of Lachesis among the Viperidae, 
may be quoted as examples. 

Of Water-snakes, some are exclusively aquatic, 
like the marine Hydrophiinse and the typical 
Acrochordinae {Acrochordus, Chersydrus) and Homa- 
lopsinae (Hipistes, Herpeton). Chersydrus and Hipistes 
occur in the sea as well as in fresh water. Many 
species of Tropidonotus (T. tessellatus and T. viperinus 
in Europe), as well as the genera Helicops, Grayia, 
Boidengerina, etc., among the Colubridae, Eunectes 
among the Boidae, Ancistrodon piscivorus among the 
Viperidae, are chiefly but not exclusively aquatic. 

Our Tropidonotus natrix stands between the 
Ground-snakes and the Water-snakes; Boas and 
Pythons are as much Water-snakes as Tree-snakes. 
As shown by these and many other examples which 
might be given, a division into categories cannot 
always be applied with precision, nor does it convey 
an expression of the natural relationships of the 
species, as was believed by many systematists of the 
last century, who appealed to such adaptations for 
the definition of families. 

A vertical pupil denotes more or less nocturnal 
habits. Nevertheless our European Vipers, which 



HABITS 93 

are provided with such a contractile pupil, are far 
from exclusively nocturnal, delighting to bask in the 
sun, and pairing and feeding in the day-time. The 
Boidse appear to be more nocturnal, but no snake is 
known to be absolutely so, and the two species of 
Coluber which have been found living in perfect 
darkness in limestone caves in the Malay Peninsula 
and China, where they feed chiefly on bats, occur 
also outside the caves, and probably never breed in 
them. 

It is often stated in books that the organs of 
locomotion for the exceedingly elongate body of 
snakes are the ribs, and these creatures have even 
been compared to Centipedes. This statement is no 
doubt true to a certain extent for slow locomotion 
on uneven ground, when the ribs and the corre- 
sponding ventral shields afford a point of support ; 
but it does not account for the rapid movements, as 
when a snake darts like an arrow in pursuit of its 
prey or to escape from an enemy. Besides, the 
winding motions are not different from those of a 
Slow-worm or Glass-snake, in which, encased as they 
are in a bony armour, the ribs cannot come into 
play at all. The action of the muscles alone is quite 
sufficient to account for the reptation of snakes, 
without the ribs having to play an essential part. 

Not only the Cobras, but several harmless snakes, 
are able to raise the anterior third of the body 
vertically, when taking up a threatening attitude in 



94 INTRODUCTION 

the presence of an enemy, at the same time widen- 
ing or inflating the region behind the head. 

Most snakes can climb, and in this case the ribs 
and ventral shields are of great assistance. The Tree- 
snakes, usually characterized by a very slender, some- 
times compressed, body, or by a prehensile tail, are 
specially adapted for twining themselves round 
branches, and in several of them the presence of a keel 
on each side of the ventral and subcaudal shields, 
accompanied by a notch corresponding to the keel, 
affords an additional help for climbing on vertical un- 
even surfaces, such as the trunks of trees. This con- 
dition of the ventral shields has a bearing on the 
extraordinary mode of locomotion with which some 
Tree-snakes [Chrysopelea, and probably also Dendro- 
phis) have long been credited by the Malays. We allude 
to the so-called Flying-snakes, remarkable for their 
habit of shooting down from trees and descending to 
the ground at an oblique angle, the body being kept 
rigid the whole time of the " flight." It has been 
observed in Chrysopelea that the ventral surface 
between the lateral keels, which may be compared 
to hinges, can be drawn in and become deeply 
concave, whilst at the same time a slight dorso- 
ventral flattening of the body takes place. During 
this muscular contraction the snake is like a piece of 
bamboo bisected longitudinally, and is buoyed up in 
such a way as to explain its parachute-like descent. 

All snakes are able to swim, and the more aquatic 



HABITS 95 

kinds may spend a few hours under the water. A 
Python molurus is known to have remained alive in a 
basket sunk for thirty-six hours in a river. The best 
adapted for aquatic life are the Hydrophiinae, or Sea- 
snakes, most of which never leave the water, and 
are quite helpless and soon die when brought on 
shore ; their body is more or less compressed 
posteriorly, and the tail oar-shaped. Sea-w/eeds and 
barnacles sometimes settle on them. Algae have also 
been observed growing on the fresh-water snake 
Herpeton tent acid 'atum. 

As regards food, Burrowing-snakes, as well as a 
few small Ground-snakes, subsist mostly on worms, 
insects, and myriopods ; Tree-snakes on lizards, 
frogs, birds and their eggs ; Water-snakes on fishes 
and batrachians. Among the other types, some 
show a predilection for mammals, others for lizards 
or snakes, whilst not a few feed indiscriminately 
upon mammals, birds, reptiles and batrachians, even 
on slugs, insects, and worms, in addition. However 
surprising, it is a fact that spiny mammals are 
occasionally eaten, spines of the Madagascar Hedge- 
hog (Ericulns) having been found in the excrements 
of a Boa madagascariensis. Even hard-shelled eggs 
and molluscs may constitute the principal or 
exclusive food of certain snakes. 

Thus, Dasypeltis eats nothing but birds' eggs, the 
shells of which are crushed in the gullet, by a special 
contrivance mentioned above (p. 80), and are soon 



96 INTRODUCTION 

after rejected through the mouth as a pellet. Other 
snakes, such as Coluber and Lioheterodon show them- 
selves partial to eggs in addition to live prey, but 
their alimentary canal does not depart from the 
normal, the eggs being broken in the stomach and 
the remains of the shells passed with the excrements. 

The Amblycephalidse subsist almost entirely on 
snails and slugs, the shells of the former being 
crushed in the anterior part of the intestine after 
their contents have been digested, and the debris are 
rejected through the vent. A small land tortoise has 
been found in the stomach of a Cobra (Naia hate) 
from Algeria. 

Snakes which take large prey secure it according 
to three methods : By catching it simply with the 
jaws, and immediately proceeding to swallow it, as 
in Tropidonotus and in some of the Constrictors when 
dealing with small animals ; by constriction, after 
having seized it with the jaws, crushing it in the 
coils of their body and thus killing it previous 
to feeding, as in the Boidse and Coluber; or by 
poisoning, by a mere stroke with the fangs, the 
result being awaited before the meal is begun, as in 
most of the Viperidae. Other poisonous snakes 
proceed according to the first method, the use of the 
venom being to reduce the struggles of the victim 
and to relax its muscles. Such snakes as are in the 
habit of previously killing their prey show little 
reluctance to accept dead food in confinement, a 



HABITS 97 

thing which others usually refuse to do ; they 
may, however, be deceived by the dead animal 
being agitated before them, and the system now 
adopted in our Zoological Gardens, of offering all 
snakes previously-killed animals, has been attended 
with comparative success. 

Some species feed almost exclusively on other 
snakes, and often manage to swallow individuals as 
large as, or even a little larger than, themselves. 
Examples are known of harmless snakes showing 
a predilection for dangerous species, to whose poison 
they are immune (see p. 71). 

As a rule snakes that eat fish will also eat batra- 
chians, but nothing higher in the scale, although 
exceptions have been reported, such as the Anaconda 
feeding on mammals, birds, reptiles, and fish, and our 
Grass-snake having taken mice and birds. Some that 
feed chiefly on lizards and snakes will occasionally eat 
also mammals, and vice versa, but rarely frogs. On 
the other hand, European Vipers accommodate them- 
selves to a more varied bill of fare, being known to 
feed on mammals, birds, reptiles, batrachians, insects, 
and slugs, and they have even been observed to eat 
voles showing signs of putrefaction. 

The enormous prey which some snakes are able 
to swallow is quite astounding. Anacondas and 
Pythons, the largest snakes, have been known to 
swallow calves and good-sized antelopes with their 
horns, animals which, even after being somewhat 
7 



9 8 INTRODUCTION 

crushed by constriction, very much exceed the 
calibre of the snake. A Python molurus 17 feet long 
is reported on good evidence to have swallowed a 
gravid Axis deer. A Grass-snake half an inch in 
diameter can manage a frog or toad three times 
that width, and a Dasypeltis of the same size a hen's 
egg. Such feats are rendered possible by the 
mobility of the jaws and palato-pterygoid arch on 
the cranium, and the elasticity of the ligaments by 
which they are attached (see above, p. 42), as well 
as by the mobility of the ribs and the absence of 
sternal apparatus, together with the great dis- 
tensibility of the skin. When a snake proceeds to 
dispose of a large prey, which, if it be a mammal or 
bird, is usually seized head-first, it pulls itself 
forward by alternate movements of the jaws, the 
maxillary and the mandibular ramus of the one side, 
and then of the other, being extended anteriorly and 
laterally, the snake at the same time producing an 
abundant salivation which renders the prey very 
slimy. Several repeated alternate movements of the 
jaws bring the head of the prey to the gullet, where 
the muscles and ribs come into play, and the two 
sides of the jaws work no longer alternately, but 
together. When once in the oesophagus, the prey 
progresses with much greater facility, and usually 
reaches the stomach in a few minutes, whilst the 
previous process of deglutition may have lasted half 
an hour. While this laborious operation is going 



HABITS 99 

on, the breathing of the snake is not impaired 
owing to a remarkable contrivance : the trachea 
can be protruded in such a manner as to bring its 
opening outside the mouth. 

In cases where the victim is eaten alive, the snake 
has to contend with its struggles, but retrogression is 
rendered impossible by the backwardly-directed 
sharp teeth with which the jaws and palate are beset. 
A frog is usually caught by one of the hind limbs and 
swallowed back-first, the long hind limbs stretching 
forwards as they fold against the body; its struggles 
are often still apparent when it has reached the 
oesophagus. Snakes when caught immediately after 
a meal are in the habit of disgorging their food, and 
it sometimes happens that a frog or toad is thus 
vomited alive. An instance is known of a naturalist 
having captured a Grass-snake and put it in a linen 
bag. On opening it a short time after, great was his 
surprise to find the snake had escaped through a 
small hole in the bag, leaving instead a living toad 
too big to pass through the hole. 

If not of too large a size, several animals will often 
be swallowed in rapid succession, after which the 
gorged snake will allow its digestive organs several 
days, or even weeks, of repose. A large Anaconda in 
the Paris Jardin des Plantes fed only thirty-six times 
in the course of seven years. Digestion is usually 
rapid in the small snakes, defecation taking place 
twenty-four to forty-eight hours after the feeding ; it 



ioo INTRODUCTION 

lasts much longer in the large Boas and Pythons. 
Thus, in the above-mentioned Anaconda it has been 
observed to take from nine to thirty-eight days. 
Even the hardest bones of birds are decomposed by 
the gastric juices, but hairs, feathers, and horny pro- 
ductions, are passed with the excrements, sometimes 
forming regular balls. It is in most cases possible to 
tell, from an inspection of the dried faeces, what a 
snake has been feeding on, hairs, feathers, beaks, 
claws, epidermal horny shields, bits of tooth-enamel, 
being found mixed with the chalky matter which 
represents the decomposed bones. As a rule there 
is but one defecation after each meal, but there are 
in addition more frequent renal dejections, consisting 
chiefly of uric acid. 

In captivity snakes show themselves capricious in 
the choice of food, one individual preferring mam- 
mals, whilst another, of the same species, will 
only take birds ; and many, although to all appear- 
ances perfectly healthy, will persist in refusing all 
food, and allow themselves to die of starvation — a 
suicide which may require months, or even years, to 
accomplish. A Rattle-snake in the menagerie of the 
Jardin des Plantes in Paris has lived two years and two 
months without taking any food, a Python sebcs nearly 
two years and a half, a Boa madagascariensis four years 
and a month. A Vipera aspis was kept for three 
years without food and without losing its vicious 
temper. Specimens thus fasting do not, as a rule, 



HABITS 101 

renew their epidermis, or do so but very rarely. Our 
Common Adder can very seldom be induced to feed 
in captivity. Other snakes may rid themselves of all 
shyness to the extent of taking food from the hand, 
or show such appetite as to seize a prey immediately 
on being released from the small box or bag in 
which they have travelled for a considerable time. 

Most snakes drink, and pretty often — not by 
lapping with the tongue, but by drawing in water 
from the mouth and immersing the anterior part of 
the head. Some are said to be fond of milk, but 
there is no foundation for the belief held by peasants, 
that they enter sheds with the object of sucking 
milk from the cows, which would be a material im- 
possibility ; their real purpose in visiting such places 
being a search for suitable dung-heaps in which to 
deposit their eggs. 

Snakes cannot be credited with much intelligence 
or educability, nor do they display any very marked 
instincts. The least stupid and most easily tamed 
are the species of the genera Coluber and Coronclla. 
There is, however, considerable difference in this 
respect between individuals of the same species. 
Most snakes, when freshly caught, defend themselves 
by biting, and some individuals retain their savage 
temper after months of captivity ; others hardly ever 
bite, even if molested. The Common Grass-snake, 
for instance, hisses loudly and takes up a very 
threatening attitude, or even pretends to snap with 



102 INTRODUCTION 

open mouth, but very seldom bites; its principal 
defensive action when caught consists in voiding a 
most repulsive secretion from its anal glands, which 
it evidently controls, as it ceases doing so when 
accustomed to being handled. The same snake also 
produces, during the spring, an oily exudation from 
the skin which has the same repulsive smell. 
Mr. H. N. Ridley has observed a Malay snake 
allied to Tropidonotus, Macropisthodon rhodomelas, to 
exude drops of a white viscid liquid from the skin of 
its neck, which is flattened out like that of a Cobra 
when in an attitude of defence, and he noticed that 
his dog, seizing the snake to worry it, foamed at the 
mouth as if he had been biting a toad. 

The hissing is produced by the rapid expulsion of 
air from the lungs through the trachea and the notch 
at the end of the mouth, which is kept shut at the 
time. Snakes provided with an epiglottis (see p. 79) 
produce a much louder hissing. Other sounds are 
produced by some snakes. Thus, the Indian and 
African Vipers of the genera Echis and Cerastes make 
a curious, prolonged, rustling noise, by rubbing the 
folds of the sides of the body against one another. 
This sound is produced by friction between the 
serrated keels of the lateral scales, which are dis- 
posed obliquely with their tips directed downwards 
and backwards ; the noise can even be repeated 
after the death of the animal, by twisting the body 
and thus rubbing or rasping these little saws against 



HABITS 103 

one another. The same thing probably takes place 
in the African genus Dasypeltis, in which we find a 
similar arrangement of the scales, though to a less 
degree. 

The best known sounding apparatus is that of the 
Rattlesnakes, described on p. 20. When alarmed, 
these snakes gather the body in a few coils or roll 
themselves up in a spiral, with the tail erect in the 
centre, and vibrating with great rapidity, whilst the 
head is ready for attack. Other snakes, such as the 
Ancistrodon and some species of Coluber and 
Zamenis, when excited, vibrate the tail in the same 
manner ; but, being deprived of the sound-producing 
apparatus, this expression of their anger does not 
attract the same attention. It is from such a habit, 
however, that the rattle must have been evolved and 
perfected, not necessarily in a Lamarckian sense, but 
through the different steps by which evolution or 
creation has proceeded ; Natura non fecit saltics, as 
Linnaeus well said. Many suggestions have been 
made as to the use of the rattle. One of them is 
that the rattling resembles the sound made by 
locusts, and serves to decoy insect-eating birds ; 
another, that it serves to call the sexes together. 
Probably it is useful to the snake as a warning to 
keep off disturbers which cannot serve as food, and 
thus prevents useless expenditure of venom, or even 
the breaking of the fangs. At any rate, it gives ex- 
pression to the snake's excitement, as does the voice 



io 4 INTRODUCTION 

in the case of many other animals, and it seems 
reasonable to suppose that it may be applied to 
different purposes. With the advent of man, this 
means of attracting attention must tend to the more 
rapid extermination of the snakes which possess it. 

Another curious behaviour is that of feigning death, 
as observed in a harmless but vicious-looking snake, 
Heterodon, often called Puff-adder in America. It 
looks more like a Viper than a harmless snake, and 
when disturbed hisses loudly and flattens out the 
anterior part of the body, much as does a Cobra, and 
pretends to strike, although it is one of the few 
snakes that never bite man. If, however, this display 
proves of no avail in frightening away the intruder, 
the snake rolls on its back and opens its mouth, and 
then lies for a time, which may exceed a quarter of 
an hour, absolutely motionless, as if dead. As soon 
as it thinks the danger over, it awakens from its 
spasm and rapidly moves off. It is the opinion of 
those who have most experience of this snake that 
this extraordinary behaviour is not to be explained 
as a convulsion or faint due to fright, but constitutes 
a deliberate trick to save its life. Individuals of the 
South African Ringhals (Sepedon hmnachates) and of 
the Common Grass-snake have also been observed to 
feign death. 

The notion that snakes fascinate their prey, 
attracting it or reducing it to immobility by a 
mysterious power in their glittering eyes, is pure 



HABITS 105 

fable. Animals placed in a cage with a snake evince 
no particular fright, and fly away when pursued, if 
not actually turning round to defend themselves. 
It is even dangerous to offer a good-sized snake a 
wild rat for food, as all keepers of menageries know. 

In cold and temperate climates snakes hibernate, 
lying more or less torpid in holes or hollow trees, 
sometimes assembled in numbers and coiled 
together in a mass. The first thing they do in 
awakening in the spring is to cast the outer coating 
of the epidermis, as described above (p. 20). Several 
exuviations take place during the period of activity, 
sometimes pretty regularly every month, sometimes 
at very irregular intervals. A few days previous to 
this operation the snake is languid and abstains from 
feeding; its skin is dull and the sight impaired by 
the opaque condition of the lid ; a day or two before 
moulting, the outer stratum of the epidermis 
becomes again transparent and the eye clear, 
through this stratum becoming detached from the 
subjacent tissue, until it is pulled off in one piece, 
by the snake rubbing itself against stones or bushes. 
The first exuviation takes place very shortly after 
birth. 

Snakes are long-lived, although the limit of dura- 
tion of life is not known in any of them. They grow 
slowly, and do not appear ever to reach sexual 
maturity until the fourth year, when they continue 
increasing in size for a long period. A Python retic- 



106 INTRODUCTION 

ulatus and an Ancistrodon piscivorus are reported to 
have lived twenty-one years in captivity in Paris. 
The young of many snakes are very secretive, and 
are not often found in the open, those that are met 
with being as a rule either new-born or approaching 
sexual maturity. 

Snakes are tenacious of life, and remarkable for 
the reflex movements which take place after they 
have been cut to pieces, the severed parts of the body 
and tail wriggling for a considerable time, and the 
head endeavouring to bite. Accounts of decapitated 
Rattlesnakes turning round and striking with their 
bloody stumps are probably not snake stories. 



CHAPTER XI 
PARASITES 

LIKE all other animals, snakes are infested with 
a multitude of vegetable and animal parasites, 
both external and internal. About 300 species of 
Ophidian parasites have been recorded ; yet our 
knowledge of them is very imperfect. Although 
some 2,000 species of snakes are known, parasites 
have not been recorded for more than 168 species, 
and in the great majority of these (102) only a single 
parasite : a tick, a hsemogregarine, or some intestinal 
worm. Owing to the more frequent opportunity of 
dissecting them, the common menagerie snakes have 
yielded better records, notwithstanding the fact that 
they usually lose most of their parasites through 
constant handling, prolonged fasting, and artificial 
surroundings. Thus, we have a list of thirteen species 
for the Indian Python molurus, and one of twenty-two 
species for the Boa constrictor. But no systematic 
search appears to have been attempted, save, perhaps, 
in the case of a few European species. 

It is interesting to notice that it was the finding of 
an Ophidian parasite which prompted Francesco 
Redi to write his famous " Observations on the Living 

107 



io8 INTRODUCTION 

Animals which are found within Living Animals." 
This work, a veritable treatise of comparative 
parasitology, published in 1684, caused the great 
naturalist, physician, and poet to be regarded as the 
father of that science. He tells us that in dissecting 
a curious dicephalous Viper a aspis, caught at Pisa, he 
found within the intestines a number of roundworms 
(Ascaris cephaloptera), and on the surface of one of 
the two lobes of the liver five cysts enclosing a small 
worm, which he rightly ascribed to the same species. 

The parasites of snakes are here enumerated by 
Dr. L. W. Sambon, in systematic order. 

Arthropoda. — Two families of the class Arach- 
nida, the Ixodidse and the Linguatulidae, furnish 
numerous species parasitic on snakes. 

Of the Ticks (Ixodidae) we find, as a rule, species 
of the genera Amblyomma and Aponomma, the 
latter genus being almost entirely confined to 
Reptiles. A single species of the genus Hcema- 
physalis (H. punctata, Can. and Franz, 1877) has 
been reported once from Viper a aspis. A few larval 
forms found on various snakes have been reported 
under the generic name Ixodes, but they probably 
belong either to A mblyomma or Aponomma. 

The Ophidian Tick-parasites, like those of 
mammals, birds, lizards, and tortoises, appear to 
be in many cases the means of transmission of 
protozoal infections from snake to snake. 

The Tongue - worms (Linguatulidae) are, with- 



PARASITES 109 

out doubt, of the greatest possible interest. Their 
systematic position has ever been a puzzle to 
zoologists, and even now is a matter of contro- 
versy. They have been looked upon as Hirudtnea 
by Winsberg (1765), Cestoda by Chabert (1787), 
Acanthocephala by Humboldt (1808), Trematoda by 
Rudolphi (1809), and Nematoda by Nordmann (1832). 
It was Van Beneden (1848) who first recognized 
their Arthropod nature, but he placed them amongst 
the Crustacea. Schubart (1853) suggested that their 
proper position is amongst the Mites (Acarina), and 
Leuckart (i860) adduced important anatomical and 
embryological evidence in support of this view, 
which was confirmed by Railliet in 1883 and by 
Sambon in 1910. 

No less than three out of the four genera of 
Linguatulids so far established are represented by 
species parasitic on snakes. They are the genera 
Porocephalus, Reighardia, and Raillietiella. 

The genus Porocephalus is of special interest, 
because some of its species, such as Porocephalus 
armillatus, a parasite of African Pythons (Python 
regius, P. sebce) and Puff-adders (Bitis arietans, 
B. nasicomis, B. gabonica), and Porocephalus monili- 
formis, a parasite of Oriental Pythons {Python 
molurus, P. reticulatus), are, in their nymphal stage, 
deadly parasites of mammals, including man. 

The genus Reighardia was established by Professor 
H. B. Ward, in 1899, for a Linguatulid of gulls and 



no INTRODUCTION 

terns, first described, in 1861, by De Filippi. In 
igio Sambon included in this genus other similarly 
structured Linguatulids from crocodiles, monitors, 
and snakes. 

The genus Raillietiella was established by Sambon 
in 1910 for a Linguatulid {Raillietiella boidengeri) of 
the African Puff-adders (Bitis arietans, B. gabonica). 
Amongst the characters of this genus is one of 
great structural and phylogenetic importance — viz., 
the position of the female sexual orifice at the 
anterior end of the abdomen, whilst in the other 
known genera it is at the posterior extremity. 

According to Prowazek, Sambon, and Laveran, the 
Ophidian Linguatulids, which live as blood-suckers 
in the air-passages of their hosts, are able to 
foster and transmit the haemogregarines of these 
hosts. 

Acanthocephala. — The early encysted stages of 
several species of Thorn-headed worms (Acantho- 
cephala), belonging to the family Echinorhynchidcu, 
have been reported from snakes belonging to very 
different genera, such as Boa, Tropidonotus, Zamenis, 
Drymobius, Xenodon, Dipsadomorphus, Oxyrhopus, 
Erythrolamprus, Diemenia, Naja, Elaps, Vipera, 
Lachesis. Their further development probably 
occurs in ophiophagous birds. Thus, Echinorhynchus 
oligacanthoides, Rud., the immature stages of which 
occur encapsuled within the body cavity of Lachesis 
lanceolatus and other neotropical snakes, when adult 



PARASITES in 

is found attached to the intestinal mucosa of Milvus 
bidcntatus. 

Nematoda. — The roundworms (Nematoda) so 
far described from snakes belong to the fami- 
lies Ascaridae, Strongylidae, Trichotrachelidae, and 
Filariidae. Some of the genera belonging to these 
families, such as Cucullanus, Nematoxys, Oxysoma, 
are as yet represented by a single species in a single 
host ; others, such as A scaris, Polydelphis, Heierakis, 
Strongylus, Diaphanocephalus, Physaloptera, Tricho- 
soma, number already several species more or less 
widely distributed. 

Eelworm infection (ascariasis) is very common in 
snakes, and not infrequently the infection is a heavy 
one; Sambon twice found over fifty specimens 
of Polydelphis in Puff-adders (Bitis arietans). This 
investigator has shown that the snake eelworms 
undergo an encysted stage of development within 
the body cavity of their hosts before migrating into 
the intestinal lumen for the purpose of fertilization 
and oviposition. Thus, Redi was quite right in 
considering the immature, encysted forms found in 
one of the livers of his double-headed Asp as 
belonging to the same species of eelworm (Ascaris 
cephaloptera) as that which the snake harboured in 
its intestine. 

Professor A. Railliet, whilst examining specimens 
of Polydelphis which had been preserved for nearly 
two months in a 3 per cent, solution of formalin, 



112 



INTRODUCTION 



found that the ova within their uterine tubes had 
undergone development, and still contained living 
embryos ; indeed, some of these hatched under the 
microscope, and moved very actively in the preserv- 
ing fluid. This is in no way surprising, because 
even after several years of preservation in formalin 
solution the embryos of other species of eelworms 
(Ascaris equorum, A. marginata) have been found in 
a living condition. 

Trematoda. — The Flukes {Trematoda) of snakes, 
so far described, belong to the following genera : 
Agamodistomum, Astiotrema, Brachylaimus, Cotylo- 
tretus, Dicroccelium, Diplodiscus, Distoma, Halipegus, 
Lecithodendrium, Metorchis, Opisthogonimus, Opis- 
thorchis, Plagiorchis, Saphedera, Telorchis, Tetracotyle, 
Zeugorchis. 

Cestoda. — Save a few larval forms (Cysticercoides, 
Piestocystis, Sparganum), the known tapeworms 
(Cestoda) of the Ophidia belong to the genera 
Bothridium and Proteocephalus. 

Protozoa. — Numerous species of Hsemogregarines 
have been described from snakes. As a rule the 
forms seen in the peripheral blood are sporonts, the 
schizogonic cycle occurring in the lungs. The 
sporonts do not greatly alter their host cells ; they 
are invariably doubled up within a more or less 
thick capsule. Some species show a marked sexual 
differentiation, others not. Tvypanosomes, Spiro- 
echaudinnicz, and Plasmodidcz have also been de- 
scribed from the blood of various snakes. 



PARASITES 113 

Within the alimentary tube have been found 
species of Trichomonas and Caryospora. 

Bacteria. — Acid-fast bacilli have been described 
in tubercular lesions found in snakes by Sibley, 
Gibbs and Shurley, Shattock, Hausemann, and 
Sambon. 

The so-called " canker," which so frequently 
develops in the oral cavity of captive snakes, is also 
a bacterial disease, due to a specific bacterium of 
thick, rod-shaped form. 

LIST OF PARASITES HITHERTO 
RECORDED FROM EUROPEAN SNAKES 

TROPIDONOTUS NATRIX, L. 
ACANTHOCEPHALA. 

Echinorhynchus incequalis, Rudolphi. 

Echinorhynchus poly acanthus, Creplin. 
Nematoda. 

Strongylus auricularis, Zeder. 

Strongylus catanensis, Rizzo. 

Trichosoma mingazzini, Rizzo. 

Oxysoma brevicaudatum, Zeder. 

Nematoxys commutatus, Rudolphi. 

Ascaris cephaloptera, Rudolphi. 
Trematoda. 

Opistorchis caudatum, Polonio. 

Dicroccelium assida, Dujardin. 

Diplodiscus conicum, Polonio. 

Tetracotyle colubri, v. Lin stow. 
8 



ii4 INTRODUCTION 

Distoma acervocalciferum, Gastaldi. 
Distoma allostomum, Diesing. 
Distoma neniatoides, Miihling. 
Saphedera naja, Rudolphi. 
Brachylaimus signatum, Dujardin. 
Telorchis ercolanii, Monticelli. 
Lecithodendrium nigrovenosum, Bellingham. 
Plagiorchis mentulatus, Rudolphi. 
Cestoda. 

Ligula panceri, Polonio. 

TROPIDONOTUS TESSELLATUS, Laur. 

Nematoda. 

Strongylus denudatus, Rudolphi. 

Physaloptera abbreviata, Rudolphi. 

Physaloptera striata, v. Linstow. 
Trematoda. 

Plagiorchis mentulatus, Rudolphi. 

TROPIDONOTUS VIPERINUS, Latr. 
ACANTHOCEPHALA. 

Echinorhynchus lobianchii, Monticelli. 
Trematoda. 

Distoma allostomum, Diesing. 

Opisthorchis caudatum, Polonio. 

Telorchis ercolanii, Monticelli. 

Astiotrema monticellii, Stossich. 
Cestoda. 

Ligula pancerii, Polonio. 



PARASITES 115 

Protozoa. 

Hamogregarina viperina, Billet. 

ZAMENIS GEMONENSIS, Laur. 
ACANTHOCEPHALA. 

Echinorhynchus cinctus, Rudolphi. 

Echinorhynchus poly acanthus, Creplin. 

Echinorhynchus heterorhynchus, Parona. 
Nematoda. 

Strongylus catanensis, Rizzo. 

Filaria parvomucronata, Rizzo. 

Trichosoma sonsinoi, Parona. 
Trematoda. 

Distoma subflavum, Sonsino. 

Brachylaimus baraldii, Sonsino. 

Saphedera naja, Rudolphi. 
Cestoda. 

Cysticercus acanthotetra, Parona. 

Cysticercoides rostratus, Mingazzini. 

COLUBER QUATUORLINEATUS, Lacep. 
ACANTHOCEPHALA. 

Echinorhynchus oligacanthus f Rudolphi. 
Nematoda. 

Ascaris cephaloptera, Rudolphi. 
Trematoda. 

Plagiorchis sauromates, Poirier. 
Protozoa. 

Hcemogregarina , sp. 



n6 INTRODUCTION 

COLUBER LONGISSIMUS, Laur. 

Protozoa. 

Hcemogregarina colubri, Borner. 

CORONELLA AUSTRIACA, Laur. 
Nematoda. 

Tricheilonema megalochihim, Diesing. 

Physaloptera colubri, Rudolphi. 
Cestoda. 

Piestocystis dithyridium, Diesing. 
Protozoa. 

Monocercomonas colubrorum, Hammersch. 

CORONELLA GIRONDICA, Daud. 

Protozoa. 

Hatnogregarina coronellce, Franca. 

VIPERA BERUS, L. 

Nematoda. 

Physaloptera dentata, v. Linstow. 
Trematoda. 

Agamodistomum viper ce, v. Linstow. 

Tetracotyle colubri, v. Linstow. 

VIPERA ASPIS, L. 

Arthropoda. 

Hcztnaphysalis punctata, Can. & Franz. 

ACANTHOCEPHALA. 

Echinorhynchus cinctus, Rudolphi. 



PARASITES 117 

Nematoda. 

Ascaris cephaloptera, Rudolphi. 

Diaphanocephalus viperce, Rudolphi. 
Protozoa. 

Caryospora simplex, Leger. 

Hczmogregarina samboni, Giordano. 

VIPERA AMMODYTES, L. 

Nematoda. 

Ascaris ammodytis, Rudolphi. 
Ascaris cephaloptera, Rudolphi. 



CHAPTER XII 
DISTRIBUTION 

REPRESENTATIVES of the order Ophidia are 
found over the whole world, with the exception 
of Iceland, Ireland, and New Zealand, between the 
Northern limit of 6y° in Europe (Vipera berus), 6o° in 
Asia (Vipera berus), and 52 in America {Tropidonotus 
ordinatus), and the Southern limit of 44 (Philodryas 
schotti). The highest altitudes reached by them are 
14,000 feet in the Himalayas (Tropidonotus baileyi), 
9,700 feet in the Alps (Vipera aspis), and 9,000 feet 
in the Andes (Liophis albiventris). They are most 
numerous between the tropics, and the number of 
species gradually diminishes to the North and South. 

For the purpose of showing the distribution of the 
principal groups, we will follow the divisions into 
families and subfamilies enumerated above (p. 4). 

Typhlopidcz. — S.E. Europe, S. Asia, Africa, Aus- 
tralia (exclusive of Tasmania), C. and S. America, 
and W. Indies. 

GlauconiidcB. — S. Asia (as far E. as Sind), Africa 
(exclusive of Madagascar), C. America (extending into 
the S. parts of N. America), S. America. 

Pythonincz. — S. Asia, Africa (exclusive of Mada- 
gascar), Australia (exclusive of Tasmania), C. America. 

118 



DISTRIBUTION 119 

Boincz. — S.E. Europe, C. and S. Asia, N. Africa, 
Madagascar, Mauritius, W. Polynesia, S.W. of 
N. America, C. and S. America, and W. Indies. 

Ilysiidtz. — S.E. Asia, S. America. 

Uropeltidcz. — India and Ceylon. 

Xenopeltidce. — S.E. Asia. 

Acrochordincz. — S.E. Asia, C. America. 

Colubvince. — The whole range of Ophidia, except 
Tasmania. 

Dasypeltincz. — Africa (exclusive of Madagascar). 

Homalopsincz. — S.E. Asia, N. Australia. 

Dipsadomorphincz. — S. Europe, C. and S. Asia, 
Africa, Australia (exclusive of Tasmania), C. America 
(extending into the S. parts of N. America), 
S. America. 

Elachistodontincz. — India. 

Hydrophiincz. — Indian and Pacific Oceans. 

Elapincz. — S. Asia, Africa (exclusive of Madagascar), 
Australia and Tasmania, Fiji Islands, C. America 
(extending into the S. parts of N. America), 
S. America. 

Amblycephalidcz. — S.E. Asia, C. and S. America. 

Viperincz. — Europe, Asia, Africa (exclusive of 
Madagascar). 

Crotalincz. — S.E. Europe, Asia, America. 

The Zoogeographical Regions into which the world 
is usually divided (Palaearctic or Europo-Asiatic, 
Oriental or Indian, Ethiopian or African, Australian, 
Nearctic or North American, Neotropical or South 



120 INTRODUCTION 

American) do not lend themselves any better than 
the ordinary divisions of physical geography to the 
study of the distribution of Snakes. Contrary to 
what we find in dealing with the Tortoises, Australia 
does not show any special affinity to South America, 
and, as in the case of the Lizards, it must be 
regarded as an impoverished extension of the Indo- 
Malay fauna; as with the Lizards, also, Europe and 
Africa hang together, whilst Madagascar stands apart, 
distinguished by many negative features and some 
points of agreement with South America (Boidse). 
There is a greater difference between the Snakes of 
Europe and those of Eastern Asia than there is 
between the latter and those of North America, 
whilst in Lizards a primary distinction must be made 
between the Old World and the New. Southern 
Asia east of Persia (the Oriental Region) is the great 
Ophidian centre, all the groups mentioned above, with 
the exception of the Dasypeltinse, having representa- 
tives within its limits, and a large and very distinct 
family, the Uropeltidse, being confined to it. The 
Pythoninse occur along with the Boinae, the Viperinae 
with the Crotalinae, and the Elapinae are represented 
by varied forms, as they are also in Africa and still 
more in Australia, where they form the overwhelm- 
ing majority, and in some parts, as well as in 
Tasmania, the exclusive Ophidian population. The 
coasts of India and Malaya are also the home of the 
great majority of the Hydrophiinse. Large genera 



DISTRIBUTION 121 

like Tropidonotus, Zamenis, and Coluber, extend over 
the Europo-Asiatic and North American regions, but 
they are equally well represented in the Oriental. 
The great difference between Madagascar and Africa 
is, as we have said, very striking. Madagascar pos- 
sesses Boidae generically identical with those of South 
America, but otherwise only Typhlopidae, Colubrinae, 
and Dipsadomorphinae ; whilst in the greater part of 
Africa the Boinae are replaced by the Pythoninae, and 
the Glauconiidas, Elapinae, and Viperinae are gener- 
ally distributed. North America agrees with Asia 
and South America in its Crotalinae, otherwise its 
Ophidian fauna is not very different from that of 
Europe, although much richer, and South America 
shares the Glauconiidae with Africa and the Ilysiidae 
with Southern Asia. South America is rich in 
Colubrinae and Dipsadomorphinae, nearly all generi- 
cally different from those of other parts of the world, 
and the Elapinae are represented by the single genus 
Elaps, with many species, two of which extend to 
the southern parts of North America. 

This rapid sketch of the principal facts of Ophidian 
distribution suffices to show how difficult it would be 
to frame geographical regions that would give expres- 
sion to these facts. Such regions would necessarily 
be very different from those adopted in dealing with 
the distribution of the other divisions of the class 
Reptilia. This is a task which need not be attempted 
on the present occasion. 



122 INTRODUCTION 

A few words as to the salient characters of 
the European fauna, which is a poor one as com- 
pared with other parts of the world. The single 
species of the genera Typhlops and Eryx must be 
regarded as outposts from South-Western Asia ; the 
single species of Ancistrodon, which extends from 
Central Asia into a very small territory to the south- 
east, is also an Asiatic type. The genera Tropido- 
notus, Z amends t Coluber, Coronella, and Contia, are 
characteristic of the Northern Hemisphere, and the 
first three are, besides, equally well represented in 
the Oriental region ; a few species of Tropidonotus are 
also found in Africa and Madagascar. Ccelopeltis, 
Macroprotodon, and Tarbophis are the northern out- 
posts of an Afro- Indian group, although, with 
the exception of the third, exclusively confined to 
the circum-Mediterranean district. The genus Vipera 
is also represented in East Africa and in Southern 
Asia, but the species V. berus is essentially a northern 
type, extending to the highest latitude reached by 
any snake, and ranging all over Northern Asia to the 
Amur and Sachalien. The same species reaches the 
greatest altitude at which any snake has been observed 
on the northern side of the Alps — viz., 9,000 feet. 

Of the twenty-eight species inhabiting Europe, 
only two are generally distributed : Tropidonotus 
natrix and Coronella austriaca. One is to be regarded 
as a northern form, although occurring locally in the 
south : Vipera berus. It is the reverse with Coluber 



DISTRIBUTION 123 

longissitnus. The others may be described as southern 
forms, two only as ranging from west to east: 
Zamenis gemonensis and Ccelopeltis monspessulana ; 
one of more central habitat: Vipera ursinii. The 
remainder may be divided into two groups — those of 
more western, and those of more eastern distribution. 
To the first group belong Tropidonotus viperinus, 
Zamenis hippocrepis, Coluber scalaris, Coronella giron- 
dica, Macroprotodon cuctdlatus, Vipera aspis and Vipera 
latastii ; to the second, Typhlops vermicidaris f Eryx 
jaculus, Tropidonotus tessellatus, Zamenis dahlii, Colu- 
ber quatuorlineatus, dione, leopardinus, Contia modesta, 
Tarbophis fallax and iberus, Vipera renardi, ammodytes, 
lebetina, and Ancistrodon halys. 

A remarkable fact in the distribution of Euro- 
pean Snakes is the altitudinal range of Vipera berus, 
V. aspis, and V. ursinii. The first being the northern- 
most snake, generally distributed in Northern 
Europe and more locally in the south, should, one 
would expect, be a mountain form in the south. 
This is so in Switzerland, where it occurs chiefly 
between 2,500 and 9,000 feet, on the northern aspect 
of the Alps, whilst V. aspis lives at altitudes below 
5,000 feet ; but on the southern aspect of the same 
chain things are reversed, and V. berus is replaced 
by V. aspis, which reaches an altitude of 9,700 feet, 
whilst the former shows a tendency to abandon the 
mountains, and has established ifself in a few locali- 
ties in the plain of North Italy. Again, in France 



124 INTRODUCTION 

V. berus is the northern and V. aspis the southern 
species, yet the latter is the only one found on the 
French side of the Pyrenees (up to 7,250 feet), whilst 
the former reappears in North- Western Spain and 
Portugal at very low altitudes, even at sea-level. 
V. ursinii is a mountain form in Italy (Abruzzi), in 
France (Basses-Alpes), and in the Balkan Peninsula 
(up to 6,800 feet); but it is restricted to the plain in 
Lower Austria and Hungary, where V. berus occurs 
only in the mountains. 

Only three species are entirely confined to Europe : 
Coluber scalaris, Vipera ursinii, and V. aspis. 

Of the species which range outside Europe, the 
following occur both in Western Asia and in North 
Africa : 

Eryx jaculus, Tropidonotus natrix, Ccelopeltis mons- 
pessulana, Vipera lebetina. 

In Western Asia and the North-East of Egypt : 
Tropidonotus tessellatus, Zamenis dahlii. 

In Western Asia : 

Typhlops vermicularis, Zamenis gemonensis, Coluber 
quatuorlineatus, C, dione, C. longissimus, C. leopardinus, 
Coronella austriaca, Contia modesta, Tarbophis fallax, 
T. iberus, Vipera renardi, V. berus, V. ammodytes, 
Ancistrodon halys. 

In North Africa : 

Macroprotodon cucullatus. 



DISTRIBUTION 125 

In North- West Africa : 

Tropidonotus viperinus, Zamenis hippocrepis, Coronella 
girondica, Vipera latastii. 

The following lists will help to elucidate the 

distribution of the snakes in the different parts of 

Europe : 

I. Scandinavia 

1. Tropidonotus natrix (as far north as 65 ). 

2. Coronella austriaca (as far north as 63 ). 

3. Vipera berus (as far north as 67 ). 

II. Great Britain 

1. Tropidonotus natrix (England and Wales, ex- 
treme south-east of Scotland). 

2. Coronella austriaca (Surrey, Berkshire, Hamp- 
shire, and Dorsetshire). 

3. Vipera berus. 

III. Belgium and Holland 

1. Tropidonotus natrix. 

2. Coronella austriaca. 

3. Vipera berus. 

IV. Germany and Denmark 

1. Tropidonotus natrix. 

2. Tropidonotus tessellatus (Middle Rhine and 
Moselle, Saxony). 

3. Coluber longissimiis (Denmark, Schlangenbad, 
Treves) . 



126 INTRODUCTION 

4. Coronella austriaca. 

5. Vipera bents. 

6. Vipera aspis (Black Forest, Lorraine). 

V. France and Switzerland, exclusive of 

Ticino 

1. Tropidonotus natrix. 

2. Tropidonotus viperinus (as far north as South 
Brittany and Fontainebleau). 

3. Zamenis gemonensis (south, locally as far north 
as the Sarthe and Aube). 

4. Coluber longissimus (locally as far north as 
South Brittany, South Normandy, and Fontaine- 
bleau). 

5. Coluber scalaris (Mediterranean Littoral), 

6. Coronella austriaca. 

7. Coronella girondica (south and west as far 
north as the Charente-Inferieure). 

8. Ccelopeltis monspessulana (Mediterranean Lit- 
toral). 

g. Vipera ursinii (Basses-Alpes). 

10. Vipera bents (as far south as the Loire basin, 
the Central Plateau, and the Alps). 

11. Vipera aspis (as far north as the Loire basin, 
Fontainebleau, and Lorraine). 

VI. Spain and Portugal 

1. Tropidonotus natrix. 

2. Tropidonotus viperinus. 



DISTRIBUTION 127 

3. Zamenis gemonensis (Catalonia). 

4. Zamenis hippocrepis (absent from the north). 

5. Coluber longissimus (Andalucia). 

6. Coluber scalaris. 

7. Coronella austriaca (north and north-west). 

8. Coronella girondica. 

9. Ccelopeltis monspessulana. 

10. Macroprotodon cucullatus (centre and south, 
Baleares). 

11. Vipera berus (north-west). 

12. Vipera aspis (Pyrenees). 

13. Vipera latastii (absent from the north). 

VII. Italy, with Ticino and Corsica 

1. Tropidonotus natrix. 

2. Tropidonotus tessellatus (as far south as Naples ; 
absent from the islands). 

3. Tropidonotus viperinus (Liguria, Piedmont, 
Ticino, Corsica, Sardinia, Sicily). 

4. Zamenis gemonensis. 

5. Zamenis hippocrepis (Sardinia). 

6. Coluber quatuorlineatus (south and Sicily). 

7. Coluber longissimus (absent from Corsica). 

8. Coluber leopardinus (south and Sicily). 

9. Coronella austriaca (absent from Corsica and 
Sardinia). 

10. Coronella girondica (absent from Corsica and 
Sardinia). 

11. Ccelopeltis monspessulana (Western Liguria, 
Sicily). 



128 INTRODUCTION 

12. Viper a ursinii (Abruzzi). 

13. Vipera berus (the Continental part only). 

14. Vipera aspis (absent from Corsica and 
Sardinia). 

15. Vipera ammodytes (Northern Venetia). 

VIII. Austria-Hungary, without Balkan 

States 

1. Tropidonotus natrix. 

2. Tropidonotus tessellatus. 

3. Zamenis gemonensis (South Tyrol, Littoral, 
South Hungary). 

4. Coluber quatuorlineatus (Istria). 

5. Coluber longissimus. 

6. Coluber leopardinus (Istria). 

7. Coronella austriaca. 

8. Coronella girondica (South Tyrol). 

9. Ccelopeltis monspessulana (Istria). 

10. Tarbophis fallax (Istria). 

11. Vipera ursinii (Lower Austria, Littoral, Hun- 
gary). 

12. Vipera berus. 

13. Vipera aspis (South Tyrol, Littoral). 

14. Vipera ammodytes (South Tyrol, Styria, 
Carinthia, Carniola, Littoral, South Hungary). 

IX. Balkan Peninsula and Archipelago 

1. Typhlops vermicularis (Greece, Turkey, Bul- 
garia). 



DISTRIBUTION 129 

2. Eryx jaculus (Greece, Turkey, Roumania). 

3. Tropidonotus natrix. 

4. Tropidonotus tessellatus. 

5. Zamenis gemonensis. 

6. Zamenis dahlii (coast of the Adriatic, Greece). 

7. Coluber quatuorlineatus. 

8. Coluber longissimus. 

9. Coluber leopardinus. 

10. Coronella austriaca. 

11. Ccelopeltis monspessulana (West Coast, Greece, 
and islands). 

12. Tarbophis fallax (West Coast, Greece and 
islands, Constantinople). 

13. Viper a ursinii (Bulgaria, Bosnia, Herzegovina, 
Montenegro). 

14. Vipera berus (Bosnia, Herzegovina, Rou- 
mania). 

15. Vipera aspis (Bosnia). 

16. Vipera ammodytes. 

17. Vipera lebetina (Cyclades). 

X. Russia 

1. Tropidonotus natrix (as far north as 6o°). 

2. Tropidonotus tessellatus (south). 

3. Zamenis gemonensis (south). 

4. Zamenis dahlii (Caucasus). 

5. Coluber quatuorlineatus (south). 

6. Coluber dione (south, between Volga and Ural) 

7. Coluber longissimus (south, Poland) 
9 



i 3 o INTRODUCTION 

8. Coluber leopardinus (Crimea). 

9. Coronella austriaca (as far north as 57 ). 

10. Contia modesta (Caucasus). 

11. Tarbophis iberus (Caucasus). 

12. Vipera renardi (south). 

13. Vipera berus (as far north as 64 ). 

14. Ancistrodon halys (south, between Volga and 
Ural). 

Without attempting anything like a complete 
bibliography, we have compiled a list of faunistic 
works and papers dealing with the snakes of Europe: 

Europe in General 

Schreiber, E. : Herpetologia Europaea. Zweite 
Auflage, Jena, 1912, 8vo. 

Steinheil, F. : Die Europaeischen Schlangen. 
Kupferdrucktafeln nach Photographien der lebenden 
Tiere. Jena, 1913, 4to. (in progress). 

Great Britain 

Bell, T. : A History of British Reptiles. 2nd 
edit., London, 1849, 8vo. 

Cook, M. C. : Our Reptiles. London, 1865, 8vo. 

Leighton, G. : The Life-History of British 
Serpents. Edinburgh and London, 1901, 8vo. 

France 

Gadeau de Kerville, H. : Faune de la Nor- 
mandie. IV. Reptiles. Paris, 1897, 8vo. 



DISTRIBUTION 131 

Lataste, F. : Essai d'une Faune Herp£tologique 
de la Gironde. Bordeaux, 1876, 8vo. 

Martin, R., et Rollinat, R.: Vertebres sauvages 
du Departement de l'lndre. Paris, 1894, 8vo. 

Switzerland 

Fatio, V. : Faune des Vertebres de la Suisse. 
III. Reptiles et Batraciens. Geneva and Basle, 

1872, 8vo. 

Spanish Peninsula 

Bosca, E. : Catalogue des Reptiles et Amphibies 
de la Peninsule Iberique et des lies Baleares (Bull. 
Soc. Zool. France, 1880). 

Italy 

Bonaparte, C. L. : Iconografia della Fauna 
Italica. II. Anfibi. Rome, 1832-1841, fol. 

Camerano, L. : Monografia degli Ofidi Italiani 
(Mem. Ace. Torin., [2] xxxix., 1888, and xli., 1891). 

Germany 

Durigen, B. : Deutschlands Amphibien und 
Reptilien. Magdeburg, 1890-1897, 8vo. 

Leydig, F. : Ueber die Einheimischen Schlan- 
gen (Abh. Senck. Ges., xiii., 1883). 

Austrian Empire 

Werner, F. : Die Reptilien und Amphibien 
Oesterreich-Ungarns und der Occupationslander. 
Vienna, 1897. 8 yo - 



1 32 INTRODUCTION 

Greece 

Bedriaga, J. de : Die Amphibien und Reptilien 
Griechenlands (Bull. Soc. Nat. Mosc, 1881). 

Bory de St. Vincent, J. B. : Expedition Scienti- 
fique de Moree. Reptiles. Paris, 1832-1835, 4to., 
fol. atlas. 

Bulgaria 

Kowatcheff, W. T. : Herpetological Fauna of 
Bulgaria. Philippopolis, 1912, 8vo. [Bulgarian 
text.] 

ROUMANIA 

Kiritzescu, C. : Contribution a la Faune Herpe- 
tologique de Roumanie. Sauriens et Ophidiens 
(Bull. Soc. Rom. Bucarest, x., 1901). 

Russia 

Nikolsky, A.: Herpetologia Rossica. St. Peters- 
burg, 1905, 4to. [Russian text.] 

Strauch, A. : Die Schlangen des Russischen 
Reichs. St. Petersburg, 1873, 4to. 



CHAPTER XIII 
SNAKES IN RELATION TO MAN 

UNDER this head, the question of poisonous 
snakes naturally occupies the first place. In 
addition to what has been said above in Chapter VI., 
dealing with the anatomical and physiological aspects 
of the subject, we have to allude to the accidents 
caused by these dangerous reptiles, and the measures 
taken to combat them. 

The enormous mortality for which snake-bite is 
responsible in India is well known. Statistics estab- 
lish the fact that an average of 20,000 human lives 
are thus lost annually : 24,264 is the official return 
for 1911. In Australia, where highly poisonous snakes 
of various genera and species abound, the fatal cases 
are likewise very numerous, though less in propor- 
tion than in South America, and no doubt also in 
Africa. In the small island of Martinique, the Fer- 
de- Lance, Lachesis lanceolatus, causes every year 
the death of about 100 human creatures. Though 
numerous in species, the poisonous snakes of Ceylon 
cause a comparatively small mortality — 200 per 
annum. 

Modern research has resulted in the discovery of 

i33 



134 INTRODUCTION 

the only effective antidote for snake-venom intoxica- 
tion : the serotherapic treatment. An animal that 
has been treated over a length of time with the 
venom of a poisonous snake, such as a Cobra, yields 
a serum which is antitoxic towards that venom ; but 
the great difficulty resides in the specificity of the 
different poisons, which often renders the use of the 
serum ineffective in countries like India and Australia, 
where several kinds of poisonous snakes occur in the 
same district (see above, p. 67). In India, where a 
special laboratory has been established for the supply 
of antivenine, at the Central Institute of Kasauli, it 
has been found impossible to obtain any venoms but 
those of the Cobra and Russell's Viper in sufficient 
quantity to immunize animals, and thus produce the 
serum necessary for dealing with the bite of the 
King Cobra, the Krait, and the Echis Viper. 

In Pondicherry the French Government places 
annually a sum of 200 rupees at the disposal of the 
director of the hospital for obtaining Cobra poison, 
the snakes, to be brought alive, being paid for to the 
natives at the rate of half a rupee to one rupee each, 
according to size and condition. Six hundred and 
fifty-three specimens were thus purchased in less 
than two years (1901-1903). The poison is utilized 
for the preparation of Calmette's antivenine, which, 
as we have said above, is only effective against cobra 
poison, and, unfortunately, useless for the cure of 
bites from other species. 



SNAKES IN RELATION TO MAN 135 

In Brazil, where the number of accidents is esti- 
mated at 19,200 per annum, and that of fatal cases 
at 4,800, over 2,000 snakes (Lachesis and Crotalus) are 
brought annually to the Serotherapic Institute of 
Batantan, in the province of S. Paolo, for the prep- 
aration of the antitoxic serum, which is given in 
exchange for the snakes. According to the latest 
report of the Institute (191 1), two serums are dis- 
tributed : the anti-crotaline (for Rattlesnake bite) 
and the anti-bothropine (for Lachesis bite) ; the 
third, the anti-elapine (for Coral-snake bite), is in 
course of preparation. 

In many countries a premium has for years been 
paid for the heads of poisonous snakes, and has led 
to the destruction of enormous numbers of them, 
without, however, resulting in a very appreciable 
diminution of the dangerous reptiles. More than 
£12,000 has been spent for this purpose in India alone ; 
the numbers destroyed in 1885 and 1886 throughout 
British India amount to 420,044 and 417,596 respec- 
tively. About forty years ago the Governor of St. 
Lucia offered a reward of 4d. for every Fer-de-Lance's 
head. But the negroes caught them alive and bred 
families of snakes for the sake of the reward, and 
thereby made what was for them a little fortune, these 
snakes bringing forth up to sixty young at a birth. 
The reward had to be abolished very soon. 

Now about the Vipers of Europe, the only really 
dangerous snakes of this part of the world. 



136 INTRODUCTION 

Although the Adder, Vtpera berus, is quite common 
in many parts of England and Scotland, accidents 
caused by its bite are rarely heard of, and cases of 
death are few and far between. It is not so, how- 
ever, on the Continent, where the same species, and 
especially its close ally, the more southern V. aspis, 
are responsible for many fatalities, due no doubt to 
the more virulent action of the venom in a warmer 
climate. 

In the French Departments Loire-Inferieure and 
Vendee, where these snakes are very plentiful, three 
or four cases of death are reported annually. From 
i860 to 1868, 370 serious accidents to man have been 
carefully recorded, 53 ending in death, not only in 
the case of children, but also of adults of all ages, 
in 10 cases within one to twenty-four hours. In 
the Puy-de-D6me cases of death are of frequent 
occurrence. In Germany and in Switzerland, 12 or 
13 percent, of the cases on record have ended fatally. 
Instances of death from the bite of the south-eastern 
V. ammodytes are also not infrequent. On the other 
hand, the bite of V. ursinii, which is but seldom 
inflicted, is not known to have ever resulted in death. 

It must be borne in mind that accidents are much 
more frequent in districts where the poorer classes 
are in the habit of going about barefoot. 

Anyhow, it is certain that Vipers are a serious danger 
in many parts of Europe, not only to man, but also 
to horses, cattle, and dogs. And it is not surprising 



SNAKES IN RELATION TO MAN 137 

that efforts have been made to reduce their numbers. 
The most efficacious means, besides the protection 
of certain animals and birds which feed on Vipers, 
appeared to be the institution of premiums to be 
paid for the heads of the dangerous snakes. By 
offering 2jd. per head, 500,000 Vipers (V. aspis) were 
destroyed from 1864 to 1890 in three French depart- 
ments, Haute-Saone, Doubs, and Jura, and in one 
district (Chaumont) of the Haute-Marne 57,045 were 
killed from 1856 to 1861 ; this gives an idea of the 
extraordinary abundance of these snakes in some 
parts of France. In the Puy-de-D6me the premium 
was fixed for a time at 5d., and one man managed to 
destroy in the course of seven years 9,175 Vipers 
(V. berus and V. aspis). A woman in the Deux- 
Sevres has made a living for many years by catching 
Vipers, the heads of which were paid to her at the 
rate of 5d. each. The average number of her cap- 
tures amounted to 2,062 per annum (mostly V. aspis). 
Around Oesnitz in Saxony, 2,140 V. berus were killed 
in 1889, and 3,335 in 1890. In a single district in 
Southern Styria the heads of 4,197 V. berus and 
7,381 V. ammodytes were sent in for the reward in 
the course of two years (1892, 1893). 

In spite of all this effort, the institution of the 
bounty has not answered expectations, and, with 
the exception of a few districts, Vipers remain as 
plentiful as ever, showing what little man can do in 
altering the equilibrium of Nature, except by inter- 



138 INTRODUCTION 

fering with the natural conditions under which 
animals live. Cultivation of the ground or destruc- 
tion by fire of the vegetation of the wilderness seems 
to be the only efficacious means of getting rid of so 
abundant and prolific a creature as the Viper. 

A word may be said, however, in defence of 
Vipers : they do a great deal of good to agriculture 
by the destruction of small rodents, on which they 
feed chiefly, and whose multiplication they serve to 
keep in check. It must be pointed out that, with the 
exception of the species of Coluber and Zamenis, other 
European snakes are to be regarded as indirectly 
injurious to agriculture, feeding as they do mainly on 
lizards or frogs and toads, which, as insectivores, 
deserve to be protected. 

Snakes are not of much economic value to man. 
Tanned skins of Boas and Pythons are utilized for 
making shoes and fancy articles, such as purses, 
pocket-books, blotters, etc., and the Siamese make 
the drum-heads of native drums out of the skins of 
Pythons and Acrochordus. To say nothing of savages, 
who seem to be partial to the flesh of large snakes, 
the peasantry in some parts of France do not disdain 
snakes as an article of food, the Grass-snake being 
occasionally served in village inns under the name of 
Anguilles de hates, or hedge-eels. 

Viper fat has for a long time been in request as an 
ointment in the case of various affections, and much 
used by quack doctors in the preparation of their 



SNAKES IN RELATION TO MAN 139 

remedies. Some forty years ago a chemist in Challans 
(Vendee) collected Vipers (V. aspis) for medicinal 
purposes, and was able to send several thousands to 
Paris in the course of a few years, thus realizing a 
considerable sum of money, but the demand has 
gradually fallen off since. 

Very frequent in the past, snake-worship is still 
prevalent in many parts of India, where the Cobra is 
held in great veneration, and is never willingly killed 
by the Hindoo. In pre-Buddhist days the gods 
were represented with a canopy of five or seven 
Cobras over them. The North African Cobra was 
sacred to the ancient Egyptians, and is profusely 
represented on the monuments and tombs ; it was 
also an emblem of the physical sun, and, as a sign of 
royal power, along with the sun's disc, formed part 
of the headdress of all solar deities. The Greeks 
and Romans also worshipped snakes, and the god of 
medicine is represented holding a snake, which is 
supposed to be Coluber longissimus, the so-called 
"iEsculapian snake " ; the occurrence at the present 
day of certain common Italian species {Zamenis 
gemonensis, Coluber longissimus, Tropidonotus tessellatus) 
in isolated localities of Central Europe, formerly 
Roman settlements, has been attributed to their 
importation for use in the temples. 

Snake-charmers have existed from the remotest 
antiquity, and are still to be found among all races 
of men, from the accomplished Indian juggler down 



140 INTRODUCTION 

to the more commonplace European snake-catcher, 
who boasts of his immunity, and of his art of attract- 
ing snakes by devices of which he has the secret. 
The Libyan Psillii of the ancient Romans have 
handed down their art to the present day, and their 
performances are to be witnessed in most of the 
towns of Egypt and Tunisia. But India above all 
lands is reputed for its snake-charmers, and the 
favourite species used by them is the Cobra, which, 
by the way in which it raises the anterior part of the 
body and expands the region behind the head, lends 
itself better than any other to the display. Con- 
stantly facing the man before him, and swaying the 
raised anterior part of the body, it seems to dance 
to the music performed by the snake-man, people 
believing it to be charmed by the sounds of the 
instrument. However, anyone sitting on the ground 
in front of a Cobra, and swaying the body from side 
to side as does the man, can obtain the same result 
without the aid of any sort of music. 

The most puzzling thing about these performances 
is how the man can thus play with impunity with so 
deadly a snake. It is a mistake to think that the 
snake is rendered harmless through the poison fangs 
having been extracted, although this subterfuge is 
frequently resorted to by the less accomplished 
jugglers. The immunity of the snake-charmer is to 
be explained by the fact that the man has submitted 
himself to a series of successive and graduated in- 



SNAKES IN RELATION TO MAN 141 

oculations of the venom, a process similar to vaccina- 
tion, which renders his blood proof against the 
venom of the particular species of snake, and that 
one only, used for his performances. 

Another deadly snake shown by the snake- 
charmers in North Africa is the Horned Viper, 
Cerastes comutus. The presence of an erect spike 
above the eye is, however, not a constant character in 
this snake, and hornless specimens are made to look 
more formidable by spines of the hedgehog being 
inserted in the proper place ; the illusion is such 
that even naturalists have been deceived by this 
trick. 

Indian snake-charmers profess to have a belief in 
the efficacy of snake-stones, or bezoar stones, as 
a remedy to be applied on the part bitten by a 
poisonous snake, a belief shared by the natives of 
many tropical countries. These stones, extracted 
from various reptiles, birds, and mammals, are 
calcareous concretions from the stomach or bladder, 
sometimes composed of superphosphate of lime, some- 
times of phosphate of ammonia or magnesia. The 
value of a bezoar stone being supposed to increase 
with its size, the larger are sold in India at very high 
prices. 

In many places a popular belief prevails that such 
stones are found in the heads of snakes. Mr. J. A. 
Bucknill, now Attorney-General at Hong-Kong, who 
spent five years in Cyprus, has informed the author 



142 INTRODUCTION 

that the Viper of the latter island, Vipera lebetina, is 
commonly believed to contain a stone which, when 
applied to the bite of a poisonous snake, quickly 
nullifies the effect ; it is also believed that, when this 
stone is allowed to stand in a glass of water and the 
water is drunk, it endows the drinker with surprising 
virility. Indeed, there was an action tried by the 
English judge at Larnaka in which the plaintiff 
claimed the return, or damages for the non-return, 
of one of these " Viper-stones " which he had lent 
for a monetary consideration to the defendant for 
the promotion of his manly vigour, and Mr. Buck- 
nill's recollection is that the plaintiff recovered £10 
for the loss. 



SYSTEMATIC ACCOUNT OF THE 
SNAKES OF EUROPE- 
First Family: TYPHLOPID.E 

SKULL compact, with short, toothless lower jaw, 
without transverse bone ; palatine and ptery- 
goid reduced and toothless ; maxillary small, loosely 
attached to lower surface of cranium and bearing a 
few small teeth ; no supratemporal, the quadrate 
articulated to the prootic ; a coronoid element in the 
lower jaw. Rudiments of a pelvic arch, reduced to 
a single bone. Body vermiform, covered with uniform 
cycloid scales ; head small, not distinct from the 
body ; mouth small, crescentic, inferior ; eyes under 
the more or less transparent head-shields, sometimes 
entirely hidden. Worm-like, smooth, shiny snakes, 
of small or very small size, the largest measuring 
little over 2 feet, of subterranean habits, or found in 
rotten trees, under stones, or in the saw-dust of saw- 
mills ; rarely appearing on the surface except when 
the ground is soaked by heavy rains. 

Inhabit the intertropical parts of the whole world, 
as well as South Africa, Southern Asia, and South- 

* For a key to the identification of the species, see above, 
p. 22. 

M3 



144 TYPHLOPIDiE 

ern Australia. One species occurs in South-Eastern 
Europe. About 120 species are known. 

Genus TYPHLOPS, Schneider 

Head with large shields ; nostril in a single or 
divided nasal. Tail extremely short. 

1. Typhlops vermicularis, Merrem 
The Greek Blind-Snake 

Form. — Slender, worm-like, the greatest diameter 
of the body 40 to 52 times in the total length. Tail 




Fig. 14 (after Sordelli) 

about as long as broad, ending in a short spine. 
Snout depressed, rounded, strongly projecting. Eyes 
distinguishable, appearing as a small black spot 
surrounded by an unpigmented circle; nostrils 
lateral. 

Head-Shields. — Rostral about one-third, or a little 
less than one-third, the width of the head, extend- 
ing on its upper surface nearly to the le. .1 of the 



PLATE I 




TVPHLOFS VERMICULARIS 
After Sordelli 





ERYX JACULUS 
A Iter Sotdelli 



9V 



TYPHLOPS 145 

eyes. Nasal incompletely divided, the cleft pro- 
ceeding from the second labial. Preocular present, 
about as broad as the ocular, in contact with the 
second and third labials. Upper head-scales feebly 
enlarged and subequal. Four upper labials. 

Scales. — Equal, 22 or 24 round the body. 

Coloration. — Brown or yellowish-brown above, 
yellowish beneath. 

Total Length. — 10 inches. A specimen from Cyprus 
is reported to measure 14 inches. 

Distribution. — This species has long been known 
from Greece, the Ionian Islands, and the Grecian 
Archipelago. It is on record from the Eli-Deren 
Pass, in Bulgaria. A specimen stated to come from 
Constantinople is preserved in the British Museum. 
The range further extends over a considerable part of 
South-Western Asia, viz., Asia Minor, Syria, Cyprus, 
Transcaucasia, Persia, Turkestan, and Afghanistan. 

Habits. — Pretty alert in its movements, this little 
snake has considerable constricting powers, and coils 
itself fast round the fingers when handled. It lives 
much after the manner of earth-worms, and if dug 
out of loam or sand a specimen must be instantly 
grasped, as it draws back with extraordinary quick- 
ness. Its food probably consists mainly of earth- 
worms and small insects. Some exotic species of 
the genus are known to feed on termites, and are often 
dug out of their nests. 

Reproduction. — No observations have been made 
10 



146 boim: 

that I am aware of, but, as some of the exotic species 
of which we know something more lay large, elon- 
gate eggs, it is probable that this species also is 
oviparous. 

Second Family : BOID^E 

Maxillary, palatine, and pterygoid bones movable; 
transverse bone present ; pterygoid extending to 
quadrate or mandible ; supratemporal present, at- 
tached scale-like to cranium, suspending quadrate ; 
prefrontal in contact with nasal ; a coronoid element 
in the lower jaw. Teeth in both jaws. Vestiges of 
pelvis and hind limbs, usually terminating, at least 
in males, ir a claw-like horny spur on each side of 
the vent. 

This family contains, besides the gigantic Boas 
and Pythons, several small more or less burrowing 
forms, among which the genus Eryx, its only 
European representative, belonging to the sub- 
family Boinse, characterized by the absence of a 
supratemporal bone and of premaxillary teeth. 
This subfamily, the largest members of which inhabit 
tropical America, is distributed over the hotter 
parts of America, Asia west of the Bay of Bengal, 
Madagascar, the Mascarene Islands, Africa north of 
the equator, Papuasia,and some islands of the South 
Pacific. The habitat of the European species is 
confined to the eastern and southern countries of 
the Mediterranean district. 

This very varied family, including terrestrial, 



ERYX 147 

arboreal, aquatic, and burrowing forms, is a compara- 
tively small one as regards the number of species, 
viz., about sixty, of which one-third pertain to the 
Pythoninae, which inhabit tropical and South Africa, 
Southern Asia, Papuasia, Australia, and Mexico. 

Genus ERYX, Daudin 

Anterior maxillary and mandibular teeth longer 
than the posterior. Head small, not distinct from 
neck, covered with small scales ; a large rostral 
shield. Eye very small, with vertical pupil. Body 
cylindrical ; scales small ; ventral shields narrow. 
Tail very short ; subcaudal shields mostly single. 

The range of this genus, embracing eight species, 
extends from South-Eastern Europe and Africa north 
of the equator to Central Asia and India. 

2. Eryx jaculus, Linnaeus 
The Javelin Sand-Boa 

Form. — Stout. Head small, not distinct from 
neck ; snout projecting beyond the mouth ; eye 
directed upwards and outwards ; a feeble mental 
groove. Tail, ending very obtusely, one-tenth to 
one-sixteenth of the total length. Anal spurs more 
or less developed, often absent in the female. 

Head-Shields. — Rostral very large and broad, with 
angular horizontal edge, followed by a pair of inter- 
nasals and a second row of two or three small shields, 



148 



BOIDM 



the rest of the upper surface of the head covered with 
scale-like shields, 5 to 8 from eye to eye across the 
vertex, 7 to 11 round the eye ; 9 to 12 upper labials, 
second or third deepest, separated from the eye by 
one or two series of scales. Nostril between the 
internasal and two nasals, the anterior of which 
sometimes fuses with the former. Two or three 
series of scales between the nasals and the eye. 






Fig. 15 (after Sordelli) 

Scales. — Smooth, feebly keeled on the posterior 
part of the body and on the tail, in 40 to 51 rows. 
Ventrals narrow, occupying about one-third of the 
ventral surface, 163 to 200 ; anal small, entire ; sub- 
caudals all or greater part single, 15 to 29. 

Coloration. — Pale greyish, reddish, or yellowish- 
brown above, with brown, purplish-brown, or black- 
ish markings, which may be very irregular or form a 
single or alternating series of large blotches or cross- 
bands on the back ; the sides with smaller spots ; 
these markings may be confluent and so large as to 



ERYX 149 

reduce the ground colour to small yellowish spots ; 
one, two, or three short, dark stripes often present on 
the nape ; a dark streak from the eye to the angle of 
the mouth ; sometimes a dark curved band from eye 
to eye across the upper surface of the snout. Lower 
parts yellowish-white, uniform or with small blackish 
spots. 

Size. — 2 \ feet is the greatest length which this 
snake is known to attain. 

Distribution. — Originally described from Lower 
Egypt, and extending westwards to Algeria, this 
Eryx has been found in Greece, in Corfu, in the 
Cyclades, in Turkey, and in Roumania. It occurs 
also in Asia Minor, in Transcaucasia, in Trans- 
caspia, in Northern Persia, and in Syria. It has 
been found at an altitude of 5,000 feet in Persia, to 
the west of Lake Urmia. A closely allied form 
(E. miliaris, Pallas), which has been confounded with 
this species, extends from Transcaspia to Turkestan 
and Afghanistan. 

The reported occurrence of this snake in Bulgaria 
is based on a specimen labelled " Bulgaria (?) " in the 
Sofia University Museum. The species is omitted 
from KovatschefPs latest list of Bulgarian Reptiles. 

Habits. — This diminutive Boid is a burrower in 
arid, sandy districts, appearing only early in the 
morning or towards dusk ; it is as a rule more 
crepuscular than nocturnal. Notwithstanding its 
rather heavy form, it is capable of very quick move- 



150 colubrim: 

ments, darting like an arrow upon its prey, which 
consists chiefly of small mammals and lizards. A 
constrictor, like all the members of the family to 
which it pertains, it crushes its prey before swallowing 
it. If given several mice at a time, it will catch and 
kill them all in succession before proceeding to feed. 
Specimens recently discovered in the Danube Valley 
in Roumania were found to live in the sand at the 
bottom of small limestone caves, going about at 
night and feeding principally on slugs. Unlike other 
snakes, it is said to lap dewdrops with its tongue. 
It is a gentle snake, seldom attempting to bite. 

Egyptian jugglers are in the habit of implanting 
the claw of a bird or small mammal in the skin of 
the head of this snake, above each eye, in order to 
give it a more formidable appearance. 

Reproduction, — Like the other species of Eryx, this 
snake is ovoviviparous, but, beyond this fact, nothing 
appears to have been observed concerning the breed- 
ing habits, although many examples have been kept 
in captivity. 

Third Family: COLUBRID.E 

Maxillary, palatine, and pterygoid bones movable ; 
transverse bone present ; pterygoid extending to 
quadrate or mandible; supratemporal present, at- 
tached scale-like to cranium, suspending quadrate ; 
prefrontal not in contact with nasal ; maxillary 
horizontal, not movable perpendicularly to the trans- 



COLUBRID.E 151 

verse bone ; no coronoid bone. Teeth in both jaws. 
No vestiges of pelvic arch. 

An enormous group, comprising the great majority 
of snakes. Divided into three parallel series : 

A. Aglypha, with all the teeth solid. 

B. Opisthoglypha, with one or more of the pos- 
terior maxillary teeth grooved. 

C. Proteroglypha, with the anterior maxillary teeth 
grooved or canaliculated. 

The third, which is not represented in Europe, 
includes some of the most deadly snakes, such as the 
Cobras, Kraits, Death-adders, etc. 

The European genera are thus distributed in the 
two other series : 

Aglypha (ColubriN;E) : Tropidonotus, Zamenis, 
Coluber, Coronella, Contia. 

Opisthoglypha (Dipsadomorphin^;) : Ccelopeltis, 
Macroprotodon, Tarbophis. 

These genera give but a feeble idea of the 
variety of forms included in this family, which 
comprises adaptations to every mode of life for which 
snakes are fitted. 

The distribution of the family coincides with that 
of the order, extending over the whole world with 
the exception of the Arctic and Antarctic regions, 
and Ireland and New Zealand, as well as most of 
the smaller islands of the Pacific Ocean. 



152 colubrim; 

Genus TROPIDONOTUS, Kuhl 

Maxillary teeth increasing in size posteriorly. 
Head more or less distinct from neck ; eye moderate 
or rather small, with round pupil. Body more or 
less elongate ; scales keeled, with apical pits. Tail 
moderate. 

This large genus, comprising about ninety species, 
and of almost cosmopolitan distribution, with the 
exception of South America and the greater part of 
Australia, may be divided into several subgenera, two 
of which are represented in Europe — Tropidonotus 
proper, with the common T. natrix, and Nevodia, 
Baird and Girard, with two closely related species 
of more thoroughly aquatic habits, T. tessellatus and 
T. viperinus. 

3. Tropidonotus natrix, Linnaeus 

[Natrix vulgaris, Laurenti ; Coluber torquatus, 

Lacepede) 

The Grass-Snake, or Ring-Snake 

Form.— Moderately slender; snout short, obtuse, 
not prominent ; eyes and nostrils lateral, the former 
moderately large. Tail four to six and a half times 
in the total length. 

Head-Shields. — Rostral broader than deep, visible 
from above. Nasal divided, very rarely semidivided. 
Internasals at least as broad as long, trapezoid, 
shorter than the prefrontals. Frontal broader than 



PLATE II 




TROl'IDONOTUS NATRIX 
After Sordelli 




T. NATRIX, VAR. CETTI1 
AJter Sordt-lli 




T. NATRIX. VAR. I'ERSA 



TROPIDONOTUS 



153 




the supraocular, once and one-third to once and a half 
as long as broad, as long as or a little shorter than 
its distance from the end of the snout, shorter than 
the parietals, not in contact with the preocular. 
Loreal deeper than long. One (rarely two) pre- 
and three (rarely two or four) postoculars. Tem- 
porals i-J-2. Upper labials seven (rarely six or 
eight), third and fourth (or fourth and fifth) entering 
the eye. Four or five lower labials in contact with 
the anterior 
chin -shields, 
which are 
shorter than 
the posterior. 

Scales with 
two apical 
pits, in nine- 
teen rows, 
strongly 

keeled on the body, of outer row smooth or faintly 
keeled. Ventral shields 157 to 181; anal divided; 
subcaudals 50 to 88. 

Coloration. — Very variable. We shall first de- 
scribe the typical form, and then allude to the 
principal varieties and individual variations with 
which we are acquainted. 

Grey, bluish-grey, olive, or brown, above, usually 
with black spots or narrow bars on the back, and 
vertical bars on the sides ; upper lip whitish or 




Fig. 16 (after Sordelli) 



154 COLUBRIDJE 

yellowish, with the sutures between the shields black ; 
the preocular, and sometimes the postoculars, yellow 
in the young ; a white, yellow, or orange collar on the 
nape, sometimes uninterrupted, more often divided 
in the middle, bordered behind by two black sub- 
triangular or crescentic blotches, which usually meet 
on the median line; the bright collar often becomes 
faint, or even entirely disappears, in large females 
(Plate II., first figure) ; belly usually checkered black 
and grey or white, more rarely grey with small black 
spots, or entirely black. Iris dark brown or reddish- 
brown, with a golden circle round the pupil. This 
is the form found in England and Central Europe 
and in some parts of Southern Europe. 

In Jersey, in the Spanish Peninsula, and in 
Cyprus, the white or yellow collar, which is always 
present in the very young, soon disappears, and so 
does usually the black collar, which is either much 
reduced or entirely absent (var. astreptophorus, 
Seoane;. Some large specimens from the Spanish 
Peninsula are uniform olive, without any markings. 

Another variation (Plate II., third figure), rare in 
France, but common in Italy, South-Eastern Europe, 
and Asia Minor (var. persa, Pallas ; bilineatus, Bibr. ; 
murorutn, Bonap.) has the collar well marked, though 
widely interrupted in the middle, and a white, yellow, 
or orange streak extends along each side of the back, 
which may bear the usual black markings in addition. 
In some specimens from Austria and Corfu (var. 



TROPIDONOTUS 155 

subfasciatus, Werner) the belly is white, with black 
bars occupying the free edge of each ventral shield. 

A very remarkable variety (var. cettti, Gene) from 
Corsica and Sardinia (Plate II., second figure) is grey 
or olive above, with the black markings confluent 
into more or less regular annuli, which are nearly 
as wide as the spaces between them ; these annuli 
are often broken up on the middle line of the back, 
and alternating; the collar is absent, or is trans- 
formed into the first annulus, and the upper surface 
of the head is more or less spotted or blotched with 
black. This pattern is most distinct in young and 
half-grown specimens ; in large examples the annuli 
may break up into spots, disposed with great sym- 
metry in transverse series. The belly is black, 
spotted with white. 

A specimen 20 inches long, from Bona, Algeria 
(Lataste collection), has the posterior half of the 
head, from between the eyes and behind the post- 
ocular shields, of an intense black, followed by the 
usual yellow and black collar ; two light dots close 
together on the parietal shields. 

Some specimens are entirely or nearly entirely 
black. In the var. picturatus, Jan, from the Caucasus, 
the upper parts are sprinkled all over with light 
dots, and the yellow collar is present ; the belly is 
grey, dotted with black, and with white spots on 
the sides. In others the body is black above, and 
checkered black and white beneath (var. scutatus, 



156 COLUBRID.E 

Pall.), or entirely black (var. ater, Eichw.). This 
melanism never appears until the second or third 
year of life, the young being marked like the typical 
form. 

Albinos have occasionally been met with, yellowish 
flesh-colour with reddish markings, and a white or 
yellow collar, the eye and the tongue red. Such an 
albino, from Horsted Keynes, Sussex, is preserved 
in the British Museum. A remarkable aberration, to 
be regarded as an imperfect albino, has been found 
in Dorsetshire, and described as uniform whitish, 
with a well-defined broad longitudinal central dorsal 
pale yellow-brown band. 

Size. — May reach a length of 6 feet 8 inches. 
Such giants, females, known from Sardinia, Sicily, 
and Istria, are, however, very exceptional, individuals 
of this species seldom exceeding a length of 4 feet. 
The largest British specimen on record, from Wales, 
is stated to measure 5 feet 10 inches. Males rarely 
exceed 3 feet. 

Monstrosity. — A dicephalous young, with the two 
well-formed heads side by side, is preserved in the 
British Museum, and several others have been 
described, one being reported to have lived for 
about a month. 

Distribution. — Tropidonotus natrix occurs all over 
Europe, with, of course, the exception of Ireland, 
as far north as the extreme south-east of Scotland, 
and the sixty-fifth degree in Scandinavia and Finland, 



TROPIDONOTUS 157 

and as high up as 7,450 feet in the Italian Alps. 
With the exception of a few districts in England and 
in Central Europe, as well as in the extreme north, 
it is common everywhere, in the north as well as 
in the south. On the Mediterranean islands it is 
absent from the Baleares and Malta. In North 
Africa it is known from Algeria and Tunisia, north 
of the Atlas, where it does not seem, however, to 
be at all common. It has a wide range in Asia, 
extending eastwards to Lake Baikal, and southwards 
to Cyprus, Asia Minor, and Northern Persia. In 
the south-east of its range, the bilineated variety 
predominates over the typical form. The melanistic 
so-called varieties are not geographically restricted, 
but occur all over the habitat of the species, though 
not recorded from England. 

Habits. — Although fond of water, and often seen 
swimming in ponds or streams or creeping by the 
water's edge, this snake is far less aquatic than its 
two congeners described hereafter ; it often occurs 
on dry chalk hills or in woods far from any water. 
It is moderately agile in its movements, and easily 
caught, on which occasions it hisses loudly and 
emits a nauseous smell from its anal glands, together 
with the renal dejections, but makes no attempt to 
bite ; exceptionally an individual may go so far as 
to strike with open mouth, but cases of this snake 
really biting are extremely rare. However, Gene 
says of the male of his Natrix cettii, " iracundum et 



158 COLUBRID.E 

mordacissimum animal." Dr. Gadow relates his 
experience with aggressive specimens which in- 
habited a swamp with a little stream to the north 
of Oporto, close to the coast. To his utter surprise, 
some of them actually made for him, swimming 
along rapidly with the head erect, about 6 inches 
above the water, and darting forwards with widely 
opened jaws ; but they did not bite. According to 
Professor Kathariner, this snake when caught has 
been observed to sham death, lying rigid and motion- 
less, with open gape. Some specimens do well in 
captivity, and are known to have lived for many 
years ; others refuse all food and die of starvation. 
After a time they become tolerably tame, and cease 
to produce the offensive odour when handled. 

The food consists of frogs and toads — the latter 
being preferred notwithstanding their poisonous 
secretion, which protects them from the attacks of 
most animals — occasionally of newts, seldom of fish ; 
these snakes are reported to have a predilection for 
tree-frogs, and to feed occasionally on mice and birds, 
but most observers agree that they will not take 
anything higher in the zoological scale than frogs. 
The prey is swallowed alive, and, if not very large, 
four or five frogs or toads are often taken in succes- 
sion ; a case is known of a snake having swallowed 
twenty very small frogs at one meal. The young 
feed on worms and batrachian larvae, in addition 
to very small frogs and toads. 



TROPIDONOTUS 159 

The Grass-snake gets on very well with the 
Adder, to whose venom it is immune. 

It has more than once been met with swimming 
in the sea, and a case is reported of one having been 
captured in the open sea twenty-five miles from the 
nearest land, no doubt carried away by the current, 
but still perfectly lively. 

The hibernating season is spent in holes in walls 
or at the root of trees, often under manure-heaps, 
and the awakening occurs in March or April, soon to 
be followed by the first exuviation and the pairing. 

Reproduction. — Pairing takes place in April or in 
May, according to the climate, and the eggs are 
laid between June and August, the young emerging 
six to ten weeks later. It is probable that a second 
pairing occasionally takes place in the autumn, as 
eggs have sometimes been found in manure-heaps 
at the end of winter. Females do not breed until 
about 2 feet long, males a little sooner. The eggs 
number 11 to 48, according to the size of the 
female, and, after being produced in a string, stick 
together in a mass, without any regularity. 

The eggs measure 1 to 1^ inches in length, and 
when newly laid are about once and a half as long 
as broad. They often contain at the time they are 
produced a more or less developed embryo. They 
are sometimes laid in recesses in walls, in heaps 
of sawdust near sawmills, under dead leaves, but 
preferably in manure, for which purpose females 



160 COLUBRIDtE 

often approach farms during the period of oviposition. 
Holes near baking ovens at the back of village houses 
are sometimes selected as breeding resorts. The 
female rolls herself up, and by violent contortions 
makes a sort of chamber in the manure, in which 
she may remain for some days after the eggs have 
been produced. It is not very unusual for several 
females to congregate for the purpose of laying, and 
as many as 1,200 eggs have been found in the same 
hole. The young on emerging has lost the um- 
bilical cord, and measures 6 to 8J inches. It often 
remains for a considerable time, sometimes until 
the following spring, in the hole or manure-heap in 
which it was born, feeding principally on worms. 
Very young specimens are never found in the water. 

4. Tropidonotus tessellatus, Laurenti 

(Coluber hydrus, Pallas) 

The Tessellated Water-Snake 

Form. — Rather slender; head rather long and 
narrow; snout obtuse, not prominent; eyes and 
nostrils directed upwards and outwards, the former 
rather small, the latter somewhat valvular. Tail 
four to six times in the total length. 

Head-Shields. — Rostral broader than deep, visible 
from above. Nasal often semidivided. Internasals 
usually as long as broad or longer, subtriangular, 
truncate in front, as long or nearly as long as the 



PLATE III 




TROPIDONOTUS TESSELLATUS 




TROPIDONOTUS V1PERINUS 
After Sordelli 




T. VIPERINUS, VAR. AUROLINEATUS 
After Sordelli 



TROPIDONOTUS 



161 




prefrontals. Frontal a little broader than the supra- 
ocular, once and a half to twice as long as broad, as 
long as or a little shorter than its distance from the 
end of the snout, shorter than the parietals, not in 
contact with the preocular. Loreal as deep as or 
deeper than long. Two (rarely one or three) pre- 
oculars, with or without a small subocular below ; 
three postoculars, often with one or two suboculars 
below. Temporals i -f- 2. Upper labials eight (rarely 
seven, nine, or 
ten), fourth or 
fourth and fifth 
(rarely third or 
fifth) entering 
the eye. Five 
(rarely four) 
lower labials in 
contact with the 
anterior chin- 
shields, which are shorter than the posterior. 

Scales with two apical pits, in nineteen rows, 
strongly keeled, of outer row smooth or feebly keeled. 
Ventrals 160 to 187; anal divided; subcaudals 
48 to 79. 

Coloration. — Olive, olive-grey, or brown above, with 

dark spots usually arranged quincuncially or forming 

narrow bars on the back (Plate III.) ; sides often with 

lighter vertical bars ; a more or less distinct /y-shaped 

dark band on the nape, sometimes produced as a 
11 




Fig. 17 



162 COLUBRID^ 

median streak to the frontal shield ; upper lip 
yellowish, with dark bars on the sutures between the 
shields. Lower parts whitish, yellow, orange, or 
red, marbled or checkered with black, or nearly 
entirely black. Iris golden, bronzy, or coppery red. 

Some specimens depart very strikingly from the 
coloration thus briefly defined. We will now 
mention the principal variations which have been 
described : Sides of body checkered with black and 
yellow or black and red (var. mbro-tnaculosus, 
Diirigen). With four dark stripes along the anterior 
part of the back (var. lineaticollis, Werner). Above 
with four light streaks in addition to the dark 
markings. Uniform grey or light brown above (var. 
concolor, Jan, hagenbecki, Werner). Uniform black or 
blackish (var. nigrescens, De Betta). The most 
remarkable variety is the var. vosseleri, Werner, 
from Asia Minor : above with small black and 
yellowish spots, beneath yellowish with three blackish 
stripes beginning at some distance from the head, 
the median much weaker than the outer ; the scales 
are less strongly keeled than in the typical form. 
There are also specimens with two very regular 
black stripes along the belly. 

A case of chlorochroism, in a specimen from Dal- 
matia, has been observed by Peracca. The snake was 
sulphur yellow with black markings ; a black band 
along the belly; iris golden. 

An imperfect albino, which has been met with 



TROPIDONOTUS 163 

several times in Dalmatia, has been described as 
var. flavescens, Werner. Yellowish-white or brownish- 
yellow above, with small blackish spots ; belly 
whitish in the middle, with a series of black spots, 
bright yellow on the sides ; eye and tongue red. 

Size. — This snake occasionally reaches a length of 
4 feet, but specimens over 3 feet are rare. The 
largest specimen in the British Museum measures 
3 feet 10 inches. 

Distribution. — The Tessellated Snake has a wide 
range in Europe and Asia. It is found south of the 
Alps, from Liguria to Naples, and eastwards, extend- 
ing northwards over the greater part of Austria- 
Hungary, and even as far as Saxony, and again re- 
appears to the west in various localities of the Middle 
Rhine district (from Bingen to Coblenz and Kreuz- 
nach, from Nassau to Lahnstein) and of the Moselle. 
From Southern Russia it extends into Siberia as far as 
the Altai, the extreme west of China, and the extreme 
north-west of India; it is also found in Asia Minor, 
Transcaucasia, Persia, Mesopotamia, Syria, and the 
neighbouring parts of Egypt. Italy and the Rhine 
constitute the western limit of its range in Europe. 
It does not ascend to any considerable altitude 
in the mountains of Europe, but it is on record from 
6,000 feet elevation in Chitral. 

Habits. — This is a far more aquatic species than 
the preceding, being seldom found in summer away 
from the water, in which it swims and dives to 



1 64 COLUBRIDJE 

perfection ; which does not prevent it from being 
equally agile on land. In accordance with these 
thoroughly aquatic habits, it feeds mostly on fish, 
although occasionally taking frogs and toads and 
their tadpoles. Small fish are swallowed in the 
water, but large ones are landed. This snake does 
not object to salt water, and it has been observed 
on the seashore near Odessa, chasing small fish, 
mostly gobies, in shallow water. Hibernation and 
pairing take place on land, and it is not until the 
latter function is accomplished that the snakes of 
this species resort to the water, which the females 
leave again for oviposition. Like the Grass-snake, 
the Tessellated Snake seldom bites. 

Reproduction. — Pairing takes place in spring, when 
large numbers have been observed to congregate for 
the purpose. As in the Viperine Snake, a second 
pairing may occur in the autumn, Dr. Werner having 
found a pair in copula on September 14, at Trebinje, 
Herzegovina, the female laying her eggs in the 
following July, which with the beginning of August 
is the time for oviposition. The eggs measure a 
little over an inch in length and two-thirds of an 
inch in width, and number 5 to 25 ; they are deposited 
under stones, in the fissures of walls and rocks, or 
under the refuse of tanneries. 



TROPIDONOTUS 



165 



5. Tropidonotus viperinus, Latreille 
The Viperine Water-Snake 

Form. — Moderately slender ; head shorter than 
in the preceding species ; snout obtuse, not pro- 
minent ; eyes and nostrils directed upwards and 
outwards, the former rather small, the latter some- 
what valvular ; tail four to six times in the total 
length. 

Head-Shields. — Rostral broader than deep, visible 
from above. Nasal usually semi-divided. Inter- 
nasals as long as 
broad or longer, 
su bt r iangular, 
truncate in front, 
as long as the pre- 
frontals. Frontal 
usually broader 
than the supra- 
ocular, once and 
a half to twice 
as long as broad, as long as or slightly longer 
than its distance from the end of the snout, shorter 
than the parietals, not in contact with the preocular. 
Loreal as deep as or a little deeper than long. One 
or two preoculars and two (rarely three) postoculars. 
Temporals 1 + 2 or 1 + 3. Upper labials seven (rarely 
eight), third and fourth (or third, fourth, or fourth 
and fifth) entering the eye. Four (rarely five) lower 





Fig. 18 (after Sordelli) 



1 66 COLUBRID.E 

labials in contact with the anterior chin-shields, 
which are usually shorter than the posterior. 

Scales with two apical pits, in twenty-one (rarely 
nineteen or twenty-three) rows, strongly keeled, of 
outer row smooth or feebly keeled. Ventrals 147 to 
164 ; anal divided ; subcaudals 46 to 72. 

Coloration. — Grey, brown, or reddish above with 
two alternating series of dark brown or black spots 
on the back, or with a black zigzag dorsal band 
(Plate III., second figure), rarely with a single series 
of black vertebral spots; a lateral series of black 
spots, usually ocellar, with yellow centres ; upper 
surface of head with dark symmetrical markings ; 
a more or less distinct dark band on the temple, and 
another on each side of the nape, often edged with 
yellow in front ; upper lip yellow, with dark bars 
on the sutures between the shields, or dark with a 
yellow spot on each shield. Lower parts yellow or 
red, checkered with black, or entirely black ; the 
black of the belly may be connected with the ocellar 
lateral spots by black vertical bars. Iris golden, 
often mixed with brown. 

A specimen from Ponte Carrega, near Genoa, 
preserved in the Genoa Museum, is remarkable as 
being of a dark olive-grey, with three series of black 
and yellow ocellar spots. It is further exceptional 
in having the scales in nineteen rows. A second 
specimen, from the same locality, with the normal 
number of scales, has some of the vertebral spots 



TROPIDONOTUS 167 

ocellar. Specimens with ocellar vertebral spots are 
found also in Sardinia and in Spain. 

As in T. natrix, there occur, in the South of 
France, in Sardinia, in the Spanish Peninsula, and 
in North Africa, specimens with two light yellow 
or reddish lines along the back (Plate III., third 
figure), in addition to the usual markings (C. auro- 
lineatus, Gervais, T. chersoides, Dumeril and Bibron). 

Melanism is rare in this species, only one specimen 
being known, from Nantes in Southern Brittany; 
uniform black, with the exception of a few white 
spots on the belly. A remarkable variety (var. 
incertus, Fatio), connecting this species with the 
preceding, occurs in Switzerland near Geneva. Not 
only is its coloration sometimes very similar to 
that of T. tessellatus, but it agrees with it in the scales 
being often disposed in nineteen rows instead of 
twenty-one, and in the presence of eight upper 
labials, fourth or third and fourth entering the eye ; 
however, the frequent presence of ocellar spots 
on the sides, and the low number of ventral shields 
(147 to 151), show that it should be referred to 
T. viperinus. 

Size. — Rarely reaches a length of 3 feet in Europe, 
the largest specimens being from Sardinia. An 
Algerian specimen 3 feet 3 inches long is on record. 

Distribution. — France as far north as Southern 
Brittany, the Forest of Fontainebleau, and the 
Department Aube, the whole of the Spanish Penin- 



1 68 COLUBRID.E 

sula and the Balearic Islands, Southern Switzer- 
land, north and south of the Alps, Liguria, Pied- 
mont, Corsica, Sardinia, and Sicily. In Africa in 
Morocco, Algeria, a,nd Tunisia, penetrating into the 
northern parts of the Sahara. 

In Liguria, Piedmont, and Ticino, T. vipevinus 
occurs alongside with T. tessellatus. It reaches an 
altitude of nearly 4,000 feet in the Alps. 

Habits. — Very much the same as in the preceding 
species, although slightly less thoroughly aquatic, 
large individuals being sometimes met with at some 
distance from water. Ponds and marshes are the 
favourite abode of the Viperine Snake, huge num- 
bers being often found on the borders, diving into 
the water when disturbed. Frogs and toads, tad- 
poles, newts, fishes, and large earthworms, are its 
principal food when adult, the young feeding chiefly 
on batrachian larvae, young fishes, and earthworms. 
A case is known of this snake having eaten a water- 
shrew {Crossopus fodiens). When a fish has been 
caught, it is usually eaten on land ; in captivity dead 
fish are rather readily accepted, provided they be 
quite fresh. Some specimens bite when handled ; 
others are as gentle as the Grass-snake. 

For hibernation, hollow trees, fissures in rocks, 
holes in the ground or in railway embankments, 
are selected, and numerous individuals sometimes 
congregate in the same retreat. In the mild winters 
of the South of Europe they remain quiet, without 



TROPIDONOTUS 169 

being torpid, and resume activity very early in the 
spring. 

In the Alemtejo, according to Gadow, when 
during the rainless and hot summer the small rivers 
have nearly dried up, these snakes collect in great 
quantities in the remaining stagnant and muddy 
pools, and, as the stock of suitable fish gets ex- 
hausted, are often reduced to a deplorably emaciated 
condition. By the month of August they have 
become so thoroughly aquatic that they cannot be 
kept alive in dry surroundings for twenty-four hours, 
apparently dying from some kind of cutaneous 
suffocation. The same observer once caught a 
Viperine Snake in a ditch whilst it was swallowing 
an eel of nearly its own length. 

Some specimens show so great a superficial re- 
semblance to the Common Adder, Viper a bents, 
which, however, being a more northern reptile, very 
seldom occurs in the same localities — that this snake 
well deserves its name Viperinns. A celebrated 
herpetologist, Constant Dumeril, was once himself 
deceived by this resemblance and bitten by a Vipera 
berus which he had picked up in the Forest of Senart, 
near Paris, believing it to be a Tropidonotus viperinus ; 
whilst, conversely, a specimen of the harmless snake 
was killed in mistake for a Viper by no less an expert 
than Dr. Viaud-Grandmarais. 

Breeding. — This snake pairs in March and April, 
and sometimes again in the autumn ; but the eggs 



170 C0LUBRID.E 

are only laid at one season, in June or July, and 

hatch in August, September, or October. The eggs, 

numbering four to twenty, are deposited in holes not 

far from water, often in abandoned galleries of voles 

or moles. The young at birth measure 4 to 6J 

inches, and soon resort to the water, where, unlike 

those of the Grass-snake, they are frequently met 

with. 

Genus ZAMENIS, Wagler 

Maxillary teeth increasing in size posteriorly, the 
two last often separated from the others by a narrow 
interspace. Head elongate, distinct from neck ; 
eye rather large, with round pupil. One or more 
subocular shields. Body much elongate ; scales 
smooth, with apical pits. Tail long. 

The species of this genus, about thirty in number, 
are distributed over Europe, North Africa, Asia, and 
North and Central America. Three inhabit Europe. 

6. Zamenis gemonensis, Laurenti 

(Coluber viridiflavus, Lacepede ; C. atrovirens, Shaw) 

The European Whip-Snake 

Form. — Slender ; snout rounded, with distinct 
canthus, moderately prominent, concave on each 
side in front of the eye. Tail three and one-third 
to four and one-third times in the total length. 

Head-Shields. — Rostral a little broader than deep, 
the portion visible from above measuring one-fourth 
to two-fifths its distance from the frontal. Frontal 









PLATE IV 




r: 






m 



Xk 



(J 



J. 

> 

< 




1! 

z 5 



8 



Z 5 



"* -. 



< 






< 

> 



</) 

Z 
H 
Z 

o 
s 

(it 
u 






\^%0k 



< 
> 



z 

Ell 

V. 



ITm 



r 



v*r 



;• 



/••«' 



V, 



ZAMENIS 171 

more or less bell-shaped, not or but little broader 
than the supraocular, once and two-thirds to twice 
as long as broad, as long as or a little longer than 
its distance from the end of the snout, a little 
shorter than the parietals. Loreal as long as deep 
or longer. One preocular (rarely two), extending to 
the upper surface of the head, but never in contact 
with the frontal ; a small subocular below the pre- 
ocular ; two postoculars (rarely three). Temporals 
2 + 2 or 2 + 3 
(rarely 1 + 2). 
Upper labials 
eight, fourth and 
fifth entering the 
eye, fifth and 
seventh deepest. 
Five lower labials 

(rarely four) in 

.,1 ,1 Fig. 19 (after Sordelli) 

c >ntact with the v ' 

anterior chin-shields, which are usually shorter than 

+ he posterior. 

Scales with two apical pits, in nineteen (rarely 
seventeen or twenty-one) rows. Ventral shields 
more or less distinctly angulate laterally, 160 to 230 
(usually under 200 in the typical form and the var. 
caspius, 190 or more in the vars. viridiflavus and 
asianus) ; anal divided ; subcaudals 87 to 131. 

Coloration. — In the typical Z. gemonensis the upper 
parts are yellowish-brown or pale olive, anteriorly 





iy 2 colubrim: 

with blackish cross-bars or numerous small black 
spots, the black scales with a yellowish shaft, the 
lower parts yellowish-white or pale yellow, rarely 
more orange ; the sides of the head are yellow, the 
shields edged with blackish. A female, 3^ feet long, 
from Levico, Trentino, preserved in the Genoa 
Museum, is uniform reddish-brown above, with mere 
traces of darker markings on the head and nape. 
There is every gradation between this form and the 
var. viridiflavus or atrovirens (Plate IV., third figure), 
which is dark green or black above, with yellow spots 
forming transverse series or bars on the anterior 
part of the body, and longitudinal streaks, following 
the series of scales, on the posterior part and on the 
tail ; the yellow sometimes predominates over the 
black, or may appear as a shaft along each dark 
scale; the preocular and postocular shields are 
yellow, the labials likewise yellow, with black spots 
or bars. The lower parts are yellow or greenish- 
white, with or without black dots, and usually with 
a series of large black spots on each side. 

Some specimens of both the typical form and the 
var. viridiflavus are entirely black or nearly black 
{Z. carbonarius, Bonaparte; Z. sardus, Suckow). In 
some localities and islands only black specimens 
occur. 

In the var. caspius, Iwan (trabalis, Pallas, Plate V., 
first figure ; persicus, Jan, Plate IV., second figure), 
from Hungary, Bosnia, Herzegovina, Corfu, Bui- 



ZAMENIS 173 

garia, Roumania, Greece, Turkey, Southern Russia, 
Northern Asia Minor, and North-West Persia, the 
upper parts are pale olive or reddish-brown, with or 
without brown or black spots, and each scale bears a 
yellowish or pale brown longitudinal streak ; there is 
often a dark longitudinal streak on the nape ; the 
belly is uniform orange or red. 

Another variety, var. asianus, Boettger, from Asia 
Minor, Rhodes. Cyprus, and Syria, has the upper 
parts brown or olive, each scale with a longitudinal 
light streak, and there are usually 'large black 
spots relieved by yellowish shafts ; the belly is red, 
spotted or dotted with black. Melanism is frequent 
in this form, such specimens being entirely black 
except on the chin and throat, which are yellow 
variegated with red. 

The very young of the typical form, as well 
as that of the var. viridiflavus, has a striking livery 
(Plate IV., first figure), the head and nape black 
with yellow markings, or olive with black-edged yellow 
markings, contrasting sharply with the pale olive-grey 
of the body ; the most conspicuous and constant of the 
yellow markings consist of a bar between the eyes, 
interrupted on the frontal shield, but sometimes con- 
tinuous with the yellow of the postoculars, five or six 
small round spots on the parietal shields, and a 
V- or W-shaped line just behind the parietals, fol- 
lowed by one or two others separating the dark 
cross-bars which may be present on the nape, and 



174 COLUBRID^: 

occasionally even continue some way down the 
anterior part of the body. This livery persists in 
some half-grown specimens. 

In young individuals from Syria (var. asianus) the 
head is not differently coloured from the olive-brown 
body, and the markings described above appear as 
mere traces ; on the other hand, the whole body has 
black and yellow spots or cross-bars above, and the 
belly is profusely marked with round black spots. 

In the new-born of the var. caspius, of which I 
have examined only one specimen, n inches long, 
from the Crimea, the head is olive-brown like the 
body, which bears dark brown spots and narrow 
cross-bars ; and there is a dark brown streak along 
the middle of the nape, as is sometimes the case in 
the typical form. The belly is unspotted. A young 
from Malta is intermediate in its markings between 
the typical form and this variety. 

The young of the so-called black variety are not 
black at birth, but similar to the normal young of the 
races to which they belong. 

The four principal forms — viridiflavus, gemonensis, 
caspius, and asianus — are so completely connected that 
I cannot regard them as more than geographical 
races or varieties. 

Size. — This handsome snake grows to a length of 
6 feet, the var. caspius even to 8 feet. I have seen a 
specimen of this variety, from Salonica, which 
measures 7§ feet. 



ZAMENIS 175 

Distribution. — From the Atlantic coast of Europe 
to South- Western Asia. The typical form, in its 
narrowest sense, inhabits the Southern Tyrol, the 
north-eastern corner of Italy, and the countries to 
the east of the Adriatic, as far as Greece and Crete. 
The specimens from France, Switzerland, Italy, 
Giglio, Montecristo, Elba, Corsica, Sardinia, Sicily, 
and Malta, are mostly referable to the form known 
as Z. viridiflavus. Farther to the east the species 
is represented by the vars. caspius and asianus, of 
which the distribution has been mentioned above. 
From Spain, this snake is only on record from Cata- 
lonia, not far from the French frontier. 

Rare or local in the north of its range (Maine-et- 
Loire, Vienne, Indre, Sarthe, Haute-Saone, Yonne, 
Aube, in France, Switzerland north of the Alps), it 
is one of the commonest snakes in Italy and on the 
borders and islands of the Adriatic, as well as on 
practically all the islands of the Mediterranean east 
of the Baleares. The highest altitudes at which it 
has been met with are 3,900 feet in the Alps, 4,500 
feet in the Balkan Peninsula. 

Habits. — The name " Whip-snake," under which 
an American representative of this genus (Z. ftagelli- 
formis) is known, like that of " Fouet " and " Lou- 
cinglant," which have been bestowed on it in some 
parts of France, expresses the quick movements with 
which, when captured, this snake lashes its long, 
slender tail, at the same time furiously biting the 



176 colubrim: 

hand that has seized it. The generic term Zamenis, 
of Greek derivation, alludes to its viciousness, which 
also accounts for its German name, " Zornnatter." 
This snake, occurring in Malta, may well have been 
the " Viper " which fastened on the hand of St. Paul. 
Some specimens have been kept for months in cap- 
tivity without losing their savage temper, hissing and 
flying with open mouth at anyone approaching the 
glass walls of their prison ; others, on the other hand, 
become quite tame in a very short time, such as one 
which I kept for nearly two years. Except when 
sunning itself on a cold early morning in the spring, 
this snake is always on the alert, and difficult to 
capture, uncoiling itself and darting away like an 
arrow at the least disturbance. It lives in preference 
among shrubs or on the edges of woods, avoiding 
damp localities, and females at least appear to have 
sedentary tastes. Lataste tells us of one, near 
Bordeaux, which he repeatedly met for over two 
years within 20 yards of the same spot, a bush 
between a wood and a meadow, without ever being 
able to capture it. 

The food of this snake is very varied, consisting of 
voles and mice, young birds which it takes from the 
nests, being a good climber on bushes and low trees, 
occasionally of frogs, but above all of other reptiles : 
lizards, slow-worms, and snakes, which it does not 
attempt to crush before deglutition. It has even 
been observed in Istria to eat locusts (Acridium 
czgyptium) and sphyngid moths. 



PLATE V 




ZAMENIS GEMONENSIS, VAE. CAS I'll S 

After Sordelli 




ZAMENIS DAHLII 
After Sordelli 




ZAMENIS HIPl'OCREPIS 
After Sordelli 



ZAMENIS 177 

Reproduction. — Eggs, laid at the end of June or 
beginning of July in a well-sheltered hole, are a little 
over twice as long as broad, and measure 1*2 to 
1*4 inches in length. The number of eggs is eight 
to fifteen according to Fatio, about a dozen accord- 
ing to Tomasini, five according to Werner. The 
pairing was observed by Schreiber at the end of May, 
the male and female seizing each other reciprocally 
by the neck with their jaws ; this mode of pairing 
must not, however, be regarded as the rule in this 
species, for in other cases observed by Schreiber and 
by Honnorat the pairs were simply entwined by 
their coils. 

7. Zamenis dahlii, Fitzinger 
Dahl's Whip-Snake 

Form. — Very slender ; head narrow, snout moder- 
ately prominent, obtuse. Tail about one-third of the 
total length. 

Head-Shields. — Rostral a little broader than deep, 
just visible from above. Frontal not or but little 
broader than the supraocular, once and two-thirds to 
once and three-fourths as long as broad, as long as 
or longer than its distance from the end of the snout, 
shorter than the parietals. Loreal longer than deep. 
One preocular, usually in contact with the frontal, 
with a subocular below it ; two postoculars. Tem- 
porals 2 + 2 or 2 + 3 (rarely 1 + 2). Upper labials 
eight or nine, fourth and fifth or fifth and sixth 
12 



178 



colubrim: 





entering the eye. Four or five lower labials in 
contact with the anterior chin-shields, which are 

shorter than the 
posterior. 

Scales with a 
single apical pit, 
very narrow, in 
nineteen rows. 
Ventral shields 
very distinctly an- 
gulate laterally, 

Fig. 20 (after Sordelli) . , 

* ' 205 to 218 ; anal 

divided ; subcaudals 98 to 132. 

Coloration. — Olive in front, with a few large black, 
white- or yellow-edged spots on each side, the anterior 
of which is sometimes confluent with its fellow and 
forms a nuchal collar, as in the specimen figured on 
Plate V. ; the greater part of the body and tail 
uniform pale olive, yellowish, or reddish above, 
yellowish-white beneath. Head uniform olive-brown 
above, the labial, preocular, and postocular shields 
yellowish-white. 

Total Length. — 3 feet, rarely nearly 4 feet. 

Distribution. — Southern Europe east of the Adriatic, 
as far north as Dalmatia, Asia Minor, Cis- and Trans- 
Caucasia, North-Western Persia, Cyprus, and Syria. 
Has also been recorded from Lower Egypt. 

Habits. — This snake is even more lively than 
Z. gemonensis, and does not stand captivity long. It 



ZAMENIS 



179 



seeks dry, bushy localities, and feeds on small lizards, 
occasionally on locusts. It does not seem to be very 
common anywhere in Europe, except perhaps in 
Dalmatia, whence most of the specimens sold by 
dealers are imported. 

Reproduction. — The pairing has been observed at 
the end of May. According to Werner, the eggs 
number usually three only, measuring ij inches by 
\ inch. 

8. Zamenis hippocrepis, Linnaeus 
The Horseshoe Whip-Snake 

Form. — Slender ; snout obtuse, feebly prominent. 
Tail one-fifth to one-fourth of the total length. 

Head-Shields. — Rostral once and one-third to once 
and a half as broad as deep, the portion visible 
from above mea- 
suring about 
one - fourth to 
one - third its 
distance from 
the frontal. 
Frontal bell - 
shaped, consid- 
erably broader 
in front than the 





Fig. 21 (after Sordelli) 



supraocular, once and one-fourth to once and a half 
as long as broad, as long as or a little longer than its 
distance from the end of the snout, shorter than the 



180 COLUBRIDJE 

parietals. Loreal longer than deep, sometimes divided 
into two. One preocular (sometimes divided into 
two), in contact with the frontal ; two postoculars ; a 
series of three or four suboculars, usually completely 
separating the eye from the labials. Temporals 
2 + 3 or 3 + 3. Eight or nine (rarely ten) upper 
labials, fifth or sixth very rarely entering the eye. 
Four lower labials in contact with the anterior chin- 
shields, which are shorter than the posterior. 

Scales with two apical pits, in twenty-five to 
twenty-nine rows, usually twenty-seven. Ventral 
shields very distinctly angulate laterally, 222 to 258; 
anal divided (rarely entire) ; subcaudals jj to 107. 

Coloration. — Brown, pale olive, reddish, yellow, or 
orange above, with a dorsal series of large dark brown, 
black-edged rhomboidal spots, often bordered with 
yellow, on each side of which is a series of smaller, 
alternating spots (Plate V.) ; these spots may become 
entirely black in the adult, and so large as to reduce 
the ground colour to a mere network or series of 
X-shaped pale lines. A dark cross-band between 
the eyes, and a y\- or horseshoe-shaped band on 
the back of the head, which may be confluent with 
an elongate spot on the nape ; a light circle often 
present in the middle between the parietal shields- 
The spots often more or less confluent into three 
longitudinal streaks on the tail. Yellow, orange, or 
red beneath, with or without black dots, but con- 
stantly with a lateral series of black spots, which 



COLUBER 181 

may be very large or unite with the spots higher up 
on the sides to form vertical bars. 

Size. — Examples 5 feet long are on record ; the 
largest examined by me measures 4 feet 3 inches. 

Distribution. — Spain and Portugal, Sardinia, Pan- 
tellaria, Morocco, Algeria, Tunisia. Does not reach 
the North of Spain nor penetrate into the Sahara. 

Habits. — This very handsome snake is as a rule 
as irascible as its European congeners. In Spain as 
well as in Algeria it is often found about the dwellings 
of man, occasionally entering houses in search of 
mice, on which it principally feeds ; it is also fond 
of birds, and, climbing with great facility, plunders 
the nests of sparrows in towns and villages. It 
must be regarded as a useful commensal of man, and 
deserving of protection. 

Reproduction. — F. Doumergue found in a hole in a 
rock near Oran, in September, the recently-laid eggs, 
five in number and as large as pigeons'. 

Genus COLUBER, Linn^us 

Maxillary teeth equal or nearly equal in length. 
Head elongate, distinct from neck ; eye moderately 
large, with round pupil. Body more or less elongate ; 
scales smooth or feebly keeled, with apical pits. 
Tail moderate or long. 

This large genus, embracing close upon fifty 
species, is represented in Europe, Asia, and North 
and tropical America. Five species in Europe. Very 



182 



COLUBRIDJE 



nearly allied to Zamenis, but distinguished principally 
by the posterior teeth of the upper jaw not being at 
all enlarged, and, further, in being, like Coronella, 
constrictors. . 

9. Coluber quatuorlineatus, Lacepede 

(Elaphis cervone, Aldrovandi ; Coluber 

quatuorradiatus, Gmelin) 

Aldrovandi's Snake 

Form. — Moderately slender. Snout obtuse, 
scarcely prominent. Tail one-sixth to one-fourth 

of the total 
length. 

Head-Shields. — 
Rostral broader 
than deep, just 
visible fr o m 
above. Frontal 
once and one- 
fourth to once 

Fig. 22 (after Sordelli) , , ir 

v and a half as 

long as broad, as long as its distance from the 
rostral, shorter than the parietals. Loreal nearly 
as long as deep, with one or two small shields 
below it. One preocular, rarely divided, with 
a subocular below it ; two or three postoculars. 
Temporals 2 + 3 or 3 + 4. Upper labials eight 
(exceptionally nine), fourth and fifth (or fifth and 
sixth) entering the eye. Four or five (rarely three) 





PLATE l'I 




COLUBER QUATUORLINEATUS 
Young, after Sordelli 




COLUBER QUATUORLINEATUS 
After IVerner 




COLUBER QUATUORLINEATUS, VAR. SAUROMATES 
After Sordelli 




COLUBER DIONE 
After Sordelli 



COLUBER 183 

lower labials in contact with the anterior chin- 
shields, which are longer than the posterior. 

Scales feebly but distinctly keeled, except on the 
outer rows, with two apical pits, in twenty-five (rarely 
twenty-three or twenty-seven) rows. Ventral shields 
not or but very obtusely angulate laterally, 195 to 
234 ; anal divided ; subcaudals 56 to go. 

Coloration. — Young (Plate VI., top) with three or 
five alternating longitudinal series of dark brown, 
black-edged spots on a yellowish, grey, or pale 
brown ground, the spots of the median series largest, 
transversely elliptical or rhomboidal; a dark streak 
across the forehead, black bars on the labial shields, 
and a black oblique streak from the eye to the angle 
of the mouth. In specimens from Italy and the 
countries bordering the Adriatic (the typical C. 
quatuorlineatus) the markings very gradually disappear 
with age, with the exception of the temporal streak, 
whilst a pair of black streaks appear along each side 
of the body, at a short distance from the head, the 
lower corresponding to the postocular streak, the 
adult being brown without spots, but four-lined 
(Plate VI., second figure). In more eastern specimens 
(C. sauromates, Pallas), which may be regarded as 
representing the original form, the markings of the 
young persist throughout life, or, if they disappear, 
they are not replaced by dark streaks (Plate VI., 
third figure). Lower parts pale yellow, closely 
spotted or marbled with brown, these markings 



1 84 COLUBRID.E 

usually disappearing in the adult, except on the tail. 
Iris dark brown. 

Size. — The largest European snake, stated to 
reach a length of 8 feet. The largest specimen ex- 
amined by me measures, however, only 4J feet. 

Distribution. — Aldrovandi's Snake inhabits Southern 
Italy and Sicily, Istria, Croatia, Dalmatia, Herzego- 
vina, Greece, and eastwards to Southern Russia, 
Transcaucasia, Asia Minor, and Persia. It has been 
observed at an altitude of 2,600 feet in Herzegovina. 

All the specimens from Roumania, Bulgaria, 
Turkey, and eastwards, belong to the var. sauromates, 
which is regarded by some authors as worthy of 
specific rank. The reported occurrence of C. quatuor- 
lineatus in various parts of France is certainly due to 
confusion with C. scalaris and C. longissimus. 

Habits. — Dry as well as marshy localities are the 
abode of this large and handsome snake, which 
often approaches the dwellings of man, attracted by 
the poultry. Comparatively slow in its movements, 
it is more easily captured than any of the other large 
Colubrids of Europe, and does well in captivity, 
where it should be provided with a tank, in which it 
will remain for hours under water. It is as good at 
swimming as at climbing. Biting readily when cap- 
tured, it becomes of gentle disposition after a short 
period of captivity. In consequence of its slow, phleg- 
matic temperament, it often allows itself to be picked 
up when surprised in liberty, but as soon as it feels the 



COLUBER 185 

grasp it turns round and defends itself. It appears 
to feed exclusively on mammals and birds, up to the 
size of a rat or dove, and will readily take dead food. 
It has a predilection for eggs, and has often been 
observed to swallow hens' eggs. 

Reproduction. — In Herzegovina pairing takes 
place from the middle of June to the middle of July, 
and the eggs are laid soon after, to hatch in Septem- 
ber or beginning of October. The eggs number six 
to sixteen, and measure 2 inches by ij inches. The 
young measure 8 to 14 inches at birth. 

10. Coluber dione, Pallas 
The Dione Snake 

Form. — Similar to the preceding. Head more 
convex, a little narrower; snout obtuse, scarcely 
prominent. Tail about one-fifth of the total length. 

Head- Shields. — Rostral broader than deep, just 
visible from above. Frontal once and one-fourth 
to once and a half as long as broad, as long as its 
distance from the end of the snout, shorter than the 
parietals. Loreal as long as deep, or a little longer 
than deep. A large preocular, with a subocular 
below it, the latter very exceptionally absent ; two 
or three postoculars. Temporals 2 + 3 or 3 + 3. 
Upper labials eight or nine (very rarely seven), 
fourth and fifth or fifth and sixth entering the eye. 
Four or five lower labials in contact with the 



i86 



COLUBRID^: 





anterior chin-shields, which are nearly as long as 

the posterior. 

Scales smooth or faintly keeled, with two apical 

pits, in twenty-five or twenty-seven (rarely twenty- 
three) rows. Ven- 
tral shields not or 
but very obtusely 
angulate laterally, 
172 to 214 ; anal 
divided ; subcau- 
dals 50 to 80. 

Coloratio n. — 
Pale brown or 

Fig. 23 (after Sordelli) . , y . 

v greyish - olive 

above, with blackish cross-lines or dark brown or 
reddish, black-edged spots, and usually two or three 
more or less distinct pale longitudinal bands ; two 
dark longitudinal stripes on the nape, usually united 
on the head and terminating on the frontal shield ; a 
curved dark cross-band from eye to eye, and another, 
oblique, from the eye to the angle of the mouth. 
Lower parts yellowish, usually dotted or spotted 
with blackish. 

Size. — Seldom exceeds a length of 3 feet. The 
largest specimen examined by me measures 37 
inches. 

Distribution. — Across temperate Asia from Asia 
Minor, Transcaucasia, and the southern border 
of the Caspian Sea, to the Amur, Corea, and China. 



COLUBER 



187 



In Europe the 
to the steppes 
Caucasus and 
figured on Plate 
Habits. — This 
ities, and is only 
Nothing more is 



habitat of this snake is restricted 
of Southern Russia, between the 
the Lower Ural. The specimen 
VI. is from Sarepta, on the Volga, 
snake frequents arid, sandy local- 
exceptionally found in small woods, 
known of its habits. 



n. Coluber longissimus, Laurenti 
{Coluber cesculapii, Lacepede ; C. flavescens, Gmelin) 

The yEsculapian Snake 

Form. — Slender. Snout obtuse, scarcely promi- 
nent ; head narrow. Tail about one-fifth to one- 
fourth of the total length. 

Head-Shields. — Rostral broader than deep, just 
visible from above. Frontal once and one-fourth 
to once and 
one-third as 
long as broad, 
as long as its 
distance from 
the rostral or 
the end of the 
snout, shorter 
than the par- 
ietals. Loreal 

as long as deep or longer than deep, 
two postoculars. Temporals 2 + 3. 





Fig. 24 



One pre- and 
Upper labials 



188 COLUBRID.E 

eight or nine, fourth and fifth or fifth and sixth enter- 
ing the eye. Four or five lower labials in contact 
with the anterior chin-shields, which are as long as 
or a little longer than the posterior. 

Scales smooth or feebly keeled on the posterior part 
of the body, with two apical pits, in twenty-three 
(rarely twenty-one) rows. Ventral shields distinctly 
angulate laterally, 212 to 248 ; anal divided ; sub- 
caudals 60 to 91. 

Coloration. — Yellowish-grey to dark olive-brown 
above, some of the scales with whitish lines on the 
margins occasionally forming a network ; sometimes 
with a yellowish vertebral stripe or with four darker 
stripes along the body (var. romanus, Suckow) ; upper 
lip, and often also a triangular patch on each side 
behind the temple, pale yellow ; a more or less dis- 
tinct dark band on the temple, and a vertical dark 
bar below the eye (Plate VII., first figure). Lower 
parts uniform pale yellow. Young (second figure) 
with dark brown dorsal spots, forming four to seven 
longitudinal series, a A-shaped black marking on the 
nape behind the yellow nuchal blotches, which are 
brighter than in the adult, a dark brown bar across 
the forehead, and a black vertical line below the 
eye; belly greyish or yellowish-olive. Iris dark grey 
or brown. Tongue pinkish-brown. 

Melanism is rare in this snake. Such specimens 
are entirely black above and beneath (var. mger, 



PLATE VII 




COLUBER LONGISSIMUS 




COLUBER LONGISSIMUS 
Young, after Sordelli 




COLUBER LEOPARDINUS 

After Sordelli 




C. LEOl'ARDINUS, VAR. QUADRILINEATl S 
After Sordelli 



COLUBER 189 

Nikolsky), or blackish - grey to black above, dark 
grey beneath (var. subgriseus, Werner), the angular 
line on each side of the belly often remaining light. 
An albino found near Vienna has been described as 
pale orange-yellow above, with small white spots ; 
pupil and tongue red. 

Size. — Grows to 6 feet. Specimens over 4J feet 
are, however, very rarely met with. 

Distribution. — Generally distributed over the 
greater part of Austria, Italy, with Sardinia and 
Sicily, and the whole of South-Eastern Europe, this 
snake has a very broken range in France, Switzer- 
land, Germany, and is found, quite isolated, as far 
north as Denmark and Poland. According to 
Segerus, quoted by Lacepede, it used to be quite 
common near Copenhagen at the end of the 
eighteenth century, but it is now much rarer. Its 
northern limit in France is in Southern Brittany, 
the Department Orne, and the Forest of Fontaine- 
bleau ; in Germany, Schlangenbad, near Wiesbaden, 
perhaps also Baden-Baden and Treves. It is on 
record from Southern Spain. Its discontinuous 
distribution in Central Europe, and its presence 
in various localities near former Roman thermal 
stations, has been ascribed to its introduction from 
Italy as an inmate of the temples erected to 
iEsculapius; but I am more inclined to look upon 
its sporadic occurrence in the North as the indica- 
tion of a once more widely distributed species 



igo COLUBRID.E 

now in process of extinction over part of its 
range. 

In Asia the iEsculapian Snake is only found in 
Transcaucasia. It occurs in the mountains as well 
as in the plain, being recorded from 5,200 feet 
altitude in the Tyrol, 3,200 feet in the Apennines. 

Habits. — The y£sculapian Snake lives in woods ; 
among shrubby vegetation ; in meadows, where it is 
often found under haystacks ; occasionally about 
old walls. It climbs well, and often ascends trees. 
Although a good swimmer, it seldom enters the 
water of its own accord. It feeds chiefly on small 
mammals, occasionally on birds and their eggs, and 
lizards. Specimens which I kept in confinement fed 
on mice only, refusing sparrows and lizards. Very 
savage when fresh caught, most individuals soon 
become tame, and like being handled by people to 
whom they are accustomed, although still resenting 
the intrusion of strangers. However, this snake 
never becomes so thoroughly domesticated as the 
Smooth Snake, and cannot be trained to take food 
from the hand, according to R. Rollinat, who has 
devoted many years to experiments on the taming 
of reptiles. This observer had no difficulty in feed- 
ing his -^Esculapian Snakes on mice and voles placed 
dead in their cage. 

This snake is particularly sensitive to cold, and 
does not emerge until late in the spring from the 
vole galleries and hollow trees which constitute its 



COLUBER igi 

winter-quarters. It also avoids excessive heat, never 
showing itself in the daytime during the hotter 
months in the South of Europe. 

Reproduction. — Pairing takes place between the 
middle of May and the middle of June. The eggs 
are laid towards the end of June or in July, in holes 
in walls or hollow trees, under moss, sometimes even 
in the dung-heaps of farms, and hatch in September. 
According to trustworthy observers, the eggs, which 
measure i J to 2 inches in length, and less than 1 inch 
in width, number only five or six, rarely up to 
eight. 

The young on emerging are highly suggestive of 
young Grass-snakes in colour and markings, as well 
as in their much less slender shape as compared with 
the adult. They measure about 5 inches, and are 
at once most ready to bite. 

12. Coluber leopardinus, Bonaparte 

(Coluber quadrilineatus, Pallas) 

The Leopard Snake 

Form. — Slender. Snout obtuse, scarcely promi- 
nent. Tail about one-fifth of the total length. 

Head-Shields. — Rostral broader than deep, just 
visible from above. Frontal once and one-third to 
once and a half as long as broad, as long as its distance 
from the end of the snout, shorter than the parietals. 
Loreal longer than deep. One pre- and two post- 
oculars. Temporals 1 + 2 or 2 + 3. Upper labials 



192 



COLUBRID.E 





eight (rarely seven), fourth and fifth (rarely third and 
fourth) entering the eye. Four or five lower labials in 
contact with the anterior chin-shields, which are 

longer than the 
posterior. 

Scales smooth, 
with two apical 
pits, in twenty- 
five or twenty- 
seven r o ws . 
Ventral shields 
rounded, not 
fig. 25 (after Sordelli) angulate later- 

ally, 222 to 260 ; anal divided ; subcaudals 68-90. 

Coloration. — Typical form (Plate VII., third figure) 
greyish or pale brown above, with one dorsal series 
of dark brown, reddish-brown, or bright red, black- 
edged transverse spots and a lateral alternating 
series of smaller black spots with or without lighter 
centres ; usually a A-shaped dark marking on the 
occiput and nape ; a crescentic black band from 
eye to eye across the prefrontal shields, an oblique 
black band from behind the eye to the angle of the 
mouth, and a black spot or vertical bar below the 
eye. Lower parts white, checkered with black, or 
nearly entirely black. Iris reddish-golden. 

In some specimens (var. quadrilineatus) the dorsal 
spots are replaced by two brown or red, black-edged 
stripes bordering a pale greyish or yellowish vertebral 



COLUBER 193 

stripe (Plate VII., fourth figure); such specimens 
are so coloured from birth. This colour variety, 
which is so strikingly different from the typical form, 
is connected with the latter by the var. schwoederi, 
Werner, in which the spots form two vertebral 
series, and the var. elsneri, Werner, in which the light 
vertebral band is broken up by dark transverse bars, 
producing a ladder-like pattern. 

Size. — Rarely exceeding a length of 3 feet. 

Distribution. — Southern Italy, Sicily, Malta, Istria, 
Dalmatia, and other parts of the Balkan Peninsula, 
Grecian islands, Crimea, Asia Minor. The altitudinal 
range does not extend beyond 1,600 feet. 

Habits. — This is not only the prettiest European 
snake as regards its markings, whether in the form 
of spots or of stripes, but also the most graceful in its 
movements. Unless compelled to fly for safety, 
there is something slow and deliberate in its be- 
haviour which is more suggestive of Coronella than of 
most other species of Coluber. It is fond of climbing, 
and if the terrarium in which it is kept be provided 
with a bush or small tree, it will spend most of the 
time gracefully coiled round the branches. Usually 
very savage when fresh caught, some specimens 
become quite tame in captivity. In Dalmatia, where 
it is not uncommon, this snake is found principally 
among prickly shrubs, in hedges, or on old walls. It 
awakens from its winter slumber later than other 
South European snakes. Although occasionally 
13 



ig 4 



COLUBRID.E 



taking lizards, its principal food consists of mammals 
and birds, which are killed before being devoured, 
the Leopard Snake being, like the other members of 
the genus Coluber, a constrictor. 

Reproduction. — According to Werner, the eggs, two 
to five in number, are deposited in midsummer; 
they are remarkably elongate : z\ inches long, f inch 
broad. 

13. Coluber scalaris, Schinz 
The Ladder Snake 

Form. — Moderately slender. Snout pointed, 
strongly projecting beyond the mouth. Tail one- 
sixth to one-fifth of the total length. 

Head-Shields. — Rostral deeper than broad, forming 
an acute angle above, wedged in between the inter- 
nasals, the portion visible from above nearly as long 

as its distance 
from the fron- 
tal. Frontal 
about once and 
one-third to 
once and a half 
as long as broad, 
as long as or 
shorter than its 
distance from 





Fig. 26 (after Sordelli) 



the end of the snout, nearly as long as the parietals. 
Loreal longer than deep. One pre- and two or three 



PLATE VII I 




COLUBER 195 

post-oculars. Temporals 2 + 3 or 2 + 4. Upper 
labials seven or eight (rarely nine), fourth or fourth 
and fifth (or fifth and sixth) entering the eye. Four 
or five lower labials in contact with the anterior chin- 
shields, which may be either longer or shorter than 
the posterior. 

Scales smooth, with two apical pits, in twenty-seven 
(rarely twenty-five or twenty-nine) rows. Ventral 
shields not angulate laterally, 201 to 220 ; anal 
divided, rarely entire ; subcaudals 48 to 68. 

Coloration. — Young yellowish-grey, or pale brown, 
above, with a series of regular H -shaped black or 
blackish-brown markings along the back, forming a 
ladder-like pattern — whence the name scalaris — and 
small black spots on the sides ; a V-shaped black 
marking on the snout, a black oblique streak from 
the eye to the angle of the mouth, and a black spot 
below the eye ; belly yellow, spotted or checkered 
with black or nearly entirely black. These dorsal 
markings disappear in the adult, and are replaced by 
a pair of brown stripes running along the back 
(Plate VIII.); the belly loses the black markings, 
and becomes uniform yellow. Iris dark brown. 

Size. — Grows to a length of 3! feet, exception- 
ally 4J feet. 

Distribution. — The Mediterranean coast of France, 
Spain and Portugal, and Minorca. Its occurrence in 
Algeria is very doubtful. 

Habits. — Not uncommon near the coast in France, 



196 COLUBRID^ 

in hedges and vineyards, often climbing on shrubs. 
In the Spanish Peninsula, according to Bosca, it is 
common in forests and on the sheltered side of 
valleys, under stones or in holes in the ground. A 
specimen I kept alive for a short time showed a more 
furious temper than I have ever witnessed in any 
snake, repeatedly flying with open mouth against the 
glass of its cage whenever I entered the room in 
which it was kept. Other specimens are reported to 
have become quite tame after a certain time. It is 
one of the quickest of European snakes, one of the 
most difficult to catch ; it is a good climber. The 
food consists of mice, birds, and lizards ; the young 
are said to occasionally eat grasshoppers. 

Reproduction. — According to J. von Fischer, the 
eggs, nine in number, are deposited twenty-five days 
after the pairing, which takes place in May or June, 
and measure about 2 inches by f inch. 

Genus CORONELLA, Laurenti 

Maxillary teeth increasing in size posteriorly. 
Head not or but slightly distinct from neck ; eye 
rather small, with round pupil. No subocular shields. 
Body moderately elongate ; scales smooth, with apical 
pits. Tail moderate. 

This genus, embracing about twenty species, is 
represented in the different parts of the Northern 
Hemisphere, extending a little beyond the Equator 
in East Africa. Two species are European. 



PLATE IX 




< 

u 

< 

< s 

< ■: 

o 



CORONELLA 



197 



14. Coronella austriaca, Laurenti 
(Coluber Iczvis, Lacepede) 
The Smooth Snake 

Form. — Moderately slender ; snout more or less 
prominent, sometimes decidedly pointed ; tail one- 
fourth (males) to one-sixth (females) of the total 
length. The considerable differences to be observed 
in the shape of the snout are merely individual, speci- 
mens with more prominent snout and a corresponding 
development of the 
rostral shield (C. 
italica, Fitz.,fitzin- 
geri, Bonap.) oc- 





Fig. 27 (after Sordelli) 



curring over the 
greater part of 
the range of the 
species. 

Head - Shields. — 
Rostral at least as 
deep as broad, more or less produced posteriorly 
between the internasals, the portion visible from 
above at least half as long (in some specimens 
quite as long) as its distance from the frontal, 
rarely separating the internasals. Frontal once and 
one-fourth to once and a half as long as broad* 
much broader than the supraocular, as long as 
or longer than its distance from the end of the 
snout, shorter than the parietals, widely separated 



198 COLUBRID.E 

from the preocular. Nasal rarely undivided ; loreal 
longer than deep. One (very rarely two) pre- and two 
postoculars. Temporals 2 + 2 or 2 + 3 (very rarely 
1 + 2). Upper labials seven (rarely eight), third and 
fourth (or fourth and fifth) entering the eye. Four 
lower labials (rarely three) in contact with the anterior 
chin-shields, which are as long as or longer than the 
anterior. 

Scales with one or two apical pits, the pit usually 
single on the back and paired on the sides, in nineteen 
(rarely twenty-one) rows.* Ventral shields 153 to 

199 ; anal divided (rarely entire) ; subcaudals 41 to 70. 
Coloration. — Grey, brown, or reddish above, with 

small blackish, dark brown, or brick-red spots usually 
disposed in pairs, sometimes forming cross-bars ; 
sometimes with one or three lighter stripes; one or two 
black dots precede on each scale the single or paired 
apical pit ; frequently two blackish, dark brown, or 
brick-red stripes on the nape, usually confluent with 
a large dark blotch on the occiput ; the top of the 
head occasionally nearly entirely blackish, especially 
in the young ; a dark streak on each side of the head, 
from the nostril to the angle of the mouth, passing 
through the eye, sometimes extending along the side 
of the neck or even of the whole body. Lower 
parts red, orange, brown, grey, or black, uniform or 
speckled or closely spotted with black and white, the 
sides often lighter (Plate IX.). 

* The only specimen with twenty-one rows I have examined 
is a male from Albano, near Rome (Genoa Museum). 



CORONELLA 199 

A colour variety, of which I have examined a single 
specimen from near Vienna, is pale brown above, 
with four black lines along the anterior part of the 
body, and two small, yellowish, dark-edged spots 
close together on the back of the head, separated by 
the suture between the parietal shields. 

Werner has described another variety, also from 
near Vienna, which resembles Coluber leopardinus, 
having two series of large, brown, dark - edged 
spots along the back, some of the spots alternating, 
others uniting across the back. Apparently very 
similar to the last variety, and also said to be 
suggestive of Coluber leopardinus, is the var. scalaris, 
Sternfeld, from Liineburg in Hanover, reddish- 
brown above, with two row r s of bright red, black- 
edged spots, partly confluent and connected across 
the spine by transverse bars producing a ladder- 
like pattern. Specimens of a uniform greyish- 
brown, without any markings, are very rare. The 
var. veithi, Schreiber, established on a single speci- 
men from Carinthia, represents a case of melanism : 
bluish-black, with the normal markings of an intense 
black. Two specimens of a " black variety " are said 
to have been found in this country, near Poole. 

Size. — Seldom exceeds a length of 2 feet, and in 
many districts, in England for instance, does not 
appear to often reach that size. The largest speci- 
men, from Austria, examined by me, measures 
25 inches ; one from Hampshire measures 24 inches. 



200 COLUBRID^: 

Distribution. — The range of the Smooth Snake 
extends over nearly the whole of Europe, as far 
north as 63 in Norway ; it becomes rare and more 
local in the south, being absent from part of Spain 
and the islands of the Mediterranean, with the 
exception of Sardinia. It is common in the hilly 
parts of Belgium, Northern and Central France, 
Germany, and Austria. In Sweden it appears to be 
restricted to the oak region. In Great Britain it 
has been found in four counties in the South of 
England : Surrey, Hampshire, Dorsetshire, and Berk- 
shire, in some parts of which it is less uncommon 
than usually supposed. Its reported occurrence in 
Dumfriesshire is the result of an error ; the snake 
figured as Coluber dumfriesiensis represents an Ameri- 
can species. In a very interesting article written for 
Science Gossip in 1888, Mr. A. L. Beldy says that 
about 1868, when Bournemouth was but a very 
small village, surrounded by large expanses of moor- 
land, Coronella austriaca was extraordinarily abundant, 
and during a hot summer examples were to be seen 
literally in scores and great numbers were killed. 
Since then, however, their numbers have gradually 
decreased. About 1880 the snake was occasionally 
found near Wellington College, Berks, and as many 
as five were captured by one person in the course of 
one year ; it is believed to be now extinct in that 
neighbourhood. From South-Eastern Europe the 
range of this species extends to South-Western Asia. 



CORONELLA 201 

The ascertained altitudinal range is 4,000 feet in 
the Alps, 6,000 feet in Bosnia, and 6,500 feet in the 
Caucasus. 

Habits. — The Smooth Snake lives on heathland, 
stony wastes, and wooded hills, showing a preference 
for dry localities. Although not infrequent on the 
Dorsetshire and Hampshire heaths, where it was first 
discovered in 1853, it was not recorded as a British 
reptile until 1859 ; it was discovered much later on 
the sandy heaths between Haslemere and Farnham, 
where it occurs in small numbers, and in Berkshire. 
These localities are likewise inhabited by the rarer 
British lizard, the Sand Lizard. Notwithstanding 
its gentle, timid appearance, this snake when fresh 
caught is usually very ready to bite; either it snaps 
angrily, or, without hissing or other warning, it 
suddenly fastens its jaws into the finger of its 
captor, even if it be gently handled. The food 
consists mostly of lizards, occasionally of slow- 
worms or small snakes, more rarely of voles or 
mice, even shrews, which are seized, constrictor-like, 
and crushed by the coils of the body. O. von 
Tomasini has observed one swallowing a Coluber 
longissimus as large as itself. 

In Central Europe this snake becomes active 
towards the end of March or beginning of April, and 
retires in September or October. It does well in 
captivity, and becomes very tame. It is one of the 
most intelligent of snakes, second to none in edu- 



202 COLUBRID.E 

cability; it can be trained to feed in the hand of 
its master. 

Reproduction. — The Smooth Snake pairs in early 
spring, and is ovoviviparous. The young, two to 
fifteen in number, are born late in August or in 
September, enveloped by a thin membrane which 
they tear immediately ; they measure 5 to 6 inches. 
Embryos 3 J or 4 inches long have the scaling and 
the characteristic markings fully developed, but the 
scales and shields much abbreviated, the former 
broader than long. Adicephalous young is preserved 
in the Bosnian Museum at Sarajev, and another 
was caught near Karlsruhe, in Germany, in 1881, 
and kept alive for sometime. According to Rollinat, 
a second autumnal pairing sometimes takes place in 
France. 

15. Coronella girondica, Daudin 

{Coluber riccioli, Metaxa) 

The Southern Smooth Snake 

Distinguished from the preceding by a somewhat 
more slender form, a more obtuse, scarcely prominent 
snout, a much lower rostral shield, which is con- 
siderably broader than deep and just visible from 
above, not penetrating between the internasals, con- 
stantly eight upper labials, fourth and fifth entering 
the eye, and the scales in twenty-one (rarely nine- 
teen or twenty-three) rows. Ventrals 170 to 200 ; 
anal divided ; subcaudals 49 to 72. 



PLATE X 




< 
u 

a 
2 -3 



•j 

Id 
Z 






o 
u 



CORONELLA 



203 



Coloration. — Brown, greyish, yellowish, or reddish 
above, with dark brown or black spots or transverse 
bars, sometimes with four dark stripes in addition ; 
dark dots in front of the apical pits as in the preceding 
species ; a pair of elongate dark spots or a U-shaped 
marking on the nape ; a dark streak from the eye 
to the angle of the mouth, and a dark cross-bar 
from eye to eye, across the prefrontal shields ; a dark 
line below the eye. 
Lower parts yellow, 
orange, or coral red, 
with large, mostly «* v 





Fig. 28 (after Sordelli) 



quadrangular black 
spots, often ar- 
ranged in chess- 
board fashion, or 
with two series of 
black spots (Plate 
X.), which may be confluent into two longitudinal 
bands. 

Total Length. — 26 inches. 

Distribution. — South of France (as far north as 
the Charente-Inferieure to the west, the Dauphine 
to the east), the whole of Spain and Portugal, 
Southern Tyrol, Italy, and Sicily. It has not been 
recorded from higher than 2,500 feet in the 
Alps. Rare in Northern Morocco and Algeria. In 
many localities in Europe it occurs alongside with 
C. austriaca. 



204 COLUBRID^E 

Habits. — All that is known to me of the habits 
of this close ally of the preceding species is derived 
from the works of Bonaparte, Gene, and Schreiber, 
and from a note by Gachet, who observed it near 
Bordeaux and described it under the name of 
Coluber rnbens. According to these authors, it 
frequents dry and rocky localities as well as old 
walls, in which it finds a refuge and a good supply 
of the lizards on which it feeds. A large specimen 
from Albano, near Rome, preserved in the Genoa 
Museum, had swallowed a full-grown Chalcides 
tridactylus. This Coronella is crepuscular, rarely 
showing itself in the daytime, leaving its retreat 
only after sunset, and has been observed to crawl 
about by moonlight. Its movements are slow, 
which accounts for crushed specimens being often 
met with on paths or roads. Contrary to the rule 
in C. austriaca, it is extremely gentle, seldom attempt- 
ing to bite. 

Reproduction.- -Whether this species is ovovivip- 
arous, like its European congener, has not, I think, 
been ascertained. All we know on this matter is 
that a female found dead on a road near Bordeaux 
by M. Lataste at the end of June contained eggs 
which showed no trace of embryos. This does not, 
however, settle the question, as the young would 
not be born until at least two months later. Accord- 
ing to Gene, pairing takes place in May, when 
specimens have been observed to congregate in 
considerable numbers. 



CONTIA 205 

Genus CONTIA, Baird and Girard 

Maxillary teeth subequal. Head not or but 
slightly distinct from neck ; eye moderate or rather 
small, with round pupil. Nasal single; no subocular 
shields. Body moderately elongate ; scales smooth, 
with apical pits. Tail moderate. 

This genus, with certain modifications in the 
above definition, is made to embrace about twenty- 
five species from South- Western Asia and Sind and 
North and Central America. One of the species 
inhabiting Asia extends into a very small part of 
Europe. 

16. Contia modesta, Martin 
The Dwarf Snake 

Form. — Moderately slender. Head small, quite 
flat above ; snout obtuse, feebly prominent. Length 
of tail four to five times in the total length. 

Head -Shields. — Rostral a little broader than deep, 
visible from above. Suture between the internasals 
as long as or a little shorter than that between the 
prefrontals. Frontal once and a half to once and 
two-thirds as long as broad, as long as or longer 
than its distance from the end of the snout, shorter 
than the parietals, as broad as or a little broader 
than the supraocular, widely separated from the 
preocular. Nostril in the middle or upper part of 
the nasal. Loreal square or longer than deep. One 



206 



COLUBRID^] 





(rarely two) pre- and two (rarely one) postoculars. 
Temporals i + 2 ; parietal sometimes nearly touch- 
ing the fifth upper labial. Upper labials seven, 
third and fourth entering the eye. Four (rarely 

five) lower labials in 
contact with the an- 
terior chin - shield ; 
posterior chin-shields 
smaller than the an- 
terior, and separated 
from each other by 
one or two rows of 
scales. 

Fig. 29 (after Sordelli) 3^ ^^ a singJe 

apical pit, in seventeen rows. Ventral shields 150 
to 191 ; anal divided; subcaudals 53 to 78. 

Coloration. — Not unlike that of a young Zamenis 
gemonensis. Greyish-olive above, uniform or each 
scale lighter in the centre. The greater part of the 
upper surface of the head behind the snout, to- 
gether with the nape, black in the young, with a 
yellow cross-bar or a pair of yellow spots between 
the eyes, the bar sometimes confluent with the 
yellow postoculars, and a horseshoe-shaped band 
of the same colour on the temples and across the 
occiput (Plate X.) ; the black of the nape again edged 
with yellow behind. More or less distinct traces 
of these markings are preserved in adult specimens. 
Upper lip yellowish, with black spots or bars on 



CCELOPELTIS 207 

the sutures between the shields. Lower parts uni- 
form white or yellowish. 

In the var. semimaculata, Boettger, from Chios, 
small dark spots are scattered over the upper parts 
of the anterior half of the body. 

Size. — This snake rarely reaches a length of 
19 inches. It is the smallest Colubrid of Europe. 

Distribution. — The Caucasus up to about 5,000 
feet, Asia Minor, Chios, Cyprus, Syria, Mesopotamia, 
and North- Western Persia. The northern slope 
of the Caucasus appears to be the only part of 
Europe included in its habitat. The British Museum 
possesses two specimens labelled as from Constanti- 
nople, but the presence of this species in European 
Turkey requires confirmation. 

A closely allied species, which has been confounded 
with C. modesia, C. collaris (Menetries), and which 
also inhabits the Caucasus without having been 
recorded from the northern slope, is distinguished 
by having the scales in fifteen rows (very rarely 
seventeen), and the posterior chin-shields in contact 
with each other. 

Habits. — Nothing is known as regards this species, 
but the North American members of the genus 
Contia are chiefly insectivorous and oviparous. 

Genus CCELOPELTIS, Wagler 

Maxillary teeth small and subequal, followed after 
a short interspace by one or two very large grooved 



208 COLUBRID^E 

fangs situated below the posterior border of the eye ; 
anterior mandibular teeth strongly enlarged. Head 
not very distinct from neck, with angular canthus 
rostralis and projecting supraocular; eye large, with 
round pupil ; nostril a crescentic slit in a single or 
divided nasal. Body elongate ; scales smooth, more 
or less distinctly grooved longitudinally in the adult, 
with apical pits. Tail moderately long. 

The range of this genus, which comprises only 
two species, extends over Southern Europe, South- 
Western Asia, and North Africa. 

17. Ccelopeltis monspessulana, Hermann 

{Natrix lacertina, Wagler ; Coluber insignitus, 

I. Geoffroy) 

The Montpellier Snake 

Form. — Slender ; head elongate, narrow, concave 
above on the snout and between the eyes; snout 
projecting, rounded, with raised canthus and con- 
cave loreal region. Tail about one-fifth to one- 
fourth of the total length. 

Head-Shields. — Rostral nearly as deep as broad, 
just visible from above. Internasals much shorter 
than the prefrontals. Frontal very narrow, twice to 
twice and a half as long as broad, its width in the 
middle not more than half that of the supraocular, 
widening in front and extending beyond the supra- 
oculars to join the preoculars, longer than its distance 
from the end of snout, as long as or a little longer 



PLATE XI 



m 




COELOPELTIS MONSPESSULANA 
After Sordelli 




MACROPROTODON CUCULLATl'S 




TARBOPHIS IBERUS 
After Sordelli 




TARBOPHIS FALLAX 



CCELOPELTIS 



209 




than the parietals. Two loreals. One preocular, the 
upper portion of which is much enlarged, and en- 
croaches upon the area occupied in other snakes by 
the prefrontal and the supraocular ; two (rarely three) 
postoculars. Temporals 2 + 3 or 4. Upper labials 
eight (rarely nine), fourth and fifth (or fifth and 
sixth) entering the eye. Four or five lower labials 
in contact with the anterior chin-shields, which are 
as long as or shorter than the posterior. 

Scales with 
single apical 
pits, in seven- 
teen or nine- 
teen rows, 
longitudinally 
grooved in the 
adult, less dis- 
tinctly in the 
young. Ven- 
tral shields 160 to 189; anal divided; subcaudals 68 
to 102. 

Coloration. — The young is elegantly marked with 
dark brown and yellowish-white on a pale brown 
ground. On the head, the principal dark markings 
usually are an oblique band on the posterior half of the 
supraocular shield, and another, or a large spot, on 
the parietal, sometimes produced backwards, and 
forming with its fellow a /^-shaped band, separated 
from a large occipital blotch by a yellowish space ; 
14 




Fig. 30 



210 C0LUBRID.E 

anterior half of the frontal shield and shields on the 
snout edged with dark brown ; a dark streak, some- 
times broken up into small spots, on the temporal 
region ; yellow spots on the pre- and post-oculars ; 
lips brown, with large, yellow, black-edged spots, or 
yellow with brown spots ; chin with three brown 
longitudinal streaks. Back with a vertebral series of 
large roundish dark spots or narrow cross-bars; 
small spots on the sides, these sometimes forming 
longitudinal series or accompanied by yellowish streaks 
or dots ; these markings often confluent into three 
longitudinal streaks on the tail. Belly pale brownish, 
greyish, or reddish, with numerous pale spots, some- 
times with a dark brown line on each side. The 
adult is greyish, reddish-brown, or olive above. Some 
specimens preserve more or less the markings of the 
young, and the dark dorsal markings (Plate XI.) may 
be edged with yellowish and ocellar in appearance 
(var. insignitus, Geoffroy) ; the belly is yellowish, with 
small dark spots which usually form longitudinal 
series, and may be confluent into streaks. A variety 
common in Dalmatia (ueumayeri, Fitzinger) is brown 
or olive above, without spots, sides with a bluish- 
grey lateral band, the scales on which are edged with 
black, the belly uniform yellow. Other specimens 
are brown or reddish, with light edges to the scales 
on the sides, or with yellowish lateral lines, or dark 
olive or dark brown above and black on the sides, 
each scale with a yellowish central spot ; in the last- 



CCELOPELTIS 211 

mentioned the second third of the back may be almost 
entirely black, and the belly dark olive-grey in the 
middle and yellowish on the sides. Iris brown, 
with a golden or coppery circle round the pupil. 

Size. — This handsome snake grows to a length of 
6J feet. Specimens 5 to 6 feet long are not un- 
common. 

Distribution. — Mediterranean coast of France and 
Western Liguria, Spain and Portugal, Sicily, Lam- 
pedusa, eastern coast of the Adriatic, Greece and 
eastern islands of the Mediterranean, Mediterranean 
coast of Asia and Sinaitic Peninsula, eastwards to 
the Caucasus and Persia, North Africa from Egypt 
to Rio de Oro. It is not known to occur above 
2,300 feet altitude in Europe. 

Habits. — A lively, swift snake, living on land and 
on low bushes, often found near human habitations. 
Some specimens are very vicious, whilst others show 
a gentle disposition after a short period of captivity. 
A specimen nearly 6 feet long, which I kept for 
some time, never attempted to bite when handled, 
and some have become so tame as to take food from 
the hand. The sense of sight appears to be better 
developed than in any other European snake. The 
food consists chiefly of mammals, even large rats 
and young rabbits, birds such as chickens, partridges, 
and quails, lizards, and other snakes, which, if of 
considerable size, are not swallowed until paralyzed 
or killed by the effect of the poison. In Eastern 



212 colubrim; 

Europe, Vipera ammodytes is said to be the principal 
enemy of Ccelopeltis, and the two snakes are conse- 
quently seldom found together in the same locality. 

Many experiments have been made on the action 
of the poison of this Opisthoglyph. Peracca and 
Deregibus, as well as, later, Phisalix, found a striking 
similarity with the symptoms of Cobra poison in 
their experiments on small animals, the suspension 
of the respiration occurring in a few minutes, the 
blood being otherwise unaffected. It has been 
stated by some authors that Ccelopeltis poison has 
little or no action on man, but a French zoologist, 
E. Taton-Baulmont, having been bitten in the index- 
finger by a four- foot-long specimen at Algiers, the 
swelling extended within thirty hours up to the 
shoulder, and was accompanied by fever and nervous 
troubles. As a rule, however, the bite of this snake 
has no poisonous effect on man, from the fact that 
the fangs conveying the venom are situated so far 
back in the mouth as not to come into action. 

Reproduction. — According to Werner, the eggs, 
four to twelve in number, are laid in July, and 
measure 2 inches in length and J inch in width. 

Genus MACROPROTODON, Guichenot 

Maxillary teeth few and very unequal in size, fourth 
and fifth or fifth and sixth enlarged and followed by 
an interspace, the two last teeth fang-like and grooved, 
situated just behind the eye ; sixth mandibular tooth 



MACROPROTODON 



213 



fang-like, and separated from the remainder by an 
interspace. Head slightly distinct from neck ; eye 
rather small, the pupil vertically elliptic or sub- 
elliptic when contracted. Body moderately elon- 
gate ; scales smooth, with apical pits. Tail moderate 
or rather short. 
A single species. 

18. Macroprotodon cucullatus, I. Geoffroy 
The False Smooth Snake 

Form. — Very similar to the Smooth Snakes, with 
which it has been confounded, but snout broader and 
very strongly de- 
pressed. Tail 
five and a half to 
six and a half 
times in the total 
length. 

Head - Shields. 
— R o st r al at 
least twice as 
broad as deep, 





Fig. 31 



not or but scarcely visible from above. Inter- 
nasals as long as or a little shorter than the pre- 
frontals. Frontal not much broader than the supra- 
ocular in the adult, once and a half to twice as 
long as broad, as long as or longer than its dis- 
tance from the end of the snout, shorter than the 
parietals. Nasal usually semidivided. Loreal once 



214 COLUBRIDJE 

and a half to twice as long as deep. One preocular, 
extending to the upper surface of the head, but not 
reaching the frontal ; two (rarely one or three) post- 
oculars. Temporals i + 2. Upper labials eight, fourth 
and fifth entering the eye, sixth usually in contact 
with the parietal. Four or five lower labials in contact 
with the anterior chin-shields, which are as long 
as or a little shorter than the posterior. 

Scales with mostly single apical pits, the pits 
sometimes paired on the sides of the body, in twenty- 
one or twenty-three rows (nineteen to twenty-five in 
North African specimens). Ventral shields 153 to 
192 ; anal divided ; subcaudals 40 to 54. 

Coloration. — Pale brown or greyish above, with 
small dark brown or blackish spots or with more or 
less distinct darker and lighter longitudinal streaks. 
Upper surface of head with dark brown vermicula- 
tions; a dark brown or black, often light-edged 
occipito-nuchal band, extending downwards to the 
gular region and produced forwards into a point to 
between the parietal shields ; a dark brown or black 
streak on each side of the head from the end of the 
snout, through the eye, to the last lower labial shield, 
traversing the four last upper labials, which are yellow- 
ish above and below the streak (Plate XL). Lower 
parts yellow or coral red, with black spots, which 
may form a tessellated pattern, two longitudinal 
series, or be so crowded as to fuse into a band along 
the middle of the belly and tail. 



MACROPROTODON 215 

The above description is taken from Spanish 
specimens (Badajos, Algeciras, Andalucia), but the 
variations are very great when we take North Africa 
into consideration. The nuchal band may be 
narrow or broken up into spots, the median of 
which sometimes forms a longitudinal streak, or 
so much enlarged as to fuse with the dark markings 
on the upper surface of the head ; in some specimens 
(from Morocco and Algeria) the upper surface of the 
head and the nape may be entirely ink black, or the 
whole head black above and beneath with the excep- 
tion of a whitish streak bordering the upper lip. 
The dark streak from the eye to the angle of the 
mouth may be absent, or reduced to a short oblique 
streak below the eye. Irrespective of the variations 
in the markings of the upper parts, the lower parts 
may be more or less spotted with black, or immacu- 
late. 

Some specimens of this small snake bear a general 
resemblance to Coronella girondica, with which Macro- 
protodon has sometimes been confounded. But a 
careful examination of its whole structure shows it 
to be more afnne to Ccelopeltis and Tarbophis, the 
other European representatives of the Opisthogly- 
phous Colubrids. 

Size. — The largest European specimen examined 
measures 17J inches. Specimens up to 22 inches 
long occur in Algeria and Tunisia. 

Distribution. — In Europe this snake is only known 



216 COLUBRIDJE 

from Spain (Estremadura, New Castille, Andalucia), 
Portugal (Alemtejo), the Balearic Islands (Majorca 
and Minorca), and the island of Lampedusa. In 
North Africa it is generally distributed from the 
north coast of Egypt to the Rio de Oro ; in Algeria 
it penetrates into the northern parts of the Sahara. 
The specimen figured on Plate XI. is from Algeciras. 

Habits. — Appear to be similar to those of Coronella 
girondica. Crepuscular in its habits, it is usually 
found under stones or in burrows in the ground* 
Unless pursued, when it darts off with great rapidity, 
its movements are slow. It is very ready to bite, 
but no experiments have been made on the effects 
of its poison. The food consists chiefly of small 
lizards. 

Reproduction. — All that is known on this head is 
that, according to Doumergue, eggs are laid in July in 
Algeria. 

Genus TARBOPHIS, Fleischmann 

Maxillary teeth few, anterior longest, gradually 
decreasing in size posteriorly, and followed, after an 
interspace, by a pair of enlarged, grooved fangs, 
situated below the posterior border of the eye ; 
anterior mandibular teeth strongly enlarged. Head 
distinct from neck ; eye moderate or rather small, 
with vertically elliptic pupil. Body moderately 
elongate ; scales smooth, oblique, with apical pits. 
Tail moderate or rather short. 

The eight species of this genus inhabit South- 



TARBOPHIS 



217 



Eastern Europe, South- Western Asia, and Africa. 
Two are dealt with here. 

ig. Tarbophis fallax, Fleischmann 

(Aihcrophis vivax, Bonaparte) 

The Cat-Snake 

Form. — Moderately slender. Head much de- 
pressed. Tail five and a half to seven times in the 
total length. 

Head-Shields. — Rostral broader than deep, just 
visible from above. Internasals shorter than the pre- 
frontals. Fron- 
t a 1 much 
broader than 
the supraocu- 
lar, once and 
one - fourth to 
once and a 
half as long as 
broad, as long 
as its distance 





Fig. 32 



from the end of the snout, shorter than the 
parietals. Nasal divided or semidivided. Loreal 
twice and a half to thrice as long as deep, entering 
the eye below the preocular, which is in contact with 
the frontal. Two (rarely three) postoculars. Tem- 
porals small, scale-like, 2 or 34-3 or 4. Upper 
labials eight (rarely seven or nine), third, fourth, and 
fifth (rarely fourth and fifth, or fourth, fifth, and 



218 COLUBRIDJE 

sixth) entering the eye. Three or four lower labials 
in contact with the anterior chin-shields ; posterior 
chin-shields very small and widely separated from 
each other by scales. 

Scales with single or paired apical pits, in nineteen 
or twenty-one rows, usually nineteen in European 
specimens. Ventral shields 186 to 222 ; anal divided ; 
subcaudals 48 to 73, 

Coloration. — Greyish above, with 40 to 57 brown 
or black spots or bars on the body ; a lateral series 
of smaller spots or vertical bars, alternating with the 
dorsals ; the first spot, on the nape, elongate, usually 
with one or three linear processes in front, extending 
on the head (Plate XI.) ; usually a dark streak on each 
side of the head, from the eye to the angle of the 
mouth. Lower parts whitish, speckled, spotted, or 
marbled with grey or brown. Iris brown, with a 
golden circle round the pupil. 

Size. — This species grows to a length of 2 feet 
10 inches. 

Distribution. — From Istriaand Dalmatiato Greece, 
the Archipelago, Constantinople, Asia Minor, Cyprus, 
and Northern Syria ; 2,600 feet appears to be its 
altitudinal limit. 

Habits. — Although to a certain extent crepuscular 
or nocturnal, the Cat-snake is often seen hunting in 
the daytime, its food consisting almost exclusively 
of lizards, rarely of small mammals. Its movements 
are rather slow. The names Katzenschlange and 



TARBOPHIS 219 

Ailurophis, translated Cat-snake, probably originated 
from the way in which this snake stalks its prey, and 
suddenly pounces upon it. According to Eiffe, the 
poison causes the death of a Lacerta vivipara in one 
minute, and P. de Grijs observed the larger Lacerta 
agilis to die in two or three minutes. As a rule 
even fresh-caught specimens allow themselves to be 
handled without attempting to bite ; some specimens, 
on the other hand, are very savage. Stony localities, 
old walls, and ruins, are the favourite abodes of this 
snake, which does well in captivity. 

Reproduction. — Seven or eight eggs are laid in 
July; they measure about ij inches in length and 
i inch in width. 

20. Tarbophis iberus, Eichwald 
The Caucasian Cat-Snake 

Very closely allied to the preceding, and differing 
from it only in the following points : Parietals 
shorter, slightly longer than the frontal, and anal 
entire. Loreal twice to twice and a half as long as 
deep. Fourth and fifth, or third, fourth, and fifth, 
labials entering the eye. Scales in nineteen or 
twenty-one rows. Ventrals 203 to 235 ; subcaudals 

54 t0 7°- 

Grey above, with 35 to 40 blackish spots on the 

body, the anterior largest and darkest ; a lateral 

series of smaller spots or vertical bars. Lower parts 



220 VIPERIDJE 

blackish, with small whitish spots and dots. Reaches 
a length of 3 J feet. 

This species inhabits the Caucasus, and, being on 
record from the northern slope (Kuban River), has 
to be included in the European fauna. It occurs 
also in Mesopotamia, a specimen from Bagdad being 
preserved in the British Museum. The young 
specimen figured on Plate XI. is stated to be from 
Constantinople. 

Nothing is known of its habits, which are probably 
the same as those of Tarbophis fallax. 



Fourth Family: VIPERID^E 

Maxillary, palatine, and pterygoid bones movable, 
the first much abbreviated, erectile perpendicularly 
to the large transverse bone, and supporting a pair 
of large canaliculated poison fangs ; mandible with- 
out coronoid bone. No vestiges of pelvic arch. 

All more or less poisonous, some being among the 
most dangerous of snakes. 

Divided into two subfamilies, each of which is 
represented by one genus in Europe : 

Viperince. — No pit on the side of the snout ; 
maxillary bone not hollowed out. 

Crotalince. — A deep pit on each side of the snout, 
between the nostril and the eye ; maxillary bone 
hollowed out above. 

The Viperince inhabit nearly the whole of Europe, 



VIPERA URSINI1 

From Zoological Society's Proceedings 



PLATE XII 





VIPERA RENARDI 
Fro»t Zoological Society's Proceedings 




VIPERA BERUS 
After Sordelli 



VIPER A 221 

Asia, and Africa ; the Crotalince are Asiatic (one 
species extending its range into a small part of 
South-Eastern Europe) and American. 

Genus VIPERA, Laurenti 

Head distinct from neck, covered with small 
shields or scales, with or without distinct frontal and 
parietal shields ; eye moderate or small, with vertical 
pupil, separated from the labial shields by scales ; 
nasal separated from the rostral by a naso-rostral. 
Body short. Scales keeled, with apical pits. Tail 
short. 

Of the eleven species of this genus, six are found in 
Europe ; two inhabit South-Western Asia, one the 
Indo-Malay region, and two Eastern Africa. 

The distinction of the European species is one of 
considerable difficulty, owing to their close relation- 
ship and the presence of intermediate forms connect- 
ing them. Matters being so, it seems curious that 
the Common Adder should have been regarded by 
so many authors as generically distinct from the 
Asp Viper, under the name of Pelias berus. It is 
highly probable that hybrids are produced in those 
districts where two species coexist, as in some parts 
of France, North Italy, and Austria. 

21. Vipera ursinii, Bonaparte 
Orsini's Viper 
Form. — Short and stout. Snout obtusely pointed, 
flat above or with the canthus slightly raised. Eye 



222 



VIPERID.E 




very small, usually smaller than the nasal shield, its 
horizontal diameter usually not exceeding its distance 
from the posterior border of the nostril, its vertical 
diameter often less than and rarely exceeding its 
distance from the mouth. Length of tail seven to 
eight times in total length in males, nine and a half 
to twelve times in females. 

Head-Shields. — Rostral as deep as broad or slightly 
deeper than broad, visible from above, in contact 

with one 
apical shield 
(rarely with 
two). Dis- 
tinct frontal 
and (usually) 
p a r i e t a 1 
shields, the 
former once 

Fig. 33 (From Proceedings of the Zoological and a half 
Society, 1893) , 

to once and 
two-thirds (rarely once and one-third) as long as 
broad, as long as its distance from the rostral 
or the end of the snout, and nearly always longer 
than the parietals ; the latter always in contact 
with the former, rarely broken up into small 
shields. Four to seven small shields on the snout 
between the canthals, of which there are two on 
each side. Supraocular well developed, extending 
posteriorly beyond the vertical of the eye, separated 




VIPERA 223 

from the frontal by one to three shields, very 
rarely in contact with it. Six to ten scales round 
the eye, usually eight or nine, the upper pre- 
ocular usually in contact with the nasal ; a single 
series of scales between the eye and the labials. 
Nasal single. Temporal scales smooth. Upper 
labials six to nine, usually seven or eight, usually 
third or third and fourth below the eye. Three 
(rarely four) lower labials in contact with the chin- 
shields, of which there is but one pair. 

Scales in nineteen (rarely twenty or twenty-one) 
rows, with two apical pits, strongly keeled on the back, 
less strongly on the sides, outer row smooth. Ventral 
shields 120 to 135 in males, 125 to 142 in females ; anal 
entire; subcaudals 30 to 37 in males, 20 to 28 in 
females. By adding the subcaudals to the ventrals in 
a hundred specimens, the total numbers are 153 to 169 
in males, 150 to 168 in females. 

Coloration. — Unlike its ally V. bents, V. ursinii 
shows no sexual differences in the coloration. The 
ground colour of the back is usually yellowish or pale 
brown, sharply defined from the darker grey or brown 
colour of the sides ; some specimens, however, are of 
an almost uniform brown ground colour. The light 
colour of the back is relieved by a series of more or 
less regular transversely oval, elliptic, or rhomboidal 
dark brown, black-edged spots, some or all of which 
may run together to form a wavy or zigzag band 
(Plate XII.). Two or three longitudinal series of dark 



224 VIPERID.E 

brown or black spots extend along the sides, the upper 
series, if present, occupying the space between the 
series of spots continued from the postocular band 
and the large dorsal spots or vertebral band, the 
lowermost following the outer row of scales. Small 
dark spots and one or two A-shaped markings are 
present on the upper surface of the head ; an oblique 
dark band proceeds from the eye to the angle of the 
mouth, and is not infrequently confluent with the 
branches of the occipital A. The rostral and the 
labial shields are uniform yellowish-white, rarely 
with a few small, blackish spots or with brown 
borders. The chin and throat are yellowish-white, 
rarely with some blackish spots. The ventral and 
subcaudal shields are black, with transverse series of 
small white spots, or grey checkered with black and 
white, or whitish with small round black spots ; the 
tail is but rarely (females) tipped with yellow. 

The form recently described as V. macrops, Mehely, 
from Bosnia and Herzegovina, is distinguished by a 
usually larger eye, the vertical diameter of which 
equals or a little exceeds its distance from the mouth, 
and the parietals are often broken up into small 
shields. The postocular dark band is often reduced, 
originating at some distance from the eye, and is not 
prolonged beyond the mouth. In this geographical 
race melanic specimens occasionally occur, which are 
dark brown or blackish above, the lower parts not 
differing from those of the typical form. 



VIPERA 225 

Size. — 20 inches appears to be the usual maximum 
size reached by this species, but, Dr. Werner informs 
me, a female 2 feet long has been found in Lower 
Austria. 

Distribution. — First discovered in Italy in the 
Abruzzi, this species has since been found in the 
Basses-Alpes, near Digne, in various parts of Hun- 
gary, in Lower Austria, on the island of Veglia in 
Istria, and in Bosnia, Bulgaria, Herzegovina, and 
Montenegro. A very broken and curious distribution, 
the more so as V. ursinii is essentially a form of the 
plain in Lower Austria and Hungary, and an alpine 
form in Italy, in France, and in the Balkan Penin- 
sula, where it only occurs between 3,000 and 6,800 
feet. In no part of its habitat does it appear ever to 
be found in company with V. berus. 

Habits. — Only a few specimens have hitherto been 
found in Italy and in France, but the species occurred 
up to a few years ago in extraordinary numbers in 
Lower Austria, in the immediate vicinity of Laxen- 
burg. The intendant of the imperial castle pays a pre- 
mium for the destruction of Vipers, and in the course 
of one year (1892) more than 1,000 specimens were 
brought to him. These snakes are found principally, 
though not exclusively, in the marshy meadows 
around the park, where they may be seen about in 
the daytime from May to September, feeding chiefly 
on lizards (Lacerta agilis), and also on small rodents. 
The lizards are swallowed as soon as seized, without 
15 



226 VIPERIDjE 

the effect of the poison being awaited as in other 
Viperid snakes. This Viper is as a rule of gentle 
disposition, allowing itself to be handled without 
attempting to bite, and village boys have been seen 
playing with them. Although occurring in such 
enormous numbers at Laxenburg, no accident from 
snake-bite has ever been heard of. The form from 
the Balkan Peninsula {V. macrops) is even more pacific 
still, and is believed never to make use of its poison 
apparatus, its food consisting of orthopterous insects. 
According to Captain Veith, who has collected a large 
number of specimens of this Southern form, the con- 
tents of the stomach as well as the excrements show 
this snake to feed exclusively on grasshoppers. On 
one occasion a big specimen showed such a swelling 
of the body as to lead to the conclusion that it had 
swallowed a mouse, but it soon after disgorged what 
proved to be a ball made up of the agglutinated 
remains of at least a hundred grasshoppers. When 
handled, this Viper hisses or even pretends to snap, 
but with closed mouth, never biting unless seriously 
hurt. The poison appears to have little effect on man. 
Reproduction. — Nothing has been published on the 
breeding habits of this species, but in a letter to 
the author, dated January 14, 1913, Herr L. 
von Kirchroth, who has examined over 4,000 
specimens since 1890, says the young are born in 
July or August, exceptionally as early as June. 
Young females bring forth from six to eight young, 



VIPERA 227 

older females from eight to eighteen ; but a large 
female from Lower Austria is reported to have con- 
tained as many as twenty-two. The length of the 
new-born young is from 5 to 6 inches, and it grows 
rapidly within the first week, probably through 
stretching out, without taking any food. 

According to Captain Veith, the form described as 
V. macrops brings forth only from three to five 
young. 

22. VlPERA RENARDI, Christoph 

Renard's Viper 

Form. — Similar to the preceding species, but snout 
more pointed, the raised canthi rostrales meeting at 
an acute angle. Eye usually as large as in V. bents, 
nearly as large as the nasal shield; its horizontal 
diameter equal to its distance from the posterior or 
anterior border of the nostril, its vertical diameter 
equal to or a little less than its distance from the 
mouth. Length of tail seven and a half to nine times 
in total length in males, eight to ten times in 
females. 

Head-Shields. — Rostral as deep as broad or a little 
deeper than broad, just visible from above, and in 
contact with a single apical shield. Distinct frontal 
and (usually) parietal shields, the former once and 
two-thirds to twice and one-third as long as broad, as 
long as or longer than its distance from the end of the 
snout, usually longer than the parietals ; the latter 



228 



VIPERID.E 



always in contact with the former, unless broken up 
into small shields. Two to six, usually three or four, 
small shields on the snout between the canthals, of 
which there are two on each side, the second broadly 
in contact with the supraocular. Supraocular well 
developed, extending posteriorly beyond the vertical 
of the eye, separated from the frontal by one to four 
shields. Nine to eleven, usually ten, scales round 
the eye, the upper preocular usually in contact with 

the nasal; either 
a single series of 
scales between 
the eye and the 
labials, or two 
series except 
under the centre 
of the eye, which 
is separated 

Fig. 34 (From Proceedings of the Zoological from the fourth 
Society, 1893) . , . _ 

labial by a 
single scale. Nasal single. Temporal scales all 
smooth, or the upper faintly keeled. Upper labials 
eight or nine, fourth or fourth and fifth below 
the eye. Four (rarely five) lower labials in contact 
with the chin-shields, of which there is but one 
pair. 

Scales in twenty-one (very rarely nineteen) rows, with 
two apical pits, strongly keeled, outer row smooth or 
feebly keeled. Ventral shields 130 to 148 in males, 





VIPERA 229 

130 to 150 in females; anal entire ; subcaudals 31 to 
37 in males, 24 to 30 in females. 

Coloration. — As in V. ursinii, the sexes are alike in 
coloration. European specimens (Plate XII.) are very 
similar to V. ursinii, except that the labial shields are 
markedly dark-edged and speckled or spotted with 
brown or black. The dorsal band or series of spots 
is dark brown, edged with blackish ; the ground 
colour of the middle of the back and of the scales of 
the two outer rows on each side is yellowish, of the 
sides (four rows of scales) greyish-brown with two or 
three series of dark brown spots ; two dark A-shaped 
markings on the head; a dark postocular streak, 
extending or not to the side of the neck. The lower 
parts are whitish or pale greyish, with blackish dots, 
of which there is a series of larger ones along each 
side of the belly. The tip of the tail is never 
yellow. 

Central Asian specimens are of a pale yellowish 
sand-colour, with a brown, dark-edged dorsal zigzag 
band or series of spots and two series of small spots 
on the sides. Belly whitish, dotted or spotted with 
black, or uniform blackish. 

Size. — 23 inches is the length of the largest 
specimen examined. 

Distribution. — In Europe V. renardi, which has 
long been confounded with V. bents, is abundant in 
the district of Uralsk, in the steppe around Sarepta, 
in Crimea, and it is also found in Cis-Caucasia and 



230 viperim: 

in Bessarabia. Its range extends far into Central 
Asia, being known from the Khirghiz steppes, the 
Emba steppes, the steppes near the Alatau, on the 
borders of the Urdshar, and in the Semipolatinsk 
district. Around Sarepta it is common in the bare 
steppe, and only exceptionally occurs in localities 
overgrown with willows and small shrubs. 

Habits. — Nothing has been published concerning 
the habits of this snake, except that it is more sensitive 
to cold than V. berus and does not appear before the 
middle of April, retiring to its winter-quarters in the 
beginning of October. The food consists of small 
mammals and lizards. 

Reproduction. — Pairs in May, and brings forth five 
to seven young in August, these young at birth 
measuring about 54 inches. 

23. Vipera berus, Linnaeus 
The Northern Viper, or Adder 

Form. — Short and stout. Snout flat above, rarely 
slightly concave, the upper contour broadly rounded 
or truncate in front, the canthus well marked, some- 
times slightly raised, the loreal region nearly vertical. 
Eye as a rule smaller in females than in males, as 
large or nearly as large as the nasal shield ; its 
vertical diameter equals or a little exceeds its dis- 
tance from the mouth. Length of tail five and a 
half to nine times in total length in males, eight to 
ten and three-quarter times in females. 



VIPERA 



231 



Head-Shields.— Rostral as deep as broad or slightly 
broader than deep, rarely once and one-third as deep 
as broad, not or but scarcely visible from above. In 
addition to the supraoculars, three large shields, the 
frontal and the parietals, are as a rule present on 
the top of the head. Frontal as long as broad or a 
little longer than broad, rarely much longer than 
broad, once and a half to twice and a half as broad 
as the supraocular, from which it is as a rule separated 
by one to four 
shields, as long 
as or a little 
shorter than its 
distance from the 
rostral, as long as 
or a little shorter 
than the parietals. 
Parietals usually 
in contact with 





Fig. 35 (after Sordelli) 



the frontal and separated from the supraoculars 
by small shields, but sometimes in contact with 
both, or separated from the frontal. Exception- 
ally, in specimens from Great Britain, Germany, 
and Austria, the parietals, or the frontal and the 
parietals, are broken up into scales, and this is more 
frequently the case in specimens from North-Western 
Spain (var. seoanei). Upper surface of snout bor- 
dered by six (rarely by five or four) small shields, viz., 
two apicals (rarely one), and on each side two can- 



232 VIPERID.E 

thals, the second of which is usually in contact with 
the supraocular; canthals very rarely united into one 
shield ; the space between these shields covered by 
four to twenty flat or convex, juxtaposed scales, 
which very exceptionally are fused into a single 
large shield. Supraocular usually extending pos- 
teriorly beyond the vertical of the eye. Six to thir- 
teen scales round the eye, usually eight to ten ; two 
or three superposed scales, rarely two vertical series 
of scales, separate the preoculars from the nasal, 
which is single. As a rule a single series of scales 
intervenes between the eye and the labials ; specimens 
with two series are of very exceptional occurrence 
(single specimens from Isle of Arran, Normandy, 
Southern Norway, and Carniola, in the British 
Museum), but there are occasionally two series 
except just below the centre of the eye. Upper labials 
six to ten, usually eight or nine ; fourth or fourth 
and fifth (rarely third and fourth) below the eye. 
Temporal scales smooth, rarely feebly keeled. Three 
or four (rarely five) lower labials in contact with 
the single pair of chin-shields. 

Scales in twenty-one (rarely nineteen or twenty- 
three) rows, with two apical pits, strongly keeled, 
those of the outer row smooth or feebly keeled. 
Ventral shields 132 to 150 (usually 137 to 147) in 
males, 132 to 158 (usually 140 to 150) in females; 
anal entire ; subcaudals 32 to 46 (usually 35 to 40) 
in males, 24 to 38 (usually 28 to 33) in females. 



VIPERA 233 

Coloration. — It is characteristic of this species, 
contrary to the rule in snakes, to present such 
marked differences of colour, according to the sexes, 
that these can be distinguished in most cases from 
that character alone. 

Whitish or pale grey specimens, with black belly 
and jet black dorsal markings (Plate XII.), are males. 
Brown and brick-red specimens, with the markings 
of a more or less dark brown or red, are females. 
There are also brown, reddish-brown or olive males 
with the markings of a deep black, and grey males 
with brown markings. A very pretty colour variety, 
which affects only females, is olive with brick-red 
band and spots. Some males, just before exuvia- 
tion, have the lower surface of a pale greyish-blue 
(Coluber cczruleus, Sheppard), with the outer ends of 
the shields black. Specimens with yellowish-white 
chin and throat, which may be tinged with red, are 
females ; males have the throat black, or whitish with 
the scales spotted or edged with black. Exceptional 
females occur (in Carniola) which in this respect re- 
semble the males. 

The markings vary considerably. Those on the back 
usually consist of a wavy or zigzag longitudinal band, 
flanked on each side with a series of spots corre- 
sponding to its sinuses ; but this band may be partly 
or even entirely broken up into rhomboidal or trans- 
versely oval spots, or, losing its indentations, form a 
straight stripe edged on each side with a yellowish 



234 VIPERID.E 

streak (as in some specimens of the var. seoanei, from 
North-Western Spain). The markings may be absent 
altogether (var. concolor, Jan), or reduced to a narrow 
straight vertebral band (Pelias dorsalis, Gray). In 
the var. seoanei the zigzag band is often replaced by 
a dark brown vertebral band, three to five scales 
wide, bordered on each side by a series of subtriangu- 
lar or crescentic black spots opposite to each other, 
as in the Pyrenean specimens of V. aspis. A pair of 
elongate dark markings are usually present on the 
back of the head, affecting the following shapes : 
J\,X#)Vi )(i By uniting together, this pair of mark- 
ings may form a A or an X. An oblique dark streak 
extends on each side from the eye to the last labial 
shields, being sometimes prolonged a short way down 
the neck. The snout and vertex may be uniform or 
bear some symmetrical dark spots, or in some males 
be entirely black, the black involving the apex of the 
A-shaped marking. The labial shields are whitish or 
yellowish, those at least which are anterior to the 
eye being more or less broadly edged with brown or 
black. 

The belly and the lower surface of the tail vary 
from grey or brown to bluish, blackish-grey, or black, 
the sides usually dotted or spotted with whitish ; 
sometimes, especially in females, the belly is dark 
grey, each shield with a white posterior border which 
is broken up by a series of small roundish black spots. 
The end of the tail is often yellowish, bright yellow, 



VIPERA 235 

or pale orange below, rarely coral red, more com- 
monly in females than in males. 

The iris is usually coppery red, more rarely golden 
suffused with brown. 

Black specimens occur, more or less frequently, all 
over the habitat of this species, and are often referred 
to as V. prester, Linnaeus. A distinction has to be 
made between individuals which are black through 
darkening of the ground colour, and such as are thus 
coloured through expansion and confluence of the 
markings. The latter are males, and among them 
we may find intermediate stages showing how this 
melanism is brought about ; in one case the black of 
the back is separated from the black of the sides by 
a narrow light brown wavy stripe, the remains of 
the ground colour. When, as in all females, and 
occasionally in males, the black is the result of a 
gradual darkening of the ground colour, the typical 
markings may still be detected under certain lights. 
Some specimens (from Schneeberg, Lower Austria) 
are black, with scattered golden dots, or of a dark 
mahogany brown speckled with yellowish. In 
nearly all the black specimens at least a few dots of 
whitish are visible on the lips, and of yellow under 
the end of the tail. 

Most of the variations enumerated above occur irre- 
spective of the geographical distribution. Two forms, 
however, deserve to be regarded as ill-defined local 
races : the var. seoanei, Lataste, from North-Western 



236 viperim: 

Spain, in which, in addition to the peculiarities of 
coloration mentioned in the description, the canthus 
rostralis is frequently more distinctly raised, and the 
frontal and parietal shields are often disintegrated 
into scales ; and the var. bosniensis, Boettger, from 
Bosnia, Carniola, and Carinthia, which is sometimes 
very suggestive of, and has been taken for, the typical 
form of V. aspis, having like it, though not at all 
constantly, two series of scales between the eye and 
the labials, and the zigzag band replaced by a series 
of dark bars across the back. The var. pseudaspis, 
Schreiber, from the plains of Sclavonia, described 
as straw yellow above, with narrow dark cross-bars, 
is hardly separable from the var. bosniensis. 

Size. — Viper a berus is said to reach very excep- 
tionally a length of 2 feet 11 inches. The largest 
specimen in the British Museum (from Belgium) 
measures 2 feet 3J inches : the largest British 
specimen 2 feet 3 inches. Both these specimens are 
females. The largest male measures 2 feet 2 inches. 

Distribution. — Vipera berus ranges over the whole 
of Northern Europe, to the extreme north of Scot- 
land, and the sixty-seventh degree in Scandinavia, 
and right across Northern Asia as far east as the 
island of Saghalien. It is generally distributed in 
Great Britain, occurring also on the Isles of Arran, 
Islay, Skye, Lewis, and Mull, rare or absent in 
some districts, common in others. Its distribution 
in Central and Southern Europe is irregular. In 



VIPERA 237 

Western France it does not extend much beyond the 
Loire to the south, only isolated captures being on 
record from the departments Vendee, Deux-Sevres, 
Vienne, and Indre. Of rare occurrence south of 
Paris, in the departments Yonne and Allier and in 
the mountains of Auvergne, it is again abundant in 
some parts of the Central Plateau. To the east it 
is recorded from the departments Aude, Haute- 
Marne, and Vosges. In Belgium, it is known from 
Flanders, Limburg, the Meuse Valley, and the 
Ardennes ; in Holland it is pretty generally distrib- 
uted in the uncultivated parts. It is spread over 
nearly the whole of the German Empire with the 
exception of the vine districts, where it is absent or 
extremely rare ; it is also very scarce in the mountains 
of the Black Forest ; it is on record from only a few 
localities in Lorraine, and has never been found in 
Alsatia. In South Germany it is rarely found below 
1,000 feet altitude. In Switzerland it is absent from 
the Jura, but occurs in the Alps chiefly between 
2,500 and 9,000 feet. To the East it extends to 
Russia, as far north as 64 , Austria- Hungary, con- 
fined to the hills in the south, and Roumania. 
In the Balkan Peninsula it occurs in the mountains 
of Bosnia, of Herzegovina, and of Bulgaria, up to 
7,000 feet. Absent from the South of France, it 
curiously reappears in the hills of the north coast of 
Spain, even at sea-level in Galicia, and in a few 
localities in North Portugal. On the southern side 



238 viperim: 

of the Alps it is much rarer than V. aspis, but it has 
established itself in a few low-lying districts in 
Lombardy, Venetia, and the neighbouring part of 
Emilia. 

Habits. — As we see from the above sketch of 
its distribution, the Adder generally avoids the 
hotter parts of Europe ; when found in the plain 
in the South, as in Italy, it dwells in marshy 
localities, and Bonaparte called it Marasso palustre 
(Marsh Viper) in opposition to his Marasso alpino, 
Vipeva ursinii. In the North, however, it usually 
selects in preference dry moors, sandy heaths, and 
hills well exposed to the sun, in which, although to 
a certain extent a nocturnal reptile, it delights to 
bask. Its food is very varied : weasels, mice, voles, 
shrews, moles, birds, lizards, slow- worms, frogs, 
salamanders, large slugs, have been found in the 
stomach, and the very young feed also on insects 
and worms. Of irascible temper as a rule, Adders 
are very ready to bite when fresh caught, but 
instances are known of their becoming quite tame 
in captivity, allowing themselves to be handled. 
As a rule they refuse food in captivity, but some 
have been known to live for as long as five years, 
being fed on lizards. Accidents from their bite, 
although seldom heard of in this country, are of 
frequent occurrence in France and in Germany, 
where many cases of fatal results on people have 
been recorded. 



VIPERA 239 

Reproduction. — Pairing takes place in April and 
May, and the young, five to twenty in number, are 
born in August or September, exceptionally as early 
as the end of July ; the young, on releasing them- 
selves from the thin, transparent membrane in which 
they are enclosed at birth, measure 6 to 8 inches. 
According to J. Geithe, a black female from Saxony 
gave birth to seventeen young, of which only one, a 
male, was black. 

It is probable that exceptionally some individuals 
pair late in the summer or in the autumn. There is 
a trustworthy record, by Eiffe, of three pregnant 
females having been caught near Hamburg on 
March 12, 1882, one of them giving birth to young 
on the following day. 

Dicephalous young have occasionally been observed. 
One 6 inches long was found crawling in a field near 
Hornburg in Germany in October, 1895, and, having 
been kept alive for some time, was observed to hiss 
and open the two mouths alternately when taking 
up a defensive attitude. Another similar monster, 
from Cornwall, is reported to have been sent alive to 
the London Zoological Gardens in 1854. 

24. Vipera aspis, Linnaeus 
The Asp Viper 

Form. — Rather more elongate than in the preced- 
ing. Snout flat above, more or less distinctly turned 
up at the end, with sharp, not or but very slightly 



240 



VIPERID^E 




raised canthus, and vertical or nearly vertical loreal 
region. Vertical diameter of the eye equal to or a 
little less than its distance from the mouth. The 
raised upper border of the transversely truncate or 
obtusely pointed extremity of the snout, coupled 
with the downward slant of the supraocular region 
and canthus rostralis, gives the head, seen from the 
side, a peculiar expression ; the eye is so oblique 
that a vertical line drawn from the posterior extremity 

of the supraocular 
shield to the lip 
usually passes 
through the eye 
or down its pos- 
terior border; but 
the extent to which 
the snout is turned 
up at the end 
varies consider- 
ably, some specimens approaching V. bents in this 
respect, others V. latastii. Length of tail five and a 
half to eight times in total length in males, seven to 
nine times in females. 

Head-Shields. — Rostral deeper than broad, its width 
two-thirds to seven-eighths its depth, extending to the 
upper edge of the snout. As a rule, with the exception 
of the large supraocular, the upper surface of the head 
is covered with small, subimbricate scales, which are 
smooth, very rarely feebly keeled, between the eyes 




Fig. 36 (after Sordelli) 



PLATE XI II 





VIPERA ASPIS 
After Calmett? 




VIPERA LATASTI1 



VIPERA 241 

and on the snout ; however, an enlarged frontal 
shield, or even a frontal and a pair of parietals, are 
sometimes present, though rarely so large as in 
a typical V. berus ; when present, the frontal is 
separated from the supraocular by one or two series 
of scales ; when the frontal is absent, four to seven 
series of scales separate the supraoculars. Upper 
surface of snout usually bordered by eight or nine 
small shields — viz., two or three apicals, in contact 
with the tip of the rostral and raised to form the 
turned-up nose, and, on each side, two canthals and 
the upper preocular, which separates the supra- 
ocular from the canthals, or three canthals ; some- 
times, however, the border is formed by six or seven 
small shields, the second canthal being in contact 
with the supraocular, as in V. bents. Supraocular 
usually with very convex outer border, not extending 
posteriorly beyond the vertical of the eye. Eight to 
thirteen scales round the eye, usually ten to twelve ; 
one or two vertical series of scales separate the pre- 
oculars from the nasal, which is single or divided, 
and often rather deeply hollowed out. As a rule 
two series of scales (very rarely three) separate the 
eye from the labials ; sometimes, however, there is 
but one scale between the eye and the fourth labial, 
the second series being incomplete. Upper labials 
nine to thirteen, usually nine to eleven, fourth and 
fifth, rarely fourth to sixth or fifth and sixth, below 

the eye. Temporal scales smooth or feebly keeled. 
16 



242 VIPERID^E 

Four (rarely five) lower labials in contact with the 
single pair of chin-shields. 

Scales in twenty-one or twenty-three (rarely nine- 
teen or twenty-five) rows, with two apical pits, 
strongly keeled, those of the outer row more or less 
distinctly keeled, rarely perfectly smooth. Ventral 
shields 134 to 158 (usually 143 to 153) in males, 
141 to 169 (usually 145 to 157) in females; anal 
entire ; subcaudals 32 to 49 (usually 37 to 45) in 
males, 30 to 43 (usually 32 to 38) in females ; the 
terminal caudal shield is sometimes shorter and less 
spine-like than in V. berus, quite obtuse in some 
specimens. 

Coloration. — Grey, greyish-brown, brown, reddish- 
brown, coppery red, or orange, is the ground colour 
in individuals from the same district; in this 
respect sexual differences are less marked than in 
the preceding species, red or copper-coloured speci- 
mens being found in both sexes, and silvery white 
specimens do not seem ever to occur. In rare cases 
markings are entirely absent. In specimens from 
the greater part of France, Italy, and the Southern 
Tyrol (see Plate XIII.), the dark brown or black 
markings on the body are mostly in the form of 
narrow cross-bars, continuous across the back or 
broken on the vertebral line and often alternating 
with each other and with similar bars on the sides, 
thus producing a pattern not unlike that frequently 
found in Tropidonotus natrix ; a narrow dark line 



VIPERA 243 

running straight or zigzag along the spine may 
connect these cross-bars, and in rare cases it is so 
broad as to produce a zigzag band similar to that of 
V. bents. In specimens from South- Western France 
and the Pyrenees, rarely in some from other parts of 
France and Italy, there is a broad dark grey or 
brown vertebral band between two series of black or 
blackish-brown spots, opposite to each other or 
alternating ; this band may be straight or wavy, some- 
times forming a regular zigzag ; there is another 
series of blackish spots or short bars lower down on 
the side, alternating with those of the dorsal series. 

The upper surface of the head may be devoid of 
any markings, or bear merely the two oblique dark 
streaks forming the branches of a A \ or a dark 
cross-bar may be present on the snout, followed or 
not by smaller spots or a pair of oblique streaks on 
the occiput ; the f\ on the back of the head may 
be united with the first cross-bar on the nape, and 
enclose a cordiform figure of the lighter ground 
colour. A light line sometimes borders the upper 
edge of the snout and the outer edge of the supra- 
ocular shield. A blackish band or a mere line 
extends obliquely from the eye to the first lateral 
spot ; below this the upper lip is whitish, yellowish, 
or pinkish, with or without dark vertical bars on the 
sutures between the labial shields. The iris is 
golden or coppery red. 

The lower parts vary as much as the upper: 



244 VIPERID.E 

sometimes black or steel blue, with or without whitish 
or reddish dots or spots, sometimes yellowish or 
pale reddish, with brown dots or marblings ; in 
some young, white with greyish dots. The throat 
is yellowish white or pale reddish, uniform or 
speckled with blackish, sometimes (males) nearly 
entirely black. The end of the tail is usually bright 
yellow or reddish, or at least with a few bright spots. 
And finally we must mention black specimens — some 
nearly black by darkening of the ground colour, 
others intensely black by enlargement of the mark- 
ings. A specimen from Piedmont, in the Turin 
Museum, shows the ground colour reduced to mere 
narrow light bars disposed in pairs. In most of 
these black specimens the chin and throat remain 
entirely or partially yellowish or reddish, and a few 
spots of the same colour are to be seen under the 
end of the tail. 

A remarkable form of V. aspis, which some herpe- 
tologists would perhaps regard as entitled to rank as 
a species, is the var. hugyi, Schinz, from Calabria 
and Sicily. It is in some respects intermediate 
between V. aspis and V. latastii. The snout is rather 
more pointed than usual in the typical form, 
often, though not constantly, more strongly turned 
up at the end, and the canthus rostralis may be 
distinctly raised. Constantly two canthal shields, 
the second in contact with the supraocular. Ventral 
shields 134 to 148 ; subcaudals 30 to 43. Pale 



VIPERA 245 

greyish, yellowish, brownish, or reddish, above, with 
a broad wavy dark brown vertebral band, edged 
with darker; this band sometimes broken up into 
transversely oval spots ; a lateral series of blackish- 
brown spots, each corresponding to the sinus of the 
dorsal band. 

Specimens so completely intermediate between 
Vipera aspis and V. bents as to render their naming 
arbitrary are known from parts of France and Italy 
where the two species coexist, and are probably to 
be regarded as hybrids. 

Size. — The largest specimen examined (St. Sever, 
Landes, in the Lataste Collection) measures 2 feet 
2.\ inches. It is a male. The largest female in the 
British Museum is 2 inches shorter. 

Distribution. — Vipera aspis is found over the whole 
of France south of a line connecting the depart- 
ments Loire-Inferieure, Orne, Seine-et-Marne, and 
Meurthe-et-Moselle, and ascends the Pyrenees to 
the altitude of 7,250 feet. In Germany it is known 
from Lorraine and the Black Forest, in Switzerland 
from the western and southern parts, up to 5,000 
feet on the northern side of the Alps. It occurs 
also in Austria, in the Southern Tyrol and in the 
Karst, and is distributed over the whole of Italy and 
Sicily, reaching an altitude of 9,700 feet in the 
Alps. Most of the specimens from the western parts 
of the Balkan Peninsula which have been referred 
to this species belong, apparently, to V. bents, var. 



246 VIPERIDJE 

bosniensis, but one from Jahorin in Bosnia, altitude 
5,650 feet, preserved in the Bosnian Museum, is pro- 
nounced by Werner to be an unquestionable V. aspis. 

Habits. — This Viper shows a predilection for hot 
and dry localities. It is both diurnal and nocturnal, 
and does not seem to wander far from its hole in a 
rock or in the earth. It is slow in its movements, 
but very irascible, and innumerable accidents, in 
some cases fatal to man, are caused yearly in many 
parts of France, where it is extremely abundant. 
Its food consists principally of small mammals, 
young birds, and lizards, but the very young eat 
insects and worms. In France it retires into its 
winter-quarters at the end of October or in Novem- 
ber, and numerous specimens often congregate in 
the same hole ; it resumes its activity towards the 
end of March or the beginning of April, sometimes 
as early as the end of February. In rare cases it 
will even leave its retreat in the middle of winter, to 
bask in the sun. In captivity it long retains its 
savage temper, and usually refuses all food. 

Reproduction. — Vipera aspis pairs in April and May ; 
the pair are entwined in each other's coils. The 
young, four to eighteen in number, but rarely more 
than ten, are born in August or September, and 
measure 7 or 8 inches. 

Several cases of dicephaly in young specimens 
have been described. 



VIPERA 



247 



25. VlPERA LATASTII, BoSCa 

Lataste's Viper 

Form. — Heavier than in the preceding. Head 
similar, but snout more pointed and loreal region 
slanting towards the lip, well visible when the head 
is viewed from above. The extent to which the 
snout is turned up at the end varies considerably, 
sometimes similar to certain specimens of V. aspis, 
sometimes forming an appendage which is only a 





Fig. 37 

little less developed than in V. ammodytes. Length of 
tail six and a half to seven and a half times in total 
length in males, seven and a half to nine times in 
females. 

Head-Shields. — Rostral once and a half to twice as 
deep as broad, nearly reaching the tip of the rostral 
wart. Upper surface of head covered with small, 
smooth or feebly keeled, subimbricate scales, among 
which a slightly enlarged frontal, or a frontal and a 



248 VIPERIDiE 

pair of parietals, may sometimes be distinguished ; 
four to seven longitudinal series of scales between 
the supraoculars, which are large, and do not as a 
rule extend posteriorly beyond the vertical of the 
eye. Five or six (rarely three) scales on the posterior 
aspect of the raised part of the snout ; two canthal 
scales on each side, the second in contact with the 
supraocular, or separated from it by the uppermost 
preocular. Eight to thirteen (usually nine to twelve) 
scales round the eye; two or three series of scales 
between the eye and the labials. Nasal hollowed 
out, entire, separated from the preoculars by one or 
two vertical series of scales ; naso-rostral sometimes 
divided into two in North African specimens. 
Temporal scales smooth or feebly keeled. Upper 
labials nine to eleven (rarely eight), fourth and fifth 
(rarely third and fourth) below the eye. Four or five 
lower labials in contact with the single pair of chin- 
shields. 

Scales in twenty-one rows, with two apical pits, 
strongly keeled, outer row smooth or feebly keeled. 
Ventral shields 125 to 146 in males, 135 to 147 in 
females ; anal entire ; subcaudals 35 to 45 in males, 
32 to 38 in females. 

Coloration. — Grey or brown above, the back often 
paler than the sides, with a broad darker, usually 
black-edged, wavy or zigzag band along the spine, and 
a lateral series of spots (Plate XIII.) ; the band some- 
times replaced by large rhombic or transversely oval 



VIPERA 249 

spots. Head with or without dark markings above, 
sometimes with two oblique dark streaks on the 
occiput ; a dark streak from behind the eye to the first 
lateral spot, sometimes originating at a considerable 
distance from the eye ; upper lip white or pale 
brown, more or less speckled or spotted with black. 
Lower parts grey, spotted with black and white, or 
blackish speckled with white, the end of the tail 
usually yellow or with yellow spots. 

Size. — This Viper is not known to exceed a length 
of 2 feet. 

Distribution. — Locally distributed over the greater 
part of Spain and Portugal, as far north as Burgos 
and Barcelona. Also found in Morocco near Tangier, 
and in Algeria near Bona and Guyotville. 

Habits. — Lataste's Viper lives in stony and arid 
districts, and also in forests. The food consists 
chiefly of small mammals, but remains of a scorpion 
have been found in the stomach of an adult, and of a 
centipede in that of a young. According to Graells, 
this Viper easily climbs low trees in search of young 
birds, five of which have been found in the stomach 
of one specimen. The bite is believed to be less 
dangerous than that of V, aspis, and rarely causes 
the death of man and domestic animals. 

26. Vipera ammodytes, Linnaeus 
The Sand-Viper, or Long-Nosed Viper 

This species may be divided into several geographi- 
cal forms. The typical form will be described first. 



250 



VIPERECME 



Form. — Short and heavy. Snout pointed, pro- 
duced into an erect, horn-like dermal appendage 
covered with scales ; canthus rostralis strong, some- 
times slightly raised, loreal region slanting more or 
less towards the lip. Vertical diameter of the eye 
less than its distance from the mouth in the adult. 
Length of tail six to nine and a half times in the total 
length in males, eight to eleven times in females. 

Head - Shields. — Rostral usually broader than 
deep. Naso-rostral (rarely divided into two) usually 

reaching the 
canthus rostra- 
lis, and extend- 
ing considerably 
higher up than 
the upper border 
of the rostral. 
Rostral append- 
age covered with 
ten to seventeen 
scales, arranged in three (rarely two or four) trans- 
verse series between the rostral shield and the apex. 
Upper surface of head covered with small smooth or 
faintly keeled, subimbricate scales, among which a 
feebly enlarged frontal shield or a frontal and a pair 
of parietals are rarely distinguishable ; when present, 
the frontal is separated from the supraocular by two 
series of scales ; on the vertex five to eight series of 
scales separate the supraoculars. Two (rarely three) 





Fig. 38 (after Sordelli) 



PLATE XIV 




VIPERA LEBETINA 
After Sordelh 




VIPERA AMMODYTES 

AJter Sordelii 




ANCISTRODON HALYS 

After Sordelii 



VIPERA 251 

canthal scales, the second separated from the supra- 
ocular by the upper preocular. Supraocular with 
very convex outer border, usually not extending 
posteriorly beyond the vertical of the eye. Ten to 
thirteen scales round the eye ; one or two vertical 
series of scales separate the preoculars from the 
nasal, which is single or rarely divided, and hollowed 
out. Two series of scales between the eye and the 
labials. Upper labials eight to twelve, usually nine 
or ten, usually fourth and fifth below the eye. 
Temporal scales smooth or feebly keeled. Four or 
five lower labials in contact with the single pair of 
chin-shields. 

Scales in twenty-one or twenty-three (rarely twenty- 
five) rows, with two apical pits, strongly keeled, those 
of the outer row smooth or feebly keeled. Ventral 
shields 143 to 161 in males, 147 to 160 in females; 
anal entire ; subcaudals 27 to 40 in males, 24 to 37 
in females. 

Coloration. — Grey, pinkish-grey, brown, yellowish- 
brown, or brick red above, with a more or less 
distinct wavy or zigzag black or brown, usually 
black-edged, band along the back, or a series of large 
rhombs connected on the median line, with or with- 
out a lateral series of dark spots ; specimens with the 
markings of an intense black are males (Plate XIV.). 
Head with or without dark markings, very variable 
in disposition, sometimes forming a A on the occiput, 
or a lyre-shaped figure confluent with the dorsal 



252 viperim: 

band ; a dark streak or broad band from the eye to 
the angle of the mouth, sometimes continued along 
the neck; dark vertical bars often present on the 
sides of the snout and under the eye ; on the lower 
lip, the dark shade, if present, broken up by light 
bars separated by two to four labial shields. Belly 
greyish or pink, powdered with brown or black, or 
dark bluish-grey, with or without black and white 
spots ; lower surface of end of tail orange or coral 
red, rarely yellow. Iris golden. Specimens with a 
straight brown vertebral band flanked with triangular 
black spots pointing outwards (var. steindachneri, 
Werner) are known from Hungary, and resemble 
certain colour varieties of V. benis and V. aspis. 
Black specimens are very rare. 

The following varieties are important geographical 
forms occurring in Europe : 

Var. montandoni, Boulenger : Naso - rostral shield 
never reaching the canthus rostralis nor the summit 
of the rostral shield, which is deeper than broad 
(once and one-seventh to once and a half) ; rostral 
appendage clad with ten to fourteen scales, in three 
(rarely two or four) transverse series between the 
rostral shield and the apex. Scales in twenty-one 
rows. Ventral shields 149 to 158; subcaudals 
30 to 38. A more or less distinct dark blotch on 
the lower lip, involving five to seven labial shields 
without complete interruption. Lower surface of 
end of tail yellow. 



VIPERA 



253 



Var. mevidionalis, Boulenger : Naso-rostral shield 
never reaching the canthus rostralis, and but rarely 
extending higher up than the upper border of the 
rostral, which is often as deep as broad or a little 
deeper than broad; rostral appendage clad with 




p IG> ^g — Side Views of Heads of V. ammodytes typica (a) and 

Var. mevidionalis (b) 






a b c 

Fig. 40 — Front Views of End of Snout, showing the Lepi- 
dosis. (From Proceedings of the Zoological Society, 1903) 

a, Form typica; b, var. mevidionalis ; c, var. montandoni 

fourteen to twenty scales, in four or five (rarely three) 
transverse series between the rostral shield and the 
apex. Supraciliary edge usually more prominent 
than in the typical form, sometimes slightly angular. 
Scales in twenty-one rows (very rarely twenty-three). 
Ventral shields 133 to 147; subcaudals 24 to 35. 



254 viperim; 

A more or less distinct dark blotch on the lower lip, 
involving five or six labial shields without interrup- 
tion. Lower surface of end of tail yellow. 

Size. — This Viper exceptionally attains a length 
of 3 feet. The largest male in the British Museum 
measures 2 feet 6 inches, the largest female 2 feet 
4 inches. In V. berus females grow to a larger size 
than males ; in this species, as in V. aspis, the reverse 
appears to be the rule. 

Hybrid. — A female specimen, presumed to be a 
hybrid between V. berus and V. ammodytes, was 
obtained by Captain Veith in 1902 in Carinthia, in 
a locality where both these species occur together. 
The shape of the head is exactly that of a typical 
V. aspis, the snout distinctly turned up at the end, 
but without wart or scaly appendage, the raised 
portion being covered by the apex of the rostral 
shield and three apical shields. The rostral shield, 
which is a little deeper than broad (5 : 4), ex- 
tends above the level of the slightly raised canthus 
rostralis, which bears two shields, the second in 
contact with the supraocular. The naso-rostral 
extends to the canthus rostralis, where it joins the 
first canthal and the lateral apical shield ; one series 
of scales between the nasal shield and the preoculars. 
On the upper surface of the head the snout is 
covered with fifteen subimbricate smooth scales, 
in addition to the canthals and apicals. A frontal 
shield and a pair of parietals are well developed, 



VIPERA 255 

although smaller than in an average V. berus ; the 
parietals are in contact with the frontal, between 
which and the supraoculars two series of scales 
intervene. The supraocular, as in V. berus, ex- 
tends backwards considerably beyond the vertical 
of the eye. Eleven scales round the eye ; two series 
of scales between the eye and the labials. Temporal 
scales smooth. Upper labials nine, fourth and fifth 
below the eye. Scales in twenty-one rows, outer 
row smooth. Ventrals 159 ; subcaudals 31. 

Grey- brown above, with a reddish -brown or 
mahogany-coloured zigzag vertebral band and a 
lateral series of paler reddish-brown spots ; temporal 
band ill-defined in front ; no markings on the upper 
surface of the head ; lips pinkish, with a few reddish- 
brown spots. Ventral shields pale brownish, finely 
speckled or powdered with blackish, and with small 
whitish spots on the free edge and reddish-brown spots 
on the sides. Tail orange red below. Iris fire 
red. 

Total length 2 feet 2 J inches ; tail 2f inches. 

Distribution. — The typical form is known from 
Northern Venetia, Austria-Hungary (Styria, Carin- 
thia, Southern Tyrol, Carniola, Illyria, Istria, Croatia, 
Slavonia, and eastward through Southern Hungary 
to Transylvania), Dalmatia, Bosnia, Herzegovina, 
Montenegro, Albania, and Servia. In the Alps up to 
1,300 feet, in the Balkan Peninsula up to 7,500 feet. 
Thevar. montandoni inhabits Roumaniaand Bulgaria. 



256 viperim: 

The var. meridionalis inhabits Greece, with the 
Archipelago, Asia Minor, and Syria. 

The specimens from Transcaucasia constitute a 
further variety (var. transcaucasiana, Boulenger), 
agreeing with the var. montandoni in the rostral 
scutellation and in the number of ventral shields 
(150 to 156), but differing in the markings on the 
back ; these consist of dark bars or alternating 
paired dark spots, as in the typical form of V. aspis. 
The dark and light markings on the lower lip are 
as in the typical V. ammodytes, and the lower surface 
of the tail is pale yellow or greenish towards the 
end. 

Habits. — Notwithstanding its name ammodytes, 
this Viper is by no means restricted to sandy 
localities ; on the contrary, it shows a predilection 
for dry stony hills with low vegetation, and has 
often been found climbing bushes. It avoids thick 
forest, but occurs on the edges of woods and in 
clearings, as well as on the borders of roads through 
woods. In the cooler regions of the mountains, 
which it ascends to a considerable altitude, it is 
essentially diurnal, leaving its retreat only when 
the sun shines ; but in warm localities it is stated 
to be principally nocturnal, appearing in numbers 
by moonlight. The length of its period of hiber- 
nation depends entirely on the climate, but when 
the winter is mild it may be seen about in mid- 
winter whenever the sun shines. The poison of 



VIPERA 257 

this Viper is stated to be more active than that 
of V. berus and V. aspis, and, as the snake is very 
common in some parts of Austria and the Balkan 
Peninsula, fatal accidents to man are frequent. 

The food consists of small mammals and birds, 
and also of lizards. V. ammodytes does much better 
in captivity than its European congeners, and takes 
food more readily. 

The hissing is louder than in V. berus and V. aspis, 
and it is often produced, on the approach of man, 
by specimens lying in such perfect concealment that 
their presence would not otherwise be suspected ; 
this habit, like the rattling of the Crotalus, is evi- 
dently detrimental to the species in its relation 
to man. 

This species is extremely abundant in some parts 
of Austria, and over 7,000 specimens were killed in 
a district of Southern Styria in the course of two 
years (1892, 1893). According to Werner, it is the 
commonest of all snakes in Bosnia and Herzegovina. 

Reproduction. — Pairing takes place in the spring, 
sooner or later according to altitudes, and the young, 
five to fourteen in number, are born in August. 

27. Vipera lebetina, Linnaeus 
The Blunt - Nosed Viper, or Kufi 

Form. — Short and heavy. Snout rounded, obtuse, 
usually with well-marked canthus, and the loreal 
region slanting towards the mouth. Eye small, its 
17 



258 



VIPERID.E 




vertical diameter less than its distance from the 
mouth in the adult. Nostril large and directed 
backwards. Length of tail six (males) to ten times 
(females) in the total length. 

Head-Shields. — Rostral as deep as broad, a little 
broader than deep, or slightly deeper than broad, 
reaching or nearly reaching the upper surface of 
the snout, and in contact with two or three apical 
shields. Upper surface of head covered with small 

sub imbricate 
scales, which 
are all more 
or less dis- 
tinctly keeled, 
or, rarely, 
smooth on the 
snout and 

forehead; 

Fig. 41 (after Sordelli) 

^ v J sevento 

twelve longitudinal series of scales between the 
eyes (supraoculars included) ; two to four canthal 
shields, of which the anterior is the largest, and 
may be regarded as a supranasal. Supraocular 
narrow, usually broken up into two or more small 
shields. Twelve to eighteen scales round the eye ; 
two or three series of scales between the eye 
and the labials ; two or three vertical series of 
scales separate the preoculars from the nasal, 
which is single and often strongly hollowed out, 




VIPERA 259 

and usually partially fused with the naso-rostral. 
Upper labials nine to twelve, usually fourth and fifth 
below the eye. Temporal scales keeled. Four or 
five lower labials in contact with the single pair of 
chin-shields. 

Scales in twenty-three to twenty-seven rows, 
usually twenty-five, with two apical pits, strongly 
keeled, those of the outer row smooth or feebly keeled. 
Ventral shields 151 to 177 in males, 153 to 180 in 
females; anal entire; subcaudals 42 to 51 in males, 
38 to 49 in females. 

Coloration. — Very variable. The typical form, 
which alone is represented in Europe, and was 
originally described from Cyprus, is grey, greyish- 
buff, or pale brown, above, with two dorsal series 
of darker spots, which may stand in pairs, alternate, 
or unite to form cross-bars, and a lateral series of 
large dark spots or bars. A more or less distinct 
dark band on each side of the head, passing through 
the eye and often extending to the neck ; a dark 
bar or triangular spot below the eye, and usually 
another below the nostril. Lower parts pinkish- 
white, powdered with grey-brown, with or without 
dark brown spots ; end of tail yellow. The ground 
colour of the young is pink or flesh-colour. In 
specimens from desert sandy regions in Asia and 
North Africa the markings may be very indistinct, 
the snake being of a nearly uniform pale buff. 

In the var. mauritanica, Guichenot, from Morocco 



2 6o viperim: 

and Algeria, the back has three series of very large 
dark brown or reddish-brown spots, separated by a 
network of the yellowish ground colour, or the 
middle series may be transformed into a wavy or 
zigzag band. The scales in this variety are usually 
in twenty-seven rows, instead of twenty-five (rarely 
twenty-three) as in the typical form. 

Size. — This species, the largest of European 
Vipers, grows to a length of 4! feet. 

Distribution. — The European habitat of V. lebetina 
is restricted to the Cyclades, where it is not un- 
common on the island of Tinos, and appears to be 
found also on Kimoli. It is common on Cyprus, 
where it is called Kufi, or Deaf Snake, and extends 
from Syria and Asia Minor through Transcaucasia, 
Mesopotamia, Persia, Northern Baluchistan, to 
Afghanistan and Cashmere. It is further found on 
the Atlas of Morocco and Algeria, near Oran and 
Bona, and in Tunisia. Its reported occurrence in 
Egypt has not been confirmed by recent investi- 
gations. 

Habits. — According to M. Doumergue — who has 
had ample opportunities of observing this Viper near 
Oran, where it is common — it is a nocturnal reptile, 
rarely moving about in the daytime. It inhabits 
rocky localities, where there is brushwood, and 
vineyards, During the day it remains sluggish 
under large stones. It is most frequently met with 
in April and May. 



ANCISTRODON 261 

On Milos, Dr. de Bedriaga observed this much- 
dreaded snake, the bite of which is probably as bad 
as that of its Indian ally, the Daboia, V, russelli, to 
occur frequently in gardens, and to crawl about 
near houses in villages after sunset. The same 
observer has noted a sort of valvular closing of 
the nostril through raising of the posterior part of 
the nasal shield when the snake prepares to strike. 

The food consists principally of mammals up to 
the size of a rabbit. 

Reproduction. — According to Doumergue's obser- 
vations in Algeria, the young, up to thirteen in 
number, are born in May and June. 

Genus ANCISTRODON, Palisot de Beauvois 

Head distinct from neck, its upper surface covered 
with large shields, as in the normal Colnbridcz ; a 
loreal pit ; eye moderate or small, with vertical 
pupil. Body moderately elongate or short. Scales 
keeled (or smooth), with apical pits. Tail moderate 
or short. 

This genus is distributed over nearly the whole 
of Asia, the eastern parts of the United States of 
America, and Mexico and Central America. One 
of the Asiatic species just penetrates into South- 
Eastern Europe, and is the sole representative of 
the Crotalinae in this part of the world. 



262 viperim: 

28. Ancistrodon halys, Pallas 
Pallas's Pit-Viper 

Form. — Moderately elongate. Head subtriangular, 
flat or slightly concave above, swollen in the tem- 
poral region, very distinct from neck ; snout rounded 
or obtusely pointed, slightly turned up at the end, 
with obtuse canthus and vertical or slightly oblique 
loreal region ; eye rather small. Tail seven to eight 
and a half times in the total length. 





Fig. 42 

Head-Shields. — Rostral as deep as broad or slightly 
broader than deep, just visible from above. A pair 
of internasals and a pair of larger prefrontals. 
Frontal as broad as the supraocular, as long as or 
a little longer than its distance from the end of the 
snout, as long as or a little shorter than the rather 
short parietals. Supraocular extending beyond the 
vertical of the posterior border of the eye. Loreal 
pit between three shields, separated from the labials. 
Nostril between two nasals, the posterior of which 



ANCISTRODON 263 

is separated from the upper preocular by a loreal. 
Two preoculars, one or two postoculars, and a sub- 
ocular. Three large lower temporals, anterior largest. 
Upper labials seven or eight, third and fourth 
largest, third entering the eye. A pair of small 
chin-shields. 

Scales sharply keeled, with two apical pits, in 
twenty-three rows. Ventral shields 149 to 174 ; anal 
entire; subcaudals 31 to 44 pairs. 

Coloration. — Pale yellowish-grey, greyish-brown, 
or reddish, sometimes greenish in young specimens, 
with transverse series of darker spots or with more or 
less regular dark bars with serrated edges across the 
back (Plate XIV.) ; these bars may be narrower than 
the interspaces between them, or so large as to cause 
the back to appear brown with light cross-bars ; the 
bars sometimes broken up on the vertebral line, and 
the two halves alternating. The sides usually paler 
and bearing two alternating longitudinal series of 
small spots, the lower of which are usually darker, 
and sometimes extend on the outer ends of the 
ventral shields. Head pale above, with a dark spot 
on the middle of the snout, a cross-bar or a pair 
of spots between the eyes, a spot or short band on 
each side of the parietal region, and a horseshoe- 
shaped band on the occiput, the branches of which are 
more or less produced on the nape ; all these mark- 
ings sometimes confluent. A broad, dark, light- 
edged band on the temple. Lips whitish, speckled 



264 VIPERID.E 

with brown. Lower parts whitish, more or less 
profusely speckled with grey or brown. The horny 
shield terminating the tail usually dark brown or 
black at the end. 

Size. — This species rarely reaches a length of 
29 inches. The largest specimen in the British 
Museum measures only 19 inches. 

Distribution. — From the north and east coasts 
of the Caspian Sea, across Central Asia to the 
Upper Yenissei, as far north as 51°. In Europe it 
is only known from two arid tracts between the 
Volga and the Ural, near the Caspian Sea, viz., the 
Saltan-Murat desert and the Induski hills. 

Habits. — Nothing has been published on the 
habits of this snake, but they are probably similar 
to those of its near and more eastern relative, 
A. blomhoffi, Schlegel, which inhabits China and 
Japan. A. blomhoffi is said to be more or less 
nocturnal, although showing a predilection for 
localities well exposed to the sun. It is ovovi- 
viparous. The symptoms of its bite, which is 
rarely fatal to man, are the same as in the Vipers. 

All the species of Ancistvodon, so far as they 
have been observed, are in the habit of raising and 
vibrating the tail, like the Rattlesnakes, when coiling 
themselves up in a defensive attitude. 



INDEX 



ACANTHOCEPHALA, IIO 

Acrochordina, 5 

Adder, 230 

iEsculapian Snake, 187 

asculapii (Coluber), 187 

A glyphs, 4, 151 

Ailurophis vivax, 217 

Albinism, 39 

Aldrovandfs Snake, 182 

Altitudinal range, 118, 123 

Amblycephalida, 5 

ammodytes (Vipera), 249 

Anal pockets, 83 

Anal shield, 17 

Ancistrodon, 261 

Ancistrodon halys, 262 

Antitoxic serums, 66 

Apical pits, 15, 75 

Arteries, 77 

Ascariasis, in 

asianns (Zamenis gemonensis, var.), 

173 
Asp Viper, 239 

aspis (Vipera), 239 

astreptophorus (Tropidonotus Jiatrix, 

var.), 154 

ater (Tropidonotus natrix, var.), 

156 
atrovwens (Coluber), 170 
aurolineatus (Coluber), 167 
austriaca (Coronella) , 197 

Bacteria, 113 
her us (Pelias), 221 
berus (Vipera), 230 
Bezoar stones, 141 
bilincatus (Tropidonotus natrix, 
var.), 154 



Blind Snake, 144 

Blood-supply, 77 

Blunt-nosed Viper, 257 

Body, 8 

BoidcB, 4, 146 

Bdna, 4, 146 

bosniensis (Vipera berus, var.), 236 

Brain, 73 

Burrowing Snakes, 91 

caruleus (Coluber), 233 
Canker, 113 
Canthus rostralis, 13 
Captivity, 100 
carbonarius (Zamenis), 172 
caspius (Zamenis gemonensis, var.) , 

172 
Cat-Snake, 217 
Caucasian Cat-Snake, 219 
cervone (Elaphis), 1 82 
Cestoda, 112 
cettii (Tropidonotus natrix, var.), 

155 

chersoides (Tropidonotus), 167 

Classification, 2 

Cobra di capello, 9 

Ccelopeltis, 207 

Coslopeltis monspessulana t 208 

collar is (Contia), 207 

Coloration, 29 

Colour, 29 

Coluber, 181 

Coluber asculapii, 187 

Coluber atrovircns, 170 

Coluber aurolineatus, 167 

Coluber cceruleus, 233 

Coluber dione, 185 

Coluber dumfriesiensis, 200 



265 



266 



INDEX 



Coluber flavescens, 187 

Coluber hydrus, 160 

Coluber insignitus, 208 

Coluber leopardinus, 191 

Coluber longissimus, 187 

Coluber quadrilineatus, 191 

Coluber quaterradiatus, 182 

Coluber quatuorlineatus, 182 

Coluber riccioli, 202 

Coluber rubens, 204 

Coluber sauromates, 183 

Coluber scalaris, 194 

Coluber torquatus, 152 

Coluber viridiflavus, 170 

Colubridcs, 4, 150 

ColubrincB, 4, 151 

concolor (Tropidonotus tessellatus, 

var.), 162 
concolor (Vipera berus, var.), 234 
Coniophis, 7 
Constriction, 96 
Contia, 205 
Contia modesta, 205 
Coral-Snakes, 32, 34, 35 
Coronella, 196 
Coronella austriaca, 197 
Coronella girondica, 202 
Coronella italica, 197 
Coryphodonts, 58 
CrotalincB, 5, 220 
Crotalon, 20 
Crotalus, 21 
cucullatus (Macroprotodon). 213 

Dahl's Whip-Snake, 177 
dahlii (Zamenis), 177 
Dasypeltina, 5 
Defecation, 99 
Definition, 1 
Deglutition, 98 
Dentition, 53 
Desert Snakes, 36 
Development, 88 
Diacranterians, 58 
Dicephalous young, 89 
Digestion, 99 
rf/0»* (Coluber), 185 
Dione Snake, 185 
Dipsadomorphina, 5, 151 



Disgorging, 99 
Distribution, 118 
Dolichosauria, 1 
dorsalis (Pelias), 234 
Drinking, 101 
dumfriesiensis (Coluber), 200 
Dwarf Snake, 205 

Economic value, 138 

Eggs, 85 

Egg-eating Snakes, 53, 80 

Egg-tooth, 89 

Elachistodonti a, 5 

Elaphis cervone, 182 

Elapince, 5 

Elaps, 35 

elsneri (Coluber leopardinus t var.), 

193 
Embryo, 88 
Epiglottis, 79 
Eryx, 147 
Eryx jaculus, 147 
European Whip-Snake, 170 
External characters, 8 
Exuviae, 20 
Exuviation, 105 
Eyes, 11, 73 

False Smooth Snake, 213 
fallax (Tarbophis), 217 
Fangs, 53 
Fascination, 104 
Fat-bodies, 81 
Faunistic works, 130 
Feigning death, 104 
fitzingeri (Coronella), 197 
flavescens (Coluber), 187 
flavescens (Tropidonotus tessellatus, 

var.), 163 
Flukes, 112 
Flying Snakes, 94 
Food, 95 
Form, 8 
Fossils, 6 

Gape, 11 
Gastrosteges, 15 
gemonensis (Zamenis), 170 
Genital organs, 82 
girondica (Coronella), 202 



INDEX 



267 



Glauconiida, 4 
Glottis, 79 
Grass Snake, 152 
Greek Blind Snake, 144 
Ground Snakes, 91 

Habits, 91 

Haemogregarines, 112 
hagenbecki (Tropidonotus tessella- 

tus, var.), 162 
halys (Ancistrodon), 261 
Head, 10 
Head-shields, 17 
Hearing, 74 
Heart, 77 
Helagras, 7 
Hemipenes, 82 
Hibernation, 105 
Hind limbs, 10 
hippocrepis (Zamenis), 179 
Hissing, 102 
Homalopsina, 5 
Hood, 9 

Horse-shoe Whip-Snake, 179 
hugyi (Vipera aspis } var.), 244 
Hybrids, 90 
Hydrophiina, 5 
hydrus (Coluber), 160 
Hyoid apparatus, 42 

iberus (Tarbophis), 219 

Iguanognathus, 54 

lly slides, 4 

Immunity, 66, 71 

incertus (Tropidonotus viperinus, 

var.), 167 
Incubation, 88 
Intelligence, 101 
insignitus (Coluber), 208 
Integument, 8 
Intestines, 79 
Iris, 37 
Isodonts, 58 
italica (Coronella) , 197 

Jacobson's organ, 73 
jaculus (Eryx), 147 
Javelin Sand-Boa, 147 



Key to the identification, 22 
Kidneys, 80 
Kufi, 257 

Labial pits, 76 

Lacertilia, 1 

lacertina (Natrix), 208 

Ladder Snake, 194 

Largest Snakes, 21 

Larynx, 79 

Lataste's Viper, 247 

latastii (Vipera), 247 

lebetina ( Vipera), 257 

Leopard Snake, 191 

leopardinus (Coluber), 191 

lineaticollis (Tropidonotus tessella- 

tus, var.), 162 
Liver, 81 

Livery of the young. 38 
Locomotion, 93 
Long-nosed Viper, 249 
longissimus (Coluber), 187 
Loreal pit, 75 
Loreal region, 12 
Lungs, 78 
Lycodonts, 58 
Lymph-hearts, 78 

Macroprotodon, 212 

Macroprotodon cucullatus } 213 

macrops (Vipera), 224 

Marine Snakes, 92, 95 

Markings, 29 

mauritanica ( Vipera lebetina i 

var.), 259 
Melanism, 39 
Mental groove, 13 
meridionalis (Vipera ammodytes, 

var.), 253 
Mimicry, 35 
modesta (Contia), 205 
monspessulana (Ccslopeltis), 208 
montandoni (Vipera ammodytes, 

var.), 252 
Montpellier Snake, 208 
Mortality from Snake-bite, 133 
Mosasauria, 1 
murorum (Tropidonotus natrix, 

var.), 154 



268 



INDEX 



Natrix lacertina, 208 

Natrix vulgaris, 152 

natrix (Tropidonotus), 152 

Neck, 9 

Nematoda, 11 1 

Nerodia, 152 

Nervous system, 73 

neumayeri (Coelopeltis monspessu- 

lana, var.), 210 
niger (Coluber longissimus, var), 

188 
nigrescens (Tropidonotus tessellatus, 

var.), 162 
Nocturnal Snakes, 93 
Northern Viper, 230 
Nostrils, 13, 74 

GEsophagus, 79 
Olfactory organ, 73 
Ophidia. 1 

Opisth glyph a, 4, 151 
Organs of a sixth sense, 75 
Orsini's Viper, 221 
Oviparous Snakes, 86 
Oviposition, 85 
Ovoviviparous Snakes, 85 

Pairing, 84 

Pallas's Pit-Viper, 262 

Parasites, 107 

Peli as berus, 221 

Pelias dorsalis, 234 

Pelvic arch, 52 

persa (Tropidonotus natrix, var.), 

154 
persicus (Zamenis gemonensis, 

var.), 172 
picturatus (Tropidonotus natrix, 

var.), 155 
Pit-Vipers, 75, 262 
Poison apparatus, 62 
Poison fangs, 55, 62 
Poison glands, 62 
Poisons, 66 
prester (Vipera), 235 
Proteroglypha, 4, 151 
Protozoa, 112 
pseudaspis (Vipera berus, var.), 

236 



Pupil, 12 
Pythonina, 4 

quadrilineatus (Coluber), 191 
quaterradiatus (Coluber), 182 
quatuorlineatus (Coluber), 182 

Rattle. 20 

Rattlesnakes. 20, 103 

Rattling, 103 

Reflex movements, 106 

Relation to man, 133 

Renard's Viper, 227 

renardi (Vipera). 227 

Replacement teeth, 55 

Reproduction, 82 

Reptation, 93 

Rep tili a, 1 

Rhiptoglossa, 1 

riccioli (Coluber), 202 

Ring-Snake, 152 

romanus (Coluber longissimus, 
var.), 188 

rubens (Coluber) , 204 

rubvo-maculosus (Tropidonotus tes- 
sellatus, var.), 162 

Rustling, 102 

Sand-Boa, 147 

Sand-Snakes, 91 

Sand-Viper, 249 

sardus (Zamenis), 172 

sauromates (Coluber), 183 

scalaris (Coluber), 194 

scalaris (Coronella austriaca, var.), 

199 
Scales, 14 • 

schwoederi (Coluber leopardinus, 

var.), 193 
scutatus (Tropidonotus natrix, 

var.), 155 
semimaculata (Contia modesta, 

var.), 207 
Sense organs, 73 
seoanei (Vipera berus, var.), 235 
Serotherapy, 66, 134 
Serum treatment. 67 
Shedding of the epidermis, 20 
Shields, 14 



INDEX 



269 



Sixth sense, 75 

Skeleton, 40 

Skull, 40 

Smallest Snakes, 21 

Smooth Snake, 197 

Snake-charmers, 139 

Snake-stones, 141 

Snake swallowing her young, 88 

Snake-worship, 139 

Snout, 12 

Solenoglyphs, 56 

Southern Smooth Snake, 202 

Squama ta, 1 

steindachneri {Viper a ammodytes, 

var.), 252 
Stomach, 79 
Subcaudal shields, 15 
subfasciatus {Tropidonotus natrix, 

var.), 155 , . • 

subgriseus {Coluber longissimus, 

var.), 189 
Symoliophis, 7 
Syncranterians, 58 

Tail. 9 

Tarbophis, 216 

Tarbophis fallax, 217 

Tarbophis iberus, 219 

Teeth, 53 

Tessellated Water-Snake, 160 

tessellatas {Tropidonotus), 160 

Thymus gland, 78 

Ticks, 108 

Tongue, 74 

Tongue-worms, 108 

torquatus {Coluber), 152 

trabalis {Zamenis gemonensis , var .), 

172 
Trachea, 78 
transcaucasiana {Viper a ammodytes, 

var.), 256 
Tree-Snakes, 36, 92, 94 
Trematoda, 112 
Tropidonotus, 152 
Tropidonotus chersoides, 167 
Tropidonotus natrix, 152 
Tropidonotus tessellatus, 160 
Tropidonotus viperinus, 165 
Typhlopida, 4, 143 



Typhlops, 144 

Typhlops vermicularis, 144 

Ureters, 81 
Uroveltida, 5 
Urosteges. 15 
ursinii {Vipera), 221 

Veins, 77 

veithi (Loronclla austriaca, var.), 

199 
Ventral shields, 15 
vermicularis {Typhlops), 144 
Vertebrae, 50 
Vertebral column, 50 

Vipera, 221 

Vipera ammodytes, 249 

Vipera aspis, 239 

Vipera berus, 230 

Vipera latastii, 247 

Vipera lebetina, 257 

Vipera macrops, 224 

Vipera pr ester , 235 

Vipera renardi, 227 

Vipera ursinii, 221 

ViperidcB, 5, 220 

Viperince, 5. 220 

Viperine Water-Snake, 165 

viperinus {Tropidonotus), 165 

viridiflavus {Coluber), 170 

Viscera, 77 

vivax {Ailurophis), 217 

vosseleri {Tropidonotus tessellatus, 

var.), 162 
vulgaris {Natrix), 152 

Warning coloration, 35, 36 
Water-Snakes, 92, 94, 160 
Whip-Snake, 170 
Worms, no 

XenopeltidcB, 5 

Zamenis, 170 

Zamenis carbonarius, 172 

Zamenis dahlii, 177 

Zamenis gemonensis, 170 

Zamenis hippocrepis, 179 

Zamenis sardus, 172 

Zoogeographical Regions, 119 



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Fiction 



25 



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Cr. Zvo. 


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A MAN 


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26 



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Fiction 



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