THE

G.A.BOULENGER

THE SNAKES OF EUROPE

UNIFORM WITH THIS VOLUME
THE LIFE OF CRUSTACEA FRESHWATER FISHES THE OX AND ITS KINDRED
BRITISH

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. LONDON

First Publisned in igiS

PREFACE

THERE
far as the

is

no work

in

the

English language
I

dealing with the Reptiles of Europe.

have
in a

therefore endeavoured to supply this desideratum, so

Snakes are concerned, by drawing up
is

concise form an account of what
characters, their distribution,

known

of their

and

their life-histories.

Professor Sordelli, of Milan, having kindly acceded
to

my

request to

reproduce some of the beautiful
for the

figures

drawn by him

work published
I

in

collaboration with the late Professor Jan under the
title

of" Iconographie G^nerale des Ophidiens,"

have
illus-

been able to supplement
trations

my

descriptions with

which leave nothing to be desired from the

point of view of accuracy.

A

few drawings have
J.

been made specially
I

for this

book by Mr.

Green.

have further to acknowledge the permission given

by the Trustees of the British Museum, the India
Office,

figures

and the Zoological Society, to reproduce a few from previous publications of which I am the
little

author.

In order to render this

book more

useful, the

account of the Snakes of Europe has been preceded

vi

THE SNAKES OF EUROPE
is

by an Introduction summarizing what
Snakes generally.
I

known

of

have purposely avoided overburdening a work of

this kind,

which aims

at concision, with bibliographilists.
I

cal references

and synonymic

am

sure

my

readers will be thankful for being spared this display
of erudition.
to trespass on

Whenever

I

have had to compile, and
is

ground that

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.
all

To

who have helped me

I

beg to tender

my

hearty thanks.

G. A. B.

CONTENTS
INTRODUCTION
CHAPTER
I.

PAGE
-

Definition and Classification

-

i

II.

External Characters— Integument
Coloration
-

-

-

8

III.

-

-

-

-

-

29

IV.

V.

VI.

Skeleton Dentition Poison Apparatus
-

-

-

-

-

40

-

-

-53
of
-

— Different
-

Kinds
-

Poisons
VII.
VIII.
IX.

-

-

-

62
Th

Nervous System— Sense Organs
Viscera

....-']']
;

Organs OF Reproduction TiON Development
;

Pairing; Oviposi-

-

-

-

82
9*

X. Habits
XI. Parasites

-

-

-

-

-

-

-

-

-

-

-

107

XII. Distribution
XIII.

-

-

-

-

-

118 i33

Snakes

in

Relation to

Man

-

-

-

SYSTEMATIC ACCOUNT OF THE SNAKES OF EUROPE
First Family:

TYPHLOPID^
-

Genus TYPHLOPS, Schneider
I.

-

144

Typhlops vermicularis, Merrem— The Greek
Blind-Snake
-

-

I44

vii

viii

THE SNAKES OF EUROPE
Second Family: BOIDiE
PAGE

Genus ERYX, Daudin 2. Eryx jaculus, Linnaeus— The

-

-

-147
-

Javelin

Sand-Boa

147

Third Family:

COLUBRID^
-

Genus TROPIDONOTUS, Kuhl NATRix, Linnaeus— The 3. TROPIDONOTUS

-

152

Grass-

Snake, or Ring-Snake
4.

-

-

-

TROPIDONOTUS

5.

The tessellatus, Laurenti Tessellated Water-Snake TROPIDONOTUS viperinus, Latreille— The Viperine Water-Snake

152

....
-

160
165

Genus ZAMENIS, Wagler
6.

-

-

170

7.

8.

Zamenis gemonensis, Laurenti The European Whip-Snake Zamenis dahlii, Fitzinger Dahl's Whip-Snake Zamenis hippocrepis, Linnaeus— The Horseshoe Whip-Snake

170
177


-

-

-

i79
181

Genus COLUBER, Linn^us Coluber quatuorlineatus, Lacepede— Aldro9.
-

vandi's
10.

Snake

11.

Coluber dione, Pallas The Dione Snake Coluber longissimus, Laurenti — The ^scula-

pian Snake12.

Coluber leopardinus, Bonaparte
ard Snake

.... -----— — The
Leop-

-

-

-

-

-

182

185

187
191

13.

Coluber

scalaris, Schinz— The Ladder Snake

194
196

Genus CORONELLA, Laurenti 14. CoRONELLA AUSTRIACA, Laurenti — The Smooth

Snake
15.

......
.
.

-

197

CORONELLA GIRONDICA, Daudin— The
Smooth Snake
.

Southern
.

202

CONTENTS
Genus CONTIA, Baird and Girard 16. Contia modesta, Martin— The Dwarf Snake
-

ix
PAGE
-

205
205

Genus CCELOPELTIS, Wagler
17.

CCELOPELTIS

monspessulana,

Montpellier Snake

-----

-

-

207

Hermann— The

208
212

Genus MACROPROTODON, Guichenot 18. Macroprotodon cucullatus, I. Geoffroy— The

False Smooth Snake

-

-

-

-

213

Genus TARBOPHIS, Fleischmann

-

-

216
217

19. TARBOPHIS fallax, Fleischmann— The Cat-Snake 20 Tareophis iberus, Eichwald— The Caucasian Cat-

Snake

.-.---

219

Fourth Family: VIPERID^E
Genus VIPERA, Laurenti
21.
-

-

-

221
221

22.
23.

24.

25.
26.

27.

ViPERA ursinii, Bonaparte— Orsini's Viper ViPERA renardi, Christoph Renard's Viper ViPERA BERUS, Linnaeus The Northern Viper, or Adder ViPERA ASPis, Linnaeus— The Asp Viper ViPERA LATASTii, Bosca Lataste's Viper ViPERA AMMODYTES, Linnaeus— The Sand- Viper, . . or Long-Nosed Viper ViPERA lebetina, Linnaeus The Blunt-Nosed
-

......
-

227

230
239
247

-

-

.

249

Viper, or Kufi

-

-

-

-

-

257
261

Genus ANCISTRODON, Palisot de Beauvois 28. ANCISTRODON HALYS, Pallas— Pallas's Pit-Viper
Index
-

-

-

262
265

-

-

-

-

-

-

LIST OF PLATES
PLATE
I.

FACING PAGE

Typhlops vermicularis, Eryx jaculus

-

144

II.

Tropidonotus natrix and
PERSA
-

vars. cettii
-

and

-

-

-152
-

III.

Tropidonotus VAR. AUROLINEATUS
VIRIDIFLAVUS
-

tessellatus, T.
-

viperinus and
-

160

IV.

Zamenis gemonensis and vars. persica and
-

-

-

-

170

V.

Zamenis gemonensis, var. caspius,
Z.

Z.

dahlii,
-

hippocrepis

-

-

-

176

VI.

Coluber quatuorlineatus and var. sauromates, C. dione
-

-

-

-

182

VII.

Coluber longissimus,

C.

leopardinus and
-

VAR. quadrilineatus
VIII.
IX.

1

88

Coluber scalaris-

.

_

.

_
-

194

Coronella austriaca

-

-

-

196 202

X.

Coronella girondica, Contia modesta

-

XI. Ccelopeltis monspessulana,

Macroprotodon
2o8
-

CUCULLATUS, TARBOPHIS IBERUS, T. FALLAX
XII.
XIII.

Vipera ursinii, v. renardi, v. berus Vipera
aspis, V. latastii
-

220

XIV. Vipera lebetina, V. ammodytes, Ancistrodon halys

-----XI

-

240

250

THE SNAKES OE EUROPE
INTRODUCTION
CHAPTER
I

DEFINITION AND CLASSIFICATION

SNAKES,
as an

Ophidia

— regarded

by some authorities

by the author as a sub-order of the order Sqtiamata, which includes besides the Lizards, Lacertilia, the Chameleons, Rhiptoglossa, and the extinct Dolichosauria and
order of the class Reptilia,

Mosasauria

— may

be

defined

as

greatly

elongate

scaly Reptiles without limbs, or with

mere vestiges
without

of the hind pair, without

movable

eyelids,

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,
at the

connected

symphysis by an
alone

elastic ligament.

The
from

latter

character

distinguishes

them
all

all

Lizards, but no single Lizard possesses

the others in combination.

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

and

for

swallowing

2

INTRODUCTION
much exceeding
skull,

prey

their

own caHbre
to

;

hence the

bones of the

on which a prehensile function
the cranium by
tissue, or articulated in

devolves, are loosely attached

ligamentous elastic

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 classification

which the more important characters, derived from the general structure, and especially
in
skull,

from the
logical.

were subordinated to the physioa

Such

system was

far

from reflecting
our knowledge

natural relationships.

Besides, as

progressed, drawing a distinction between poisonous

and harmless snakes became more and more
cult, so

diffi-

many

snakes previously regarded as harmless

proving to be poisonous in various degrees

at least

enough
subsist,

to paralyze the small prey
if

on which they

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
left

is

out of

consideration.

Further, in this as in

many
for

other groups of the animal kingdom, external

characters do

not

furnish

trustworthy indications
the
definitions

higher

divisions,

and

of

the

DEFINITION
families are
logical

AND CLASSIFICATION

3

therefore based exclusively

on osteo-

For those who wish to name snakes with facility, the key which concludes the
characters.

chapter on External Characters
this defect,

will,

however, remedy
all

and

suffice for

the identification of

the European species without any reference to their

anatomy.

Many

attempts

have
but

been
the

made

to

furnish an easy criterion for the distinction of
less

harm-

from

poisonous

snakes,

characters

hitherto suggested with this object can only be applied successfully to the small
tives in a limited area.
it

number
in

of representa-

Thus,
all

Southern Australia

might be stated that
dangerous to

snakes showing the regular

nine large shields on the upper surface of the head
are

man, whilst
;

those with
in

small

shields or scales are harmless

but 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.
those

Only

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
may sometimes
feel

even they

embarrassed, unless
the

the dentition

be examined;

mistakes which

have occasionally been made by some experienced
herpetologists are proof sufficient of the fallacy of

external characters for this purpose.

The Ophidia
first,

are divided into nine families, the

third, seventh,

and ninth

of

which have repre-

sentatives in
I.

Europe
(ectopterygoid)

No

transverse

bone

;

pterygoid
;

not extending to quadrate or mandible

no supra-

temporal
Maxillary

;

nasals
;

in

contact with

prefrontals

coronoid present
loosely

vestiges of pelvis.
to

attached
;

lower

surface

of

cranium, toothed
pelvic

lower jaw edentulous; a single
i.

bone

Typhlopid^.
toothless

Maxillary bordering mouth, forming a suture with
premaxillary, prefrontal,

and

frontal,

pubis and ischium present, latter forming a symphysis
II.
2.

Glauconiid^.

Transverse bone present

;

both jaws toothed.
in contact

A. Coronoid present; nasals
frontals.
I.

with pre-

Vestiges of pelvis
large,

;

supratemporal present.
BoiDiE.

Supratemporal

suspending quadrate
3.

(Subfamilies

:

Pythoninc^, Boince.)
in the cranial wall
4.

Supratemporal small, intercalated

Ilysiid^.

DEFINITION
2.

AND CLASSIFICATION
;

5

No

vestiges of pelvis

supratemporal absent
5.

Uropeltid^.
reaching

B. Coronoid absent
1.

;

supratemporal present.
;

Maxillary

horizontal

pterygoids

quadrate or mandible.
Nasals in contact with prefrontals
6.

Xenopeltid^.
7.

N asals
Three
dincB^

not in contact with prefrontals

COLUBRID^.

series:

A.

Aglypha {suhisimilies:
DasypeltincB)
;

A cwchor;

Cohibrinc^,

B.

Opisthoglypha

{HomalopsincB, Dipsadomorphince, ElachistodontincB)

C. Proteroglypha {HydrophiincEf Elapince).
2.

Maxillary horizontal, converging posteriorly

towards palatine; pterygoid not reaching
quadrate or mandible
3.

8.

Amblycephalid.e.

Maxillary vertically erectile perpendicularly to
transverse bone
rate or
;

pterygoid reaching quadg.

mandible
:

VlPERID^.

(Subfamilies

Vtperin^s, Crotalince.)

The

technical

terms

employed

in

the

above

synopsis will be found explained and illustrated by
figures in the chapter

on the Skeleton.
affinities

No
cator
;

serial

arrangement can express the

of the various groups as conceived by the classifi-

diagram therefore follows to show the author's views as to their interrelationships, and
a

possibly

their

phylogeny.

Leaving

aside

the

Typhlopidai 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
Viperidce

may have been

derived.
Amblycephalidse

Colubridas opisthoglyphae
I

Colubridas proteroglyphae
I
I

Uropeltidae
Ilysiidae
!

Xenopeltidae
\

Colubrid?e aglyphas
I

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 remains of which

the

Lower Eocene upwards,

can be identified with some measure of certainty,
being either non-poisonous types {Boidce,
Ilysiidco^

Palc^ophiidcF, Coluhridce) or Vipendce (Viperines

from

the Miocene of France and Germany, Crotalines from

the Miocene of North America).

The

vertebrse from

the Puerco Eocene of America, on the limit between
the Cretaceous and Eocene periods, described as the

DEFINITION AND CLASSIFICATION
approach the Lacertilian type.

7

oldest snake remains, Helagras, Cope, are stated to

Whether
Marsh, from

the

vertebrae

named

Syrnoliophis^

Sauvage, from the chalk of France, and Coniophis,
the

Laramie Cretaceous

of

North

America, are Ophidian, as claimed by their describers, or Dolichosaurian,

cannot be decided without

further material.

CHAPTER

II

EXTERNAL CHARACTERS— INTEGUMENT

THE form comparatively
more or
filiform, in others.

varies enormously, worm-like in some,

short

and heavy, elongate and

less slender, or

extremely gracile and almost

In this respect our

common Grass-

snake occupies a central position, and for this reason
is

termed a moderately slender form, anything above
standard being described as
or
elongate.

or below this

paratively short
stoutest

Our

comshortest and

European Snakes are the Vipers, especially Vipera ursinii ; our longest and slenderest, the Coluber and Zamenis, especially Zamenis dahlii. These
extremes in both directions
passed
are,

however,
as

far

sur-

by

many

exotic

snakes,

we

find

on

comparing, for instance, one of the African Puffadders
(Bitis),

with certain Oxybelis and Leptognathus

from Tropical America.

The body may be some-

what

rigid, as in

some burrowing and ground snakes,
or extremely flexible, as in
in the

not unlike in appearance to our Slow- worm and other
limbless Lizards
;

many

Pythons and Boas and
This
flexibility

Tree-snakes generally.

may

be accompanied by a vertical

compression of the body in relation with an arboreal
existence, whilst sluggish snakes, such as
8

most of the

EXTERNAL CHARACTERS
Viperidse,

9

the body,

may be remarkable for the flattening of which they may further increase when
appearance

basking in the sun or in order to assume a more
formidable

on

the

approach
possessed by

of

an

enemy.

This power of flattening out the whole or
is

the anterior part of the body

many

snakes, poisonous as well as harmless, and reaches
its

highest degree in the Cobras of India and Africa,

the expanded anterior part being
*'

hood,"

from

the

known as the Cobra di Portuguese name
'*

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
In some of these Sea-snakes
**

the body compressed.

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
gests a limbless Plesiosaur.
tail,

and sug-

The
versely

tail,

the part of the body behind the transvent,
is

cleft

most frequently about one;

fourth or one-fifth of the total length

but

it

may

be
in

much
one

shorter, even reduced to a

mere stump, as
snake,
as
in

the Typhlops, or, at the opposite extreme, enter for
half in

the length

of the

the

African Xemirophis.
to a fine point
;

This organ

may

taper gradually

or end abruptly, as

if

mutilated
see in

;

or

terminate in a horny spine, such as
of the Typhlops

we

some

or in the Australian Death-adder,

10

INTRODUCTION
like

AcanthophiSf or in a series of horny segments which
are vibrated

a

rattle,

as

in

the

well-known

Crotalus of America, to

which we
is

shall refer again at

the end of this chapter.

In some of the burrowing
obliquely truncated,

Uropeltidse, the very short tail

with indurated shields above, and acts as a trowel.

And,

finally,

the marine snakes of the

subfamily

Hydrophiinse are distinguished by a strongly compressed,
outline.
tail is

oar

-

shaped

tail,

with

rounded

vertical

In a few forms, arboreal or aquatic, the
less prehensile.

more or

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
mentioned further on
in

of Bold snakes, to be
description

the

of

the

skeleton, terminate in a claw-like

horny spur, which
distinctly,
in

appears on each side of the vent in the male, and

sometimes
female.

also,

though

less

the

These spurs are probably of use in facilitating the pairing, an explanation which appears the more plausible from the fact that the snakes provided 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
Although never actually compressed, except
rostral region,
it

ii
in the

may

be very narrow and
it

elongate,

whilst in the opposite extreme

may

be strongly

depressed,

and so broad behind as
the
anterior
is

to be abruptly

defined

from

part of the body,

or

" neck."

This feature
that

very marked in some of

the Viperidae, and this has given rise to the incorrect
generalization

poisonous

snakes
a

are

distinflat

guished from the harmless by

broad and

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 Glauconiidae aside
for the present,

snakes have a wide gape,

cleft far

beyond the

vertical of the eyes, with,
in front for the

when

closed,

one or two notches
is in

passage of the

protrusible, bifid tongue.

In most snakes this chink

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
free

from minute to enormous, are usually
surrounding shields, and

from the
a trans-

may move under

parent cap like a watch-glass, which appears to

12

INTRODUCTION
The view
as

represent the lower eyelid of Lizards.
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
in

transparent disc appearing like a small

window

the movable lower eyelid, gradually increasing in
size so as to

occupy the whole of the lower

eyelid,

which
upper

finally
lid

and

becomes fused with the rudimentary loses its mobility. In Ilysia and in

most of the Uropeltidse, the transparent disc over the eye is confluent with a thick horny shield of
which
it

occupies the middle.
is

The

pupil

usually circular or vertical,
it

rarely

horizontal.

In some forms

is

difficult to decide elliptic
;

whether
like the

it is

round or vertically

in others,

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 contraction 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, depressed or compressed, sometimes projecting strongly

beyond the mouth, turned up at the end, or terminating in one {Langaha) or two (Herpeton) long scaly dermal appendages. In some burrowing forms
it

is

provided with a more or

less

trenchant hori-

zontal or vertical edge.
(loreal region)

When

the sides of the snout

form an angle with the upper surface.

EXTERNAL CHARACTERS
the angle
is

13

termed the ''canthus

rostralis,"

which

may be intensified by the loreal region being concave. The deep pits which are sometimes present on
the Hps 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 Typhlopidae, the head passes gradually into
the vermiform body, and the small

mouth

is

situated
;

on the under surface of the projecting snout head so resembles the extremely short
tail,

the

and the

mouth
which

is
is

so similar in shape

and position

to the vent,

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.
It is

There

is

no mental groove.
abbreviated

much

the

same

with the Glauconiidse, which have, however, a some-

what

less

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
be provided
with a sharp cutting horizontal

may
edge.

14

INTRODUCTION
Snakes are covered with epidermal folds in the form and shields, the shape and arrangement of
affords important characters for their classifica-

of scales

which
tion.

Dermal
scales

ossifications are absent.

The

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
scales,
in

and

also with the vertebras.
is

number with the series of The body of the
uniformly covered

Typhlopidae and Glauconiidge

with polished, closely adherent, rounded, overlapping,
sub-equal scales, without even an indication of ventral shields.

In

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 hexagonal,

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
the body, as
obliquely,
is

15

ning in longitudinal series parallel with the axis of
the rule, they are sometimes disposed
in

and among those

which we meet with

this peculiarity several
in

genera are further remarkable

having some of the oblique lateral scales furnished
keel, to

with a serrated
in

which we

shall again allude

the chapter on Habits,

when

dealing with the

rustling sounds

produced by certain snakes.

The

number
Boa)
;

of longitudinal series of scales on the body

varies from 10 {Herpetodryas) to nearly 100 {Python,
in

the

European species

modesta) to 50 (Eryx jacuhis).

from 17 {Contia The scales are some-

times furnished near the end with one or two shallow
impressions, termed ''apical pits," which afford indications for the distinction of genera

and species
is

;

unless

of a lighter or darker colour, as

often the case,

these pits are not always easy to see, except in a

strong light and with the aid of a powerful magnifying glass.

The
they

ventral

shields,

also

called " gastrosteges,"
;

usually occupy the whole width of the belly

but

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
Colubrids.

in the posterior border,

corresponding to

the keel, as in several of the more arboreal genera of

The

shields under the

tail,

termed subin certain

caudals or ''urosteges," are sometimes similar to the
ventrals, but

more

often disposed in pairs

;

i6

INTRODUCTION
and the others paired.
is

species or individuals
single,

some of the subcaudals are When the number of
is

subcaudals

given in the descriptions, each pair

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,
sexes.

and sometimes also

for the distinction of

The subcaudals
in
is

are nearly always
is

much

fewer

than the ventrals, but the difference
great

often not so
tail

the males as in the females, the

of
It

which
is

usually shorter in proportion to the body.
in

noteworthy that

many

species,

if

the

number

of

subcaudals (C.) be added to that of the ventrals
the total
is

(V.),

nearly the

same

in the

male as

in the

female, however
differ

much
way

the respective numbers

may

when taken

separately.

The

following figures

may

be given by

of example, taken

from British

specimens
Coronella austriaca
:

S
?

V. 154

;

C.

V. 165; C. 48 V. 138; C. V. 144; C.

Vipera herns:

^
?

58=212 = 213 35 = 173
29=173

Although
to hold

this rule is
all

does not apply at

to

by no means universal, and some species, it will be found
cases,

good

in

many

and

is

of interest

in
in

showing that the changes that have taken place

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.

i

In dealing with

certain species

— of Vipers,
two

for instance

it is

impor-

tant, for systematic purposes, to

keep the counts of

shields distinct for the

sexes.

The
is

shield

which covers the

vent, the jjnal^hield,

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-

and number of which furnish some of the most important characters for the distinction of genera and species. These head-shields belong to
portions,

two primarily

different types,

from each of which

all

further modifications

may

be regarded as derived by

alteration in shape or by disintegration.

type

is

that

The first shown by the Typhlopidse and Glauis

coniidse,

which

explained by the figure on the

next page.

The rostral, which is usually the
shields, extends to the

largest of the head-

upper surface of the head, of
the greater part.

which

it

may occupy

In

the

Glauconiidae, the ocular usually borders the mouth.

As may be seen by a comparison of the
is

first

figure

with the second, the arrangement of the head-shields
essentially different

from that which prevails

in the

Colubrids and the majority of other snakes.

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
second.
loreal
first

row, three in the

The internasals and the temporals, and the and the preocular, are sometimes absent, and

the prefrontal or the internasal
or

may

be single.

One

two large shields are in rare cases present behind the parietals, and are called occipital.

A

breaking up into smaller shields takes place in
snakes.

many

In the

Pythons, for instance, the

Fig.

I

Head of
;

Typhlops hraminiis. British India ")
;

(From "Fauna
n,
;

of

/,

Frontal

ip, interparietal p, parietal •,po, preocular;

I,

labial

;

nasal
y,

;

0,
;

/>;/,

prefrontal

rostral

'50,

ocular supra-

ocular.

frontal
cleft,

be divided into two by a longitudinal and separated from the prefrontals by small
In

may

shields.

some
size,

Vipers,

such

as

V.

hems and

V.

iirsinii, in

which the

frontal

and

parietals,

though

reduced

in

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
snakes
the
rostral are

19

small shield or shields behind the termed " apical," and those on the upper

edge of the snout are termed " canthals."

The

shield
is

which, in Vipers, separates the rostral from the nasal

I

pro

Fig.

2— Head

of Zamenis

ventrimaculatus.

(From " Fauna

of

British India ")
^5,

Chin-shields (anterior)
in,

internasal
;

;

^,

loreal
;

pto,

m, mental n, nasal postocular ; r, rostral; temporals (first row) t' t,
;

chin-shields (posterior) /, frontal labial (upper) la\ labial (lower) p, parietal pf, prefrontal pro, preocular
;

cs',
;

;

la,

;

;

;

sbo,
,

subocular so, supraocular temporals (second row) y, first
; ;

ventral.

called " naso- rostral."

to the scaly dermal

Allusion has been made above appendages which terminate the

snout in certain genera.

Some Viperidae are furnished

20

INTRODUCTION
their sinistral

with horn-like erect spines above the eyes or at the

end of the snout, which add greatly to
appearance.

The

periodical shedding of the outer layer of the

epidermis in

a

single
is

piece,

including

even

the

covering of the eye,
peculiarities of

one of the most striking
to

snakes, although paralleled in the

Lizards
British

of

the

family Anguidae,
belongs.

which
inside

our

Slow-worm
at

The
is

skin

becomes
out

detached

the
tail,
is

lips,

and

turned

from head to

without any sort of laceration

when

the snake

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
to stretching.

reptile,

owing
is

In Sea-snakes the epidermis
is

cast

piecemeal, and sloughing
in ordinary snakes.

a longer operation than

In
*'

Rattlesnakes

each

piece
tail

of

the

rattle,

or

crotalon," in which the

terminates, represents

a retained portion of the sloughed epidermis.

This

remarkable appendage looks
rings, but
it

like a

number

of horny

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
together

fits,

thus keeping the
the

pieces

without

impairing

mobility

necessary to produce the rattling sound for which

INTEGUMENT
the apparatus
is

21

intended.
is

At each exuviation one
added.

bell-shaped horny piece

The number
to the

of

segments in the

rattle

is,
;

therefore, not an index to

age, as formerly believed

nor

is it

number

of

exuviations, for w^hilst 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 been
six

have

six pieces to the rattle if there

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 show^s

whether

it

was formed

at birth or at
in the

any

later period,

no growth taking place

horny

tissue.

So

far as trustworthy records are concerned, the

largest snakes known, the Malay Python reticulatus and the South American Anaconda, Eunectes murinuSy

reach a length of 25 to 30
skins

feet. Measurements of 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
skin,
it is

from such a stretched
of

necessary to deduct the interstitial spaces
scales,

showing between the
snake known

and about one-fourth

the scale to allow for the overlap.
is

The

smallest

4 inches long {Glaucoma disstmilis).
quatuorlineatus)
;

The
is

largest

European snake (Coluber

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 inferior; body vermiform, covered with uniform scales above and beneath vent close to the end of the body, the extremely short tail
;

small,

;

ending
II.

in a

small spine
small,

Typhlops vermicularis.

Eyes very
surface

with vertical pupil;
covered with
small

upper
scales

of

head

ventral shields
tail short,

much narrower than
;

the body

ending obtusely
;

subcaudals single, or
feebly keeled, in

mostly single
40 to 50 rows
III.

scales

smooth or

Eryx jaculus.

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
I.

;

anal shield usually divided.

Dorsal scales strongly keeled, with paired
apical pits
a.
;

a single anterior temporal.
;

Nostrils lateral

internasals broadly trun-

cate in front.

Scales in 19 rows;
oculars
;

normally

i

pre-

usually 7 upper labials, third
;

and 3 postand fourth
subcaudals

entering the eye

ventrals 157-181

;

50-88

Tropidonotus natrix.

EXTERNAL CHARACTERS
b.

23
internasals

Nostrils

directed

upwards

;

much narrowed
Scales in 19 rows
oculars
; ;

in front.

normally 2 pre- and 3 or 4 postusually suboculars sometimes present
;

8 upper

labials, fourth or fourth
;

and

fifth

entering

the eye; ventrals 160-187

subcaudals 48-79
Tropidonohis
tessellaUcs.

Scales in
pre-

21 (rarely 19 or 23) rows

;

normally

i or 2

third

and 2 postoculars; usually 7 upper labials, ventrals and fourth entering the eye
;

147-164; subcaudals 46-72.
2.

Tropidonotus viperinus.
or

Dorsal

scales

smooth
loreal.

feebly

keeled;

normally a single
a.

Two
porals

or three

superposed
;

anterior
nostril

tem-

(very rarely one)

usually

between two nasals.
a.

A

subocular below the preocular.

* Scales smooth, in 17 or 19 rows.

Tw^o upper labials entering the eye
in contact

;

preocular not

with the frontal

;

scales with

two apical
angulate

pits

;

ventrals

more

or

less

distinctly

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
ally,

ventrals very distinctly angulate later;

205-218

subcaudals 98-132.
in 23 to 29

Zamenis

dahlii,

^^ Scales

rows (usually 25 or
pits.

27), with two apical

24

INTRODUCTION
labials

Upper
series

usually separated from the eye by a

of suboculars;
;

preocular in

contact with

the

frontal

scales

smooth

;

ventrals
;

very dis-

tinctly

angulate

laterally,

222-258

subcaudals
hippocrepis.

77-107

Zamenis
with the frontal
ventrals

Two

upper labials entering the eye; preocular not
;

in contact

scales feebly but dis-

tinctly

keeled;

not

angulate

laterally,

195-234; subcaudals 56-90. Coluber quatuorlineatus.

Two
in

upper labials entering the eye
contact

;

preocular not

with the frontal
;

;

scales

smooth or

faintly keeled

ventrals not or but very obtusely

angulate laterally, 172-214; subcaudals 50-80
Coluber dione.
/3.

No

subocular

;

scales smooth, or faintly

keeled on the posterior part of the body. * Ventrals more than 200 scales with
;

two apical
Snout obtuse;
ally,

pits.

rostral broader

than deep; scales

in

21 or 23 rows; ventrals distinctly angulate later-

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

Cohiber leopardmns.
;

Snout pointed, strongly projecting
than broad, wedged
scales in 25 to 29
ally,

rostral

deeper

in
;

between the internasals
ventrals not angulate later-

rows

201-220

;

subcaudals 48-68.

Coluber scalaris.

EXTERNAL CHARACTERS
mostly with a single apical
Rostral at least as deep as broad, often
pit.

25

^^ Ventrals not more than 200; scales

wedged

in

between the internasals
third
(rarely 21)

;

usually 7 upper labials,
;

and fourth entering the eye
rows;
ventrals

scales in 19

153-199;

subcaudals

41-70
Rostral broader than deep
fourth
(rarely
;

Coronella austriaca.

usually 8 upper labials,
scales in

and

fifth

entering the eye;
ventrals

21

19 or 23) rows;

170-200;

sub-

caudals 49-72
b.

Coronella givondica.
;

A

single anterior temporal

nostril

in

a

single

nasal;

scales

smooth, with single

apical pits, in 17 rows; ventrals 150-191

subcaudals 53-78
3.

Contia modesta,

Scales longitudinally grooved in the adult, in

two strongly marked;
17 or 19 rows
;

loreals

;

canthus rostralis
very

frontal

narrow, in
;

contact with the preocular; ventrals 160-189

subcaudals 68-102

...

Co^lopeltismonspessulana,

B. Pupil vertical or vertically subelliptic (sometimes appearing round in Macroprotodon).
I.

Scales smooth, mostly with single apical pits

;

upper surface of head with nine large shields.
Frontal ij 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 cucullatns.

26

INTRODUCTION
much
shorter than the
parietals
loreal

Frontal il to ij times as long as broad,
;

entering the

eye; scales oblique, in 19 or 21 rows; ventrals

186-222; anal divided

;

subcaudals 48-73
Tarbophis fallax.
as broad, nearly

Frontal

ij to i^ times as long
;

as long as the parietals

loreal entering the eye

scales oblique, in ig or 21

rows

;

ventrals 203-235

;

anal entire; subcaudals 54-70
2.

...

Tarbophis iberus.
pits
;

Scales

keeled, with

two apical
nostril
if

anal

shield entire.
a.

No

pit

between the
from
eye
the

and the eye;
present
;

upper head-shields small,
separated
rostral
labials
a.
;

nasal

rostral

by a
the

naso-

separated

from
at the

upper
supra-

by suboculars.

Snout not turned up
ocular
usually

end

;

extending

posteriorly

beyond the

vertical of the posterior border

of the eye; frontal and parietal shields

usually well developed
series of scales

;

usually a single

between the eye and the
flat

upper

labials.

Snout

obtusely pointed,
;

above, or with the

canthus slightly raised

rostral usually in contact
;

with a single apical shield, rarely with two

6 to g

upper

labials, usually 7

or 8

;

scales in ig rows,

rarely 21; ventrals:

^

120-135, ? 125-142
Viper a ursinii.

EXTERNAL CHARACTERS
Snout pointed, with raised canthus
tact with a single apical
labials;
cT
;

27

rostral in con-

shield

;

8

or g
;

upper

scales

in 21

rows, rarely ig

ventrals

130-148, ? 130-150

Vipera renardi.
flat

Snout truncate or broadly rounded,
with slightly raised canthus
;

above or
contact
;

rostral

in

with two apical shields, rarely with one

8 or g

upper labials; scales
ventrals:
13,

in 21 rows, rarely ig or 23;
?

c^

132-150,

132-158
less

Vipera berus.

Snout usually more or

turned up at

the end or produced into a scaly dermal

appendage
posteriorly
terior

;

supraocular not extending
vertical of the pos;

beyond the

border of the eye
often

frontal

and

parietals

absent or very

small

2 or 3 series of scales

between the eye and
;

the upper labials

;

g to 13 upper labials

scales in 21 or 23 rows, rarely ig 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: (^134-158, ? I4i-i6g.

Vipera aspis.

Snout simply turned up or produced into a small
appendage, the raised portion with 5 or 6 (rarely 3) scales rostral ij to 2 times as deep as broad
;

ventrals: ^125-146, ? 135-147

...

Vipera

laiastii.

Snout produced into an appendage covered with
10 to 20 scales
of the
rostral
;

rostral not reaching the
S'

summit
133-161,

appendage; ventrals:

$135-163

Vipera ammodytes.

28
7.

INTRODUCTION
Snout not turned up
small shields
at the

end

;

supra-

ocular narrow or broken up into several
;

upper

with small, usually keeled scales
three series of scales
the upper labials
usually 25
; ;

head two or between the eye and
surface
of
;

scales in 23 to 27 rows,
:

ventrals

S

151-177, ? 153-

180
b.

Vipera lehetina.

A

pit

between the

nostril

and the eye;

upper surface of head with g large shields nasal in contact with the rostral; third

upper
23

labial
;

entering the eye; scales in

rows

ventrals

149-174

;

subcaudals
halys.

31-44

A ncistrodon

CHAPTER
IN

III

COLORATION
dealing with the coloration,

we have

first

to

distinguish between the colour
is

and the markings.

The former

very often

highly variable

among

snakes of the same species, to say nothing of the

changes which
after exuviation

may
;

take place with age or with

the condition of the individuals, whether before or
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
latter afford

Common

Viper.

The
often

more important characters, and
indications for the

furnish valuable
species
;

distinction of
is

but even the disposition of the markings

subject to great individual variations,

more

likely to

mislead than to help the inexperienced student in
the discrimination of species.
advisable to resort in the
first

It is therefore

always

instance to structural

characters for the purpose of specific identification,

and to

fall

back on coloration only as a means of

If we were to be guided by colour and markings alone, how could we believe that an

confirmation.

adult

four-lined

Coluber quatuorlineatus
29

is

of the

30

INTRODUCTION
species
;

same

as

the
yet,

sauromates

and

if

handsomely spotted Coluber we compare the young of
find

these two snakes

we

them

to

be absolutely

identical in their markings, and, in the absence of

any structural
species, of
It is

differences,

we

are forced to conclude

that they only represent two forms of the same

which the

latter is the

more

primitive.

nevertheless a fact that, with a few excep-

tions, the

markings, however variable they
certain fundamental

may

be,

are

reducible to

patterns

to

which the innumerable variations may be traced back, and their derivation followed and scientifically
explained.

Let us consider,

for instance,
its

another
:

species of Coluber, highly variable in

markings

C. leopardinus, of

which the typical form, so called
first

from having been the
primitive.
First,

described and named,
the

is

not by any means to be regarded as

most

we must

take for granted that the markings
of
spots,

of

all

such snakes, whether consisting

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)
(L)
;

;

(2)

Dorso-lateral

(DL)

;

(3)
first

Lateral
starts

(4)

Ventrolateral (VL),
line of the head,
;

The
and
is

from the middle

con-

tinued along the spine

the second occupies the
third,

space between the

first

and

which originates

at the tip of the snout, passes

through the eye, and

COLORATION
is

31

continued on the temple and along the side of the
;

body

the fourth follows the lower

lip,

and extends
in

along each side of the belly.

Bearing this

mind,

we

find

that

the variety of C. leopardinus

named
is

schwoederi, with a vertebral series of paired spots,

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
into a single row,

D

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
(see

produces

the

variety

named

qtiadrilineatus

Plate VII.).
series

In this particular instance, the paired

D

has fused into a single streak on the head,
series

and the

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
from

snakes show an interocular band extending

lip to lip,

through the eyes, across the
In others the lateral stripe
of the

inter-

orbital region.

L may
branch

bifurcate

in

front

eye,

an

upper

extending

across

the

snout,

through

transverse

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
fusion of series

D

a

W-shaped

figure.

The

pattern of markings on the
is,

upper surface of the head
complicated, and hence

however, often very
of explanation.

difficult

32

INTRODUCTION

As a second example of the derivation of patterns, we may mention Vipera aspis, which varies enormously 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 transverse bars
;

all

these are derived from

the paired
fuse

spots of series D.

Each

pair of spots

may

and

form transversely oval or
the spots

elliptical spots or bars, or

may assume an

alternate disposition from

which, through confluence, the zigzag or smuous

band

results.

Thus, spotted and striped patterns
to

may
first

be

traced

a

common

origin,

however
at

fundamental the difference between them appears
sight.
If

the elements of the four series, D,
unite transversely with each other,

DL,
and

L, and

VL,

also with the spots

on the ventral surface, we
as

obtain

ringed

forms

such

the

Coral-snakes.

That the black nuchal collar of our common Grasssnake 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 vipermus, and Vipera berus. Although it sometimes happens that a definite
system of markings prevails throughout a genus,

COLORATION
Elaps, this
is

33

such as the annulate form in the South American
far

from being universally the case;
or individuals
of the

many
same

closely allied species,
species,

may

be distinguished by very different
the

patterns.

Even on

same individual we may

find

two opposite types of markings without any transition, as in two Central American species of widely
different genera, Polyodontophis annulatus

and Zamenh
is

mexicanus, in which the anterior part of the body

annulate
striped.

or

barred,

and

the

rest

longitudinally

It is also

a remarkable fact that very often 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 correspond in number on the two sides. In the handsome South American Lachesis alternatus, 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 alterbut some may lie opposite nating, as is the rule each other and back to back, this being due to to the fact that the numbers of the markings do not In one specimen I correspond on the two sides. twenty-four of these markings on the left side, count and twenty-seven on the right. This shows that
two
sides of the
;

34

INTRODUCTION
number
In
fact,

great importance cannot be attached to the

of the markings, for systematic purposes.
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.
bilateral

The

which we have alluded produces the chess-board arrangement of the ventral

asymmetry

to

spots in

many snakes. Among the markings which
some Colubrids, of a
pair of light

call for investigation

as to their
in

meaning, we must allude to the presence,
small, light, dark-edged spot,

or of a

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
Moth.

?

At present

it

is

as

inexplicable as the lugubrious
of the Death's-head

emblem on

the thorax

It

cannot be suggested

that
for

mark intended to terrify intruders, when the Cobra is at rest the hood is folded, and
it is

a warning

the characteristic marking
as soon as
faces its
it

is

not displayed

;

whilst
it

is

aroused, and the hood expanded,
in

enemy

such a way that the spectacle, or

ocellus, is not to

be seen.

First

among

the most brilliantly coloured snakes,

COLORATION
of

35

which there are many, stand the Coral-snakes,

ElapSy of America, mostly annulate with red, yellow

or white,

and

black.

This striking coloration obtains

also in diverse harmless snakes inhabiting the

same
been

part of the world,

and

this coincidence has

adduced
for

in favour of the theory of

mimicry, corre-

lated with that of natural selection,

which accounts
a

the resemblance as being of advantage to
is

harmless species, which
notorious for
its

thus mistaken for one

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
aulicuSf

the Krait, Btmgarus cc^ruleus, and Lycodon
in

India, the

Pit-viper, Ancistrodon
pulverulentus,

himathe

layamcs,

and Psammodynastes

in

Himalayas and Assam, are good examples of such
cases.

On

the other hand, there are equally striking

what one would regard as mimics if is no better case of general resemblance between a poisonous and a harmless snake than we find in the Indian Cobra and the Cohiber corais of tropical America, where Cobras are absent, or between a Viper and the Boid Enygms asper, from New Guinea, where
instances of

they only occurred together; thus, there

no Vipers

exist.

Without attempting
account for

any suggestion to the similarity of markings which prevails
to offer

36

INTRODUCTION
may be drawn
predominance of longitudinal dark and light Indo-Malayan representatives of the

in certain parts of the world, attention

to the

sftipes in the

American Elaps, shared by many innocuous snakes of similar form inhabiting the same region, and
to the striped tails

common
if

to

various

Colubrids

of Madagascar, as

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

We

may

point out at the
in

resemblance
Batrachian

and black markings. same time the very marked form and coloration between the
glutinosus,

Uropeltid Melanophidium bilineatum, and the Apodal
Ichthyophis

both

occurring

together in Southern India.

The
brid,

colour of snakes often harmonizes with their

surroundings.

Thus,

many

Tree-snakes, Boid, Colugreen,
like

or Viperid, are

of a bright

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
districts the desert individuals are paler,

^^ without or

with

less distinct

markings, as compared to the^r
In addition to

fellows
their

among

other surroundings.

markings, some snakes are adorned with a
gloss*,

metallic iridescent

due to a

fine striation of

the scales.

The
red,

iris is

often metallic, gold, bronze, or copper-

and the black streaks of the head sometimes
it.

extend over

Although, unlike
to

many
their

lizards,

snakes are unable

rapidly

alter

semblance of
neck or body

this
;

some produce a phenomenon when inflating their
colours,
is

this

due to the presence of dark
in the

and

light

markings or of a bright pigment

interstitial skin,

which
in the

is

not seen

when

the scales

overlap.

Thus,

Indian Tree-snake Dryophis

mycterizans the skin
scales

between the green or brown

in the anterior part of the body is black and white, producing a striped pattern when the the skin of the same region is neck is inflated bright vermilion in the Malay Tvopidonotus submini attis ; many more examples could be quoted. The spectacle marking on the hood of the Indian
;

Cobra involves the
skin.

scales as well as the interstitial

As a

rule there are

no sexual differences
in

in colour.

Yet these are so marked
nized

our

Common
is,

Adder that

the sex of a specimen can nearly always be recog-

hy the coloration.

This

however, the

38

INTRODUCTION
genus to which the Adder
is

exception, even in the
belongs.

A

nuptial dress

unknown

in snakes.

A special
than
its
is

livery for the

but very often the new-born
parents,

young is rather exceptional, is more vividly coloured

and

in

many

black varieties the

young

similar to the typical form.

Some green

Tree-Boids (Chondropython and Corallus caninus) are
not green, but yellowish, cream-colour, or pinkish,

when young,
spots,

the green appearing around the white

which are the remains of the ground colour, and
Pit-viper Lachesis
is

gradually spreading over the whole body. Conversely,
the young of a variety of the
waglerij

common

in the

Malay Peninsula,
African

green,

and the adult black and yellow.
Grayia ornata, a

In the young of

West

Water-snake,

the

markings of the young are to those of the adult
like positive

bars, forked

and negative in photography, the white on the sides, which extend across the
light

black back of the former being gradually trans-

formed into black bars on a
latter;
in

ground

in the

such a case

it

is

impossible to decide

whether the dark or the light parts are to be considered as the ground colour.

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

COLORATION
Melanism
affects
all
is

39

frequent in
in

snakes, and sometimes

individuals

seems undesirable to bestow varietal
aberrations, as
is

same locality. It names on such so frequently done by systematists,
the
in

any more than we should

the case of albinos.
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

Melanism may be produced

prolonged sojourn

in spirits.

Some-

times, as in Zamenis gemonensisy the uniform black

colour appears only as the snake approaches the
adult
livery.

condition,

the

young

having

the

normal

Partial albinism terized
still.

is

rare; perfect albinism, characin the eye, rarer

by absence of black pigment
in

Cases have been observed, among European
Tropidonotus natrix

species,

and

tessellattis,

in

Coluber longissimuSj and in Coronella aiistriaca.

CHAPTER

IV

SKELETON

THE
frontals

typical

Ophidian skull

is

characterized by

a solidly ossified brain-case, with the distinct

and the united

parietals extending

to the basisphenoid,

which

is

large

downwards and produced
ossified,

forward into a rostrum extending to the ethmoidal
region.

The
The

nasal region

is less

completely

and the paired nasals are often attached only at their
base.
occipital condyle
is

either trilobate

and

formed by the basioccipital and the exoccipitals, or a
simple knob formed by the basioccipital
occipital
is
;

the supra-

excluded from the foramen

magnum.

The

basioccipital

may
is

bear a strong, curved ventral
(in

process or hypapophysis

the Vipers).

The

prefrontal

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
it

between

Pythons a supraorbital and the prefrontal.
is

is

intercalated

The
The

premaxillary

single

and

small,

and as a

rule connected with the maxillary only

paired

vomer

is

narrow.
40

The

palatine

by ligament. and

SKELETON
;:;'-/•

41
pro ste

sor

-P

pT

cpg

p

<j'

CO,

sce>

st& CO,

^g;

Fig. 3

— Skull

of Python amethystiniis. (From British Catalogue of Snakes)
;

Museum

an,

Angular; ar, articular; bo, basioccipital cor, coronoid c.a, columella auris (stapes)
;
;

bs,
<f,

;

basisphenoid dentary eo, ex;
;

occipital ; epg, ectopterygoid (transverse) /, frontal m, maxillary nasal pm, premaxillary 71, //, palatine p^ parietal pro, prootic ptf, postfrontal pg, pterygoid pyf, prefrontal sor, supraorbital so, supraoccipital sp, splenial q, quadrate
; ;
;

;

;

;

;

;

;

;

;

ste,

supratemporal

;

tu,

turbinal

;

v,

vomer.

42
pterygoid
of the
are

INTRODUCTION
elongate
the latter

and

parallel

to

the axis

skull,

diverging behind and ex-

tending to the quadrate or to the articular extremity the pterygoid is connected with of the mandible
;

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
(often

is

usually large

and elongate, and
In rare cases
is

attached to the cranium through the supratemporal

regarded as the squamosal).

{Miodon, Polemon) the transverse bone
articulates with

forked,

and

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

the calibre of the mouth. For the same reason, the rami of the lower jaw, which consist of dentary, splenial, angular, and articular
elements, with the addition of a coronoid in the

much exceeding

Boidae and a few other small families, are connected
at the

symphysis by a very extensible
is

elastic ligament.
cartila-

The hyoid apparatus
ginous
filaments

reduced to a pair of

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 (PsammophiSf

SKELETON
CcelopeUis)\ the maxillary

43

may
as

be

much

abbreviated
;

and movable
pterygoids

vertically,

in

the Viperidae

the

may
;

taper

and

converge

posteriorly,

without any connexion with the quadrate, as in the

Amblycephalidae

the supratemporal

may

be

much

Fig. 4

Skull of Typhlops

lumhricalis.

(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

tremely large

;

the prefrontals

may be may join
;

short or exin a

median

suture in front of the frontals
freely

the dentary

may

be

movable, and detached

from the articular

posteriorly.

44

INTRODUCTION
The
deviation

from the

normal

type

is

much

greater
like

still when we consider the degraded, wormmembers of the families Typhlopidse (Fig. 4,

p. 43)

and Glauconiidae

(Fig. 5), in

which the

skull

is

very compact and the maxillary

much

reduced.

In

j>ro

Fig. 5

Skull of

Glauconia macvolepis. (From British Catalogue of Snakes)

Museum

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
The
principal modifications

45
of the skull in the
as
in

European genera may be
following synopsis
I.

contrasted

the

Quadrate articulating with the cranium, supratemporal absent; mandible much shorter than the
skull,

with coronoid bone
prn.

;

maxillary small, on
pTTV

ptf.

\

Fig, 6

Skull of

Tropidonotus natrix. (From British Catalogue of Snakes)

Museum

Lettering of the bones as in Fig. 3

lower aspect of cranium
ing to quadrate
;

;

pterygoids not extend-

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.

46

INTRODUCTION
.

A

Mandible with coronoid bone
contact with frontals and

;

nasals in suturai
;

prefrontals

trans-

verse bone short, not projecting

much beyond
as

cranium

;

maxillary

not

half

long

as

Fig. 7

Skull of Zamenis

gemonensis.

(From

British

Museum

Catalogue of Snakes)

mandible, which
occiput)

is

not longer than skull (to

Eryx.
;

B,

No
I.

coronoid bone

nasals isolated.

Maxillary elongate, not movable vertically.
a.

Maxillary half as long as mandible.
skull, projecting far

Supratemporal half as long as

SKELETON
beyond cranium
skull
;

47

mandible much longer

than

Tropidonotus.
skull, projecting

Supratemporal not half as long as
far

beyond cranium

;

mandible much longer than
Zamenis.

skull

Fig.

8— Skull

of Coluber longissimiis. (From British Catalogue of Snakes)

Museum

Supratemporal not half as long as
but
slightly

skull, projecting

beyond

cranium

;

mandible
skull,

much

longer than skull

Coluber.

Supratemporal not half as long as
jecting
skull

not pro-

beyond cranium

;

mandible not longer than
Coronella, Contia.

48
h.

INTRODUCTION
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

(From British Coronella austriaca. Catalogue of Snakes)

Museum

much

longer than quadrate, nearly straight in front
;

of prefrontal

a large vacuity between the frontal
Ccelopeltis,
;

bones and the basisphenoid

Quadrate not longer than supratemporal
little

maxillary

longer than quadrate, strongly curved in front

of prefrontal

Macroproiodon.

SKELETON
Quadrate
little
;

49

longer than supratemporal maxillary longer than quadrate, nearly straight in front
Tarbophis.
p//'

of prefrontal

f

pi-f

p pro

Fig. io

— Skull

of Vipera lebetina. (From British Catalogue of Snakes)

Museum

Lettering of the bones as in Fig. 3

2.

Maxillary

much

abbreviated
half

and
skull

erectile

supratemporal

not

as long

as
;

skull

mandible much longer than
cipital with a strong process.
4

basioc-

50

INTRODUCTION
Vipera.

Maxillary bone solid

Maxillary bone hollowed out

A ncistrodon.

The The

vertebrae

number 130
ursinii) to

to 500

in the

European

forms 147 {Vipera
vertebral

330 {Coluber leopardinus).

column

consists of an atlas (com;

posed of two vertebrae) without ribs
caudal vertebrae,
all

numerous

pre-

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
processes

ribless caudal vertebrae

with

simple transverse processes.
transverse

When

bifid,

the ribs or

have the branches regularly
usual cup-and-ball articulation,
elliptic

superposed.

The centra have the

with the nearly hemispherical or transversely

condyle at the back (procoelous vertebrae), whilst the
neural arch
is

provided with additional articular sur-

faces in the form of preflattened,

and post-zygapophyses, broad, and overlapping, and of a pair of anterior
fitting

wedge-shaped processes called zygosphene,
just

into a pair of corresponding concavities, zygantrum,

below the base of the neural spine.

Thus the
by eight

vertebrae of snakes articulate with each other

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
and-mortice " in carpentry.

51
vertebrae

The precaudal

have a more or
plate-like above,

less

high neural spine which, as a

rare exception (XenopJwlis),

may

be expanded and
trans-

and short or moderately long

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.

Fig. II Posterior Precaudal Vertebra of Lioheterodon (A) and Helerodon (B). (From British Museum Catalogue of Snakes)
a,

Back view

;

b,

lower view

;

c,

side view.

or

haemapophyses, which are sometimes continued

throughout (Fig. 11, 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
pelvic are

arch, but remains of the
Typhlopidce,
Ilysiidce.

found

in

the

the

Glauconiidce,
first
?)

the BoidcB,

and

the

In the

these vestiges are reduced to a
;

single

bone (ilium

on each side

in the

second they
latter form-

consist of ilium, pubis,

and ischium, the
is

ing a ventral symphysis, and a rudimentary femur
whilst in the third there

a long ilium, attached to
bifurcate transverse pro-

the lower branch of the
cess

first

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
the most generalized INshow the nearest approachsnakes — those which to hzards — 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
including

is

devoid of teeth in the great majority,

all

European representatives, of the Ophidia.
is

In the egg-eating snakes of the genera Dasypeltis

and Elachistodon the dentition
in

very

much

reduced,

accordance with the peculiar regime, and this
is

deficiency

compensated by the development on
(hypapophyses) directed forwards, and

some

of the anterior thoracic vertebrae of long, tooth-

like processes

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
shell
its

contents are

lost,

the

being afterwards rejected through the mouth in

the form of a pellet.
53

54

INTRODUCTION
With the exception
of the worm-like Typhlopidae,

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

Viperidse, the teeth in the

jaws as well as on the
series;

palate

form

single

longitudinal

they are

elongate, conical, with or without a sharp posterior

edge,

more or
function,
is

less recurved, acutely pointed,

some-

times needle-like, and directed backwards, as behoves
their

which, in addition to

attack and

defence,

to prevent the retrogression of the prey

in the act of

prehension and deglutition.

A

notable

exception occurs in the genus Iguanognathiis, from

Sumatra,

all

the

teeth

having

spatulate

crowns

ribbed along the outer side.

Unfortunately, nothing

known The teeth
is

as to the food of this remarkable snake.

are coated with a thin layer of enamel.

It

was

held, for a time, that the glossy outer coating

only due to a denser structure of the dentine.
all living

was As in

Reptiles with the exception of the Crocodiles,
are

the teeth

not implanted in true sockets, but

simply ankylosed to the bone on which, when detached, 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
lost,

or feeding,

some of the teeth are frequently

and

are readily replaced by others lying in reserve in the

DENTITION
gum
at the inner side,

55
fixed to the

and becoming

bone

soon after a vacancy occurs.
so that in a snake like our

Such replacement

teeth,

of different grades of development, form several series,

mouth may contain

four

common Tropidonotus times as many teeth as

the
are

functional, without reckoning different earlier stages

of tooth germs which escape ordinary observation,

being placed vertically one above the other.

Three types of
mediate
step,

teeth,

connected by every inter:

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
It varies in

wound

the secretion of a poison gland.

depth according to the species, and
ing glass.

may

be so slight

as to escape detection without a very strong magnify-

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
is

condition the so-called ''perforated" fang

derived

through the complete fusion of the borders of the
tooth,
at

and the

obliteration of the line of union except

each extremity.

The

structure of such a fang

may
all

be best understood by imagining a tooth, lined

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
line in
is

median

such a way that the inner wall of
reality

the tooth

in

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
less

in

more or
pean

some cases may likewise show a distinct groove. Such may also be

present on the teeth of the lower jaw, as in the EuroCcelopeltis, in
is

some specimens of which a

faint

groove

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
erectile maxillary bone,

much

abbreviated and
teeth.

which bears no other

The Proteroglyphs

(Cobras, Coral-snakes, Sea-snakes)

andthe Solenoglyphs (Vipers, Pit-vipers, Rattlesnakes)

may be regarded as the diverging extremes in the development of the poison apparatus, both culminating in forms with tubular fangs, the former as derived directly from the

Aglyphs (harmless snakes),
insertion of the poison
really
is

the latter from the Opisthoglyphs, Hkewise evolved out
of the Aglyphs.

That the
is

fangs of the Viperidae

on the posterior
evident from the
position,
in

extremity of the maxillary bone
condition
of the

bone

in

its

recumbent

especially in the

African

Viper, Caiisus, which

DENTITION
brid type than our

57

several respects departs less markedly from the Colu-

European Vipers.
in

The poison

fangs of the Viperidae appear to be

movable, folding
This, however,

the

erected, or even thrust
is

mouth when at rest, and forward, when ready to act.

simply due to the mobility of the

maxillary bone, to which they are ankylosed as in
all

other snakes.

There are normally two equallyand
side by side, to

developed fangs, close together

each maxillary, followed by several replacement fangs
loosely attached behind them, usually in
four.

two

series of

When

the two fangs are

in situ,

they of course

both function
is

in the act of biting,

although only one
;

in relation

with the single poison duct
only one fang

often,

howthe

ever, there

is

in position, either

right or the

left,

the place of the other being indifilled

cated by a shallow socket which will soon be

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
bone

dentition of the snakes in which the maxillary
is

not
:

movable

vertically

falls

under three
all solid

divisions

the Aglyphs, in which the teeth are

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
principal

canaHculated teeth are situated in front, followed or
not by solid teeth.

Beyond these three

divisions, the dentition furnishes
for the classification,

important characters

although that importance has

sometimes been over-estimated.

The

maxillary teeth

maybe

equal in length (Isodonts), or the anterior the

longer (Lycodonts), or the posterior the longer, increasing gradually in size (Coryphodonts) or abruptly,

without (Syncranterians) or with a diastema, or break,
in front of are,

them (Diacranterians).
in

These categories
to

however, so completely connected as to preclude
use

their

taxonomy beyond helping
of maxillary teeth

define

genera.
relative

and the proportions and disposition of the mandibas follows,

The number

ular teeth also afford useful generic characters.

The European genera may be arranged
according to the dentition
I.
:

Teeth few, disposed
upper jaw only

in a transverse series in the

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
2.

Eryx,

Maxillary teeth equal, or increasing in size
posteriorly.

DENTITION
a.

59
;

Mandibular teeth 17 to 30
15 to 22.

maxillary teeth

Posterior maxillary teeth longest

;

mandibular teeth
Tropidonotus.

subequal, more than 20
Posterior maxillary teeth longest
;

mandibular teeth
Zamenis.

not more than 20, posterior smallest
Maxillary teeth subequal
posterior smallest
b.
;

mandibular teeth 20 to 25,
Coluber.
15,

Mandibular teeth 14 or
lary teeth 12 to 15.

subequal

;

maxil-

Maxillary teeth increasing in size Maxillary teeth subequal
B.

Coronella.

Coniia.

One

or

two enlarged grooved fangs behind the

series of solid maxillary teeth.

14 to 17 subequal sohd maxillary teeth, forming a

continuous

series

;

21

to

23

mandibular teeth,
Ccelopeltis.
fifth

anterior strongly enlarged

9 to II soHd maxillary teeth, fourth and

or fifth

and sixth enlarged, followed by an interspace
sixth

mandibular tooth fang-like, followed by an
Macroprotodon.

interspace

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

mandibular teeth strongly enlarged
C. Maxillary

Tarbophis.

with only two large canaliculated

fangs side by side, one of which
ing
;

may

be miss-

anterior mandibular teeth longest
Vipera, Ancistrodon.

6o

INTRODUCTION
In counting the teeth for the purpose of using this

number, more are missing but their place is indicated by the shallow pits in which their base was implanted, the overkey, care
it
;

must be taken

to ascertain the full

as

frequently happens that one or

FiG.

12

Maxillary and Mandible of
monspessulana
;

(a)

TarbopJiis
cucullatua.

fallax

{b) Coelopeltis

British

Museum

Macroprotodon Catalogue of Snakes)
(c)

(From

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

which are in reserve on the inner side of the jaws or behind the tubular fangs are not taken into
teeth

consideration,

the numbers

given

being those of

DENTITION
functional teeth only.

6

Although as a rule the teeth
on a specimen preserved
lips

can be counted
spirit,

easily,

in

and gums with the finger, it is sometimes necessary to remove and clean the bones of the jaws, an operation which
by simply pushing aside the
does not require

much

skill.

CHAPTER

VI

POISON APPARATUS— DIFFERENT KINDS OF POISONS

THE

gland which secretes the poison

is

a modifi-

cation of the parotid sahvary gland of other

Vertebrates, and

is

usually situated on each side of

the head below and behind the eye, invested in a

muscular sheath.
in

It is
is

provided with large alveoli

which the venom

stored before being conveyed

by a duct

to the base of the channelled or tubular
it is

fang through which

ejected.

In the Vipers, which furnish examples of the

most highly developed poison apparatus, although
inferior to
is

some in its toxic' effects, the poison gland and in intimate relation with the masseter or temporal muscle, consisting of two bands,
very large

the superior arising from behind the eye, the inferior

extending from the gland to the mandible.
to be powerfully wrung,
into the duct.

When

the snake bites, the jaws close up, causing the gland

and the poison pressed out

From

the anterior extremity of the

gland the duct passes, below the eye and above the
maxillary bone, where
orifice of the
it

makes a bend,
62

to the basal
(p.

poison fang, described above

55),

POISON APPARATUS

63

which is ensheathed in a thick fold of mucous By means of the membrane, the vagina dentis.
movable maxillary bone
prefrontal,
{supra, p. 49)

hinged to the

and connected with the tranverse bone set in action by the opening of the mouth, the tubular fang is erected
which
is

pushed forward by muscles

ff

^

e d

<:

a^

Fig. 13

Poison Apparatus of Rattlesnake: Venom Gland AND Muscles (Lateral View). (After Duvernoy)
;

a,

Venom

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

;

;

;

;

and the poison discharged through the
in

distal orifice

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
as the maxillary

bone

in these

snakes

elongate,

and not or but

slightly

and more or less movable vertically,
;

is

64

INTRODUCTION
its

the poison duct runs above the latter, making a bend

only at

anterior extremity,

and the tranverse bone

has not the same action on the erection of the fangs.

Otherwise the mechanism
the

is

the same.

In the Opisthoglyphous Colubrids, with grooved
teeth
situated
at

posterior

extremity of
of

the

maxilla,

a small

posterior

portion

the upper

labial or salivary

gland

is

converted into a poison-

secreting
colour,

organ, distinguished

by a

light

yellow

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
duct
is

;

the

not in direct connexion with the groove, but

the two communicate through the mediation of the
cavity enclosed by the folds of

mucous membrane
teeth,

surrounding the tooth, and united in front.

The

reserve

or

successional

which
are in

are

always present just behind or on the side of the
functional fang of
all

venomous snakes,
It

no

way connected with the duct until called upon to
replace a fang that has been lost.
otherwise,
since

could not be
a

the

duct

would

require

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
its

occupy exactly the same position as

predecessor.

Two
brids,

genera, Doliophis

among

the Elapine Colu-

and

Caiisus
for

among

the Viperids, are highly
its

remarkable

having the poison gland and

duct

POISON APPARATUS
body and terminating
of the
in front of the heart.

65

of a great length, extending along each side of the

Instead

muscles of the temporal region serving to
is

press out the poison into the duct, this action

performed by those of the side of the body.

When
the skin,

biting, a Viperid

snake merely

strikes, dis-

charging the venom the

moment

the fangs penetrate

and then immediately leaves go. A Proteroglyph or Opisthoglyph, on the contrary, closes
its

jaws

like a

dog on the part
is

bitten, often holding

on firmly

for a considerable time.

The

poison, which

mostly a clear limpid
colour,

fluid of

a pale straw or

amber

sometimes
matter,
is

with a

certain

exhausted after

more rarely greenish, amount of suspended 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.
that

The

fact

some

of these deadly snakes

when

irritated are

in the habit of shooting poison

from the mouth, at a

distance
at a

of

4 to
face,

8

feet,

even apparently aiming

man's

has been too often witnessed in

India and Malaya, and especially in Africa, from the

days of the ancient Egyptians,
subsist as to their being

for

any doubt to
this faculty,
is

endowed with

but the mechanism by which this action

produced
all

has not been satisfactorily explained.
bility,
5

In
the

probaof

the

poison

escapes

from

sheath

66

INTRODUCTION

mucous membrane surrounding the base of the fangs, and is mixed with ordinary saHva, the membranes of the mouth perhaps acting as lips, in which
case the term " spitting " would not be incorrect.
spitting,

The

which may take place three or four times
has been observed to be preceded by
If reaching

in succession,

some chewing movements of the jaws.
tion of the cornea

the eye, the poisonous fluid causes severe inflamma-

and conjunctiva, but no more washed away at once. Snake poisons is a subject which has always attracted much attention, and which has made great
serious results
if

progress within the last quarter of a century, especially as regards the defensive reaction

by which
effect

the blood

may

be rendered proof against their

by processes similar to vaccination
serotherapy.

—antipoisonous

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
supplying
their

anti-

dotes

;

that, in addition to the poison glands, snakes

possess other glands

blood with

substances antagonistic to the poison, such as also
exist in various

animals refractory to snake poison,

the

hedgehog
is

and

the

mungoose

for

instance.

Unfortunately, the specificity of the different snake
poisons

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
scutatus,

European
to

in

Australia
of

who had become immune
deadly
Notechis

the

poison

the

manipulating

these

snakes

with

was under the impression that
tended
also

to

other species,

and immunity exwhen bitten by a
impunity,
his

Denisonia super ba, an allied Elapine, died the following day.

In India, the serum prepared with the

venom

without

Naia tripudians has been found to be on the poison of Naia bungarus, the two species of Bungarus, and the Vipers Vipera Echis carinatus, and Lachesis graminetis. nisselli, Vipera russelli serum is without effect on Colubrine In venoms, and on those of Echis and Lachesis. Brazil, serum prepared with the venom of Lachesis
of
effect

lanceolatus

has

proved

to

be without

action

on

Crotaliis 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
India,

— at

least

as

regards

countries

like

where, different kinds of poisonous snakes
it

occurring together,

is

sometimes impossible to
for

know by which

the bite has been inflicted.

Chemistry teaches that snake venoms consist

the most part of solutions of modified proteids, and
all

attempts to separate the toxic principles from

68

INTRODUCTION
proteids

such

have

hitherto

been

unsuccessful.

Accordingly, at the present time

we must
in

regard
special

such toxic principles as residing

some

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,
is

Pseudechis, Notechis, Acanthophis)

mainly on the

nervous system, respiratory paralysis being quickly

produced by bringing the poison into contact with
the
central
;

nervous
the

respiration

pain

mechanism which controls and local swelling which
Echis,
Lachesis,

follow a bite are not usually severe.

Viper

poison

{Viper a,

Crotalus)

acts more on the vascular system, bringing about

coagulation

of

the

blood
its

and

clotting

of

the

pulmonary
system
tion
is

arteries;

action

on
effect

the

nervous

not great, no individual groupof nerve cells

appears to be picked out, and the
is

upon
is

respira-

not so direct

;

the influence upon the circulation

explains the great depression which
of Viperine poisoning.
severe,

a

The

pain of the

symptom wound is

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
The
bite
is

69

immediately followed by local pain of a
;

burning character
discoloured,
prostration,
diarrhoea,
usual.
slight

the limb soon swells and becomes

and within one to three hours great
accompanied by vomiting, and often
in.

sets

Cold,

clammy

perspiration

is

The

pulse becomes extremely feeble,

and
In

dyspncea and restlessness

may
in

be seen.

severe cases,

which occur mostly

children, the

pulse
cold
;

may become
the patient

imperceptible and the extremities

may

pass into

coma.

In from

twelve to twenty-four hours these severe constitutional

symptoms

usually pass off; but in the
discoloration

mean-

time the swelling and

have spread

enormously.

The limb becomes phlegmonous, and
somewhat suddenly, but death may
severe

occasionally suppurates. Within a few days recovery
usually occurs
result

from

the

depression

or

from
are

the

secondary
infrequent

effects

of suppuration.
in

That cases of
not
is

death, in adults as well as
in

children,

some parts of the
all

Continent

mentioned

in the last chapter of this Introduction.

The

bite

of

the

Proteroglyphous Colubrids,

even of the smallest and gentlest, such as the Blaps
or Coral-snakes,
is,

so far as

known, deadly

to

man.

The

Viperidse differ

much among

themselves in the

venom. Some, such as the Indian Viper a russelli and Echis carinatuSy the American Ancistrodon, Crotalus, Lachesis mtitus and lanceolatus,
toxicity of their

the African Causus, Bitis, and Cerastes, cause fatal

70
results unless a

INTRODUCTION
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.
bite of the

The

larger

European Vipers may be very
results, especially

dangerous, and followed by fatal
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

although very

seem to be possessed of a very virulent poison, and, common in some parts of AustriaHungary,
is

not

known

to have ever caused a serious

accident.
It is
is

noteworthy that the

size of the poison fangs

no relation to the virulence of the venom. The comparatively innocent Indo-Malay Lachesis
in

alluded to above have enormous fangs, whilst the
smallest fangs are found in the most justly dreaded

of

all

snakes, the Hydrophids.
is

Little

known

of the physiology of the poison of

the Opisthoglyphous Colubrids, except that in most
cases
it

approximates to that of the Proteroglyphs.

Experiments
rhinuSf

on

Ccelopeltis,

Psammophis,

TrimeroDryophis,

Dipsadomorphus,
Hypsirhina,

Trimorphodon,

Tarbophis,

and

Cerberus,

have shown
rapidly

these snakes to be possessed of a specific poison,

small

mammals,

lizards,

or
in

fish,

being

paralyzed and succumbing

a very short time,

whilst others {Eteirodipsas, Ithycyphiis) do not

seem

POISONS
to be appreciably
easily affected
least in

71
it is

venomous.

Man,

true, is

not

by the

bite of these snakes, since, at

most of those which have a long maxillary bone, the grooved fangs are placed too far back to
inflict

a

wound under ordinary

circumstances.
case

There

are,

however, exceptions.

A

was reported a
Dispholidtis

few years ago of a

man

in

South Africa nearly dying

as a result of the bite of the
typus,

Boomslang,

the symptoms, carefully recorded, being those

characteristic of Viperine poisoning, an
fact

important

to

oppose to the conclusions, based on the
experiments

physiological

on
from
the

Ccelopeltis,

which

appeared to disprove the theory that the Viperidse

may

have

been

derived

Opisthoglyphous
secretion

Colubrids.

Experiments made with
parotid gland of

of

the

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 harmless

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
brazili,

Coronella getiila

and the Brazilian Rhachidelus

72

INTRODUCTION
same districts, and which they are able overpower and feed upon. The Cribo, Spilotes
is

are proof against the poison of the Crotalines which

frequent the
to

variabilis^

the
it

enemy

of the Fer-de-lance in St.

Lucia, and

is

said that in their encounters the

Cribo

is

invariably the victor.

Repeated experiSnake, Tropidonotus

ments have shown our
natrix, not to

Common

be affected by the bite of Viper a her us
being due to the presence, in the

and

V. aspis, this

blood

of

the

harmless snake, of toxic principles
parotid

secreted

by the

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 feedmg on snakes, are known
to be

immune

to an ordinary dose of snake poison

;

whether the pig may be considered so is still uncertain, although it is well known that, owing to its
subcutaneous layer of
impunity.
fat,
it

is

often bitten with
quercinus)

The Garden Dormouse (Myoxus
been

has

recently

added to the

list

of animals

refractory to Viper poison.

CHAPTER

VII

NERVOUS SYSTEM— SENSE ORGANS

THE

brain

consists of

optic lobes,

tory lobes

;

It small and of very oblong shape. smooth cerebral hemispheres, small a still smaller cerebellum, and long olfacthe pineal body is not accompanied by a
is

parietal organ.
is

The

spinal accessory cranial nerve
is

absent,

and the sympathetic system
(p.

but feebly

developed.

The

eyes have been noticed above

12).

When
indepen-

normally developed they are susceptible of a slight

movement under the transparent

disc, quite

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
is

olfactory organ proper

is little

developed, but

accompanied by an accessory organ, Jacobson's

organ, consisting of a pair of pediculate, cup-shaped

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
sacs,

the

mouth

just

in

front

of 73

the choanae.

As

this

74

INTRODUCTION
nerves,
its

organ, richly provided with

communicates

with the inside of the mouth,
smell the prey as
glutition.
it

function

may

be to

passes through previous to de-

Snakes cannot be credited with a keen
it

sense of smell, although undoubtedly guided by

during the nuptial period.
In the more thoroughly aquatic snakes, the nostril

may
trils

be closed,
tissue,

when

respiration

is

suspended, by a

spongy

which acts as a stopper, and such nosis

are called "valvular," although a valve
;

not, in

the strict sense, present

when

the animal breathes,

the nostril

is

opened by a compression, through special
In

muscles, of the cavernous tissue.

some Sand-

snakes the narial opening
crescentic
slit.

may

be reduced to a

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), wath a fibrous or cartilaginous pad at
the outer end, extends from the fenestra ovalis in the

cranium to the quadrate, but
ing forms the stapes
fenestra ovalis.
is

in the

degraded burrow-

a small bony plate closing the

With one exception {Eryx jacuhcs, which
Schreiber to lap like a lizard), the tongue
for

is

said

by

is

not used

drinking or for the prehension or gustation of
is

food, nor for hissing, but

a tactile organ protruded
It is

on any object the snake wishes to probe.

slender

and deeply

bifid at the end,

smooth, very protractile,

SENSE ORGANS
with

75

often quite to the length of the head, and furnished

many

sensory corpuscules.

It is
is

darted and

vibrated on the least excitement, and

usually looked

upon by the ignorant
it is it

as a

**

sting."

In most snakes
;

much pigmented, dark brown
flesh-coloured or bright

or black

in a

few
is

is

red.

The tongue
glottis

entirely retractile into a sheath

below the

and

opening

in front of

it

;

it

is

always withdrawn into

the sheath

when

the snake bites or feeds.
in

Other organs, which,

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 Colubridse. 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 headshields,

this

being particularly noticeable in

the

Typhlopidse.

The

large

and deep

pit situated

between the

nostril

and the eye

(loreal pit) in the Crotaline

Viperidse

—whence
of Mexico

the

name
is

Pit-vipers, or that of

"cuatro

naricas" which

bestowed on them by the Spaniards

is

divided into two chambers: an outer

76

INTRODUCTION
orifice,

with large external

and an

inner, rather

more
inner

posterior in position

and occupying an excavation on
maxillary bone.

the outer face of the
walls of these

The

chambers are very thin and membraminute opening
this partition is

nous, and form a partition separating the two, except
for the presence of a
;

stretched across the hollow of the maxillary bone like

the

membrane

of a drum,

and

is

supplied with bloodcells of

vessels

and nerves, the

latter

terminating in

variable form.

The

use of the organ, thus situated

at the base of the poison fang,

and therefore

in close
is

proximity to the sphincter of the poison duct,

still

unknown.
Several of the Boidse, such as Python and Corallus,

have deep

pits in

some

of the upper

and lower

labial
;

shields, or also

on each side of the
all

rostral shield

these problematic organs are in
sensory.

probability also

CHAPTER

VIII

VISCERA

IN

most snakes there

is

a very marked asymmetry

of the viscera

and
left.

their blood-supply, the organs

of the right side being anterior to, as well as larger

than, those of the

The

heart in most cases

is

situated between the

and the anterior fourth of the body it may be much farther back, beyond the anterior third, in Doliophis, Platurus, and some Viperidas and
anterior seventh

Amblycephalidae, in the middle in Chersydnis.

It is

of

rather elongate form, enclosed in a pericardium in

which
Three
off

it

lies

freely,

auricles,

and a

arteries

and has a sinus venosus, two by a septum. leave the ventricle, the pulmonary and
single ventricle divided

two systemic arches.

The

right systemic arch gives
in

the carotid artery, which

common

Grass-snake

two, or in others be

many snakes, the for instance, may branch into double from its origin. The
is

anterior abdominal vein

double

in

most snakes, some Boidse, and conveys blood from the
single in
liver.

ventral body-wall to the

The caudal

vein

is

continued as the renal portal.

Veins which have

been regarded as remains of the two posterior cardinal
77

78

INTRODUCTION
some
of the

of lower Vertebrates have been found in
Boidae.

The

bifurcate transverse processes of the vertebrae

at the limit

between the body and

tail

enclose the
or
less

lymph-hearts, which are large and

more

elongate, metamerically divided into several
bers, the right often

chammore developed than the left.
on each side of the trachea,
is

The thymus gland
line, close to

lies

near the heart, and the thyroid gland

in the

middle

the base of the carotid artery.
is

The trachea
complete

long,

and the tracheal rings may be

in front

plete throughout.

and incomplete behind, or incomThe 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
In most of the Boidse
left

nearly to the cloacal region.

there are two well-developed lungs, the

shorter

than the
front,

right.

The lung has highly

cellular walls in

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 Typhlopidse

some of the

Boidae, Colubridse,

and Viperidae, as well as in and Amblycephalidae, and provided with
air cells,

the posterior end or the greater part of the trachea

may have

its

wall enlarged

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
it is

79
it

not a prolongation of the true lung, nor does
left

represent the missing

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
to produce,

species oi Coluber {Pityophis), said

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 Epiglottuphis
is

has been proposed.

This epiglottis

a narrow, thin

flap, erect in front of the glottis;

it is

not hinged, and
to cover the

therefore not

capable of falling

down

opening of the windpipe during the process of swallowing, its function evidently

being to increase the sound
air

produced by the escape of the

from the windpipe.

The larynx
bands of

is

represented

by two longitudinal
;

cartilage, united

by transverse bands

it is

extremely long in some snakes {Leptognathus),

The
gate,

oesophagus, which

may

be extremely elon-

sometimes measuring almost one-third of the
sac-like
itself

digestive canal, passes into the tubular or

stomach, often with thickened walls, which

gradually or abruptly merges into the narrower intestine.

The windings

of the

small intestine are

connected by ligamentous

tissue,

and enclosed

in

8o
a

INTRODUCTION
common
sheath of peritoneum.
In
several

of

the Glyphodont Water-snakes (Homalopsinae and

Hydrophiinse), 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 reticulate appearance, followed farther

down by

transverse

and then longitudinal
intestine
at
is

folds.

In these snakes the

abruptly constricted behind the stomach,
shells are

which point the

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
is

contents reaching the stomach, the
in front of the latter, at the

oesophagus

narrowed

point where the tooth-like ventral processes of the
vertebrae project

in order to aid in this function, after
shell is rejected

and pierce the wall of the cesophagus which the broken

through the mouth.

The more

or less elongate, feebly-lobed kidneys are

placed in the posterior part of the body, often extending 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
The suprarenal bodies are narrow and
on the renal veins or on the vena cava
inferior.

8i
elongate, placed

The
which

ureters leave the hind ends of the kidneys,

and

open through the side-walls of the cloaca on a papilla
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
fifth to

is

usually long and narrow, measuring one-

one-fourth the length of the body, on the right

side of the alimentary canal,

commencing just behind
It is

the heart or farther back.
in Chersydrus.
It is

exceptionally short

sometimes divided by transverse
extremity
is

furrows.
left

Its posterior

bilobate,

and the

lobe usually extends beyond the right, although

the reverse has been observed in
gall-bladder,
its

some snakes. The which may be absent, is remarkable for distance from the liver. The pancreas, elongate
is

but comparatively small,
the
left

located near the spleen, on

side of the alimentary canal, at a considerable

distance from the liver.

into a

The peritoneal part of the body-cavity is subdivided number of spaces or coelomic compartments

enclosed in serous capsules
tino-genital, a gastric

viz.,

a posterior or intesside,

on the

left

and a pair
lobes.

round the

liver,

corresponding to

its

two

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
elongate.
mally,

genital glands are situated anterior to the

kidneys, the right extending farther forward

and often larger than the

left.

The

testes

are

The

vas deferens

is

closely folded proxi-

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
opens into the cloaca
;

is

above the rectum and

this vaginal

chamber may be

more or

less

completely divided into two.
are provided with a pair of intromittent

The males

organs, or hemipenes, each connected with one of

the caudal vertebrae by a muscle {retractor penis) which
often exceeds
it

in length.

drical or club-shaped

surface divided

into

These organs are cylinand hollow, with the inner numerous cavities and beset
also with hard spines, of

with

papillae,

and usually

which those towards the apex may be greatly developed, folded against the walls, and directed
towards the extremity.

Such spines are absent
82

in

ORGANS OF REPRODUCTION
the snakes

83
of

provided with claw-like rudiments

hind

limbs.

The

cavities

of

the

hemipenis are

connected by a branch with the dorsal artery, and
it

is

the organ

by a flow of blood into them that erection of Each hemipenis is is accomplished.

lodged in a cavity on each side of the base of the
tail;

when protruded
spines
vagina,

it

turns inside out, and the

inner surface becomes

the outer, the papillae and
to

erected
in the

serving

maintain

a

firm hold

from which the organ cannot be
It

withdrawn except by invagination.
penis
walls.

has been

observed that the presence of spines on the hemiis

associated

with
is

much tougher
along
its

vaginal
entire

The organ
the

grooved

length,

groove

being the stdcus

spermaticus,

which,

when

the edges of the two hemipenes meet,
fellow a canal to convey the
;

forms with

its

semen

into the oviduct

this sulcus

may

be bifurcate, as in

the Viperids and

some Colubrids.
tail,

Anal pockets, secretory organs on each side of the
vent and lodged in the base of the
females, to be the

seem, in
;

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
active during the breeding season.
is

more

A

Viper-

catcher in France

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
autumn.
full

again

at

the end

of

summer

or

in

After hibernation the testes of the males

are rather voluminous,

and the sperm-ducts are often

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
then

hers.

In the Vipers the bodies of the pairing
are

individuals

completely entwined.
bring the

The male
orifices

endeavours to

two anal

together,

and when he has succeeded

in getting the

female to distend her cloacal opening, the intromittent 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
tions

last

a whole day.

Several copula-

may

take place at intervals of a few days.

Many
season,

snakes are gregarious during the breeding

and great numbers of males have been seen
the
females,

wriggling round
coils

forming with
with

their

huge lumps or an entangled mass

like a ball.

The

more

or

less

prehensile

tail

which
in

thoroughly aquatic snakes, such as Hydrophis and
Acrochordus, are

provided,

is

no doubt of use

PAIRING
facilitating

85
it

the pairing,

when

has to take place
pair

in the water.

Our European Water-snakes

on

land.

During the rutting season a
base of the male's
tail

slight pressure

on the

the hemipenes, and so
spine of the reptile.

may cause the protrusion of may a violent blow on the
more

Thus, recently killed specimens

of our Adder, with the organs everted, have

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 which the hemipenes are

normally everted, have been described by zoologists,

who should have known

better, as

examples showing
their

external vestiges of limbs.

The spermatozoa soon make
oviducts,
in

way up

the

which the

ripe

ova have previously
after,

descended, or which gradually descend shortly
these ducts
are usually

becoming dilated

in consequence.

There

more eggs

in the right

than

in the left

oviduct, although the reverse has occasionally been

observed.

Some
their

snakes lay eggs shortly after impregnation,
;

or a few weeks later

in others the

young undergo
capsule,

development within the oviducts, each envelin
is

oped

a thin, transparent,
torn

membranous
or

which

immediately before

immediately

after parturition,

such species being termed " ovovi-

viparous."

Just before oviposition the female curves

86
the base of the
cloacal opening.

INTRODUCTION
tail

upwards, in order to extend the
eggs are
all

The

produced together,
fluid

usually at intervals of a few minutes, and generally

adhere to one another by means of a sticky
secreted

by the oviducts, thus forming a clump.
snakes the young are born in

In

ovoviviparous

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
is

or less developed
said to produce

embryos, and Coronella punctata
time required for the eggs of
thus

thin-shelled eggs which hatch in less than half the

under the most favourable circumstances.
ovoviviparity.

American congeners There is almost every degree between oviparity and
its

These two modes of parturition bear no
the natural affinities of snakes.
Coronella aiistriaca
is

relation to

Thus, the European
its

ovoviviparous, and
;

North

American congeners are oviparous
it is

whilst, curiously,

the inverse in the genus Tropidonotiis. It was long

believed to be an invariable rule for the Viperidse to

bring forth live young, the

from

this

well-known peculiarity, but

name Viper being derived 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 Hydrophiinae, are ovoviviparous,

and thus dispensed from going on land

for

parturition.

OVIPOSITION
The
yolk entirely
or, if
fills

87
;

the eggshell
it is

there

is

no

albumen,

any exists,

so

much reduced
of lime,

as to

easily escape observation.

The

eggshell in oviparous

species contains

a small

amount

and

is

not hard, but tough and parchment-like, white or

yellowish
surface
is

;

it

is

usually smooth, but in Pythons

its

studded with minute pores, and in the
constrictor
it

American Zamenis

is

rough,

as

if

sprinkled over with loose grains of

salt.

The shape
It

varies from a short oval to a long ellipse.

has

been observed

in

some snakes

that

the eggs, on

leaving the cloaca, are of an elongate shape, suggestive of a short cigar,

and immediately

after

assume

a more oval form.

After they have been laid, the
size, espe-

eggs absorb moisture and thus increase in
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 much according to the
a higher

young of one brood varies species, and also according to
or a larger size than smaller
species.

the age of the mother, large females usually producing

number and of specimens of the same
our Tropidonotus,
15
to

Our European
;

Zamenis, Coluber, and Coronella produce only 2 to 15

48

;

our Vipers, 3 to 22.
as the

Among
A bastor

exotics

we may mention,
up
to 47

most

prolific,

Bitis nasicornis,

young; Tropidonotus fasciatus,

erythrogr animus,
;

and Farancia abacura, 50
Vipera
russelli,

Lachesis lanceolatus, 60

63

;

Boa

con-

88
strictor,
;

INTRODUCTION
Pseudaspis Tropidonotus ordinatus, 78 64 Python molurus, nearly 100 eggs.
;

cana, 80

;

The
of

eggs are deposited in holes without any sort

nest,

under
cases
for

moss
of
it

or

decomposing
or
in

leaves,

in

accumulations
In

saw-dust,

manure-heaps.

many

has been observed that the female

remains

some time with her eggs or young, and

Pythons a sort of incubation takes place, the female remaining coiled in a spiral over the mass an increase of several of eggs for six to eight weeks
in the large
;

degrees in her temperature at that period has been
ascertained by experiments conducted with every
possible care, a remarkable
so-called
'*

fact in

the case of a

cold-blooded" animal.
reports of

The numerous
stantiated

young snakes seeking
evidence, and,

refuge in their mother's gullet have not been sub-

by satisfactory
it

scientific

although

is

perhaps wise to say that the question
it

remains an open one,

may

be mentioned that, in

Europe

at least, trained observers

who have devoted
abundant, have

special attention to the habits of Vipers, in districts

where these
never

reptiles are exceedingly

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.
is is

The embryo
before birth
it

closely coiled

up in a spiral. Just distinguished by a large, convex

DEVELOPMENT
tively

89

head, with large, prominent eyes, and a comparashort

body,

the scales and ventral
life.

shields

being
is

much

shorter than later in

The umbihcus

situated in the posterior part of the body, from

six to ten

times as far from the head as from the
after birth the

vent.
visible,

Long
and

umbilical

slit

remains

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
indistinct,

is

much
the

reduced, and

sometimes
species.

very

in

ovoviviparous
is

The
it,

function of the egg-tooth

to cut

through the tough eggshell.
has
left

This, after the
slits in
its

young

shows one or several
if

anterior

extremity, cut as clean as

with a sharp knife.
is

The
shed

egg-tooth becomes loose soon after birth, and

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
time
;

live

for

a

short

there are even records of a three-headed snake,

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
stated to have been seen at

immediately after birth resent
ing,

all

interference, hiss-

snapping, or puffing themselves up, after the

go

INTRODUCTION
their parents.

manner of
birth

The

first

shedding of the

outer coating of the epidermis follows soon after
;

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
aspis,

inter-

mediate between

Vipera

herus

and V.

and
been

between

V.

hems

and

V.

ammodytes,

have

assumed, with

much

probability, to be hybrids.

CHAPTER X
HABITS

SNAKES
mode

may
of

be

grouped, according to their
in
five

life,

principal

categories,

gradually merging into each other, or two of them

not infrequently found combined in one and the

same

species.

These categories are
Burrowing-snakes,

:

— Ground-snakes,
Tree- snakes, and

Sand-snakes,

Water-snakes.

Ground-snakes may be defined as living above
ground, and only occasionally climbing bushes or
entering
Coronella
type,

the

w^ater.

Among European
and
Zamenis

genera,
this

and Vipera are perfect examples of
Coluber

whilst

approach

the

Tree-snakes in often
trees.

ascending

bushes, or

even

Sand-snakes
sand, in

are

adapted for living

on

loose

which they seek concealment.

Such are
the

Lytorhynchus

and

some Psammophis

among

Colubridse, Cerastes

among
live

the Viperidae.

Eryx conand

nects this category with the next.

Burrowing-snakes
as in

chiefly underground,

often have the visual organ atrophied in consequence,

Typhlops

;

all

the Typhlopidse, Glauconiidse,
91

92

INTRODUCTION
this category; the Viperid
is

and Uropeltidae, belong to
Atractaspis
also a

burrowing type.
life

Tree-snakes spend the greater part of their

on

among the Boidae, Dendrophis and Dendraspis among the Colubridse, Athens and various species of Lachesis among the Viperida^, may be quoted as examples. Of Water-snakes, some are exclusively aquatic,
bushes or trees.
Coralhis
like

the

marine

Hydrophiinae

and

the

typical

Acrochordinas (Acyochordus, Chersydrus) and
lopsinas (Hipistes, Herpeton).

Homa-

Chersydrus and Hipistes

occur in the sea as well as in fresh water.
species of Tropidonotus {T. tessellatus
in Europe), as well as the

Many

and

T. viperinus

genera Helicops^ Grayia,
the Colubridae, Eunectes

Botdengerina, etc.,

among

among
Our

the Boidae, Ancistrodon piscivorus

among

the

Viperidae, are chiefly but not exclusively aquatic.

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
of the

an expression of the natural relationships of the
species, as

was believed by many systematists

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
from exclusively nocturnal, delighting to bask
sun,

93

are provided with such a contractile pupil, are far
in the

and pairing and feeding in the day-time. The Boidae 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
of

locomotion for the exceedingly elongate body
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
but

ribs

and the

corre;

sponding ventral shields afford a point of support
it

does not account for the rapid movements, as
darts like an arrow in pursuit of
its

when a snake

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,
several harmless snakes,

without the ribs having to play an essential part.

Not only the Cobras, but
are able
vertically,

to

raise the anterior

third

of the

body

when taking up

a threatening attitude in

94

INTRODUCTION
same time widen-

the presence of an enemy, at the

ing or inflating the region behind the head.

Most snakes can chmb, and in this case the ribs and ventral shields are of great assistance. The Treesnakes, usually characterized by a very slender, some-

times compressed, body, or by a prehensile
specially

tail,

are

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 uneven 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 Dendrophis)

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
It

the ground at an oblique angle, the body being kept
rigid the

whole time of the
in

''

flight."

has been
surface

observed

Chrysopelea
lateral keels,

that the ventral

between the
to

which may be compared
in

hinges,

can be drawn

and become
place.
is like is

deeply

concave, whilst at the same time a slight dorsoventral flattening of the
this

body takes and

During
in

muscular contraction the snake
longitudinally,

a piece of

bamboo bisected such a way as to

buoyed up

explain

its

parachute-like descent.

All snakes are able to swim,

and the more aquatic

HABITS
kinds

95

may spend
is

a few hours under the water.

A
in a

Python molunis

known

to

have remained alive

basket sunk for thirty-six hours in a river.

The

best

adapted

for aquatic Hfe are the

Hydrophiinse, or Sea-

which are quite helpless and their body is shore posteriorly, and the tail
snakes, 'most of
;

never leave the water, and

soon die when brought on

more or
oar-shaped.

less

compressed

Sea-weeds and
Algae have also

barnacles sometimes settle on them.

been observed growing on the fresh-water snake
Herpeton tentaculatum.

As regards
insects,

food, Burrowing-snakes, as well as a

few small Ground-snakes, subsist mostly on worms,

and myriopods and

;

Tree-snakes
;

on

lizards, fishes

frogs, birds

their eggs

Water-snakes on
the

and batrachians.

Among

other types,

some

show a
or

predilection for

mammals, others

for lizards

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
of a

(Ericuhis) having been found in the excrements

Boa

tnadagascariensis.

Even hard-shelled eggs
the
principal

and

molluscs

may

constitute

or

exclusive food of certain snakes.

Thus, Dasypeltis eats nothing but

birds* eggs, the

which are crushed in the contrivance mentioned above (p.
shells of

gullet,

80),

by a special and are soon

96
after rejected

INTRODUCTION
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
snails

subsist
shells

almost entirely on

and

slugs,

the

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 haie)

from Algeria.

Snakes which take large prey secure
to three

it

according

methods

:

By
in

catching

it

simply with the
it,

jaws, and immediately proceeding to swallow
in Tropidonotus

as

and
it

some

of the Constrictors
;

when
after

dealing with small animals

by constriction,
it

having seized
coils

with the jaws, crushing
killing
it

in the

of their body and thus
in

previous

to feeding, as

the

Boidae and Coluber; or by

poisoning,
result being

by a mere stroke with the fangs, the awaited before the meal is begun, as in
Other
first

most of the Viperidae.
proceed according to the

poisonous

snakes

method, the use of the
as are in the
little

venom being
and
habit

to reduce the struggles of the victim

to relax its muscles.

Such snakes
their
in

of previously

killing

prey show

reluctance to accept

dead food

confinement, a

HABITS
thing

97
refuse
to

which others

usually

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
large
as,

species

feed almost

exclusively

on other

snakes, and often
or

manage
little

to swallow individuals as

even a

larger than, themselves.

Examples
they are

are

known

of harmless

snakes showing

a predilection for dangerous species, to whose poison

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.
feed chiefly on lizards
also

Some that

and snakes will occasionally eat
but rarely frogs.

mammals, and

vice versa,

On

the other hand, European Vipers
selves to a

more varied bill of feed on mammals, birds, reptiles, batrachians, insects, and slugs, and they have even been observed to eat
voles

accommodate themfare, being known to

showing signs of putrefaction.
are able

The enormous prey which some snakes
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,
7

even after being somewhat

g8
crushed

INTRODUCTION
by
constriction,

very

much exceed

the

calibre of the snake.
is

A

Python molurus 17 feet long

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,
egg.

and a
feats

Dasypeltis of the

same

size a hen's

Such

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
as
sternal

(see above, p. 42), as well

by the mobility of the
apparatus,

ribs

and the absence of
the
great
dis-

together

with

tensibility of the skin.

When

a snake proceeds to
if it

dispose of a large prey, which,
bird,
is

be a
it

mammal
pulls

or

usually

seized

head-first,

itself

forward by alternate movements of the jaws, the
maxillary and the mandibular ramus of the one side,

and then of the
laterally, the

other, being extended anteriorly

and

snake at the same time producing an

abundant salivation which renders the prey very
slimy.

Several repeated alternate

movements
gullet,

of the

jaws bring the head of the prey to the
the muscles and ribs
sides of the

where

come

into play,

and the two

together.

jaws work no longer alternately, but When once in the oesophagus, the prey

progresses with

much

greater facility,

and usually
lasted half
is

reaches the stomach in a few minutes, whilst the
previous process of deglutition

may have

an hour.

While

this laborious operation

going

HABITS
on,

99
is

the breathing
to

of the

snake

not
:

impaired

owing

a remarkable contrivance

the trachea
its

can be protruded in such a manner as to bring opening outside the mouth.
In cases where the victim
is

eaten aHve, the snake
is

has to contend with

its

struggles, but retrogression

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;
are often
still

its

struggles

apparent when

it

has reached the
after

oesophagus.

Snakes when caught immediately

a meal are in the habit of disgorging their food, and
it

sometimes happens that a frog or toad
alive.

is

thus

vomited
bag.

An
it

instance

is

known

of a naturalist
it

having captured a Grass-snake and put

in a linen

On

opening

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
organs several

be swallowed in rapid succession, after which the

gorged snake

will allow its digestive

days, or even weeks, of repose.

A

large

Anaconda
is

in

the Paris Jardin des Plantes fed only thirty-six times
in the course of

seven years.

Digestion

usually

rapid in the small snakes, defecation taking place

twenty-four to forty-eight hours after the feeding

;

it

100
lasts

INTRODUCTION
much
in the

longer in the large Boas and Pythons.

Thus,

above-mentioned Anaconda
of birds are

it

has been

observed to take from nine to thirty-eight days.

Even the hardest bones

decomposed by

the gastric juices, but hairs, feathers, and horny productions, are passed with the excrements, sometimes

forming regular
tell,

balls.

It is in

most cases possible to
faeces, w^hat

from an inspection of the dried

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
is

decomposed bones.

As a

rule there

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
mals,
whilst

mam-

another,
;

only take birds

of the same species, will and many, although to all appearall

ances perfectly healthy, will persist in refusing
food,

suicide

and allow themselves to die of starvation 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 sehce nearly two years and a half, s. Boa madagascariensis four years and a month. A Viper a aspis was kept for three years without food and without losing its vicious
temper.

Specimens thus fasting do not, as a

rule,

HABITS
renew their epidermis, or do so but very
rarely.

loi

Our
all

Common Adder
in captivity.

can very seldom be induced to feed

Other snakes

may

rid

themselves of

shyness to the extent of taking food from the hand,
or

show such

appetite as to seize a prey immediately
in

on being released from the small box or bag

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.
there
is

Some

are said to be fond of milk, but
for the belief held

no foundation

by peasants,

that they enter sheds with

the object of sucking

milk from the cows, which would be a material impossibility
;

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

marked instincts. The least stupid and most easily tamed are the species of the genera Coluber and Coronella. 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
or educability, nor do they display any very
;

bite,

even

if

molested.
hisses

The Common
takes

Grass-snake,

for

instance,

loudly and

up a very

threatening attitude, or even pretends to snap with

102

INTRODUCTION
its
^

open mouth, but very seldom bites;
defensive action

principal

when caught
it

consists in voiding a
its

most repulsive secretion from
it

anal glands, which

evidently controls, as

ceases doing so

when
also

accustomed to being handled.
the
skin

The same snake
repulsive

produces, during the spring, an oily exudation from

which

has

the

same

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
in

is

flattened out like that of a

Cobra
at the

when

an attitude of defence, and he noticed that
snake to worry
it,

his dog, seizing the

foamed

mouth The
air

as

if

he had been biting a toad.
is

hissing

produced by the rapid expulsion of
kept shut at the
(see p. 79)

from the lungs through the trachea and the notch

at the

end of the mouth, which

is

time.

Snakes provided with an epiglottis

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.
by
friction

This sound

is

produced

between the
dis-

serrated keels of the lateral scales,

which are

posed obliquely with their tips directed downwards

and backwards
after the

;

the

noise

can even be repeated

death of the animal, by twisting the body
little

and thus rubbing or rasping these

saws against

HABITS
one another.
in the African

103

The same

thing probably takes place

genus Dasypeltis, in which we find a

similar arrangement of the scales, though to a less

degree.

The

best

known sounding apparatus
p.

is

that of the

Rattlesnakes, described on

20.

When
tail

alarmed,

these snakes gather the body in a few coils or roll

themselves up in a
centre,

spiral,

with the

erect in the

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,
sense, but

however, that the rattle must have been evolved and
perfected, not necessarily in a

Lamarckian

through the different steps by which evolution or
creation has proceeded
;

Natura non

fecit salttis, as

Linnaeus well said.

Many

suggestions have been

made

as to the use of the rattle.

One
sound

of

that the rattling resembles
locusts,

the

them is made by
birds;

and serves
it
it

to

decoy insect-eating

another, that

serves to call the sexes together.

Probably
keep

is

useful to the snake as a

warning to

which cannot serve as food, and thus prevents useless expenditure of venom, or even the breaking of the fangs. At any rate, it gives exoff disturbers

pression to the snake's excitement, as does the voice

104
in the case of

INTRODUCTION
many
other animals, and
it

it

seems

reasonable to suppose that
different purposes.

may

be applied to

With

the advent of man, this

means of

attracting attention

must tend

to the

more
it.

rapid extermination of the snakes which possess

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
If,

as does a Cobra,
is

and

pretends to strike, although

it

one of the few

snakes that never bite man.
the snake rolls on

however, this display

proves of no avail in frightening
its

away the
its

intruder,

back and opens

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

those

spasm and rapidly moves off. It is the opinion of who have most experience of this snake that
behaviour
is

this extraordinary

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 hcemachates) and of the Common Grass-snake have also been observed to
feign death.

The notion
attracting
it

that

snakes
it

fascinate to

their

prey,

or

reducing

immobility by a
is

mysterious power in their glittering eyes,

pure

HABITS
fable.

105

Animals placed
fright,

in a

cage with a snake evince

no particular
It is

and fly away when pursued, if not actually turning round to defend themselves.
even dangerous to
all

offer a

good-sized snake a

wild rat for food, as

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
first

and

coiled

together in a mass.

The

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. this operation the

A

few days previous to
sight impaired

snake
is

is

languid and abstains from

feeding;

its

skin

dull

and the
;

by

the opaque condition of the lid

a day or two before
of

moulting,

the

outer

stratum

the

epidermis

becomes

again

through this

and the eye clear, stratum becoming detached from the
transparent
it

subjacent tissue, until

is

pulled off in one piece,
against stones or bushes.

by the snake rubbing

itself

The

first

exuviation takes place very shortly after

birth.

Snakes are long-lived, although the limit of duration of
life is

not

known

in

any of them.
to

They grow

slowly,

and do not appear ever

reach sexual

maturity until the fourth year,

when they continue

increasing in size for a long period.

A

Python

retic-

io6
ulatus

INTRODUCTION
and an Ancistrodon
of
piscivorus are reported to
in captivity in

have lived twenty-one years

Paris.

The young

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.
life, and remarkable for 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

Snakes are tenacious of

the reflex

head endeavouring to

bite.

Accounts of decapitated
stories.

Rattlesnakes turning round and striking with their

bloody stumps are probably not snake

CHAPTER

XI

PARASITES

L
and

IKE

all

other animals, snakes are infested with

a multitude of vegetable

both external and internal.

and animal parasites, 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,
in the great majority of these (102) only a single
:

parasite

a tick, a haemogregarine, or
to the

some

intestinal

worm.

Owing

more frequent opportunity of
menagerie snakes have

dissecting them, the

common

yielded better records, notwithstanding the fact that

they usually lose

most of their parasites through
artificial

constant handling, prolonged fasting, and
surroundings. Thus, we have a
for the Indian Python mohcrus,
list

of thirteen species

species for the

Boa

constrictor.

and one of twenty-two But no systematic

search appears to have been attempted, save, perhaps,
in the case of

a few European species.
notice that
it

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

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
Dr. L.

parasites of snakes are here enumerated by

W. Sambon, in Arthropoda. — Two
the Ixodidse

systematic order.
families of the class Arach-

nida,

and the Linguatulidae, furnish
on snakes.

numerous species

parasitic

Of
of
latter

the Ticks (Ixodidae)

the

genera

we find, as a rule, species Amhlyomma and Apojtomma, the
almost
species
entirely

genus being

confined

to

Reptiles.
physalis

A

single

of the

genus

Hmnahas

(H, punctatay

Can. and Franz,
aspis.

1877)

been reported once from Vipera

A

few larval

forms found on various snakes have been reported

under the generic name Ixodes, but they probably
belong either to

Amhlyomma
lizards,

or

Aponomma,
like

The Ophidian mammals, birds,
be
in

Tick-parasites,

those

of

and tortoises, appear to many cases the means of transmission of
-

protozoal infections from snake to snake.

The Tongue worms

(Linguatulidae)

are,

with-

PARASITES
out doubt, of the greatest possible interest.
systematic
zoologists,
versy.

log

Their
to

position

has ever

been

a

puzzle

and even now is a matter of controThey have been looked upon as Hirudinea
(1765),

by Winsberg

Cestoda by Chabert (1787),

Acanthocephala by Humboldt (1808), Trematoda by
Rudolphi (1809), and Nematoda by Nordmann (1832).
It

their

was Van Beneden (1848) who first recognized Arthropod nature, but he placed them amongst

Schubart (1853) suggested that their proper position is amongst the Mites (Acarina), and
the Crustacea.

Leuckart (i860) adduced important anatomical and
embryological evidence
in

support
Railliet
in

of

this

view,

which was confirmed by

1883 and by

Sambon

in 1910.

No
species

less

than three out of the four genera of
far established

Linguatulids so
parasitic

are represented by

on snakes. Reighardia, and PorocephaluSy

They
is

are the genera

Railltetiella.

The genus
armillatuSy
regiuSf

Porocephalus
its

of

special

interest,

because some of

species, such

as Porocephalus

a parasite of African Pythons {Python
sebce)

P.

and
of

Puff-adders

(Bitis

arietans^

B. nasicornis, B. gahonica), and Porocephalus moniliformis,

a

parasite

Oriental

Pythons

{Python

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
moluruSf P. reticulatus), are, in their

no
terns,
first

INTRODUCTION
described, in
1861, by

De

Filippi.

In

igio

Sambon

included in this genus other similarly
monitors,

structured Linguatulids from crocodiles,

and snakes.

The genus
in

Raillietiella

was established by Sambon
{Raillietiella botdengeri) of

igio

for a

Linguatulid

the African Puff-adders {Bitis arietans, B. gabonica).

Amongst
the

the characters of this genus

is

one of

great structural and phylogenetic importance
position

viz.,

of

the female sexual

orifice

at

the

anterior end

of the
it is

abdomen, whilst

in

the other

known genera

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.
several species of
cephala),

— The early encysted

stages of

Thorn-headed worms (Acantho-

belonging to the family Echinorhynchidce,

have been reported from snakes belonging to very
different genera,

such as Boa, Tropidonotus, Zamenis,
Dipsadomorphus,
Oxyrhopus,
Vipera,

Drymobius,

Xenodon,

Erythrolamprus,
Lachesis.

Diemenia,
further

Naja,

Elaps,

Their

development

probably

occurs in ophiophagous birds.
oligacanthoides, Rud., the

Thus, Echinorhynchus

immature stages of which

occur encapsuled within the body cavity of Lachesis
lanceolatus

and other neotropical snakes, when adult

PARASITES
is

III

found attached to the intestinal mucosa of Milvus

bidentatus.

Nematoda.
far
lies

— The
from

roundworms
snakes

(Nematoda)
to

so

described

belong

the

fami-

Ascaridae, Strongylidse, Trichotrachelidae,

and

Filariidae.

Some

of the genera belonging to these

families,

such as Cucullanus, Neniatoxys^ Oxysoma,

are as yet represented

by a single species

in

a single

host

;

others, such as
.

A scaris,

Polydelphis, Heiercikis,

StrongyluSj

DiaphanocephaluSf

Pkysaloptera,

Tricholess

soma,

number already

several species

more or

widely distributed.

Eelworm
one;

infection (ascariasis)

is

very

common
is

in

snakes, and not infrequently the infection

a heavy

Sambon

twice

found
that

over

fifty

specimens

of Polydelphis in Puff-adders {Bitis arietans).

This

investigator has

shown

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.
one of the
livers

Thus, Redi was quite right
of
his

in

considering the immature, encysted forms found in

double-headed Asp

as

belonging to the same species of eelworm (Ascaris
cephaloptera) as that
its intestine.

which the snake harboured

in

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

some of these hatched under the microscope, and moved very actively in the preserv;

indeed,

ing fluid.

This

is

in

no way surprising, because
in

even after several years of preservation
solution the
{Ascaris

formalin

embryos of other species of eelworms equorum, A. marginata) have been found in

a living condition.

Trematoda.
so
far

—The

Flukes (Trematoda) of snakes,
:

described, belong to the following genera
Astiotrema,

Agamodistomtim,
tretus,

Brachylaimus,

Cotylo-

Dicroccelium, Diplodiscus, Distomay Halipegus,
Metorchis,

Lecithodendrium,

Opisthogonimus,

Opis-

thorchis, Plagiorchis, Saphedera, Telorchis, Tetracotyle,

Zeugorchis.

Cestoda.
Piestocystis,

— Save a few larval forms
Sparganum),
the

(Cysticercoides,

the

known
to

tapeworms
the

(Cestoda)

of

Ophidia

belong

genera

Bothridium and Proteocephahis,

Protozoa.

— Numerous species of Hsemogregarines
As a
rule the

have been described from snakes.
schizogonic

forms seen in the peripheral blood are sporonts, the
cycle

occurring in

the lungs.
;

The
they

sporonts do not greatly alter their host cells
are invariably doubled
thick capsule.
differentiation,

up within a more or less Some species show a marked sexual
others
not.

Trypanosomes,

Spiro-

echaudinnicBy

and Plasmodidce have also been de-

scribed from the blood of various snakes.

PARASITES
Within
the

113

alimentary tube have been

found

species of Triclwmonas

and Caryospora.
baciUi have been described

Bacteria.
in

—Acid-fast
Shurley,

tubercular lesions found in

snakes by Sibley,

Gibbs and Sambon.

Shattock,

Hausemann, and
so

The

so-called

" canker,"

which

frequently
is

develops in the oral cavity of captive snakes,

also

a bacterial disease, due to a specific bacterium of
thick, rod-shaped form.

LIST OF PARASITES HITHERTO RECORDED FROM EUROPEAN SNAKES

TROPIDONOTUS NATRIX,
ACANTHOCEPHALA.
Echinorhynchus
incequalisy

L.

Rudolphi.

Echinorhynchus polyacanthus, Creplin.

Nematoda.
Strongylus auricularisy Zeder.
Strongylus catanensis, Rizzo.

Trichosoma mingazzini, Rizzo.

Oxysoma brevicaudattim, Zeder.
Nematoxys commutatus, Rudolphi.
Ascaris cephaloptera, Rudolphi.

Trematoda.
Opisthorchis caudatum, Polonio.

Dicroccelium assula, Dujardin.
Diplodiscus conicum, Polonio.
Tetracotyle colubri, v. Linstow.
8

114

INTRODUCTION
Distoma acervocalciferum, Gastaldi.

Distoma allostomum, Diesing.
Distoma nematoides, Miihling.
Saphedera naja, Rudolphi.

Brachylaimus sig7tatum, Dujardin.
Telorchis ercolanii, Monticelli.

Lecithodendrium nigrovenosum^ Bellingham.
Plagiorchis mentulatus, Rudolphi.

Cestoda.
Ligula pancerif Polonio.

TROPIDONOTUS TESSELLATUS,
Nematoda.
Strongylus denudatuSf Rudolphi.

Laur.

Physaloptera ahhreviata, Rudolphi.
Physaloptera striata, v. Linstow.

Trematoda.
Plagiorchis mentulatus, Rudolphi.

TROPIDONOTUS VIPERINUS,
ACANTHOCEPHALA.
<

Latr.

Echinorhynchus lobianchii, Monticelli.

Trematoda.
Distoma allostomum, Diesing.
Opisthorchis caudatum, Polonio.
Telorchis ercolanii, Monticelli.

Astiotrema monticellii, Stossich.

Cestoda.
Ligula pancerii, Polonio.

PARASITES
Protozoa.
Hcemogregarina viperina.
Billet.

115

ZAMENIS GEMONENSIS,
ACANTHOCEPHALA.
Echinorhynchus
cincUtSf

Laur.

Rudolphi.

Echinorhynchus polyacanthis, Creplin.
Echinorhynchus heterorhynchusy Parona.

Nematoda.
Strongylus catanensis, Rizzo.

Filaria parvomticronata, Rizzo.

Trichosoma sonsinoi, Parona.

Trematoda.
Distoma subflavum, Sonsino.

<

Brachylaimtis baraldii, Sonsino.

Saphedera naja, Rudolphi.

Cestoda.
Cysticercns acanthotetvay Parona.
Cysticercoides rostratuSf Mingazzini.

COLUBER QUATUORLINEATUS,
ACANTHOCEPHALA.

Lacep.

Echinorhynchus oligacanthns, Rudolphi.

Nematoda.
Ascaris cephaloptera^ Rudolphi.

Trematoda.
Plagiorchis sauromates, Poirier.

Protozoa.
HcBmogregarinUf sp.

ii6

INTRODUCTION
COLUBER LONGISSIMUS,
Laur.

Protozoa.
HcBmogregarina
colubri,

Borner.
Laur.

CORONELLA AUSTRIACA,
Nematoda.

Tricheilonema megalochilum, Diesing.
Physaloptera colubri, Rudolphi.

Cestoda.
Piestocystis dithyriditim, Diesing.

Protozoa.
Monocercomonas cohibrorum, Hammersch.

CORONELLA GIRONDICA,
Protozoa.
Hcemogregarina
coronellce,

Baud.

Fran9a.

VIPERA BERUS,
Nematoda.

L.

Physaloptera dentata, v. Linstow.

Trematoda. Agamodistomum

viperc^, v.

Linstow.

Tetracotyle colubri, v. Linstow.

VIPERA
Arthropoda.

ASPIS,

L.

Hcemaphysalis punctata. Can.

&

Franz.

Acanthocephala.
Echinorhynchiis cinctus, Rudolphi.

PARASITES
Nematoda.
Ascaris cephaloptera, Rudolphi.

117

Diaphanocephahis

vipercBy

Rudolphi.

Protozoa.
Caryospora simplex^ Leger.

HcBmogregarina samboni, Giordano.

VIPERA AMMODYTES,
Nematoda.

L.

Ascaris ammodytis, Rudolphi.

Ascaris cephaloptera, Rudolphi.

CHAPTER
REPRESENTATIVES
of Iceland, Ireland,

XII

DISTRIBUTION
of the order Ophidia are

found over the whole world, with the exception

and
in

New

Zealand, between the

Northern limit of 67°
Asia (Vipera
ordinatus),
schotti).

berus),

Europe {Viper a herus), 60° in and 52° in America (Tropidonotus
limit of 44° {Philodryas

and the Southern

The

highest altitudes reached by

them

are

14,000 feet in the Himalayas {Tropidonotus baileyi),

9,700 feet in the Alps {Vipera
in the

aspis),

and g,ooo

feet

Andes {Liophis numerous between the

albiventris).

tropics,

They are most and the number of
North and South.
into

species gradually diminishes to the

For the purpose
principal groups,
families

of

showing the distribution of the
will follow the divisions

we

and subfamilies enumerated above

(p. 4).

TyphlopidcB,

— S.E. — S.
S.

Europe, S. Asia, Africa, AusTasmania), C. and S. America,
Asia

tralia (exclusive of

and

W.

Indies.
(as far

GlauconiidcB.

E. as Sind), Africa

(exclusive of Madagascar), C. America (extending into

the S. parts of N. America), S. America.
Pythomnce.

Asia,

Africa (exclusive of

Mada-

gascar), Australia (exclusive of Tasmania), C. America.
118

DISTRIBUTION
Boince.

119

— S.E.

Europe, C. and

S. Asia, N. Africa,

Madagascar,
Ily slides.

Mauritius,

W.

Polynesia,

S.W.

of

N. America, C. and S. America, and

W.

Indies.

— S.E. Asia, S. America. — India and Ceylon. XenopelUdcB, — S.E. Asia. Acrochordince. — S.E. Asia, C. America. —The whole range of Ophidia,
Uropeltidee.

Cohibrince.

except

Tasmania.
DasypeltincB.

Homalopsince.

— Africa (exclusive of Madagascar). — S.E. Asia, N. Australia. DipsadomorphincB, — S. Europe, C. and S. Asia,
America
into

Africa, Australia (exclusive of Tasmania), C.

(extending
S.

the

S.

parts

of

N.

America),

America.
ElachistodontincB.

HydrophiincB.
ElapincB.

— India. — Indian and Pacific Oceans.
Madagascar),
America),
the
S.

S. Asia, Africa (exclusive of

Australia and Tasmania, Fiji Islands, C. America

(extending
S.

into

parts

of

N.
S.

America.
AmhlycephalidcB.
Viperince,

— S.E. Asia, C. and — Europe, Asia, Africa
into

America.
of

(exclusive

Madagascar).
CroialincE,

— S.E. Europe, Asia, America.
which the world

The Zoogeographical Regions
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.
w^hat

Contrary to

we find
as
in

in dealing with the Tortoises, Australia

does not show any special affinity to South America,
and,
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 (Boidae).

There

is

a greater difference between the Snakes of
of

Eastern Asia than there is and those of North America, whilst in Lizards a primary distinction must be made between the Old World and the New. Southern

Europe and those

between the

latter

Asia east of Persia (the Oriental Region)

is

the great

Ophidian centre,

all

the groups mentioned above, with

the exception of the Dasypeltinse, having representatives within its limits,

and a

large

and very

distinct

family, the Uropeltidae, being confined to

it.

The

Pythoninse occur along with the Boinae, the Viperinge with the Crotalinse, and the Elapinse are represented

by varied forms, as they are also in Africa and still more in Australia, where they form the overwhelming 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
like

121

and Coluber, extend over the Europo-Asiatic and North American regions, but
Tropidonottis, Zamenis,

they are equally well represented in the Oriental.

The
is,

great difference between Madagascar

and Africa

as

we have

said, very striking.

sesses Boidas generically identical with those of

Madagascar posSouth

America, but otherwise only Typhlopidse, Colubrinae,

and Dipsadomorphinae

;

whilst in the greater part of

Africa the Boinse are replaced by the Pythoninse, and

the Glauconiidae, Elapinse, and Viperinse are generally distributed.

North America agrees with Asia

and South America in its Crotalinse, 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
cally different

all

generi-

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

sion to these facts.

would give expresSuch regions would necessarily
in dealing

be very different from those adopted

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
few words
as to

A

the

salient
is

characters

of

the European fauna, which

a poor one as

comsingle

pared with other parts of the world.
species
of the genera Typhlops

The

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 southeast, is also

an Asiatic type.
Coluber,

The genera
and

Tropido-

notus, ZameniSy

Coronella,

Contia, are

characteristic of the Northern Hemisphere,
first

and the

three are, besides, equally well represented in
;

the Oriental region
also

a few species of Tropidonotus are

found

in Africa

and Madagascar.
group,

Ccelopeltis,

Macroprotodon, and Tarbophis are the northern outposts
of

an

Afro -Indian

although, with

the exception of the third, exclusively confined to the circum-Mediterranean district.
is

The genus
and
in

Vipera

also represented in East Africa
is

Southern

Asia, but the species V. berus

essentially a northern

type, extending to the highest latitude reached

by

any snake, and ranging

all

over Northern Asia to the

Amur and

Sachalien.

greatest altitude at

The same species reaches the which any snake has been observed

on the northern side of the Alps
only

viz.,

g,ooo feet.

Of the twenty-eight species inhabiting Europe, Tropidonotus two are generally distributed 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
longisstmus.

123

The others may be
only
as

described as southern
east:

forms, two

ranging from west to

Zamenis gemonensis
remainder

and

Ccelopeltis

monspessulana

one of more central habitat:

Vipera ursinii.

The

may be

divided into two groups
of

—those of
viperimis,

more western, and those

more eastern

distribution.

To
dica,

the

first

group belong Tropidonotus

Zamenis

hippocrepis, Coluber scalariSy Coronella giron-

Macroprotodon cucullatus, Vipera aspis and Vipera
to the second, Typhlops vermicularis^

latastii;

Eryx

jaculus, Tropidonotus tessellatus,

Zamenis

dahlii, Colu-

ber quatuorlineahis, dione, leopardinus, Contia modesta,

T arbophis fallax 2ind
lebetina,

iberus,

Vipera renardi, armnodytesy

and Ancistrodon
is

halys.

A
V.

remarkable fact in the distribution of Eurothe altitudinal range of Vipera berus,
ursiftii.

pean Snakes
aspis,

and V.

The

first

being the northernin

most

snake,

generally

distributed

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 g,ooo feet, on the northern aspect
of the Alps, whilst F. aspis lives at altitudes below

5,000 feet

;

but on the southern aspect of the same
is

chain things are reversed, and V. berus

replaced

by V.

which reaches an altitude of 9,700 feet, whilst the former shows a tendency to abandon the
aspis,

mountains, and has established
ties in

itself in

a few locali-

the plain of North Italy.

Again, in France

124
V. berus
is

INTRODUCTION
the northern and V, aspis the southern
is

species, yet the latter

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
in the

France (Basses-Alpes), and
(up to 6,800 feet)
;

Balkan Peninsula
V. herus occurs

but

it is

restricted to the plain in

Lower Austria and Hungary, where
only in the mountains.

Only three species are entirely confined to Europe: Coluber scalaris, Vipera ursinii, and V. aspis.
which range outside Europe, the following occur both in Western Asia and in North
species

Of the

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. longissiimcs, C. leopardinus,

Coronella austriaca, Contia modesta, Tarbophis fallax,

T. iberusy

Vipera renardi,

V,

berus,

V.

aminodytes,

Ancistrodon halys.

In North Africa

Macroprotodon cucullatus.

DISTRIBUTION
In North- West Africa

125

Tropidonotus viperinus, Zamenis htppocrepis, Coronella
girondicaf Vipera latastii.

The
Europe

following

lists

will

help to elucidate

the

distribution of the snakes in the different parts of

I.

Scandinavia
65**).

1.

Tropidonotus natrix (as far north as
Coronella austriaca (as far north

2.
3.

as 63°).

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,
3.

and Dorsetshire).
Vipera berus.
III.

Belgium and Holland

1.

Tropidonotus natrix,
Coronella austriaca.

2.

3.

Vipera berus.

IV.
1.

Germany and Denmark
tessellatus

Tropidonotus natrix.

2.

Tropidonotus

(Middle

Rhine and

Moselle, Saxony).
3.

Coluber longissimus (Denmark, Schlangenbad,

Treves)

126
4.
5.

INTRODUCTION
Coronella aiistriaca.

Vipera herus. Vipera aspis (Black Forest, Lorraine).

6.

V.

France and Switzerland, exclusive of
TiCINO
Tropidonotus natrix,

1.

2.

Tropidonotus viperinus (as far north as South

Brittany and Fontainebleau).
3.

Zamenis gemonensis (south,

locally as far north

as the Sarthe
4.

and Aube).
north
as

Coluber longissimus (locally as far

South Brittany, South Normandy, and Fontainebleau).
5.

Coluber scalaris (Mediterranean Littoral),
Coronella austriaca.

6.

7.

Coronella girondica

(south

and west

as far

north as the Charente-Inferieure).
8.

Ccelopeltis

monspessulana

(Mediterranean Lit-

toral).
9.

Vipera ursinii (Basses-Alpes).
Vipera berus (as far south as the Loire basin,

10.

the Central Plateau, and the Alps).
11.

Vipera aspis (as far north as the Loire basin,

Fontainebleau, and Lorraine).

VI. Spain
1. 2.

and Portugal

Tropidonotus natrix,

Tropidonotus viperinus.

DISTRIBUTION
3. 4.
5.

127

Zamenis gemonensis (Catalonia). Zamenis hippocrepis (absent from the north).
Coluber longissimus (Andalucia).

6.

Coluber scalaris.
(north and north-west).

7. Coronella austriaca
8.

Coronella girondica.
Ccelopeltis monspessulana.

g.

10.

Macroprotodon

cucullatus

(centre

and south,

Baleares).
11.
12.

Vipera berus (north-west).

Vipera aspis (Pyrenees).
Vipera
latastii

13.

(absent from the north).

VII. Italy,
1.

with Ticino and Corsica

Tropidonotus natrix, Tropidonotus tessellatus (as far south as Naples

2.

absent from the islands).
3.

Tropidonotus

viperinus

(Liguria,

Piedmont,

Ticino, Corsica, Sardinia, Sicily).
4. 5.

Zamenis gemonensis,

Zamenis

hippocrepis (Sardinia).

6.
7.

Coluber quatuorlineatus (south and Sicily). Coluber longissimus (absent from Corsica).
Coluber leopardinus (south and Sicily).
Coronella
austriaca (absent

8.
9.

from Corsica and
from Corsica and

Sardinia).
10. Coronella girondica (absent

Sardinia).
11.
Sicily).
Ccelopeltis

monspessulana

(Western

Liguria,

128
12. 13.
14.

INTRODUCTION
Viper a ursinii (Abruzzi).

Vipera berus (the Continental part only).
Vipera
aspis

(absent

from

Corsica

and

Sardinia).
15.

Vipera ammodytes (Northern Venetia).

VIII. Austria-Hungary,

without Balkan

States
1. 2.

Tropidonotus natrix.

Tropidonotus

tessellatus.

3.

Zamenis gemonensis

(South

Tyrol, Littoral,

South Hungary).
4.
5.

Coluber quatuorlineatus (Istria).

Coluber longissimus.
Coluber leopardinus (Istria).

6.

7. Coronella austriaca.
8.

Coronella girondica (South Tyrol),
Ccelopeltis

g.

monspessulana (Istria).

10. Tarbophis fallax (Istria).
11.

Vipera ursinii (Lower Austria, Littoral,

Hun-

gary).
12. 13.

Vipera berus.

Vipera aspis (South Tyrol, Littoral).

14. Vipera

ammodytes

(South

Tyrol,

Styria,

Carinthia, Carniola, Littoral, South Hungary).

IX. Balkan Peninsula and Archipelago
I.

Typhlops vermicularis

(Greece, Turkey,

Bul-

garia).

DISTRIBUTION
2.
3.

129

Eryx jacuhis (Greece, Turkey, Roumania).
Tropidonotus natrix.

4. Tropidonotus tessellatus,
5.
6.
7.

Zamenis gemonensis.
Zamenis dahlii (coast
Coluber longissimus, Coluber leopardinus.
of the Adriatic, Greece).

Coluber quatuorlineatus,

8.
g.

10. Coronella austriaca.
11. Ccelopeltis monspessulana

(West Coast, Greece,
Coast,

and

islands).

12. Tarbophis

fallax

(West

Greece

and

islands, Constantinople).
13.

Vipera ursinii (Bulgaria, Bosnia, Herzegovina,

Montenegro).
14.

Vipera

berus

(Bosnia,

Herzegovina,

Rou-

mania).
15.
16.
17.

Vipera aspis (Bosnia).
Vipera ammodytes. Vipera lebetina (Cyclades).

X. Russia
1.

Tropidonotus natrix (as far north as 60°).
Tropidonotus tessellatus (south).

2.

3.
4.
5.

Zamenis gemonensis (south). Zamenis dahlii (Caucasus).
Coluber quatuorlineatus (south). Coluber dione (south, between Volga and Ural)

6.

7.

Coluber longissimus (south, Poland)
9

130
8.
g.

INTRODUCTION
Coluber leopardinus (Crimea).
Coronella austriaca (as far north as 57°).

10. Contia modesta (Caucasus).
11. 12.

Tarbophis iberus (Caucasus).
Viper a renardi (south). Viper a berus (as far north as 64°).

13.

14. Ancistrodon halys (south, between Volga and

Ural).

Without attempting anything
bibliography,

like

a

complete

we have compiled

a

list

of faunistic

works and papers dealing with the snakes of Europe

Europe
ScHREiBER, E.
Steinheil, F.
Tiere.
:

in

General
Zweite

Herpetologia Europsea.

Auflage, Jena, 1912, 8vo.
:

Die Europaeischen

Schlangen.

Kupferdrucktafeln nach Photographien der lebenden
Jena, 19 13, 4to. (in progress).

Great Britain
edit.,

Bell, T. A History of British Reptiles. 2nd 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.
mandie.
IV. Reptiles.

:

Faune de

la

Nor-

Paris, 1897, 8vo.

DISTRIBUTION
Lataste, F.
de
la
:

131

Essai d'une Faune Herp^tologique

Gironde.

Bordeaux, 1876, 8vo.

Martin, R., et Rollinat, R.: Vert6bres sauvages du Departement de I'lndre. Paris, 1894, 8vo.

Switzerland
Fatio,
III.

Faune des Vertebres de la Geneva and Reptiles et Batraciens.
V.
:

Suisse.

Basle,

1872, 8vo.

Spanish Peninsula
Bosca, E.
de
la
:

Catalogue des Reptiles

et

Amphibies

Peninsule Iberique et des lies Baleares (Bull.

Soc. Zool. France, 1880).

Italy

Bonaparte,
Italica.

C.

L.

:

Iconografia

della
fol.

Fauna
Italiani

II. Anlibi.
:

Rome, 1832-1841,

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

1891).

Germany
DiJRiGEN,
Reptilien.

B.

:

Deutschlands

Amphibien

und

Leydig, F.

Magdeburg, 1890-1897, 8vo. Ueber die Einheimischen Schlan:

gen (Abh. Senck. Ges.,

xiii.,

1883).

Austrian Empire

Werner,

F.

:

Die

Reptilien

und

Amphibien

Oesterreich-Ungarns
Vienna, 1897, 8vo.

und

der

Occupationslander.

132

INTRODUCTION
Greece

Bedriaga,

J.

DE

:

Die Amphibien und Reptilien
B.

Griechenlands (Bull. Soc. Nat. Mosc, 1881).

BoRY DE
fol.

St. Vincent,

J.

:

Expedition Scienti-

fique de Moree.
atlas.

Reptiles.

Paris, 1832-1835, 4to.,

Bulgaria

KowATCHEFF, W. T.
Bulgaria.
text.]

:

Herpetological Fauna of
I912,
8vo.

Philippopolis,

[Bulgarian

ROUMANIA
KiRiTZESCU, C.
tologique
(Bull. Soc.
:

Contribution a

la

Faune Herpeet

de

Roumanie.
Bucarest,

Sauriens
x.,

Ophidiens

Rom.

1901).

Russia
NiKOLSKY, A.: Herpetologia Rossica.
burg, 1905, 4to.
St. Peters-

[Russian
:

text.]

Strauch, a.
Reichs.

Die

Schlangen

des

Russischen

St. Petersburg, 1873, 4to.

CHAPTER

XIII

SNAKES IN RELATION TO MAN

UNDER
addition to

this head,

the question of poisonous
first

snakes naturally occupies the

place.

In

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
lish the fact that

mortality for which snake-bite
is

is

responsible in India

well known.

Statistics establives

an average of 20,000 human
:

are thus lost annually
for 191 1.

24,264

is

the

official

return

In Australia, where highly poisonous snakes

of various genera and species abound, the fatal cases
are likewise very numerous, though less in proportion than in
Africa.

South America, and no doubt also
lanceolatus,

in

In the small island of Martinique, the FerLachesis

de- Lance,

causes

every year

the death of about 100

human
small

creatures.

Though
per

numerous

in species, the

poisonous snakes of Ceylon
mortality

cause a comparatively

— 200

annum. Modern research has

resulted in the discovery of
133

134

INTRODUCTION
the serotherapic treatment.

the only effective antidote for snake-venom intoxication
:

An animal

that

has been treated over a length of time with the

venom

of a poisonous snake, such as a Cobra, yields
is

a serum which

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
King Cobra, the
In

for

dealing with the bite of the

Krait,

and the Echis Viper.

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.
fifty-three

Six hundred and
in less

specimens were thus purchased

than two years (1901-1903).
as

The

poison

is

utilized

for the preparation of Calmette's antivenine,

which,

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
In Brazil, where the

135

mated

at ig,20o per

at 4,800,

number of accidents is estiannum, and that of fatal cases over 2,000 snakes {Lachesis and Crotalus) are

brought annually to the Serotherapic Institute of
Batantan, in the province of S. Paolo, for the preparation of the antitoxic serum, which
is

given in
latest
dis-

exchange
tributed

for the snakes.

According to the

report of the Institute (igii),
:

two serums are
Lachesis
bite)

the anti-crotaline (for
(for

Rattlesnake bite)
;

and the anti-bothropine
third, the anti-elapine

the

(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,
in

without, however, resulting

a very appreciable

diminution of the dangerous

reptiles.

More than

in India alone ;f 12,000 has been spent for this purpose

the numbers destroyed in 1885 and 1886 throughout
British India
tively.

amount to 420,044 and 417,596 respecAbout forty years ago the Governor of St.
alive

Lucia offered a reward of 4d. for every Fer-de-Lance's
head.

But the negroes caught them

families of snakes for the sake of the reward,

and bred 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
bents, is quite

Although the Adder, Vipera
in

common

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, however, on the Continent, where the same species, and especially its close ally, the more southern V. aspis,
parts of
are responsible for

many

many

fatalities,

due no doubt to
in a

the more virulent action of the
climate.

venom

warmer

In the French Departments Loire-Inferieure and

Vendee, where these snakes are very
i860 to 1868, 370 serious accidents to

plentiful, three

or four cases of death are reported annually.

From

man have been
all

carefully recorded, 53 ending in death, not only in

the case of children, but also of adults of
in 10 cases within

ages,

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
is

the other

hand, the bite of V.
inflicted, is
It

tirsinii,

which

but seldom

not

known
in

to have ever resulted in death.

must be borne

mind

that accidents are

much

more frequent

in districts

where the poorer classes

are in the habit of going about barefoot.

Anyhow, it
in

is

certain that Vipers are a serious danger

many

parts of Europe, not only to

man, but also
not surprising

to horses, cattle,

and dogs.

And

it is

SNAKES IN RELATION TO
that efforts have been

MAN

137

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.
offering 2^6.. per head, 500,000 Vipers {V. aspis)

By
were

destroyed from 1864 to 1890 in three French departments, Haute-Saone, Doubs, and Jura, and in one
district

(Chaumont) of the Haute-Marne 57,045 were
;

killed

from 1856 to 1861
In the

this gives

an idea of the

extraordinary abundance of these snakes in
parts of France.

was

fixed for a time at

some Puy-de-D6me the premium 5d,, and one man managed to

destroy in the course of seven years 9,175 Vipers
{V.

hems and F.

aspis).

A woman
many

in

the Deux-

Sevres has

made

a living for

years by catching

Vipers, the heads of which were paid to her at the

The average number of tures amounted to 2,062 per annum (mostly
rate of 5d. each.

her capV. 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. hems 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
fering

INTRODUCTION
with
live.
fire

the

natural

conditions under which

animals
tion

by

ground or destrucof the vegetation of the wilderness seems
Cultivation of the

to be the only efficacious

means

of getting rid of so

abundant and
Vipers

prolific a creature as the Viper.

A word may
:

be said, however, in

defence

of

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
It

to

keep

in check.

must be pointed out

that,

with the

exception of the species of Cohiher and Zamenis, other

European snakes are
lizards or frogs

to be regarded as indirectly

injurious to agriculture, feeding as they do mainly on

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 making shoes and fancy articles, such
pocket-books, blotters,
etc.,

utilized for

as purses,

and the Siamese make the drum-heads of native drums out of the skins of Pythons 3.nd A crochordtts. 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
Anguilles de hates, or hedge-eels.

name

of

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
remedies.

139

Some

forty years ago a chemist in Challans

(Vendee) collected Vipers (F. asph) 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
is

gradually fallen off since.

Very frequent
prevalent in

in the past,

snake-worship

still
is

many

parts of India, where the Cobra
is

held in great veneration, and

never willingly killed

by the Hindoo.

In pre-Buddhist days the gods
five

were represented with a canopy of

or

seven

Cobras over them.
represented on the
also an

The North

African Cobra was
is

sacred to the ancient Egyptians, and

profusely
;

monuments and tombs

it

was

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 "vEsculapian 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
to

Roman

settlements, has

been

attributed

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
to the

INTRODUCTION
more commonplace European snake-catcher,
boasts of his immunity, and of his art of attract-

who

ing snakes by devices of which he has the secret.

The Libyan handed down

PsilHi

of

the

ancient

Romans have
in

their art to the present day,

performances are to be witnessed

and their most of the
all

towns of Egypt and Tunisia.
lands
is

But India above

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
it

raised anterior part of the body,
to the

seems to dance
of

music performed by the snake-man, people
it

believing

to be

charmed by the sounds
sitting

the

instrument.
in front of

However, anyone

on the ground
result

a Cobra, and swaying the body from side

to side as does the

man, can obtain the same

without the

.aid

of any sort of music.

is

The most puzzling thing about these performances how the man can thus play with impunity with so
It
is

deadly a snake.

a mistake to think that the

snake

is

rendered harmless through the poison fangs
is

having been extracted, although this subterfuge
frequently resorted
jugglers.

to

by the

less

accomplished
is

The immunity

of the snake-charmer

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
tion,

141

oculations of the venom, a process similar to vaccina-

which renders

his

blood proof

against

the

venom

of the particular species of snake, and that
for his performances.

one only, used

Another
charmers
in

deadly

snake
Africa

shown
is

by

the

snakeViper,

North

the Horned
of an

Cerastes cornutus.

The presence

erect spike
in

above the eye
this snake,

is,

however, not a constant character

and hornless specimens are made more formidable by spines of the hedgehog being
to look

inserted in the proper place;

the illusion

is

such
this

that even
trick.

naturalists

have been deceived by

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
from

tropical

countries.
reptiles,

various

birds,

These stones, extracted and mammals, are

calcareous concretions from the stomach or bladder,

sometimes composed of superphosphate of lime, sometimes of phosphate of

ammonia

or magnesia.

The

value of a bezoar stone being supposed to increase

with
In

its size,

the larger are sold in India at very high

prices.

many

places a popular belief prevails that such

stones are found in the heads of snakes.
Bucknill,

Mr.

J.

A.

now Attorney-General

at

Hong-Kong, who

spent

five

years in Cyprus, has informed the author

142

INTRODUCTION
is

that the Viper of the latter island, Vipera lebetina,

commonly

believed to contain a stone which,

when
this

applied to the bite of a poisonous snake, quickly
nullifies the effect
;

it is

also believed that,

when

stone

is

allowed to stand in a glass of water and the
drunk,
it

water

is

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 "
for a

which he had

lent
for

monetary consideration to the defendant

the promotion of his
nill's

manly

vigour,

and Mr. Buck;f lo

recollection

is

that the plaintiff recovered

for the loss.

SYSTEMATIC ACCOUNT OF THE SNAKES OF EUROPEFIRST Family:

TYPHLOPID^
;

SKULL compact, with short, toothless lower jaw,
without transverse bone
goid reduced and toothless
;

palatine

and ptery-

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
to

lower jaw.

Rudiments of a pelvic arch, reduced

a single bone.
cycloid scales
;
;

Body vermiform, covered with uniform
head small, not distinct from the
;

the

body mouth small, crescentic, inferior eyes under more or less transparent head-shields, sometimes
Worm-like, smooth, shiny snakes,
of subterranean habits, or found in

entirely hidden.

of small or very small size, the largest measuring
little

over 2

feet,

rotten trees, under stones, or in the saw-dust of sawmills
;

rarely appearing
is

on the surface except when

the ground

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
ern Australia.

TYPHLOPIDiE
One
species occurs in South-Eastern

Europe.

About 120 species are known.
Genus TYPHLOPS, Schneider

Head with
divided nasal.

large shields

;

nostril

in

a single or

Tail extremely short.

I.

Typhlops vermicularis, Merrem The Greek Blind-Snake

Form.
of the

— Slender, worm-like, the

greatest diameter

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.
distinguishable,

Eyes
spot

appearing as a

small black
circle;

surrounded
lateral.

by

an

unpigmented

nostrils

Head-Shields.
less

— Rostral about one-third, or

a

little

than one-third, the width of the head, extendits

ing on

upper surface nearly to the level of the

PLATE

I

TVPHLOFS \'ERMICULARIS
After Sordelli

£RYX JACULUS
After SoKielli

TYPHLOPS
eyes.

145
cleft

Nasal incompletely divided, the
labial.

pro-

ceeding from the second

Preocular present,

about as broad as the ocular, in contact with the

second and third

labials.

enlarged and subequal.
Scales.

Upper head-scales Four upper labials.

feebly

Coloration.

— Equal, 22 or 24 round the body. — Brown or yellowish-brown

above,

yellowish beneath.
Total Length.
is

— 10 inches.

A specimen

from Cyprus

reported to measure 14 inches.
Distribution.

— This
It is

species has long been

known

from Greece, the Ionian Islands, and the Grecian
Archipelago.

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
viz.,

South-Western Asia,
Habits.

Asia Minor, Syria, Cyprus,

Transcaucasia, Persia, Turkestan, and Afghanistan.

— Pretty

alert in its

movements,

this little

snake has considerable constricting powers, and coils

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
itself fast

grasped, as
ness.
Its

draws back with extraordinary quickfood probably consists mainly of earthit

worms and
dug out of
10

small insects.

Some

exotic species of

the genus are

known to

feed on termites, and are often

their nests.

Reprodtiction.

— No

observations have been

made

146
that
I

BOIBM
am
aware
of,

but, as

some

of the exotic species

of which

we know something more
it

lay large, elonis

gate eggs,
oviparous.

is

probable that this species also

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
in the lower jaw.
;

a coronoid element

Teeth

in

both jaws.

Vestiges of

pelvis

in males, in a claw-like

and hind limbs, usually terminating, at least horny spur on each side of

the vent.

This family contains, besides the gigantic Boas

and Pythons,
forms,

several small

more

or less burrowing
its

among which

the

genus Eryx,
belonging
to

only
sub-

European

representative,

the

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
Pacific.

The

confined to

some islands of the South European species is the eastern and southern countries of
habitat of the

the Mediterranean district.

This

very

varied

family,

including

terrestrial.

ERYX
arboreal, aquatic,
tively small
viz.,

147
is

and burrowing forms,

a compara-

one as regards the number of species,
of which one-third pertain to the

about

sixty,

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.
neck,
shield.

Head

small, not distinct

from

covered with small scales;

a large

rostral

Eye very
; ;

small, with vertical pupil.
;

Body
narrow.

cylindrical

scales small

ventral shields

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
small,

Form,
neck
;

— Stout.
Tail,

Head

not

distinct
;

from
eye

snout

projecting

beyond the mouth
;

directed upwards

and outwards

a feeble mental

groove.

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
row

very large and broad, with

angular horizontal edge, followed by a pair of internasals and a second
of

two or three small

shields,

148

BOIDiE

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
of scales.

9 to 12 upper labials,

second or third deepest, separated from the eye by

one or two
internasal

series

Nostril between

the

and two

nasals, the

anterior of which

sometimes fuses with the former.
series of scales

Two

or three
eye.

between the nasals and the

Fig. 15 (after Sordelli)

Scales.

— Smooth,

feebly keeled

on the posterior

part of the body and on the

40 to 51 rows. Ventrals narrow, occupying about one-third of the
tail, in

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 blackish markings, which may be very irregular or form a
single or alternating series of large blotches or cross-

bands on the back
these markings

;

the sides with smaller spots

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
the

;

a dark streak from the eye to the angle of
;

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.— 2i feet

is

the greatest length which this

snake

is

known

to attain.

Distribution.

— Originally
in

described

from

Lower

Egypt, and extending westwards to Algeria, this

Eryx has been found
also
in

Greece, in Corfu, in the
It

Cyclades, in Turkey, and in Roumania.

occurs

Asia Minor, in Transcaucasia, in Trans-

caspia, in

Northern Persia, and
at

in Syria.

It

has

been found

an altitude of 5,000

feet in Persia, to
allied

the west of Lake
{E. miliaris, Pallas),
this species, extends

Urmia.

A

closely

form

which has been confounded with
from Transcaspia to Turkestan

and Afghanistan.

The
is

reported occurrence of this snake in Bulgaria
'*

based on a specimen labelled

Bulgaria
species

(?)

" in the

Sofia University

Museum.

The

is

omitted

from KovatschefPs
Habits,
arid,

latest Hst of

Bulgarian Reptiles.
is

—This

diminutive Boid

a burrower in

sandy districts, morning or towards dusk;
rather heavy form,

appearing only early in the
it

is

as

a rule more
its

crepuscular than nocturnal.
it is

Notwithstanding

capable of very quick move-

150

COLUBRIDiE
its

ments, darting like an arrow upon
consists chiefly of small
constrictor, like all the

prey,

which

mammals and
members
its

lizards.

A

of the family to

which
it.

it

pertains,

it

crushes

prey before swallowing
it

If given several

mice

at a time,

will catch

and

kill

them

all in

succession before proceeding to feed.
in the

Specimens recently discovered
in

Danube Valley

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

dewdrops with its tongue. is a gentle snake, seldom attempting to bite. Egyptian jugglers are in the habit of implanting
it

is

said to lap

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,
many examples have been

this

snake

is

ovoviviparous, but, beyond this fact, nothing

appears to have been observed concerning the breeding habits, although
in captivity.

kept

Third Family:
Maxillary, palatine,

COLUBRIDiE
;

and pterygoid bones movable
pterygoid

transverse

bone present

extending
present,

to
at-

quadrate or mandible; supratemporal

tached scale-like to cranium, suspending quadrate
prefrontal

not

in

contact

with nasal

;

maxillary

horizontal, not

movable perpendicularly

to the trans-

COLUBRID.E
verse bone
;

151
in

no coronoid bone.

Teeth

both jaws.

No vestiges of pelvic arch. An enormous group, comprising the
of snakes.

great majority
:

Divided into three parallel series
all

A. Aglypha, with

the teeth solid.

B. Opisthoglypha, with one or more of the posterior maxillary teeth grooved.

C. Proteroglypha, with the anterior maxillary teeth

grooved or canaliculated.

The

third,

which
of the

is

not represented in Europe,

includes

some

most deadly snakes, such as the
etc.

Cobras, Kraits, Death-adders,

The European genera
two other
series
:

are thus distributed in the

Aglypha (CoLUBRiNiE)
Coluber, Coronella, Contia.

:

Tropidonohis,

Zamenis,

Opisthoglypha (Dipsadomorphin^)

:

Co^lopeltis,

M acroprotodon,
These
variety

Tarbophis.

genera
of

give

but

a
in

feeble
this

idea
family,

of

the

forms

included

which
which

comprises adaptations to every
snakes are
fitted.

mode of life

for

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

COLUBRm.E
Genus TROPIDONOTUS, Kuhl

Maxillary teeth

increasing

in

size
;

posteriorly.

Head more

or less distinct from neck

eye moderate
or

or rather small, with round pupil.
less elongate
;

Body more

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
in

of

which are represented

Europe
itatrix,

Tropidonotus

proper, with the

common

T.

and Nerodia,

Baird and Girard, with two closely related species
of

more thoroughly aquatic

habits, T. tessellatus

and

T. viperinus.

3.

Tropidonotus natrix, Linnaeus
;

{Matrix vulgaris, Laurenti

Coluber torquatus,

Lacepede)

The Grass-Snake,
Form.

or Ring-Snake

— Moderately
;

slender; snout short, obtuse,

not prominent

eyes and nostrils lateral, the former
Tail four to six

moderately large.
in the total length.

and a half times

Head-Shields.

— Rostral
as

broader than deep, visible

from above.

Nasal divided, very rarely semidivided.

Internasals at least

broad as long, trapezoid,
Frontal broader than

shorter than the prefrontals.

TROPIDONOTUS
as long as broad, as long as or a
its
little

153

the supraocular, once and one-third to once and a half
shorter than

distance from the end of the snout, shorter than
in

the parietals, not

contact with the preocular.

Loreal deeper than long.

One
four)

(rarely two) pre-

and three

(rarely

two or

postoculars.

Temor

porals 1-I-2.
eight), third

Upper
and fourth
five

labials

seven

(rarely six
fifth)

(or fourth

and

entering

the eye.
the

Four or

lower labials in contact with

anterior

chin -shields,

which
shorter

are
than

the posterior.
Scales with

two apical
pits, in

nine-

teen

rows,
Fig. 16 (after Sordelli)

strongly
keeled.

keeled on the body, of outer

row smooth or

faintly

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
olive, or

which we are acquainted.
Grey, bluish-grey,
bars on

brown, above, usually
upper
whitish
or

with black spots or narrow bars on the back, and
vertical

the sides

;

lip

154

COLUBRID.E
;

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 subline; the bright collar often
in large

triangular or crescentic blotches, which usually meet

on the median
faint, or

becomes
females

even entirely disappears,
first figure)
;

(Plate II.,

belly usually checkered black

and grey or white, more
spots, or entirely black.

rarely grey with small black
Iris

dark brown or reddish-

brown, with a golden
is

circle

round the

pupil.

This

the form found in England and Central Europe
in

and

some parts of Southern Europe.
Jersey,
in

In

the

Spanish

Peninsula,

and
is

in

Cyprus, the white or yellow
does usually the black

collar,

which

always

present in the very young, soon disappears, and so
collar,

which
(var.

is

either

much

reduced
Seoane;.

or

entirely

absent

astreptophorus,

Some

large specimens from the Spanish

Peninsula are uniform ohve, without any markings.

Another variation (Plate
France, but

II.,

third figure), rare in

common

in Italy,

South- Eastern Europe,
;

and Asia Minor

(var. persa, Pallas

bilineatus, Bibr.

mtiromm, 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

in addition.

some specimens from Austria and Corfu

(var.

TROPIDONOTUS
suhfasciatusj

155

Werner) the belly

is

white, with black

bars occupying the free edge of each ventral shield.

A

very remarkable variety (var.
II.,

cettii,

Gene) from

Corsica and Sardinia (Plate

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

are often broken

them these annuli up on the middle line of the back,

and alternating; the collar is absent, or is transformed into the first annulus, and the upper surface
of the head
black.
is

more or
is
;

less spotted or

blotched with

This pattern

most

distinct in

young and

half-grown specimens

in large

examples the annuli
belly

may

break up into spots, disposed with great symin

metry

transverse series.

The

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 postocular shields, of an intense black, followed by the

usual yellow and black collar

;

two

light dots close

together on the parietal shields.

Some specimens
black.

are entirely or

nearly entirely

In the var. piduratus, Jan, from the Caucasus,
all

the upper parts are sprinkled
dots, and the yellow collar
is

over with light
;

present

the belly

is

grey, dotted with black,

the sides.

In others the body

and with white spots on is black above, and
beneath
(var.

checkered black and white

scutatus,

156

COLUBRID^

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
flesh-colour with reddish markings,

v^ith,

yellowish

yellow collar,
albino,

and a white or the eye and the tongue red. Such an
is

from Horsted Keynes, Sussex,

preserved
to

in the British

Museum.

A remarkable aberration,

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
is

British specimen on record, from Wales,

stated to measure 5 feet 10 inches.
feet.

Males rarely

exceed 3

Monstrosity.

—A

dicephalous young, with the two
side, is

well-formed heads side by
British

preserved in the

Museum, and
one

several

others
to

have

been
for

described,

being reported

have lived
occurs

about a month.
Distribution.

— Tropidonotus

natrix

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,

PLATE

11

TROl'IUONOTUS NATKIX
After SordelU

T.

NATRIX. VAR. CETl
AJter Sordelli

T.

NATRIX, VAR. I'ERSA

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
Africa
it

and Malta.

In

North

is

known from
where
it

Algeria and Tunisia, north

of the Atlas,

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.
but occur

The

melanistic

so-called varieties are not geographically restricted,
all

over the habitat of the species, though

not recorded from England.
Habits.

— Although
in

fond of water, and often seen

swimming

ponds or streams or creeping by the
is

water's edge, this snake

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.

says of the male of his

However, Gen6 Matrix cettii, " iracundum et

158

COLUBRID.E
Dr.

mordacissimum animal."
experience with
habited a
aggressive

Gadow

relates his

specimens

which

in-

swamp
them

with a

little

stream to the north

of Oporto, close to the coast.

To

his utter surprise,

some

of

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 motionless,

captivity,
;

Some specimens do well in 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
with open gape.
to produce the offensive odour

when handled.
and toads

The food
being
secretion,

consists of frogs

— the

latter

preferred

notwithstanding

their

poisonous

which protects them from the attacks of

most animals
tree-frogs,

—occasionally of newts, seldom of

fish

;

these snakes are reported to have a predilection for

and to feed occasionally on mice and
in the zoological scale
alive,

birds,

but most observers agree that they will not take

anything higher

than frogs.

The prey
sion
;

is

swallowed

and,

if

not very large,

four or five frogs or toads are often taken in succes-

a case

is

known

of a snake having swallowed

twenty very small frogs at one meal.
feed on

The young
in addition

worms and batrachian
and toads.

larvae,

to very small frogs

TROPIDONOTUS
The Grass-snake
It

159

gets
it

on
is

very well with the

Adder, to whose venom

immune.

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.
is

The hibernating season

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
later.

takes place in April or in

laid

May, according to the climate, and the eggs are between June and August, the young emerging
weeks
It is

six to ten

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.
feet long,

Females do not breed
little

until

about 2

males a

sooner.

The eggs
the

number

11

to

48, according to the size of

female, and, after being produced in a string, stick

together in a mass, without any regularity.

The eggs measure i to i^ 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

i6o

COLUBRID.E
at the

often approach farms during the period of oviposition.

Holes near baking ovens
are

back of village houses
resorts.

sometimes selected as breeding
rolls herself up,

The

and by violent contortions chamber in the manure, in which she may remain for some days after the eggs have
female

makes a

sort of

been produced.

It is

not very unusual for several the same
the
It

females to congregate for the purpose of laying, and
as

many as 1,200 eggs have been found in hole. The young on emerging has lost
bilical cord,

umoften

and measures 6

to 8J inches.

remains for a considerable time, sometimes until
the following spring, in the hole or manure-heap in
it was born, feeding principally on worms. Very young specimens are never found in the water.

which

4.

Tropidonotus tessellatus, Laurenti
{Coluber hydrus, Pallas)

The
Form.
narrow;

Tessellated Water-Snake
slender
;

— Rather

head

rather

long and

snout obtuse,

nostrils directed

not prominent; eyes and 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
Internasals

from above.

Nasal often semidivided.

usually as long as broad or longer, subtriangular,

truncate in front, as long or nearly as long as the

PLATE

III

TROl'IDONOTUS TESSELLATUS

'ROPIDONOTUS Vll'ERINUS
After SardHli

V.

VIPERINUS, VAR. AUROLINEATUS
After Sordelli

TROPIDONOTUS
prefrontals.

i6i

Frontal a

little

broader than the supradistance from the
parietals, not in

ocular, once

and a half
little

to twice as long as broad, as
its

long as or a

shorter than

end of the snout, shorter than the
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
or

1

+ 2.

Upper labials

eight (rarely

seven, nine,
ten),

fourth

or

fourth

and

fifth

(rarely third or
fifth)

entering

the eye.

Five

(rarely four)
lower labials in
contact with the
anterior
shields,

chin-

Fig. 17

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-shaped produced as a dark band on the nape, sometimes
a
distinct
II

more or less

1

62
streak
to

COUJERIBM
the frontal
shield
;

median
shields.

upper

lip

yellowish, with dark bars on the sutures between the

Lower

parts whitish,

yellow, orange,

or

red, marbled or checkered with black, or nearly

entirely black.

Iris

golden, bronzy, or coppery red.

Some specimens
coloration

depart very strikingly from the
defined.

thus

briefly

We

will

now

mention the principal variations which have been Sides of body checkered with black and described
:

yellow

or

black

and

red

(var.

rubro-maculosus,

Diirigen).

With

four dark stripes along the anterior
lineaticollis,

part of the back (var.

Werner).
to

Above

with four light
markings.

streaks

in

addition
light

the dark

concolor, Jan, hagenbecki,

brown above (var. Uniform black or The most blackish (var. nigrescens, De Betta). remarkable variety is the var. vosseleriy Werner, above with small black and from Asia Minor
Uniform grey or
Werner).
:

yellowish spots, beneath yellowish with three blackish
stripes beginning at

some distance from the head,
the outer
;

the median

much weaker than

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
several times in
var. flavescens,

163

Dalmatia, has been described as
Yellowish-white or brownishsmall
blackish
spots
;

Werner.
with

yellow

above,

belly

whitish in the middle, with a series of black spots,
bright yellow on the sides
Si^e.
;

eye and tongue red.

This snake occasionally reaches a length of
the British

4

feet,

but specimens over 3 feet are rare.

largest

specimen

in

The Museum measures

3 feet 10 inches.

Distribution.

— The

Tessellated Snake has a wide
It is

range in Europe and Asia.

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 w^est 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.
constitute the western limit of
It

Italy
its

and the Rhine
in

range

Europe.
altitude

does not ascend

to

any considerable
it is

in the

mountains of Europe, but

on record from
species than

6,000 feet elevation in Chitral.
Habits.

—This

is

a far

more aquatic
it

the preceding, being seldom found in

summer away

from the water,

in

which

swims and dives to

i64
perfection
;

COLUBRID^
which does not prevent it from being In accordance with these
it

equally agile on land.

thoroughly aquatic habits,

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
fish,

on the seashore near Odessa, chasing small
mostly gobies, in shallow water.
pairing take place on land, and
latter function
is
it

Hibernation and
is

not until the

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 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,
the purpose.

Herzegovina, the female laying
the time for oviposition.

her

eggs in the

following July, which with the beginning of August
is

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
5.

165
Latreille

Tropidonotus viperinus,

The Viperine Water-Snake
Form.
in

— Moderately
eyes

slender;

head shorter than

the preceding species;

snout obtuse, not prodirected

minent;

and
tail

nostrils

upwards and

outwards, the former rather small, the latter some-

what valvular;
length.

four to six times in the total

Head-Shields.— Rostra.1 broader than deep, visible

from above.
broad
or

Nasal

usually

semi-divided.

Inter-

nasals as long as
longer,

su bt riangular,
truncate in front,
as long as the prefrontals.

Frontal

usually

broader
supra-

than
ocular,

the

once
to

and
twice
Fig. 18 (after Sordelli)

a
as

half

long as broad, as
its

long

as

or

slightly

longer

than

distance from the end of the snout, shorter

than the parietals, not in contact with the preocular.
Loreal as deep as or a
or two preoculars
little

deeper than long.

One

and two + 2 or i +3. Upper labials seven (rarely eight), third and fourth (or third, fourth, or fourth and fifth) entering the eye. Four (rarely five) lower
(rarely three) postoculars.

Temporals

i

i66
labials

COLUBRIDJE
in

contact with the

anterior

chin-shields,

which are usually shorter than the
Scales with

posterior.

two apical

pits, in

twenty-one (rarely
Ventrals 147 to

nineteen or twenty-three) rows, strongly keeled, of
outer row smooth or feebly keeled.

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,

spots; a lateral series

of black
;

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
shields, or

bars

on the sutures between the
yellow spot on each shield.
red,

dark with a

Lower

parts yellow or
;

checkered with black, or entirely black

the

black of the belly
lateral

may

be connected with the ocellar
Iris

spots by black vertical bars.

golden,

often

mixed with brown.
specimen from Ponte
Carrega,
is

A

near Genoa,

preserved in the

Genoa Museum,
It
is

remarkable as

being of a dark olive-grey, with three series of black

and yellow
in

ocellar spots.
in

further exceptional

having the scales

nineteen rows.

A

second

specimen, from the same locality, with the normal

number

of scales, has

some

of the vertebral spots

TROPIDONOTUS
ocellar.

167

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

As
in

in

T. natrix,

there

occur,

in

the

South of

France, in Sardinia, in the Spanish Peninsula, and

North Africa, specimens with two

light yellow

or reddish lines along the back
figure), in addition to

(Plate III., third

the usual markings (C. auro-

lineatusy 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
it

that of T.

tessellatus,

but

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
presence
of
ocellar

however, the frequent

spots

on the
(147

sides,

to

and the low number of ventral shields 151), show that it should be referred to

T. viperinus.

Size.

— Rarely reaches a length of 3 feet in Europe,
specimens
being from
Sardinia.
is

the largest

An

Algerian specimen 3 feet 3 inches long
Distribution.

on record.
Southern
the

— France
Forest

as of

far

north

as

Brittany,

the

Fontainebleau, and

Department Aube, the whole of the Spanish Penin-

1 68

COLUBRIDJE
and the Balearic
Islands, Southern Switzer-

sula

land, north and south of the Alps, Liguria, Pied-

mont, Corsica, Sardinia, and
northern parts of the Sahara.
In
Liguria,

Sicily.

In Africa in

Morocco, Algeria, and Tunisia, penetrating into the
Piedmont, and Ticino, T. viperimis
fessellatus.

occurs alongside with T,

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
poles,

when

disturbed.

Frogs and toads, tadits

newts, fishes, and large earthworms, are

principal food

when
larvae,

adult, the

on batrachian

young feeding chiefly young fishes, and earthworms.

A

case

is

known

of this snake having eaten a waterfodiens).

shrew {Crossopus
caught,
fish
it is

When
;

a

fish

has been

usually eaten on land

in captivity

dead

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, 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
holes in

TROPIDONOTUS
spring.

169

being torpid, and resume activity very early in the

In

the

Alemtejo,

according

to

Gadow,

when

during the rainless and hot

summer

the small rivers
in great

have nearly dried up, these snakes collect
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 tv^enty-four hours,

apparently dying
suffocation.

from

some kind
observer
it

of

cutaneous
a

The same

once caught

Viperine Snake in a ditch whilst

was swallowing

an

eel of nearly its own length. Some specimens show so great

a superficial re-

semblance to the

Common
same

Adder,

Vipera
reptile,

hems,

which, however, being a more northern

very

seldom occurs
well

in the
its

localities

that this

snake

deserves

name

Viperinus.

A

celebrated

was once himself deceived by this resemblance and bitten by a Vipera berus which he had picked up in the Forest of Senart,
herpetologist, Constant Dumeril,

near Paris, believing
whilst, conversely, a

it

to be a Tropidonotus viperinus

;

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,
in the

and sometimes again

autumn

;

but the eggs

170

COLUBRID^
in

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

hatch

August, September, or October.

The

eggs,

numbering four
far

to twenty, are deposited in holes not
in

from water, often

abandoned
at

galleries of voles

or moles.
inches,

The young

birth

measure 4 to 6J

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
shields.

elongate,

distinct

from

neck

eye rather large, with round pupil.

One

or
;

more
scales

subocular

Body much elongate
pits.

smooth, with apical

Tail long.

The

species of this genus, about thirty in number,

are distributed over Europe, North Africa, Asia, and

North and Central America.
6.

Three inhabit Europe.

Zamenis gemonensis, Laurenti
;

(Coluber viridiflavusy Lacepede

C. atrovirens,

Shaw)

The European Whip-Snake
Form.

— Slender

;

snout

rounded,

with

distinct

canthus, moderately prominent,
side in front of the eye.

concave on each

Tail three and one-third
in the total length.
little

to four

and one-third times

Head-Shields.

— Rostral a

broader than deep,

the portion visible from above measuring one-fourth
to two-fifths its distance

from the

frontal.

Frontal

z

$

n-iKAivH

ZAMENIS
more or
less bell-shaped,

171
little

not or but

broader

than the supraocular, once and two-thirds to twice
as long as broad, as 'long as or a
its

little

longer than
a
little

distance from

the end of the snout,

shorter than the parietals.
or longer.

Loreal as long as deep

One

preocular (rarely two), extending to

the upper surface of the head, but never in contact

with the frontal
ocular
;

;

a small subocular below the pre(rarely three).

two postoculars 2 or 2 + 3 2 + (rarely i + 2).

Temporals

Upper
fifth

labials

eight, fourth

and

entering the

eye, fifth
seventh

and

deepest.

Five lower labials
(rarely
four)
in
Fig. 19 (after Sordelli)

contact with the
anterior chin-shields,

which are usually shorter than
pits,

the posterior.
Scales with

two apical
twenty-one)

in

nineteen (rarely

seventeen

or

rows.

Ventral shields

more

or less distinctly angulate laterally, i6o to 230

(usually under 200 in the typical form
caspius,
I go
;

and the

var.

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

172

COLUBRIDJE

with blackish cross-bars or numerous small black
spots, the black scales with a yellowish shaft, the

lower parts yellowish-white

dl"

pale yellow, rarely

more orange

;

the sides of the head are yellow, the

shields edged with blackish.

A

female, 3J feet long,
in

from Levico, Trentino, preserved

the

Genoa

Museum,
There
which
is

is

uniform reddish-brown above, with mere

traces of darker markings

on the head and nape.

every gradation between this form and the

var. viridiflavus or atrovirens (Plate IV., third figure),
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
scale
;

may

appear as a shaft along each dark
postocular
shields

the

preocular and

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
and the
In

a series of large black spots on each side.

Some specimens
var.

of both the typical form

viridiflavus are entirely black

or nearly black

(Z. carbonarius, Bonaparte; Z. sardiis, Suckow).

some
occur.

localities

and islands only black specimens
{irabalis, Pallas,

In the var. caspius, Iwan
first

Plate V.,
figure),

figure

;

persicus, Jan, Plate IV.,

second

from

Hungary, Bosnia,

Herzegovina, Corfu, Bui-

ZAMENIS
garia,

173

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
belly
is

the

uniform orange or red.
variety, var. asianus, Boettger,

Another
parts
light

from Asia

Minor, Rhodes, Cyprus, and Syria, has the upper

brown or
streak,

olive,

each scale with a longitudinal
are

and there

usually
;

large

black
is

spots relieved by yellowish shafts

the belly
is

red,

spotted or dotted with black.
in this form,

Melanism

frequent

such specimens being entirely black

except on the chin and throat, which are yellow
variegated with red.

The
(Plate

very young

of

the

typical

form,

as well

as that of the var. viridiflavus, has a striking livery
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
fol-

V- or W-shaped

line just

behind the parietals,

lowed by one or two others separating the dark cross-bars which may be present on the nape, and

174
occasionally

COLUBRID^
even
continue

some way down the
This livery persists
in

anterior part of the body.

some half-grown specimens. In young individuals from Syria
head
is

(var. asianus)

the

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, ii inches long,
;

from the Crimea, the head
cross-bars

is

olive-brown like the

body, which bears dark brown spots
;

and narrow
in

and there

is

a dark brown streak along
is

the middle of the nape, as
the typical form.

sometimes the case
is

The

belly

unspotted.
its

A young

from Malta

is

intermediate in

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
caspncs,
I

four principal forms

viridiflavuSy gemonensis,

and asianus

—are so completely connected that
to a length of
I

cannot regard them as more than geographical

races or varieties.
Sijse.

—This
of

handsome snake grows
feet.

6

feet,

the var. caspius even to 8
this

have seen a

specimen

variety,

from Salonica,

which

measures 7f

feet.

ZAMENIS
Distribution.

175

— From the

Atlantic coast of

Europe
its

to

South- Western Asia.

The

typical form, in

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

by the vars. caspius and asianus, of is represented which the distribution has been mentioned above. From Spain, this snake is only on record from Catalonia, not far

from the French

frontier.

Rare or
Aube,
is

local in the north of its range (Maine-et-

Loire, Vienne, Indre, Sarthe, Haute-Saone,
in

Yonne,
it

France, Switzerland north of the Alps),

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
feet in the

with are 3,900 feet in the Alps, 4,500

Balkan Peninsula.

Habits.

— The

name

*'

Whip-snake," under which
(Z. flagelli*'

an American representative of this genus
formis)
is

known,

like that of

Fouet " and " Lou-

which have been bestowed on it in some parts of France, expresses the quick movements with
cinglant,"

which,
slender

when
tail,

captured, this snake lashes
at the

its

long,

same time

furiously biting the

176

COLUBRID.E
it.

hand that has seized
also accounts for its

The

generic term Zamenis,
its

of Greek derivation, alludes to

viciousness,
*'

which

This snake, occurring the " Viper " which fastened on the hand of St. Paul.

German name, Zornnatter." in Malta, may well have been
kept for months in cap-

Some specimens have been
tivity

without losing their savage temper, hissing and

flying with

open mouth

at
;

anyone approaching the
others,

glass walls of their prison

on the other hand,
such as one

become
which
this
I

quite

tame

in a very short time,

kept for nearly two years.
itself
is

Except when
in the spring,

sunning

on a cold early morning
always on the

snake

capture, uncoiling itself

alert, and difficult to and darting away like an It lives in

arrow at the

least disturbance.

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
years within

for over

two

20 yards of the same spot, a bush

between a wood and a meadow, without ever being
able to capture
it.

The
voles

food of this snake

is

very varied, consisting of

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
it

lizards,

slow-worms, and snakes, which
Istria

does not
has even

attempt to crush before deglutition.
been observed in
c^gyptium) to
eat

It

locusts

{Acridium

and sphyngid moths.

PLATE V

ZAMENIS GEMONENSIS, VAF
After Sordelli

ZAMENIS DAHLI
After Sordelli

ZAMENIS HIPl'OCREFIS
A/ter Sordelli

ZAMENIS
Reproduction.

177
the end of June or
little

— Eggs,

laid at

beginning of July in a well-sheltered hole, are a
over twice as long as broad, and
1*4 inches in length.

measure
of eggs
is

i*2 to

The number

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 moderTail about one-third of the

ately prominent, obtuse.
total length.

Head-Shields.

— Rostral a

little

broader than deep,
little

just visible from above.

Frontal not or but

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
porals 2
eight

+

2 or 2

or
12

nine,

two postoculars. Tem+ 2). Upper labials fourth and fifth or fifth and sixth
it
;

+

3 (rarely i

178
entering the
eye.

COLUBRIDtE
Four or
five

lower

labials

in

contact with the anterior chin-shields, which are
shorter

than
with

the

posterior.
Scales

a
pit,

single

apical

very

narrow,

in

nineteen
Ventral

rows.
shields

very distinctly angulate
Fig. 20 (after Sordelli)

laterally,
;

205 to 218

anal

divided

;

subcaudals 98 to 132.

Coloration.

— Olive in front, with a few large black,
sometimes confluent with
collar, as in the
its

white- or yellow-edged spots on each side, the anterior
of which
is

fellow

and
tail

forms a nuchal
Plate

specimen figured on

V.

;

the greater part

of the
or

body and
reddish

uniform

pale olive, yellowish,

above,

yellowish-white beneath.

Head uniform

olive-brown

above, the labial, preocular, and postocular shields
yellowish-white.
Total Length.
Distribution.

rarely nearly 4 —3 — Southern Europe east of the Adriatic,
feet,
feet.

as far north as Dalmatia, Asia Minor, Cis- and Trans-

Caucasia, North- Western Persia, Cyprus, and Syria.

Has

also been recorded from

Habits.

—This

snake

is

Lower Egypt. even more lively than
It

Z. gemonensiSf and does not stand captivity long.

ZAMENIS
occasionally on locusts.
It

179

seeks dry, bushy localities, and feeds on small lizards,

does not seem to be very

common anywhere
dealers are imported.

in

Europe, except perhaps in

Dalmatia, whence most of the specimens sold by

Reproduction,

—The

pairing has been observed at
to

the end of May.

According

Werner, the eggs

number
J inch.

usually three only, measuring ij inches by

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 measuring

about
to
its

one

-

fourth

one - third
distance

from
-

the

frontal.
bell

Frontal

shaped, considerably
in front

broader

than the

Fig. aMafterSordelli)

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

i8o
parietals.

COLUBRID^
Loreal longer than deep, sometimes divided

into two.

One

preocular (sometimes divided into

two)

,

in contact

with the frontal
eye

;

two postoculars

;

a

series of three or four suboculars, usually completely

separating
2

the

from the
nine

labials.

Temporals
ten)

+

3

or

3

+

3.

Eight or

(rarely

upper

labials, fifth or

sixth very rarely entering the eye.

Four lower labials in contact with the anterior chinshields, which are shorter than the posterior.
Scales

with

two
rows,

apical

pits,

in

twenty-five to

twenty-nine

usually twenty-seven.

Ventral

shields very distinctly angulate laterally, 222 to 258

anal divided (rarely entire)
Coloration.

;

subcaudals yy to 107.

— 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.)
entirely black in the adult,

;

these spots

may become
as to reduce

and so large

the ground colour to a mere network or series of

X-shaped pale
the eyes, and

lines.

A

dark

cross-band between

a

y^-

or horseshoe-shaped

band on
often

the back of the head, which

may

be confluent with
light circle

an elongate spot on the nape; a

present in the middle between the parietal shields-

The

spots often

more or

less
tail.

confluent into three

longitudinal streaks on the

Yellow, orange, or

red beneath, with or without black dots, but constantly with a lateral series of black spots,

which

COLUBER
may
on the sides to form vertical bars.
Size.

i8i

be very large or unite with the spots higher up

— Examples

5 feet

long are on record

;

the

examined by me measures 4 feet 3 inches. Spain and Portugal, Sardinia, PanDoes not reach tellaria, Morocco, Algeria, Tunisia.
largest
Distvihutio7i.

the North of Spain nor penetrate into the Sahara.
Habits.

— This
its

very handsome snake

is

as a rule

as irascible as

European congeners.
it is

In Spain as

well as in Algeria

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
Reproduction.

as a useful

commensal of man, and
in a hole in a

deserving of protection.

— F.

Doumergue found
as large as pigeons'.

rock near Oran, in September, the recently-laid eggs,
five in

number and

Genus COLUBER, Linn^us
Maxillary teeth equal or nearly equal in length.

Head
large,

elongate, distinct from neck

;

eye moderately
;

Body more or less elongate 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
with round pupil.
scales

l82
nearly allied to

COLUBRID.E

Zamenk, 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, Lacep^de
(Elaphis cervone, Aldrovandi
;

Coluber

quatuorradiaUis, Gmelin)

Aldrovandi's Snake

Form.

Moderately
Tail

slender.

Snout

obtuse,

scarcely prominent.

one-sixth to one-fourth

of the total
length.

Head-Shields.

Rostral

broader
just

than
above.

deep,

visible from
Frontal

once and onefourth to once
Fig. 22 (after Sordelli)
j
V,

If

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
it.

below

One
2

preocular,
it
;

rarely

divided,

with

a subocular

below

Temporals

+3

or

3

two + 4.

or

three

postoculars.
labials

(exceptionally nine), fourth
sixth) entering the eye.

Upper and fifth Four or five

eight

(or fifth

and

(rarely three)

PLATE

VI

COLUBER QUATUORLINEATUS
Young, after Sordelli

COLUBER QUATUORLINEATUS
After IVerner

COLUBER QUATUORLINEATUS, VAR. SAUROMATES
After SordelU

COLUBER DIONE
After Sordelli

COLUBER
lower labials in
shields,

183
the anterior chinposterior.

contact

with

which are longer than the

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
five

;

anal divided

;

subcaudals 56 to 90.
VI., top) with three or

Coloration.

— Young (Plate
on
a

alternating longitudinal series of dark brown,

black-edged spots

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
the

Italy

and the
C.

countries

bordering

Adriatic (the typical

qiiatuorlineatus)

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
Pallas),

spots,

but

four-lined

(Plate VI., second figure).

In more eastern specimens

which may be regarded as representing the original form, the markings of the young persist throughout life, or, if they disappear,
satiroinates,

(C

they are not replaced by dark streaks (Plate VI.,
third
figure).

Lower

parts

pale

yellow,

closely

spotted or marbled

with

brown, these

markings

i84

COLUBRID^
tail.

usually disappearing in the adult, except on the
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 4I feet. Distribution. Aldrovandi's Snake inhabits Southern Italy and Sicily, Istria, Croatia, Dalmatia, Herzego-

vina,

Greece, and eastwards to Southern
It

Russia,

Transcaucasia, Asia Minor, and Persia.

has been

observed at an altitude of 2,600
All

feet in

Herzegovina.
Bulgaria,

the

specimens

from

Roumania,

Turkey, and eastwards, belong to the var. sauromates,

which

is

regarded by some authors as worthy of

specific rank.

The reported occurrence
France
is

of C. quatuor-

lineatus in various parts of

certainly due to

confusion with C. scalaris and C. longissimus.

marshy localities are the and handsome snake, which often approaches the dwellings of man, attracted by
Habits.

— Dry
this

as well as

abode of

large

the poultry.
it is

Comparatively slow

in its

movements,

more
it

easily captured than

any of the other large

Colubrids of Europe, and does well in captivity,

where
will

should be provided with a tank, in which
It is as

it

remain for hours under water.
as at climbing.

good

at

swimming
tured,
it

Biting readily

when

cap-

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 feeJs the

COLUBER
grasp
it

185
itself.

turns round and defends

It

appears

to feed exclusively on
size of a rat or dove,
It

mammals and

birds,

up to the

and

will readily take

dead food.

has a predilection for eggs, and has often been

observed to swallow hens' eggs.
Reprodtwtion.

— In
laid

Herzegovina
soon

pairing

takes

place from the middle of June to the middle of July,

and the eggs are
to sixteen,

after, to

hatch in Septem-

ber or beginning of October.

and measure

2 inches

The eggs number six by ij inches. The

young measure

8 to 14 inches at birth.

10.

Coluber dione,

Pallas

The Dione Snake
Form,
convex,

— Similar
a
little

to

the preceding.

Head more

narrower;

snout obtuse, scarcely

prominent.

Tail about one-fifth of the total length.

Head- Shields.
visible

— Rostral

broader

than

deep, just

from above.

Frontal once and one-fourth
its

to once

and a half as long as broad, as long as
Loreal as long as deep, or a
little

distance from the end of the snout, shorter than the
parietals.

longer

than deep.

A

large

preocular, with

a subocular

below
or

it,

the latter very exceptionally absent
postoculars.
eight
or

three

Temporals
nine
(very

2

+

3

two or 3 + 3.
;

Upper

labials

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
pits, in

smooth or

faintly keeled, with

two apical
Ven-

twenty-five or twenty-seven (rarely twentythree) rows.

tral shields not or

but very obtusely

angulate laterally,
172 to 214
divided
;

;

anal

subcau-

dals 50 to 80.

Coloration.
Pale
Fig. 23 (after Sordelli)


or

brown
-

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.
spotted

Lower

parts yellowish, usually dotted

or

with blackish.
S^'^^.— Seldom exceeds a length of 3 feet.
largest

The
37

specimen examined

by

me measures

inches.
Distribution,

—Across

temperate

Asia from Asia

Minor, Transcaucasia,
of the Caspian Sea, to

and the southern border the Amur, Corea, and China.

COLUBER
In Europe the habitat of this snake
to
is

187
restricted

the

steppes of

Southern Russia, between the

Caucasus
Habits,

and

the

Lower
is

Ural.

The specimen

figured on Plate

VL

from Sarepta, on the Volga,

ities,

snake frequents arid, sandy localand is only exceptionally found in small woods, Nothing more is known of its habits.
II.

— This

Coluber longissimus, Laurenti
Lacepede
;

{Coluber c^sculapii,

C. flavescens,

Gmelin)

The ^sculapian Snake
Form.

— Slender.

Snout obtuse, scarcely promiTail about one-fifth to one-

nent; head narrow.

fourth of the total length.
Head-Shields.
visible

— Rostral

broader

than deep, just

from above.
as

Frontal once and one-fourth

to

once and

one-third

long as broad,
as long as
its

distance from

the rostral or

the end of the
snout, shorter

than the parietals.

Fig. 24

Loreal

as long as deep or longer than deep,

One

pre- and
labials

two postoculars.

Temporals

2

+ 3.

Upper

i88

COLUBRlDiE

and fifth or fifth and sixth enterFour or five lower labials in contact with the anterior chin-shields, which are as long as
eight or nine, fourth

ing the eye.

or a

little

longer than the posterior.
feebly keeled on the posterior part

Scales

smooth or

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
stripes along the
lip,

;

sometimes
upper

with a yellowish vertebral stripe or with four darker

body

(var.

romamts, Suckow)

;

and often

also a triangular patch
;

on each side
less dis-

behind the temple, pale yellow
tinct

a

more or

dark band on the temple, and a vertical dark
first figure).

bar below the eye (Plate VII.,
parts uniform pale yellow.

Lower
figure)

Young (second

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
Iris

the forehead, and a black vertical line below the

eye

;

belly greyish or yellowish-olive.

dark grey

or brown.

Tongue pinkish-brown.
is

Melanism
are entirely

rare in this snake.

Such specimens

black above and beneath (var. niger,

PLATE

VII

COLUBER LONGISSIMUS
YoiDig, after Sordellt

COLUBER LEOPARDINUS
After Sordellt

C.

LEOPARDINUS, VAR. QU ADRILINEATUS
After Sordellt

COLUBER
Nikolsky), or blackish
-

189

grey to black above, dark

grey beneath
line

(var. stcbgriseus,

Werner), the angular
light.

on each side of the belly often remaining

An

albino found near Vienna has been described as

pale orange-yellow above, with small white spots

pupil

and tongue

red.
feet

Size.

are,

Specimens over 4J however, very rarely met with.
to 6 feet.

— Grows
part

Distribution.

— Generally

distributed

over

the

greater
Sicily,

of Austria,

Italy,

with Sardinia and
this

and the whole of South-Eastern Europe,
is

snake has a very broken range in France, Switzerland,

Germany, and
as

found, quite isolated, as far

north

Denmark and Poland.
it

According

to

Segerus, quoted by Lacepede,

used to be quite
the

common

near

Copenhagen
it

at

end

of

the
Its

eighteenth century, but

is

now much
in

rarer.

northern hmit in France

is

Southern Brittany,

the Department Orne, and the Forest of Fontaine-

Germany, Schlangenbad, near Wiesbaden, perhaps also Baden-Baden and Treves. It is on
bleau
;

in

record

from

Southern
in

Spain.

Its

discontinuous
its

distribution
in

Central

Europe, and

presence

various localities

near former
its

Roman

thermal

stations, has

been ascribed to

introduction from
erected
to

Italy

as

an

inmate
I

of

the

temples

iEsculapius; but
its

am more

inclined to look

upon

sporadic occurrence in the North as the indicaof

tion

a

once more

widely

distributed

species

igo

COLUBRID^
in

now
range.

process

of

extinction

over

part

of

its

In Asia the ^Esculapian Snake

is

only found in

Transcaucasia.
as in

It

occurs in the mountains as well

the plain,

being recorded

from

5,200

feet

altitude in the Tyrol, 3,200 feet in the Apennines.

Habits. The ^sculapian Snake lives in woods among shrubby vegetation in meadows, where it is
; ;

often found

under
It

haystacks

;

occasionally about

old walls.

climbs well, and often ascends trees.
it

Although a good swimmer,
water of
its

seldom enters the

own

accord.

It feeds chiefly

on small

mammals,
lizards.

occasionally on birds and their eggs, and
I

Specimens which

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
still

whom

they are accustomed, although
strangers.

resenting

the intrusion of

However,

this

snake

never becomes so thoroughly domesticated as the

Smooth Snake, and cannot be
devoted

trained to take food

from the hand, according to R. Rollinat, who has many years to experiments on the taming
This observer had no
difficulty in feed-

of reptiles.

ing his ^sculapian Snakes on mice and voles placed

dead

in their cage.
is

This snake
vole galleries

particularly sensitive to cold,

and
its

does not emerge until late in the spring from the

and hollow

trees

which constitute

COLUBER
winter-quarters.
It also

igi

avoids excessive heat, never

showing

itself

in

the

daytime during the hotter
takes

months

in the

South of Europe.

Reproduction,

— Pairing

place

between the

middle of
in walls or

May and

the middle of June.

The

eggs

are laid towards the end of June or in July, in holes

hollow trees, under moss, sometimes even dung-heaps of farms, and hatch in September. According to trustworthy observers, the eggs, which
in the

measure i J to
in

2 inches in length,
five

and

less

than

i

inch

width,

number only

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
5

the adult.
at

They measure about
bite.

inches,

and are

once most ready to
12.

Coluber leopardinus, Bonaparte
{Coluber quadrilineatuSy Pallas)

The Leopard Snake
Form.
nent.

— Slender.

Snout obtuse, scarcely promibroader than deep,
just

Tail about one-fifth of the total length.

Head-Shields.
visible

— Rostral

Frontal once and one-third to once and a half as long as broad, as long as its distance

from above.

from the end of the snout, shorter than the parietals. Loreal longer than deep. One pre- and two postoculars.

Temporals

i

+

2 or 2

+

3.

Upper

labials

192

COLUBRII)^

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

longer than the
posterior.
Scales smooth,

with two apical
pits, in

twenty-

five

or twenty-

seven

rows.
not
later-

Ventral shields

rounded,
Fig. 25 (after SordelH)

angulate
;

ally,

222 to 260

;

anal divided

subcaudals 68-go.

Coloration.

— Typical form (Plate VII., third figure)
brown above, with one dorsal
spots
series

greyish or pale

of dark brown, reddish-brown, or bright red, black-

edged transverse
centres

and a

lateral

alternating

series of smaller black spots
;

with or without lighter

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
eye.

vertical bar

below the

Lower

parts white, checkered with black, or
Iris

nearly entirely black.

reddish-golden.

In some specimens (var. quadriltneatus) the dorsal
spots are replaced by two

brown or

red, black-edged

stripes bordering a pale greyish or yellowish vertebral

COLUBER
stripe

193

(Plate VII., fourth figure);

such

specimens

are so coloured

from

birth.

This colour variety,

which
is

is

so strikingly different from the typical form,

connected with the latter by the var. schwoederi,

Werner,
series,

which the spots form two vertebral and the var. elsneri, Werner, in which the light
in

vertebral

band

is

broken up by dark transverse bars,

producing a ladder-like pattern.
Size.

Distribution.

— Rarely exceeding a length of 3 — Southern Malta,
feet.

Italy, Sicily,

Istria,

Dalmatia, and other parts of the Balkan Peninsula,

Grecian islands, Crimea, Asia Minor. The altitudinal
range does not extend beyond 1,600
Habits.
feet.

— This

is

not only the prettiest European

snake as regards

its

markings, whether in the form
its

of spots or of stripes, but also the most graceful in

movements.
there
is

Unless compelled to

fly

for safety,

something slow and deliberate
is

in

its

be-

haviour which

more suggestive

of Coronella than of

most other species of Coluber.

It is
it is

fond of climbing,
kept be provided

and

if

the terrarium in which
tree,
it

with a bush or small

will

spend most of the
Usually

time gracefully coiled round the branches.
very

savage
quite

when
tame

fresh

caught,

some specimens
found principally
It

become
it is

in captivity.

In Dalmatia, where
is

not

uncommon,

this snake

among

prickly shrubs, in hedges, or on old walls.

awakens from its winter slumber later than other Although occasionally South European snakes.
13

194
taking lizards,
its

COLUBRIDiE
principal food consists of
killed

mammals
of

and

birds,

which are

before being devoured,

the Leopard Snake being, like the other

members

the genus Coluber^ a constrictor.
Reproduction.
to five in

—According to Werner, the eggs, two
in

number, are deposited
:

midsummer

they are remarkably elongate
broad.
13,

2^ inches long, f inch

Coluber

scalaris, Schinz

The Ladder Snake
Form,

— Moderately

slender.

Snout

pointed,

strongly projecting beyond the mouth.
sixth to one-fifth of the total length.

Tail one-

Head-Shields.

— Rostral deeper than broad, forming
in

an acute angle above, wedged
nasals, the portion visible

between the
as

inter-

from above nearly as long
its

distance

from the frontal.

Frontal

about once and

one-third to
once and a half
as long as broad, as
F,a..6(afterSordem)

long

as

or
its

shorter than

^.^^^^^^

^^^^^

the end of the snout, nearly as long as the parietals.

Loreal longer than deep.

One

pre-

and two or three

PLATE

VIII

"^
\
^^-^5^

COLUBER
post-oculars.

195

Temporals

2

+

3 or 2

+

4.

Upper
Four

labials seven or eight (rarely nine), fourth or fourth

and

fifth (or fifth

and

sixth) entering the eye.

or five lower labials in contact with the anterior chinshields,

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, rarety entire
Coloration.

subcaudals 48 to 68.
pale brown,

—Young yellowish-grey, or
— whence
the
;

above, with a series of regular

H -shaped
name

black or

blackish-brown markings along the back, forming a
ladder-like pattern
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.

markings disappear
a pair of

in the adult,

These dorsal and are replaced by

brown

stripes

running along the back
dark brown.
feet,

(Plate VIII.); the belly loses the black markings,

and becomes uniform yellow.
Size.

Iris

— Grows

to

a length of 3J

exception-

ally 4^ feet.

Distribution.

— The Mediterranean coast of France,
Its

Spain and Portugal, and Minorca.
Algeria
is

occurrence in

very doubtful.

Habits.

— Not uncommon near the coast in France,

196
in

COLUBRID^
it

hedges and vineyards, often climbing on shrubs.
is

In the Spanish Peninsula, according to Bosca,

common
valleys,

in

forests

and on

the

sheltered side of

under stones or
I

in holes in the

ground.

A
any

specimen
furious

kept alive for a short time showed a more
I

temper than

have ever witnessed

in

snake, repeatedly flying with open
glass of its cage
it

mouth

against the

whenever 1 entered the room in 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
which
;

;

are said to occasionally eat grasshoppers.
Reproduction.
eggs, nine in

According to J. von Fischer, the number, are deposited twenty-five days

which takes place in May or June, and measure about 2 inches by f inch.
after the pairing,

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
pits.

;

scales smooth, with apical

Tail moderate.
is

This genus, embracing about twenty species,

represented in the different parts of the Northern

Hemisphere, extending a
in

East Africa.

Two

beyond the Equator species are European.
little

PLATE IX

CORONELLA
14.

197

CoRONELLA AUSTRIACA, Laurenti
(Cohcber
Icevis,

Lacepede)

The Smooth Snake
Form.
fourth
length.
in the

— Moderately
to

slender

;

snout more or
;

less

prominent, sometimes decidedly pointed
(males)

tail

one-

one-sixth

(females)

of the total

The considerable

differences to be observed

shape of the snout are merely individual, speciw^ith

mens

more prominent snout and a corresponding
(C.

development of the
rostral

shield

italica, Fitz.ffitzin-

geri,

Bonap.) ocover
part

curring
greater

the
of

the

range of the

species.

Head - Shields.


Fig. 27 (after Sordelli)

Rostral at least as

deep as broad, more or

less

produced posteriorly
visible

between the internasals, the portion
above
quite
at

from

least

half
as

as
its

long

(in

some specimens
the
frontal,

as

long)

distance from

rarely separating the internasals.

Frontal once and
as

one-fourth to once and a half as long

broad,

much broader than
or longer

the

supraocular,

as

long as

than

its

distance from

the end of the

snout, shorter than the parietals, vi^idely separated

ig8

COLUBRIDiE
Nasal rarely undivided
(very rarely two) pre2 or 2
;

from the preocular.
longer than deep.
postoculars.
I

loreal
tv^o

One

and

+

2).

Temporals 2 + Upper labials seven
fifth)

+

3 (very rarely

(rarely eight), third

fourth (or fourth and

entering the eye.

and Four

lower labials (rarely three) in contact with the anterior
chin-shields,
anterior.
Scales with

which are as long as or longer than the
one or two apical
rows.^
pits,

the pit usually

single

on the back and paired on the

sides, in nineteen

(rarely twenty-one)

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
stripes;

sometimes with one or three lighter
apical pit

one or two

black dots precede on each scale the single or paired
;

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
of the

nostril to the angle of the

mouth, passing

through the eye, sometimes extending along the side 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 a male from Albano, near Rome (Genoa Museum).

is

CORONELLA
A
colour variety, of which
I

igg

have examined a single
is

specimen from near Vienna,

pale

brown above,

with four black Imes 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
having

variety, also

from

near Vienna, which resembles Coluber leopardinus,

two

series

of

large,

brown,

dark

-

edged

spots along the back,

some

of the spots alternating,

others uniting across the back.
similar
to

Apparently very
also

the

last

variety,

and
is

said

to

be

suggestive of Coluber leopardinus,
Sternfeld,

the var. scalaris,
reddish-

from

Llineburg

in

Hanover,

brown above, with two rows of bright red, blackedged spots, partly confluent and connected across
the spine by transverse bars producing a ladder-

Specimens of a uniform greyishbrown, without any markings, are very rare. The
like

pattern.

var. veithi, Schreiber, established

on a single

speci-

men from
black.
to

Carinthia, represents a case of melanism

bluish-black, with the normal markings of an intense Two specimens of a " black variety " are said

have been found
Size.

in this country, near Poole.
feet,

— Seldom
districts,

exceeds a length of 2
in

and

in

many

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
Distribution.

COLUBRIDJE

— The

range of

the

Smooth Snake

extends over nearly the whole of Europe, as far
north as 63° in Norway
local in

becomes rare and more the south, being absent from part of Spain
;

it

and
parts

the islands of

the
It

Mediterranean, with the
is

exception of Sardinia.
of Belgium,

common
it

in

the hilly

Northern and Central
In

France,

Germany, and
restricted

Austria.

Sweden

appears to be
it

to the oak region.

In Great Britain
in the

has been found in four counties

South of

England
shire, in

:

Surrey, Hampshire, Dorsetshire, and Berkparts of which
Its
it

some

is

less

uncommon
;

than usually supposed.
Dumfriesshire
is

reported occurrence in
the snake

the result of an error

figured as Coluber dumfriesiensis represents an Ameri-

can species.
Science Gossip

In a very interesting article written for
in

1888, Mr. A. L. Beldy says that

about 1868, when

Bournemouth was but a very

small village, surrounded by large expanses of moor-

was extraordinarily abundant, and during a hot summer examples were to be seen literally in scores and great numbers were killed.
land, Coronella anstriaca

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

neighbourhood.

now extinct in that From South-Eastern Europe the

range of this species extends to South-Western Asia.

CORONELLA
The
ascertained
altitudinal

201
is

range

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 Although not infrequent on the for dry localities. Dorsetshire and Hampshire heaths, where it was first

was not recorded as a British it was discovered much later on reptile until 1859 the sandy heaths between Haslemere and Farnham, where it occurs in small numbers, and in Berkshire. These localities are likewise inhabited by the rarer
discovered in 1853,
;

it

British lizard, the
its gentle,

Sand Lizard.

Notwithstanding

timid appearance, this snake

when
it

fresh

caught
angrily,

is

usually very ready to bite; either

snaps
it

or,

without
its

hissing

or other warning,

suddenly fastens
captor, even
consists
if
it

jaws into

the

finger

of

its

be gently handled.

The

food

mostly of lizards,
or

occasionally of slowrarely

worms

small

snakes,

more

of voles

or

mice, even shrews, which are seized, constrictor-like,

O. von and crushed by the coils of the body. Tomasini has observed one swallowing a Coluber
longissimtcs as large as itself.

In

Central

Europe

this

snake

becomes active
It

towards the end of March or beginning of April, and
retires in

captivity,

September or October. and becomes very tame.

does well in
is

It

one of the
in edu-

most

intelligent of snakes,

second to none

202
cability;
its
it

COLUBRID.E
can be trained to feed
in

the hand of

master.

Reproduction.
spring,
fifteen

—The

and
in

is

ovoviviparous.

Smooth Snake pairs in early The young, two to
late in

number, are born

August or

in

September, enveloped by a thin membrane which
they measure 5 to 6 inches. Embryos 3J or 4 inches long have the scaling and the characteristic markings fully developed, but the
;

they tear immediately

scales

and

shields

much

abbreviated,

the former
is

broader than long.
in

A dicephalous young
at

preserved

the

Bosnian

Museum

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 considerably broader than deep and just visible from
above, not penetrating between the internasals, constantly eight upper labials, fourth

and

fifth

entering

the eye, and the scales in twenty-one (rarely nineteen or twenty-three) rows.

Ventrals 170 to 200

;

anal divided

;

subcaudals 49 to 72.

FLATE

X

CORONELLA
Coloration.

203
or reddish

— Brown, greyish, yellowish,
in the

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
species
;

preceding

a pair of elongate dark spots or a U-shaped
;

marking on the nape
to

a dark streak from the eye

the angle of the mouth, and a dark cross-bar
eye, across the prefrontal shields
;

from eye to
line

a dark

below the eye.
yellow,

Lower parts
with
large,

orange, or coral red,

mostly

quadrangular black

spots, oft en
ranged
board
in

ar-

chessor
Fig. 28 (after Sordelli)

fashion,

with two series of

111 black
X.),

spots

,

(Plate

/-rj,

.

which may be confluent into two longitudinal

bands.

— 26 inches. Distribution. — South of France
Total Length.

(as far

north as

the Charente-Inferieure to the west, the Dauphine
to

the east),

the whole
Italy,

of

Spain

and Portugal,
It

Southern Tyrol,
recorded
Alps.

and

Sicily.

has not been
feet

from
in

higher

than

2,500

in

the

Rare

Northern Morocco and

Algeria.

In

many

localities in

Europe

it

occurs alongside with

C. austriaca.

204
Habits.

COLUBRID^

—All

that

is

known

to

me

of the
is

habits

of this close ally of the preceding species

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
nibens.

According

to

these

authors,

it

frequents dry and rocky localities as well as
walls, in

old

which

it

finds a refuge
it

and a good supply

of the lizards on

which

feeds.

A

large specimen
in the

from Albano, near Rome, preserved
tridactylus.

Genoa
rarely
retreat

Museum, had swallowed a full-grown
This
in

Chalcides

Coronella

is

crepuscular,
its

showing
about

itself

the daytime, leaving

only after sunset, and has been observed to crawl

by moonlight.
for

Its

movements

are

slow,

which accounts
in C. austriacay

crushed specimens being often

met with on paths or roads.
it is

Contrary to the rule

extremely gentle, seldom attempt-

ing to bite.
Reproduction.
arous, like
its

— Whether
All

this species

is

ovovivipI

European congener, has

not,

think,
is

been ascertained.

we know on

this

matter

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
ing to
later.

Accord-

Gene, pairing takes place
been observed
to

in

May, when

specimens have

congregate in

considerable numbers.

CONTIA
Genus CONTIA, Baird and Girard
Maxillary
teeth

205

subequal.
;

Head
;

not

or

but

slightly distinct

from neck

eye moderate or rather

small, with round pupil.
shields.

Nasal single
;

no subocular

Body moderately elongate
pits.

scales smooth,

with apical

Tail moderate.
certain
to

This genus, with

modifications in the

above

definition, is

made

embrace about twenty-

five species

from South-Western Asia and Sind and

North and Central America.
Europe.
16.

One

of the species

inhabiting Asia extends into a very small part of

CoNTiA MODESTA, Martin

The Dwarf Snake
Form.
flat

— Moderately
;

slender.

Head

small, quite

above

snout obtuse, feebly prominent.

Length

of

tail

four to five times in the total length.

Head-Shields.
visible

— Rostral a
little

little

broader than deep,

from above.

Suture between the internasals
shorter than that between the

as long as or a
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
little

than the parietals, as broad as or a
preocular.

broader

than the supraocular, widely separated from the
Nostril in the middle or upper part of

the nasal.

Loreal square or longer than deep.

One

206
(rarely two) pre-

COLUBRID^
and two
;

(rarely one) postoculars.

Temporals
ing the
third

i

+2

parietal
labial.

sometimes nearly touchlabials

Upper and fourth entering the eye.
fifth

upper

seven,
(rarely
in

Four

five)

lower labials

contact with the anterior

chin

-

shield

;

posterior chin-shields

smaller than the anterior,

and separated

from each other by
one or two rows of
scales.
Fig. 29 (after Sordelli)

^^^^^^ ^.^j^ ^ ^j^^j^

apical pit, in seventeen rows.
to 191
;

Ventral shields 150

anal divided

;

subcaudals 53 to 78.
above, uniform
or each

Coloration.

— Not

unlike that of a young Zamenis

gemonensis.

Greyish-olive

scale lighter in the centre.

The

greater part of the
to-

upper surface of the head behind the snout,

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
of the

horseshoe-shaped band

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
the sutures between the shields.

207

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.
feet,

Caucasus up to about 5,000 Asia Minor, Chios, Cyprus, Syria, Mesopotamia,

—The

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 Constantinople, but the presence of this species in

European

Turkey requires confirmation. A closely allied species, which has been confounded with C. modesta, C. collaris (Menetries), and which
also

inhabits

the

Caucasus without having been
is

recorded from the northern slope,

distinguished

by having the scales
seventeen),

in fifteen rows (very rarely and the posterior chin-shields in contact

with each other.
Habits.

— Nothing

is

known

as regards this species,

American members of the genus Contia are chiefly insectivorous and oviparous.
but the North

Genus CCELOPELTIS, Wagler
Maxillary teeth small and subequal, followed after
a short interspace by one or

two very large grooved

2o8

COLUBRID^
Head
with

fangs situated below the posterior border of the eye
anterior mandibular teeth strongly enlarged.

not very distinct from neck, with angular canthus
rostralis

and projecting supraocular; eye
;

large,

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.

which comprises only two species, extends over Southern Europe, Southwestern Asia, and North Africa.
of this genus,
17.

The range

Cgelopeltis monspessulana, Hermann

{Natrix lacertina,
I.

Wagler

;

Coluber insignitus,

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.
just visible

— Rostral

nearly as deep

as

broad,
shorter

from above.

Internasals

much
width

than the prefrontals.

Frontal very narrow, twice to
its

twice and a half as long as broad,

in the

middle not more than half that of the supraocular, widening in front and extending beyond the supraoculars to join the preoculars, longer than
its

distance

from the end of snout, as long as or a

little

longer

PLATE XI

COELOPELTIS MONSPESSULAN
After Sordelli

MACROPROTODON CUCUI.LATUS

TARBOPHIS

IBERU.'

After Sordelli

TARBOPHIS FALLAX

CCELOPELTIS
than the parietals.

209

Two

loreals.

One

preocular, the

upper portion of which is much enlarged, and encroaches upon the area occupied in other snakes by
the prefrontal and the supraocular
postoculars.

eight

(rarely

two (rarely three) Temporals 2 + 3 or 4. Upper labials 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
apical

single
pits, in

seven(l
I I

teen

or nine-

11

/<Kl<»>

teen rows,
longitudinally

grooved

in the

adult, less distinctly in

the
Fig. 30

young.

Ven160 to i8g
anal divided
;

tral shields

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
its

forming with

fellow a /^-shaped band, separated

from a large occipital blotch by a yellowish space
14

210

COLUBRID.E
a dark streak, some-

anterior half of the frontal shield and shields on the

snout edged with dark brown

;

times broken up into small spots, on the temporal
region
lips
;

yellow spots on the pre- and post-oculars
large, yellow,
;

brown, with

black-edged spots, or

yellow with brown spots
longitudinal streaks.
large

chin with three brown
vertebral series of

Back with a
spots or

roundish dark

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
tail.

longitudinal streaks on the
greyish, or reddish, with

Belly pale brownish,
spots,
side.

numerous pale

some-

times with a dark brown line on each
adult
is

greyish, reddish-brown, or olive above.
less the

The Some

specimens preserve more or

markings of the

young, and the dark dorsal markings (Plate XI.)

may

be edged with yellowish and ocellar in appearance
(var. insignituSy

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 {neumayeri, 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.
are

Other specimens
lines, or

brown

or reddish, with light edges to the scales

on the

sides, or

with yellowish lateral

dark
sides,

olive or

dark brown above and black on the
;

each scale with a yellowish central spot

in the last-

CCELOPELTIS
and the

211

mentioned the second third of the back may be almost
entirely black,
belly dark olive-grey in the
sides.
Iris

middle and yellowish on the

brown,

with a golden or coppery circle round the pupil.
Size.

—This handsome
Specimens

snake grows to a length of
long are not un-

6J

feet.

5 to 6 feet

common.
Distribution.

— Mediterranean

coast of France and
Sicily,

Western Liguria, Spain and Portugal,

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.
Habits.

It

is

not

known

to occur above

2,300 feet altitude in Europe.

—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

COLUBRIDtE
is

Europe, Vtpera ammodytes

said to be the principal

enemy

and the two snakes are consequently seldom found together in the same locality. Many experiments have been made on the action of the poison of this Opisthoglyph. Peracca and
of Ccelopeltis,

Deregibus, as well

as, later, Phisalix,

found a striking
in

similarity with the
their experiments

symptoms

of

Cobra poison

on small animals, the suspension
unaffected.

of the respiration occurring in a few minutes, the

blood

being

otherwise

It

has

been

stated by
little

some authors that Ccelopeltis poison has or no action on man, but a French zoologist,
in the index-

E. Taton-Baulmont, having been bitten
finger

by a four-foot-long specimen

at

Algiers, the

swelling
shoulder,
troubles.

extended within thirty hours up to the

and was accompanied by As a rule, however, the
effect

fever

and nervous
snake
fact that

bite of this

has no poisonous

on man, from the

the fangs conveying the

back

in the

mouth
in

as

venom are situated not to come into action.
to
laid
in

so far

Reproduction.

—According
in length

Werner, the eggs,
July,

four to twelve

number, are

and

measure 2 inches

and J inch

in width.

Genus

MACROPROTODON,
and
sixth enlarged

Guichenot
size, fourth

Maxillary teeth few and very unequal in

and

fifth

or fifth

an interspace, the two

last teeth fang-like
;

and followed by and grooved,

situated just behind the eye

sixth mandibular tooth

MACROPROTODON
fang-like,

213

and separated from the remainder by an

interspace.

Head

slightly distinct

from neck

;

eye

rather small,
elliptic

the pupil vertically elliptic
contracted.

or subelon-

when

Body moderately

gate

;

scales smooth, with apical pits.

Tail moderate

or rather short.

A

single species.

18.

MACROPROTODON cucuLLATUS, I. The False Smooth Snake

Geoffroy

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.

— Rostral
least

Head -

Shields.

at
as
Fig. 31

twice
as

broad

deep,
scarcely
visible
little

not or but

from

above.

Inter-

nasals as long as or a
frontals.

shorter than the pre-

Frontal not

much broader than

the supra-

ocular in

the adult, once and a half to twice as
as long as or longer than
its

long as broad,

dis-

tance from the end of the snout, shorter than the
parietals.

Nasal usually semidivided.

Loreal once

214

COLUBRID^

and a half to twice as long as deep. One preocular, extending to the upper surface of the head, but not
reaching the frontal
oculars.
;

two

(rarely

one or three) posteight, fourth

Temporals

I

+

2.

Upper labials

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.
single

Scales

with mostly

apical

pits,

the
in

pits

sometimes paired on the sides of the body,

twentyin

one or twenty-three rows (nineteen to twenty-five

North African specimens).
192
;

Ventral shields 153 to
greyish above,

anal divided

;

subcaudals 40 to 54.

Coloration,

— Pale

brown

or

with
or

small dark

brown

or blackish spots or with

more

less distinct

darker and lighter longitudinal streaks.

Upper
tions;

surface of head with dark

brown vermiculaoften

a

dark

brown

or

black,

light-edged
to the

occipito-nuchal band, extending

downwards

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 yellowish

above and below the streak (Plate XL).
a tessellated
pattern,

Lower

parts yellow or coral red, with black spots, which

may form
series, or

be so crowded as to fuse
tail.

two longitudinal into a band along

the middle of the belly and

MACROPROTODON
The above
description
is

215

taken

from Spanish

specimens (Badajos, Algeciras, Andalucia), but the
variations are very great
into

consideration.

when we take North Africa 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 exception of a

whitish streak

bordering the upper

lip.

The dark streak from mouth may be absent,
streak below the eye.
in the

the eye to the angle of the
or reduced to a short oblique

Irrespective of the variations

markings of the upper parts, the lower parts
less

may
late.

be more or

spotted with black, or immacu-

Some specimens

of this small snake bear a general

resemblance to Coronella girondica, with which Macroprotodon has sometimes been confounded.
careful examination of
to be
its

But a
it

whole structure shows

more

affine 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

2i6

COLUBRIDJ^:

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
it
;

in Algeria

penetrates into the northern parts of the Sahara.
figured on Plate XI.
is

The specimen
Habits.
girondica.

from Algeciras.
usually

—Appear to be similar to those of Coronella
Crepuscular in
in
its

habits,
in

it

is

found under stones or
Unless pursued, when
its
it

burrows

the ground*

darts off with great rapidity,
It
is

movements
its

are slow.

very ready to bite,

but no experiments have been
of poison.

made on

the effects

The

food consists chiefly of small

lizards.

Reprodnctio7t.

— All that

that, according to

is known on this head Doumergue, eggs are laid in July

is

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,
posterior

situated

below the
from neck

border of the eye

anterior mandibular teeth strongij^^nlarged.
distinct
;

Head

eye moderate or rather small,
pupil.

with

vertically

elliptic

Body moderately

elongate;

scales smooth, oblique, with apical pits.

Tail moderate or rather short.

The

eight species of this genus

inhabit

South-

TARBOPHIS
Two
are dealt with here.
ig.

217

Eastern Europe, South- Western Asia, and Africa.

Tarbophis fallax, Fleischmann
{Aihirophis vivax, Bonaparte)

The Cat-Snake
Form.
pressed.

— Moderately

slender.

Head

much

de-

Tail five and a half to seven times in the

total length.

Head-Shields.
visible

— Rostral

broader than

deep,

just

from above.
Fron-

Internasals shorter than the pre-

frontals.
t

a

1

much
supraocu-

broader than
the
lar,

once and
to

one - fourth

once and

a

half as long as

broad, as long
Fig. 32

as

its

distance
of

from the end
parietals.

the

snout,

shorter

than

the

Nasal divided or semidivided.

Loreal

twice and a half to thrice as long as deep, entering
the eye below the preocular, which the frontal.
porals
is in

contact with

Two

(rarely three) postoculars.

Tem-

small,

scale-like,

2

or 3

+3

or

labials eight (rarely seven or nine), third,
fifth (rarely

Upper 4. fourth, and
fifth,

fourth and

fifth,

or fourth,

and

2i8

COLUBRID.^
Three or four lower
labials

sixth) entering the eye.
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 i86 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
usually a dark streak on each

with one or three linear processes in front, extending

on the head (Plate XI.)

;

side of the head, from the eye to the angle of the

mouth.

Lower

parts whitish, speckled, spotted, or
Iris

marbled with grey or brown.
golden circle round the pupil.
Size.

brown, with a

—This

species grows to a length of 2 feet

to inches.
Distribution.

— From Istriaand Dalmatiato Greece,
;

the Archipelago, Constantinople, Asia Minor, Cyprus,

and Northern Syria
altitudinal limit.

2^00

feet

appears to be

its

Habits.

— Although
its

to a certain extent crepuscular
is

or nocturnal, the Cat-snake

often seen hunting in

the daytime,

food consisting almost exclusively

of lizards, rarely of small

mammals.

Its

movements

are rather

slow.

The names Katzenschlange and

TARBOPHIS
from the way
in

219

Ailurophis, translated Cat-snake, probably originated

which

this
it.

snake stalks

its

prey,

and

suddenly pounces upon

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,
Stony
localities,

on the other hand, are very savage.
snake, which does well in captivity.
Reproduction.

old walls, and ruins, are the favourite abodes of this

— Seven

or eight
iJ

eggs

are

laid

in

July

;

they measure about

inches in length and

i inch in width.

20.

Tarbophis iberus, Eichwald

The Caucasian Cat-Snake
Very closely
from
it

allied to the preceding,

and
:

differing

only in

the

following

points

Parietals

shorter, slightly longer than the frontal,
entire.

and anal

Loreal twice to twice and a half as long as

deep.
labials

Fourth and
entering

fifth,

or third, fourth,

and

fifth,

the

eye.

Scales

in
;

nineteen

or

twenty-one rows.

Ventrals 203 to 235

subcaudals

54 to 70. 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

VIPERID-E
and
dots.

blackish, with ?:"all whitish spots

Reaches

a length oi j-

i^et.

This species inhabits the Caucasus, and, being on
record from the northern slope ^_Kuban River), has
to be included in
also in

the European fauna.

It

occurs

Mes::
in

::::.mia. a

specimen from Bagdad being

preserved

the

British

Museum.
is

The young

specimen figured on Plate XI.
Constantinople.

stated to be from

Nothing

is

known

of

its

habits,

which are probably

the same as those of Tarhophis 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 daagerous of snakes. Dmded into two subfamilies, each of which is rep :':r-:.ted by one genus in Europe:
r.-.
TT.

.:r.
--'
':

— No
:r.e

pit

on

the

side

of

the

snout;

not hollowed out.
ci-rp pit

,

,:.

—A
'

on each side of the snout,
;

between the
h'..._
-'
,

nostril
-e.

and the eye

maxillar}-

bone

-

.^. ,.,..,

.r.r.aoit

nearly the whole of Europe,

PLATE

XII

'^W^W^^^^^'
-^^^^;^^^^:^-J^'
VIPERA CRSIXII Fr: rn Z; r'.i^ical Society's Proaeli'iss

\IPERA RENARDI

VIPERA BERUS
A/ter Scrdeiii

VIPERA
Asia,

221

and Africa

;

the Crotalince are Asiatic (one
its

species extending

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
parietal shields
pupil,
;

and

eye moderate or small, with vertical
;

separated from the labial shields by scales

nasal separated from the rostral by a naso-rostral.

Body
short.

short.

Scales keeled, with apical pits.

Tail

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 relationship and the presence of intermediate forms connect;

ing them.

Matters being

the
so

Common
many

so, it seems curious that Adder should have been regarded by

authors as generically distinct from the
Pelias herns.
It
is

Asp Viper, under the name of
districts

highly probable that hybrids are produced in those

where two species

coexist, as in

some parts

of France, North Italy,
21.

and Austria.

ViPERA

URSiNii, Bonaparte

Orsini's Viper

Form.
flat

— Short and

stout.

Snout obtusely pointed,
raised.

above or with the canthus slightly

Eye

222

VIPERID.E
its

very small, usually smaller than the nasal shield,
horizontal diameter usually not exceeding
its

distance

from the posterior border of the
diameter often
less

nostril, its vertical
its

than and rarely exceeding

distance from the mouth.

Length of

tail

seven to

eight times in total length in males, nine
to twelve times in females.

and a half

Head-Shields.

— Rostral as deep as broad or slightly
with
apical
(rarely

deeper than broad, visible from above, in contact

one
shield

with

two).
tinct

Disfrontal

and
p a

(usually)
r
i

e

t

a

1

shields,

the

former
Fig. ^^

once

(From Proceedings of the Zoological
Society, 1893)

and

a

half
and

to once

two-thirds (rarely once and

one-third) as

long as
rostral

broad, as long as

its

distance from

the

or the end of the snout,

and nearly always longer
latter

than
with

the
the

parietals

;

the

always

in

contact

former,

rarely

broken

up

into

small

shields.

Four

to seven small shields

on the snout

each

between the canthals, of which there are two on side. Supraocular well developed, extending

posteriorly

beyond the

vertical of the eye, separated

VIPERA
from the
the
frontal

223
three
shields,

by one
it.

to

very

rarely in contact with
eye,

Six to ten scales
nine,

round
pre-

usually

eight

or

the

upper
;

ocular usually in contact with the nasal
series

a single
labials.

of scales

between the eye and the

Nasal single.
labials six to

Temporal

scales

smooth.

Upper
Three

nine, usually seven or eight, usually

third

or

third

and fourth below the
is

eye.

(rarely four) lower labials in contact with the chinshields, of

which there

but one pair.

Scales

in

nineteen (rarely twenty or twenty-one)
pits,

rows, with two apical

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
entire; subcaudals

anal

females.

By

30 to 37 in males, 20 to 28 in 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. berus,

V.

itrsinii

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
light

an almost uniform brown ground colour.
colour of the back
is

The

relieved

by a

series of

more or
of which

less regular transversely oval, elliptic, or

rhomboidal

dark brown, black-edged spots, some or

all

may

run together to form a wavy or zigzag band

(Plate XII.).

Two

or three longitudinal series of dark

224

VIPERIDJE
sides, the

brown or black spots extend along the
series, if present,

upper

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
labial

is

not infrequently confluent with the

branches of the occipital A.
shields

The

rostral

and the

are

uniform

yellowish-white, rarely

with a few small, blackish spots or with brown
borders.

The

chin and throat are yellowish-white,
spots.

rarely with

some blackish

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
tail is
;

the

but rarely (females) tipped with yellow.
recently^described as V. macrops, Mehely,
is

The form

from Bosnia and Herzegovina,
equals or a

distinguished by a

usually larger eye, the vertical diameter of which
little

exceeds

its

distance from the mouth,

and the
shields.

parietals are often broken

up

into small

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
Size.

225

— 20 inches appears to be the usual maximum
feet

size

reached by this species, but, Dr. Werner informs
long has been found in

me, a female 2
Austria.
Distribution.

Lower
the

— First

discovered

in

Italy in

Abruzzi, this species has since

been found

in the

Basses-Alpes, near Digne, in various parts of
gary, in
Istria,

Hunin

Lower
in

Austria,

on the island of Veglia

and

Bosnia, Bulgaria, Herzegovina, and

Montenegro.
plain in

A very broken and curious distribution,
is

the more so as V. ursinii

essentially a

form of the
alpine

Lower Austria and Hungary, and an
it

form
sula,
feet.

in Italy, in

France, and in the Balkan Peninhabitat does

where

only occurs between 3,000 and 6,800
its
it

In no part of

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-

and in the course 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
for the destruction of Vipers,

mium

of one year (1892)

the daytime from

May

to September, feeding chiefly

on

lizards {Lacerta agilis), lizards are
15

and

also

on small rodents.

The

swallowed as soon as seized, without

226

VIPERID^
This Viper
itself
is

the effect of the poison being awaited as in other

Viperid snakes.
disposition,

as a rule of gentle

allowing
bite,

to be handled without

and village boys have been seen Although occurring in such playing with them. 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
attempting to
still,

and

is

believed never to
its

make use

of

its

poison

apparatus,

food consisting of orthopterous insects.

According to Captain Veith,

who has collected a large

number

of specimens of this Southern form, the con-

stomach as well as the excrements show On this snake to feed exclusively on grasshoppers. one occasion a big specimen showed such a swelling of the body as to lead to the conclusion that it had
tents of the

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

Reproduction.

to have little effect on man. Nothing has been published on the

breeding habits of this species, but in a letter to
the
author,

dated

January
has

14,

1913,

Herr

L.

von

examined over 4,000 specimens since 1890, says the young are born in
Kirchroth,
or

who

July

August,

exceptionally as early as June.
forth from six to eight young,

Young females bring

VIPERA
older

227
;

females from eight to eighteen
is

but a large

female from Lower Austria
tained as

reported to have con-

many

as twenty-two.
is

The

length of the

new-born young
rapidly

within

the

from 5 to 6 inches, and it grows 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.

ViPERA RENARDI, Chrlstoph
Renard's Viper

Form. Similar more pointed, the
an acute angle.

to the preceding species, but snout

raised canthi rostrales meeting at

Eye
its

usually as large as in F. bencs,
its

nearly as large as the nasal shield;

horizontal

diameter equal to
equal to or a

distance from the posterior or
its
its

anterior border of the nostril,
little less

vertical diameter

than

distance from the

mouth.
in
total

Length of
length in

tail

seven and a half to nine times
eight
to

males,

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^
in contact

always

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
in contact

side,

the second broadly

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
or

the eye and the
labials,

two

series except

under the centre
ofthe eye, which
is
Fig. 34

separated

(From Proceedings

by a 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
1
1

of the Zoological Society, 1893)

from the fourth
1

u a b

a

1

u

1

single

scale.

Nasal

single.

pair.

Scales in

twenty-one (very rarely nineteen) rows, with
pits,

two apical

strongly keeled, outer

row smooth or

feebly keeled.

Ventral shields 130 to 148 in males.

VIPERA
37 in males, 24 to 30 in females.
Coloration.

229

130 to 150 in females; anal entire; subcaudals 31 to

— 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
is

or black.

The

dorsal

band or

series of spots
;

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.
of which there
side

The lower

parts are whitish or pale greyish, with blackish dots,
is

a series of larger ones along each

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
on the
Size.

series of spots

and two

series of small spots

sides.

Belly whitish, dotted or spotted with

black, or uniform blackish.

— 23

inches

is

the

length

of the

largest

specimen examined.

which has long been confounded with V. berus, is abundant in the district of Uralsk, in the steppe around Sarepta, in Crimea, and it is also found in Cis- Caucasia and
Distribution,

— In

Europe V.

renardi,

230
in

VIPERIDJE
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

Around Sarepta it is common in the bare steppe, and only exceptionally occurs in localities overgrown with willows and small shrubs.
district.

Habits.

— Nothing

has been published concerning
it is more sensitive and does not appear before the

the habits of this snake, except that
to cold than V. berus

middle of April, retiring to
beginning of October.

its

winter-quarters in the
consists of small

The food
in

mammals and
to seven

lizards.

Reproduction.

— Pairs
in

young

May, and brings forth five August, these young at birth

measuring about 5i inches.
x,^
23.

ViPERA BERUS, Linnseus
Viper, or Adder

The Northern
Form.
slightly concave, the

— Short and stout.

Snout

flat

above, rarely

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 as

large or nearly 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
Head-Shields.

231

— Rostral as deep as broad or slightly
In

broader than deep, rarely once and one-third as deep
as broad, not or but scarcely visible from above.

addition to the supraoculars, three large shields, the
frontal

and the

parietals, are as a rule present

on

the top of the head.
little

Frontal as long as broad or a

longer than broad, rarely

much

longer than

broad, once and a half to tv/ice and a half as broad
as the supraocular,

from which

it is

as a rule separated

by one
shields,

to

four

as

long
little

as

or

a

shorter

than

its

distance from the
rostral, as

long as
shorter

or a

little

than the parietals.
Parietals usually
in

contact with

Fig. 35 (after Sordelli)

the frontal and

separated

from
the

the supraoculars
in

by small
both, or
ally,

shields,

but sometimes

contact with

separated

from

frontal.

Exception-

and Austria, the

Germany, 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 borin

specimens

from Great

Britain,

parietals, or the frontal

dered by

six (rarely

by

five or four)

small shields,
side

viz.,

two apicals

(rarely one),

and on each

two can-

232
thals, the

VIPERIDJE
second of which
is

usually in contact with

the supraocular; canthals very rarely united into one
shield
;

the space between these shields covered by
fl^t

four to twenty

or

convex, juxtaposed scales,
single

which very exceptionally are fused into a
large
shield.

Supraocular usually extending pos-

beyond the vertical of the eye. Six to thirteen scales round the eye, usually eight to ten two or three superposed scales, rarely two vertical series
teriorly
;

of

scales,
is

separate the preoculars from the nasal,
single.

which

As a

rule a single series of scales
;

intervenes between the eye and the labials

specimens

with two series are of very exceptional occurrence

from Isle of Arran, Normandy, Norway, and Carniola, in the British Museum), but there are occasionally two series except just below the centre of the eye. Upper labials
(single specimens

Southern

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
Coloration.

233
species,

It

is

characteristic of this

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.
olive males

There are also brown, reddish-brown or
with brown markings.

with the markings of a deep black, and grey males

A

very pretty colour variety,
is

which
tion,

affects

only females,

olive with

brick-red

band and
{Coluber

spots.

Some

males, just before exuvia-

have the lower surface of a pale greyish-blue
cceruleiis,

Sheppard), with the outer ends of

the shields black.

Specimens with yellowish-white

chin and throat, which
females
;

may

be tinged with red, are

males have the throat black, or whitish with
Exceptional
(in

the scales spotted or edged with black.

females occur

Carniola) which in this respect re-

semble the males.
usually consist of a

The markings vary considerably. Those on the back wavy or zigzag longitudinal band,
its

flanked on each side with a series of spots corre-

sponding to

sinuses

;

but this band

may

be partly

or even entirely broken up into rhomboidal or transversely oval spots, or, losing
straight stripe edged
its

indentations, form a

on each side with a yellowish

234
streak (as in

VIPERID.E

some specimens of the var. seoanei^ from North-Western Spain). The markings may be absent
band
(Pelias dorsalis, Gray).

altogether (var. concolor, Jan), or reduced to a narrow
straight vertebral

In

the var. seoanei the zigzag
a dark

band

is

often replaced by
to five scales

brown vertebral band, three

wide, bordered on each side by a series of subtriangular 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
/\,)^»)^^)(i
ings

By
a

uniting together, this pair of markor an X.

may form

A

An

oblique dark streak
last labial

extends on each side from the eye to the
shields, being

sometimes prolonged a short way down
vertex

the neck.

The snout and

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
to the

yellowish, those at least

which are anterior

eye being more or less broadly edged with brown or
black.

and the lower surface of the tail vary from grey or brown to bluish, blackish-grey, or black,
belly

The

the sides usually dotted or spotted with whitish

sometimes, especially in females, the belly

is

dark

grey, each shield with a white posterior border
is

which

broken up by a

series of small roundish black spots.
is

The end

of the tail

often yellowish, bright yellow,

VIPERA
or pale orange below, rarely coral red,

235

more com-

monly The

in females
iris is

than

in males.

usually coppery red,

more

rarely golden

suffused with brown.

Black specimens occur, more or
to as V. prestev, Linnaeus.

less frequently, all

over the habitat of this species, and are often referred

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
is

latter are males,

and among them
showing how
this

we may
the back

find intermediate stages

melanism

brought about

;

in

one case the black of
the remains of

is

separated from the black of the sides by

a narrow light

brown wavy

stripe,

the ground colour.

When,

as in all females,
is

and

occasionally in males, the black

the result of a

gradual darkening of the ground colour, the typical

markings

may

still

be detected under certain lights.

Some specimens
mahogany
nearly
all

(from Schneeberg,

Lower

Austria)

are black, with scattered golden dots, or of a dark

brown

speckled

with

yellowish.

In

the black specimens at least a few dots of
lips,

whitish are visible on the

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.

seoaiiei,

Lataste, from North-Western

236

VIPERID^

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
of,

var. bosniensiSf

Boettger, from

very suggestive

is sometimes and has been taken for, the typical form of V. aspiSf 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

Bosnia, Carniola, and Carinthia, which

of dark bars across the back.

The

var. pseudaspts,

Schreiber,

from the plains of Sclavonia, described

as straw yellow above, with narrow dark cross-bars,
is

hardly separable from the var. bosniensis.
Size,

— Vipera
in

berus

is

said to reach very excep-

tionally a length of 2 feet 11 inches.

The

largest

specimen
specimen 2
females.

the
feet

British
3J

Museum
:

(from Belgium)

measures 2

feet 3

inches the largest British inches. Both these specimens are
male measures
2 feet 2 inches.

The

Distribution.

— Viper a

largest

berus ranges over the

whole

of Northern Europe, to the extreme north of Scotland,

and the sixty-seventh degree
of Saghalien.
It is

in
far

Scandinavia,
east as the

and right across Northern Asia as
island

generally distributed in
Isles of

Great Britain, occurring also on the
Islay,

Arran,
in

Skye, Lewis, and
districts,

Mull,

rare

or absent

some

common

in others.
is

Its distribution

in Central

and Southern Europe

irregular.

In

VIPERA
Western France
it

237

does not extend

much beyond

the

Loire to the south, only isolated captures being on
record from the departments Vendee, Deux-Sevres,

Vienne, and
Paris, in the

Indre.

Of

rare occurrence south of
in

departments Yonne and Allier and
it is

the mountains of Auvergne,

again abundant in

some
is

parts of the Central Plateau.

To
is

the east

it

recorded from the departments Aude,
In Belgium,
the
it it

Haute-

Marne, and Vosges.
Flanders,

known from
and
the

Limburg,
;

Meuse
is

Valley,

Ardennes

in

Holland

pretty generally distribIt
is

uted in the uncultivated parts.
nearly the whole of the

spread over
the

German Empire with
it is

exception of the vine districts, where

absent or

extremely rare

;

it is

also very scarce in the
;

mountains

of the Black Forest

it is

localities in Lorraine,

on record from only a few and has never been found in
it is

Alsatia.

In South

Germany

rarely found below
it is

1,000 feet altitude.

In Switzerland

absent from

the Jura, but occurs in the Alps chiefly between
2,500 and 9,000
feet.

To
the
it

the East

it

extends to

Russia, as far north as 64°, Austria-Hungary, confined to the
hills

in

south,

and Roumania.

In the Balkan Peninsula

occurs in the mountains

of Bosnia, of Herzegovina,

7,000

feet.

Absent
at

from, the

and of Bulgaria, up to South of France, it
Galicia,

curiously reappears in the hills of the north coast of

Spain, even
localities in

sea-level

in

and

in a

few

North Portugal.

On

the southern side

238
of the Alps
it is

VIPERIDiE

much
in

rarer than V. aspis, but

it

has

established

itself

a few low-lying districts in

Lombardy, Venetia, and the neighbouring part of
Emilia.
Habits.
its

— As

we

see

from the

above

sketch

of

Adder generally avoids the hotter parts of Europe when found in the plain
distribution,

the

;

in

the

South, as in Italy,

it

dwells
it

in

marshy
alpino,

localities,

and Bonaparte

called

Marasso palustre

(Marsh Viper)
Vipeva ursinii.

in opposition to his

Marasso
it

In the North, however,

usually

selects in preference dry moors,
hills

sandy heaths, and
delights to

well exposed to the sun, in which, although to
it

a certain extent a nocturnal reptile,
bask.
Its

food

is

very varied

:

weasels, mice, voles,

shrews,

moles,

birds,

lizards,

slow-worms,

frogs,

salamanHers, large slugs, have been found in the

stomach, and the very young feed also on insects

and worms.
are

Of

irascible
bite

temper as a

rule,

Adders
but

very ready to

when

fresh

caught,

instances are
in captivity,

known

of their becoming quite

tame

allowing themselves to

be

handled.

As a

rule they refuse food in captivity, but
live for as

some
bite,

have been known to

long as

five years,

being fed on lizards.

Accidents from their

although seldom heard of in this country, are of
frequent

occurrence in France and in Germany,
fatal

where many cases of
been recorded.

results

on people have

VIPERA

239

Pairing takes place in April and Reproduction. May, and the young, five to twenty in number, are born in August or September, exceptionally as early
as the end of July
selves
;

the young, on releasing them-

from the

thin, transparent

membrane

in w^hich

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
been

or in the autumn.

There

is

a trustworthy record, by Eiffe, of three pregnant
females

having
12,

caught

near

Hamburg on
birth to

March

1882, one of

them giving

young

on the following day.
Bicephalous young have occasionally been observed.

One

6 inches long
in

was found crawling

in a field

near

Hornburg

Germany
for

in October, 1895, and,

having

some time, was observed to hiss and open the two mouths alternately when taking up a defensive attitude. Another similar monster,
been kept alive

from Cornwall,
the

is

reported to have been sent alive to
in 1854.

London Zoological Gardens
24.

ViPERA

ASPis, Linnaeus

The Asp Viper
Form.
ing.

— Rather more elongate than
flat

in the preced-

Snout

above, more or less distinctly turned

up

at the end,

with sharp, not or but very slightly

240
raised canthus,
region.
little

VIPERID^
and
its

vertical or nearly vertical loreal

Vertical diameter of the eye equal to or a

less

than

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
side,

rostralis, gives the head,
;

seen from the
is

a peculiar expression

the eye

so oblique

that a vertical line

drawn from the

posterior extremity

of the supraocular
shield

to

the

lip

usually
through
or

passes
the
its

eye
pos-

down

terior border; but

the extent to which
the snout
is

turned

up
Fig. 36 (after Sordelli) ' ^
^

at

the

end

^-^ vanes

^^^^;^^^ consider-

ably,

respect, others V. latastii.

some specimens approaching V. berus in this 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

its

width

two-thirds to seven-eighths

depth, extending to the
rule,

upper edge of the snout.

As a

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

PLATE

XIII

VIPERA ASPIS
After Calnuite.

VIPERA LATASTII

VIPERA
and on the snout
shield, or
;

241

however, an enlarged frontal

even a frontal and a pair of parietals, are
in
is

sometimes present, though rarely so large as
a typical V. hems;

when

present,

the frontal

separated from the supraocular by one or two series
of scales
series
;

when

the frontal

is

absent, four to seven

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
times, however, the border
is

someseven

formed by
herus.

six or

small shields, the second canthal being in contact

with the supraocular, as in V.

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 preis

oculars from the nasal, which

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

the eye.
i6

Temporal

scales

and sixth, below smooth or feebly keeled.

242

VIPERID.E
(rarely five) lower labials in contact

Four

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
153)

or less

distinctly keeled, rarely perfectly smooth.

Ventral
in males,
;

shields 134 to 158

(usually

143 to

141 to 169 (usually 145 to 157) in females
entire;

anal

subcaudals 32 to 49 (usually ^y to 45) in the males, 30 to 43 (usually 32 to 38) in females
;

terminal caudal shield
spine-like

is

sometimes shorter and
quite obtuse in

less

than in V.

beriis,

some

specimens.
Coloration.

— Grey,
from

greyish-brown, brown, reddishis

brown, copper}^ red, or orange,
in

the ground colour
district
;

individuals

the

same

in

this
in

respect sexual differences are less

marked than

the preceding species, red or copper-coloured speci-

mens being found

in

both sexes, and silvery white
In rare cases
In specimens from

specimens do not seem ever to occur.

markings are entirely absent.

the greater part of France, Italy, and the Southern

Tyrol (see Plate XIII.), the dark brown or black
in the form of narrow cross-bars, continuous across the back or broken on the vertebral line and often alternating

markings on the body are mostly

with each other and with similar bars on the sides,
thus producing a pattern not unlike that frequently

found in Tropidonotus

tiatrix

;

a narrow dark line

VIPERA
running straight or zigzag
along the
spine

243

may

connect these cross-bars, and in rare cases it is so broad as to produce a zigzag band similar to that of In specimens from South-Western France V. bcriis. and the P}Tenees, rarely in some from other parts of

France and

Italy,

there

is

a broad dark grey or
series of black or

brown

vertebral

band between two
opposite

blackish-brown spots,
alternating
;

to

each

other

or

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

A

on the back of the head may
cross-bar on the nape,
of the
lighter

be united with the

first

and

enclose a cordiform
colour.

figure

ground

A

light

line

sometimes borders the upper

edge of the snout and the outer edge of the supraocular shield.

A

blackish

band or a mere
first

line

extends obliquely from the eye to the
spot
;

lateral

below

this the

upper

lip is whitish, yellowish,

or pinkish, with or without dark vertical bars on the

sutures

between

the

labial

shields.

The
the

iris

is

golden or copper}* red.

The lower

parts

var}'

as

much

as

upper;

244

VIPERID^
steel blue,

sometimes black or

with or without whitish

or reddish dots or spots, sometimes yellowish or pale reddish, with brown dots or marbhngs; in

some young, white with greyish
is

dots.

The

throat
or

yellowish white

or

pale

reddish,

uniform

speckled with blackish, sometimes
entirely black.

(males)
is

nearly

The end

of the tail

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 markings.

A

specimen from Piedmont,

in

the Turin
to

Museum, shows the ground colour reduced
narrow light bars disposed in pairs.

mere

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 F. aspis, which some herpetologists would perhaps regard as entitled to rank as a species, is the var. hugyi, Schinz, from Calabria

and

Sicily.

It

is

in

some
usual

respects

intermediate
is

between
often,

F. aspis

and F.
than

latastii.

The snout
the

rather

more pointed
up
at

in

typical

form,

though not constantly, more strongly turned the end, and the canthus rostralis may be
Constantly two canthal shields,
Ventral
Pale

distinctly raised.

the second in contact with the supraocular.
shields

134

to

148

;

subcaudals 30 to 43.

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
arbitrary are

completely intermediate between

Vipera a^pis and V, hems as to render their

naming

known from

parts of France and Italy

where the two species
be regarded as hybrids.
Size.

coexist,

and are probably to
(St.

—The

largest

specimen examined

Sever,

Landes,

in the
It is

Lataste Collection) measures 2 feet a male.
is

2J inches.
British

The

largest female in the

Museum

Distribution.

— Vipera

2 inches shorter.
aspis is

found over the whole

of France south of a line connecting the depart-

ments Loire-Inf^rieure, Orne, Seine-et-Marne, and
Meurthe-et-Moselle, and ascends the
the altitude of 7,250
feet.

Pyrenees to
it

In

Germany

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
in the

also in Austria, in the Southern Tyrol

and

Karst,
Sicily,

and

is

distributed over the whole of Italy

and
the

reaching an altitude of 9,700 feet in

Alps.
of the

Most

of the specimens from the western parts

Balkan Peninsula which have been referred

to this species belong, apparently, to V. berus, var.

246
bosniensis,

VIPERID^
but one from Jahorin in Bosnia, altitude
feet,

preserved in the Bosnian Museum, is pronounced 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
5,650

rock or in the earth.

It is

slow in

its

movements,
in

but very irascible, and innumerable accidents, in

some cases
Its

fatal to

man, are caused yearly
it

many

parts of France, where

is

extremely abundant.
small

food consists
birds,

principally of
lizards,

mammals,
its

young
insects

and

but the very young eat
it

and worms.

In

France

retires into

winter-quarters at the end of October or in
ber,

Novem-

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.
will

In rare cases

it

even leave

its retreat in the middle of winter, to

bask in the sun.

In captivity

it

long retains
all food.

its

savage temper, and usually refuses
Reproduction.

— Vipera
in

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

August or September, and

measure 7 or 8 inches.
Several cases of dicephaly in

young specimens

have been described.

VIPERA
25.

247

ViPERA LATASTII, BoSCa
Lataste's Viper

Form.

— Heavier

than

in

the

preceding.

Head

similar, but

snout more pointed and loreal region
lip,

slanting towards the
is

well visible

when
to

viewed from above.
is

The

extent

the head which the

snout

turned up at the end varies considerably,
aspis,

sometimes similar to certain specimens of V.

sometimes forming an appendage which

is

only a

Fig. 37
little less

developed than

in V.

tail six

and a half

to seven

ammodytes. Length of 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
scales,

deep as broad, nearly reaching the tip of the rostral Upper surface of head covered with small, wart.

smooth or feebly keeled, subimbricate which a slightly enlarged frontal, or a

among
and a

frontal

248
pair of parietals,

VIPERIDiE

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
eye.

beyond the

vertical of the

Five or

six (rarely three) scales

on the posterior
;

aspect of the raised part of the snout
scales

two canthal
with the

on each

side, the

second

in contact
it

supraocular, or separated from
preocular.
scales

by the uppermost
series of scales

Eight to thirteen (usually nine to twelve)
Nasal hollowed

round the eye; two or three
labials.

between the eye and the
out, entire, separated

from the preoculars by one or
;

naso-rostral sometimes 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
vertical series of scales

two

divided

into

lower labials in contact with the single pair of chinshields.
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 anal entire subcaudals 35 to 45 in males, females
;
;

32 to 38 in females.
Coloration.

— Grey
wavy

or

brown above, the back

often

paler than the sides, with a broad darker, usually

black-edged,

or zigzag

a lateral series of spots (Plate XIII.)
times replaced by large

band along the spine, and the band somerhombic or transversely oval
;

VIPERA
spots.

249

Head with
;

or without dark markings above,

sometimes with two obhque 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
less

upper

lip

white or pale

brown, more or

speckled or spotted with black.

Lower

parts grey, spotted with black

and white, or
of the tail

blackish speckled with white, the end
usually yellow or with yellow spots.
Size.

— This Viper

is

not

known

to exceed a length

of 2 feet.
Distribution.

— Locally distributed
Bona and
Viper
in

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

Guyotville.

Habits.
districts,

— Lataste's
and also

lives in

stony and arid
consists

forests.

The food
of

chiefly of small

mammals, but remains
in the

of a scorpion

have been found

stomach

an adult, and of a

centipede in that of a young.
this Viper easily climbs
birds, five of

According to Graells,
in search of

low trees
bite

which have been found

in

young the stomach

of one specimen.

The

is

believed to be less

dangerous than that of V.
the death of
26.

aspis,

and

rarely causes

man and

domestic animals.

ViPERA AMMODYTES, Linnaeus
or

The Sand-Viper,
This species
cal forms.

Long-Nosed Viper

may be divided into several geographiThe typical form will be described first.

250

VIPERID^

Form.

— Short

and heavy.
erect,
;

Snout pointed, pro-

duced into an

horn-like dermal

appendage

covered with scales

canthus rostralis strong, someVertical diameter of the eye

times slightly raised, loreal region slanting more or
less

towards the
than
its

lip.

less

distance from the

mouth

in the adult.
in the total

Length

of tail six to nine

and a half times
usually

length in males, eight to eleven times in females.

Head
deep.

-

Shields.

— Rostral

broader

than

Naso-rostral (rarely divided into two) usually

reaching the
canthus rostralis,

and extend-

ing considerably

higher up than
the upper border
of the
rostral.

Rostral append-

age covered with
Fig. 38 (after Sordelli) ^
^

ten to seventeen

,

scales,

arranged in three (rarely two or four) trans-

verse series between the rostral shield

and the apex.
a

Upper

surface of head covered with small smooth or

faintly keeled, subimbricate scales,

among which
;

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)

PLATE XIV

VIPERA LEBETINA
After SordelU

VIPERA AMMODYTES
A/ter Sordelli

ANCISTRODON HALYS
After SordelU

VIPERA
ocular by the

251

canthal scales, the second separated from the supra-

upper preocular.

Supraocular with
not

very convex outer border,
posteriorly

usually

extending

beyond the vertical of the eye. Ten to thirteen scales round the eye one or two vertical
;

series of scales

separate

the preoculars

nasal,
out.

which

is

single or rarely divided,

from the and hollowed

Two

series of scales

between the eye and the

labials.

Upper
usually
scales

labials eight to twelve, usually nine

or

ten,

Temporal
five

fourth and fifth below the eye. smooth or feebly keeled. Four or
in

lower labials

contact with the single pair of

chin-shields.
Scales in twenty-one or twenty-three (rarely twentyfive)

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
in females.

;

subcaudals 27 to 40

in males,

24 to 37

Coloration.

— Grey, pinkish-grey, brown, yellowishless

brown, or brick red above, with a more or
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 without 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,
the dorsal

or a lyre-shaped figure confluent with

252

VIPERID^
;

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
lip,

and under the eye
if

;

on the lower

the dark shade,

present, broken

up by

light

bars separated by two to four labial shields.
greyish or pink, powdered with

Belly

dark bluish-grey,
spots
;

brown or black, or with or without black and white
tail

lower surface of end of
Iris

orange or coral
a

red, rarely yellow.

golden.

Specimens with
(var.

straight

brown

vertebral

band flanked with triangular
steindachneri,

black

spots pointing

outwards

Werner) are known from Hungary, and resemble
certain

colour varieties of V.
rare.

berus

and V.

aspis.

Black specimens are very

The

following varieties are important geographical
in

forms occurring

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 halfj

rostral

appendage clad with ten
(rarely

to fourteen scales, in three

two or
Ventral

four) transverse series

between the
subcaudals

rostral shield

and the apex.
shields

Scales in twenty-one
to

rows.

149

158

;

30 to 38.
the lower

A more
lip,

or less distinct dark blotch on

involving five to seven labial shields

without complete interruption.

Lower

surface of

end of

tail

yellow.

VIPERA
Var. mevidionalis, Boulenger
:

253
Naso-rostral shield

never reaching the canthus rostrahs, and but rarely extending higher up than the upper border of the
rostral,

which

is

often as deep as broad or a
rostral

little

deeper than broad;

appendage clad with

Fig. 39

Side Views of Heads of V. ammodytes Var. meridionalis {b)

typica

{a)

and

CI

d

c

Fig. 40 Front Views of End of Snout, showing the Lepi(From Proceedings of the Zoological Society, 1903) Dosis.
a,

Form

typica;

b,

var. meridionalis

;

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

VIPEKID.E
or less distinct dark blotch on the lower
lip,

A more
tion.
Size.

involving five or six labial shields without interrup-

Lower

surface of end of

tail

yellow.

—This
The

Viper exceptionally attains a length
largest

of 3 feet.

male

in the British

Museum

measures
4 inches.

2 feet 6 inches, the largest

female 2 feet

In V. berus females grow to a larger size
;

than males
Hybrid.

in this species, as in V. aspts, the reverse
rule.

appears to be the

—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
is

shields.

The

rostral shield,
(5
:

which

a

little

deeper

than

broad

4),

ex-

tends above the level of the slightly raised canthus
rostralis,

which bears two
with
the

shields,

the second in
naso-rostral
it
;

contact

supraocular.

The

extends to the canthus rostralis, where
first

joins the

canthal and the lateral apical shield

one

series

of scales between the nasal shield

and the preoculars.
is

On

the

upper surface of the head the snout
with
fifteen

covered

subimbricate smooth scales,

in addition to the canthals

and

apicals.

A

frontal

shield

and a pair of

parietals are well developed,

VIPERA
although smaller than
in

255

an average V. hems; the
with the frontal, between
series

parietals are in contact

which and the supraoculars two
intervene.

of scales
berus,

The

supraocular,

as

in

F.

ex-

tends backwards considerably beyond the vertical
of the eye.

Eleven scales round the eye

;

of scales between the eye
scales smooth.

and the

labials.

two series Temporal
and
fifth

Upper

labials nine, fourth

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
with a few reddish-

surface of the head

lips pinkish,

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
red.

sides.

Tail

orange red below.

Iris

fire

Total length 2 feet 2 J inches
Distribution.

;

tail

—The

2f inches.

typical

form

is

known from

Northern Venetia, Austria-Hungary (Styria, Carinthia,

Southern Tyrol, Carniola,

Illyria, Istria, Croatia,

Slavonia, and eastward through Southern
to Transylvania), Dalmatia, Bosnia,

Hungary

Herzegovina,

Montenegro, Albania, and Servia.
1,300
feet, in

In the Alps up to
feet.

the Balkan Peninsula up to 7,500

The var. montandoni inhabits Roumania and

Bulgaria.

256

VIPERID^
var.

The

meridionalis

inhabits

Greece,

with

the

Archipelago, Asia Minor, and Syria.

The specimens from Transcaucasia
further
variety
(var.

constitute a

transcaucasiana,

Boulenger),

agreeing w^ith the var.
scutellation

montandoni in the rostral

and

in

the

number

of ventral shields

(150 to 156), but differing in the

markings on the
or
alternating
aspis.

back

;

these

consist

of

dark bars

paired dark spots, as in the typical form of V.

The dark and
of the tail
is

light

markings on the

low^er lip are

as in the typical V. ammodytes,

and the lower surface

pale yellow or greenish towards the

end.
Habits.
this

— Notwithstanding
is

its

name

ammodytes^
to

Viper
;

by no
hills

means
it

restricted

sandy

localities

on the contrary,

shows a predilection
It

for

dry stony

with low vegetation, and has
avoids thick

often been found climbing bushes.
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,
it

which

ascends to a considerable altitude,
diurnal, leaving
;

it

is

essentially

its

retreat only
it

when
stated

the sun shines

but in

warm

localities

is

to be principally nocturnal, appearing in

numbers

by moonlight.
the winter

The
mild
it

length of

its

period of hiber-

nation depends entirely on the climate, but
is

when

may

be seen about in mid-

winter whenever the sun shines.

The

poison of

VIPERA
this

257

Viper

is

of V. berus

common

in

stated to be more active than that and V. aspis, and, as the snake is very some parts of Austria and the Balkan

Peninsula, fatal accidents to

The food
and
in captivity

consists of small

also of lizards.

V.

man are frequent. mammals and birds, ammodytes does much better

than

its

European congeners, and takes

food more readily.

The hissing is louder than in F. herus and V. aspis, and it is often produced, on the approach of man, by specimens lying in such perfect concealment that
their presence w^ould not otherwise be suspected
this habit, like the rattling of the Crotalus,
is

evi-

dently detrimental to
to

the

species

in

its

relation

man.
This species

is extremely abundant in some parts and over 7,000 specimens were killed in a district of Southern Styria in the course of two According to Werner, it is the years (1892, 1893). commonest of all snakes in Bosnia and Herzegovina.

of Austria,

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, Llnnaeus
Viper, or Kufi

The Blunt - Nosed
Form.

— Short and heavy.

Snout rounded, obtuse,

usually with well-marked canthus, and the loreal

region slanting towards the mouth.
17

Eye

small, its

258
vertical

VIPERID^
diameter
in the
less

than

its

distance from the
large

mouth

adult.

Nostril
tail six

and directed

backwards.

Length of

(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
or

all

more
dis-

less

tinctly keeled,

rarely, smooth on the
or,

snout and forehead;
i i^,

41 (after Sordelli)

sevento
of
;

twelve longitudinal
eyes

series

scales

between the
four

(supraoculars included)

two
is

to

canthal

shields, of

which the anterior
as

the largest, and

may

be

regarded

a

supranasal.

Supraocular

narrow, usually broken up into two or more small
shields.

Twelve

to eighteen scales

round the eye
eye
of
series

two or three series of scales and the labials two or three
;

between the
vertical

scales

separate
is

the

preoculars

from

the

nasal,
out.

which

single

and often strongly hollowed

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
is

variable.

The
is

typical

form,

which alone
buff,

represented in

Europe, and was
grey, greyish
series

originally des'cribed

from Cyprus,

or pale brown, above, with

two dorsal

of darker spots,

which may stand

in pairs, alternate,

or unite to form cross-bars,
large dark spots or bars.

and a

lateral series of

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
is

tail

yellow.

The ground
In

colour of the young

pink or flesh-colour.

specimens from desert sandy regions in Asia and

North Africa the markings may be very
the snake being of a nearly uniform pale

indistinct,

buff.

In the var. mauritajtica, Guichenot, from Morocco

26o

VIPERID^
Algeria, the back has three series of very large

and

dark brown or reddish-brown spots, separated by a

network of the yellowish ground colour, or the
middle series
zigzag band.
in

may The

be transformed into a wavy or
scales in this variety are usually

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 4J feet.
Distribution.
is

— The' European habitat of V. lebetina
where
it

restricted to the Cyclades,

is

not un-

common on
where
it

the island of Tinos, and appears to be
It is

found also on Kimoli.
is

common on

Cyprus,

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
investi-

Bona, and
gations.

in Tunisia.

Its reported

occurrence in

Egypt has not been confirmed by recent
Habits.

—According
it is

to

M. Doumergue
this

—who

has

had ample opportunities of observing
Oran, where
rarely

Viper near

common

it is

a nocturnal reptile,

It inhabits moving about in the daytime. 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
On
Milos, Dr. de Bedriaga observed this
is

261

muchto

dreaded snake, the bite of which

probably as bad
russelli,

as that of its Indian ally, the Daboia, V.

occur frequently in gardens, and to crawl about
near houses in villages
observer has noted
a
after

sunset.

The same
closing of

sort

of valvular

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
in

to

Doumergue's obserto thirteen
in

vations in

Algeria,

the young, up

number, are born

May and

June.

Genus ANCISTRODON, Palisot de Beauvois

Head
loreal

distinct

from neck,

its

upper surface covered
;

with large shields, as in the normal Coluhridcd
pit
;

a

eye moderate

or

small, with

vertical

pupil.

Body moderately elongate

or short.

Scales

keeled (or smooth), with apical
or short.

pits.

Tail moderate

This genus

is

distributed over nearly the whole

of Asia, the eastern parts of the United States of

America, and Mexico and Central America.
of the Asiatic species just penetrates

One

into South-

Eastern Europe, and

is

the sole representative of

the Crotalinse in this part of the world.

262
28.

VIPERID.E
Ancistrodon halys,
Pallas's Pit- Viper

Pallas

Form.
flat

— Moderately elongate.

Head subtriangular,
;

or slightly concave above, swollen in the tem-

poral region, very distinct from neck or obtusely pointed, slightly turned

snout rounded

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.—^Kostral as deep as broad or slightly

broader than deep, just visible from above.
of

A

pair

internasals

and a pair
its

of

larger

prefrontals.

Frontal as broad as the supraocular, as long as or
a
little

longer than

distance from the end of the
little

snout, as long as or a

shorter than the rather

short parietals.

Supraocular extending beyond the
Loreal
labials.

vertical of the posterior border of the eye.
pit

between three

shields, separated

from the

Nostril between

two

nasals, the posterior of

which

ANCISTRODON
is

263

separated from the upper preocular by a loreal.
preoculars, one or

Two

two postoculars, and a sublargest.

ocular.

Three large lower temporals, anterior
labials

Upper
largest,

seven

or

eight,

third

and fourth
of small

third entering

the eye.

A

pair

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

with transverse series
less regular

young specimens, of darker spots or with more or

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
ings

less

produced on the nape

;

all

these markdark, light-

sometimes confluent.

A

broad,

edged band on the temple.

Lips whitish, speckled

264
with brown.

VIPERID.E
Lower
parts whitish,

more

or less

profusely speckled with grey or brown.

The horny
brown
or

shield terminating the tail usually dark

black at the end.
Size.

— This

species

rarely

reaches a length of
in

29 inches.

The

largest

specimen

the

British

Museum measures only From Distribution,

ig inches.

the north and east

coasts

of the

Caspian Sea, across Central Asia to
Yenissei, as far north as 51°.

the
it

Upper
is

In Europe

only

known from two

arid tracts

between the
viz.,

Volga and the Ural, near the Caspian Sea,
Saltan-Murat desert and the Induski
Habits,
hills.

the

— Nothing
its

has

been published

on

the

habits of this snake, but they are probably similar

more eastern relative, A, blomhoffi, Schlegel, which inhabits China and A. blomhoffi is said to be more or less Japan. nocturnal, although showing a predilection for
to

those of

near and

localities

well

exposed to the sun.
of
its

It

is

ovoviis

viparous.

The symptoms

bite,

which
as

rarely fatal to
All

man, are the same as

in the Vipers.
far

the species of

Ancistrodon, so

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
Acrochordince, 5

Adder, 230 iEsculapian Snake, 187
asciilapii {Coluber),

187

151 Ailurophis vivax, 217
4,

Aglypha,

Blind Snake, 144 Blood-supply, 77 Blunt-nosed Viper, 257 Body, 8 Boidcs, 4, 146 Boina, 4, 146
bosniensis {Vipera berus, var.),

236

Albinism, 39 Aldrovandi's Snake, 182 Altitudinal range, 118, 123

Brain, 73

Burrowing Snakes, gi
caruleus {Coluber), 233

A mhlycephalidcB,
Anal
shield, 17

5

ammodytes [Vipera), 249 Anal pockets, 83
Ancistrodon, 261 Ancistrodon halys, 262 Antitoxic serums, 66

Canker, 113
rostralis, 13 Captivity, 100 carbonarius {Zamenis), 172 caspim {Zamenis gemonensis, var.)

Canthus

Apical

pits, 15,

75

Arteries, 77 Ascariasis, iii asiamis [Zamenis gemonensis, var.),

172 Cat-Snake, 217 Caucasian Cat-Snake, 219
cervone {Elaphis), 182

Cestoda, 112
cettii

173 Asp Viper, 239
aspis {Vipera),
var.), ater

{Tropidonotus natrix, var.),

155
chersoides {Tropidonotus), 167 Classification, 2

239

astreptophorus {Tropidonotus natrix,

154
var.),

Cobra

di capello, 9

{Tropidonotus natrix,

Ccelopeltis,

207 208

156
atrovirens {Coluber), 170 aurolineatus {Coluber), iGy austriaca {Coronella), 197

Caelopeltis monspessulana ,

coUaris {Contia), 207

Coloration, 29 Colour, 29
Coluber, 181

Bacteria, 113
berus {Pelias), 221 berus {Vipera), 230

Bezoar stones, 141
bilineatus

{Tropidonotus

natrix,

var.), 154

Coluber Coluber Coluber Coluber Coluber Coluber

asculapii, 187 atrovirens, 170

aurolineatus, 167
caruleus, 233 dione, 185

dumfriesiensis,

200

265

266
Coluber Jiavescens, 187 Coluber hydrus, 160 Coluber itisignitus, 208 Coluber leopardinus, 191 Coluber lotigissimus, 187 Coluber quadrilineatus, 191 Coluber quaterradiatiis, 182 Coluber quatuorlineatus 182 Coluber riccioli^ 202 Coluber rubens, 204 Coluber sauromates, 183 Coluber scalaris, 194 Coluber torquatus, 152 Coluber viridiflavus, 170 Colubrida, 4, 150
,

INDEX
Disgorging, 99
Distribution, 118
Dolichosauria,
i

dorsalis {Pelias),

234

Drinking, loi
dumfriesieyisis [Coluber),

200

Dwarf Snake, 205
Economic
value, 138

Eggs, 85 Egg- eating Snakes, 53, 80 Egg-tooth, 89
Elachistodotiti a, 5 Elaphis cervone, 182

CohibrincB, 4, 151 concolor {Tropidonotus

tessellatus,

Elapina, 5 Elaps, 35 ^/s«^n {Coluber leopardinuSf var.),

var.), 162 concolor [Vipera berus, var.),

193

234

Embryo, 88
Epiglottis, 79

Coniophis, 7 Constriction, 96 Contia, 205 Contia modesta, 205

Eryx, 147

Eryx jaculus, 14^ European Whip-Snake, 170
External characters, 8 Exuviae, 20 Exuviation, 105 Eyes, II. 73
fallax {Tarbophis)

Coral-Snakes, 32, 34, 35 Coronella, 196 Coronella austriaca, 197
Coronella girondica, 202 Coronella italica, 197
CrotalincB, 5,

Coryphodonts, 58 220

Crotalon, 20 Crotalus, 21
cucullatus [Macroprotodon). 'zi^

False Smooth Snake, 213 217 Fangs, 55 Fascination, 104
,

Dahl's Whip-Snake, 177
^a/i/n (Zamenis), 177 DasypeltincB, 5

Fat-bodies, 81 Faunistic works, 130 Feigning death, 104
fitzingeri {Coronella),

Jiavescens {Coluber),

197 187

jiavescens {Tropidonotus tessellatus,

Defecation, 99
Definition, i Deglutition, 98 Dentition, 53

var.), 163 Flukes, 112

Desert Snakes, 36

Flying Snakes, 94 Food, 95

Form,
Gape,

Development, 88
Diacranterians, 58

8 Fossils, 6
II

Dicephalous young, 89 Digestion, 99
rf7"o«^

{Coluber), 185
5,

Dione Snake, 185
Dipsadomorphina,
151

Gastrosteges, 15 gemonensis {Zamenis), 170 Genital organs, 82
girondica {Coronella), 202

INDEX
Glaxiconiida^ 4 Glottis, 79

267
to the identification, 22

Key

Grass Snake, 152 Greek Blind Snake, 144

Kidneys, 80 Kufi, 257
Labial
pits,
i

Ground Snakes,

91

76
208

Lacerttlia,

Habits, 91 HEemogregarines, 112
Jiagenbecki

lacertina [Matrix),

Ladder Snake, 194
tessella-

{Tropidonotiis

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

Largest Snakes, 21 Larynx, 79 Lataste's Viper, 247 latastii [Viper a), 247 lebetina [Vipera), 257 Leopard Snake, 191
leopar dimes [Coluber), 191
lineaticollis
tus, var.),

[Tropidonotus 162

tessella-

Liver, 81

Hissing, 102 Homalopsina, 5 Hood, 9

Livery of the young. 38 Locomotion, 93 Long-nosed Viper, 249
longissitmis [Coluber),

187

Horse-shoe Whip-Snake, 179 Imgyi [Vipera aspis, var.), 244 Hybrids, 90
Hydyophiina, 5 hydrus [Coluber), 160 Hyoid apparatus, 42
iberiis

Loreal pit, 75 Loreal region, 12 Lungs, 78 Lycodonts, 58 Lymph-hearts, 78
Macroprotodon, 212 MacYoprotodon cuctdlatus, 213 macrops [ Vipera), 224 Marine Snakes, 92, 95 Markings, 29 lebetina^ mauritanica [ Vipera
var.), 259

[Tarbophis), 219

Iguanognathus, 54 Ilysiida, 4 Immunity, 66, 71
incevtus

[Tropidonotiis 4,viperimis,

van), 167 Incubation, 88 Intelligence, loi
insignitus [Coluber),

Melanism, 39 Mental groove, 13
mendionalis
[Vipera

ammodytes,

208

Integument, 8
Intestines, 79
Iris,

253 Mimicry, 35
var.),

37 Isodonts, 58
italica [Coronella),

modesta [Contia), 205 monspessulana [Coelopeltis) 208 ammodytes, montandoni [ Vipera
,

197

jacxilus [Eryx),

Jacobson's organ, 73 147 Javelin Sand-Boa, 147

var.), 252 Montpellier Snake, 208 Mortality from Snake-bite, 133

murorum
var.),

Mosasauria, 1 [Tropidonotus

natrix,

154

268
Natrix Matrix
lacertina,

INDEX
vulgaris,

208 152
,

Pupil, 12 Pythonina, 4

natrix {Tropidonotus)

152
quadrilineatus (Coluber), 191 quaterradiatus (Coluber), 182 quatuorlineatus (Coluber), 182
mo7ispessuvar.),

Neck, 9

Nematoda,

11

Neyodia, 152

Nervous system, 73
neumayeri
niger
[Ccelopeltis

Rattle, 20

lana, var.),

210

(Coluber longissimus,

188
nigrescens {Tropidonotus tessellatus,
var,), 162

Rattlesnakes, 20, 103 Rattling, 103 Reflex movements, 106

Nocturnal Snakes, 93 Northern Viper, 230
Nostrils, 13, 74

Relation to man, 133 Renard's Viper, 227 renardi (Vipera). 227

Replacement

teeth, 55

Reproduction, 82 Reptation, 93
Reptilia, i

(Esophagus, 79 Olfactory organ, 73
Ophidia.
i

Rhiptoglossa,

i

Opist/ioglypha, 4, 151

Organs of a

sixth sense, 75 Orsini's Viper, 221

202 Ring-Snake, 152 romanus longissimus, (Coluber
riccioli (Coluber),

var.),

188
tes-

Oviparous Snakes, 86
Oviposition, 85

rubens (Coluber), 204 riibro-maculosus (Tropidonotus
sellatus, var.),

Ovoviviparous Snakes, 85
Pairing, 84 Pallas's Pit- Viper, 262 Parasites, 107 Pelias berus, 221
Pclias dorsalis, 234

162

Rustling, 102

Sand-Boa, 147 Sand-Snakes, 91 Sand-Viper, 249
sardus (Zamenis), 172 sauromates (Coluber), 183
scalaris (Coluber), 194 scalar is (Coronella austriaca, var.),

Pelvic arch, 52
persa (Tropidonotus natrix, var.)^

154
persicus
picttiratus

(Zamenis

gemonensis,
natrix,

199
Scales, 14 schwoederi
(Coluber
leopardimis,
natrix,

var.), 172

(Tropidonotus

van). 155
Pit-Vipers, 75, 262 Poison apparatus, 62 Poison fangs, 55, 62 Poison glands, 62 Poisons, 66
prester (Vipera), 235 Proteroglypha, 4, 151

var.), 193 scutatus (Tropidonotus var.), 155

semimacidata
var.),

(Contia

modesta,

207

Sense organs, 73
seoanei (Vipera berus, var.),

Protozoa, 112
pseudaspis

[Vipera

berus,

var.),

235 Serotherapy, 66, 134 Serum treatment, 67 Shedding of the epidermis, 20
Shields, 14

236

INDEX
Sixth sense, 75 Skeleton, 40 Skull, 40 Smallest Snakes, 21
Typhlops, 144 Typhlops vermicularis, 144

269

Ureters, 81
Uroycltida, 5

Smooth Snake, 197
Snake-charmers, 139
Snake-stones, 141

Urosteges. 15
ursinii (Vipera),

221

Snake swallowing her young, 88
Snake-worship, 139 Snout, 12 Solenoglyphs, 56 Southern Smooth Snake, 202 Squamata, i
steindachneri

Veins, 77
veithi

[Loronella austriaca, var.),

199 Ventral shields, 15
vermicularis (Typhlops), 144

{Vipera

ammodytes,

Vertebrae, 50

252 Stomach, 79
var.),

Subcaudal

shields, 15

subfasciatus [Tropidonotiis natrix, var.), 155
subgriseus
var.), 189

{Colubey

lojigissimus,

Symoliophis, 7

Syncranterians, 58
Tail, 9

Tar bop his, 216
Tavbophis fallax, 217 Tarbophis iberus, 219 Teeth, 53 Tessellated Water-Snake, 160 tessellatas {Tropidonotns), 160 Thymus gland, 78 Ticks, 108

Vertebral column, 50 Vipera, 221 Vipera ammodytes, 249 Vipera aspis, 239 Vipera berus, 230 Vipera latastii, 247 Vipera lebetina, 257 Vipera ma crops, 224 Vipera prester, 235 Vipera renardi, 227 Vipera ursinii, 221 Viperida, 5, 220 Viperince, 5, 220 Viperine Water-Snake, 165 viperinus (Tropidonotus), 165 viridiflavus (Colubey), 170 Viscera, 77 vivax (Aibirophis), 217
vosseleri

(Tropidonotus

tessellatus,

Tongue, 74 Tongue-worms, 108
torquatus {Coluber), 152 trabalis {Zamenis gemo7iensis var.),
,

var.), 162 vulgaris (Natrix), 152

Warning

172

Trachea, 78
transcaucasiana [Vipera ammodytes,
var.),

Water- Snakes, 92, Whip-Snake, 170

coloration, 35, 36 94, 160

Worms, no
Xenopeltida, 5

256

Tree-Snakes,- 36, 92, 94

Trematoda, 112
Tropidonotiis, 152 Tropidonotus chersoides, 167 Tropidonotus natrix, 152 Tropidonotus tessellatus, 160 Tropidonotus viperinus, 165 Typhlopida, 4, 143

Zamenis, 170

Zamenis carbonarius, 172 Zamenis dahlii, 177 Zamenis gemonensis, 170 Zamenis hippocrepis, 179 Zamenis sardus, 172 Zoogeographical Regions, 119

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