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THE

RAY SOCIETY.

INSTITUTED MDCCCXLIY.

This volume is issued to the Subscribers to the Ray Society for

the Year 1S96.

LONDON

MT>cccxevir.

THE

TAILLESS BATRACHIANS

OB'

EUROPE.

BY

G. A. BOULENGER, F.R.S.

PART I.

LONDON:
PRINTED FOR THE RAY SOCIETY.

JIDCCCXCVII.

l'UINTED BY ADLAKD AND SON,
BARTHOLOMEW CLOSE, EC, AND 20 HANOVER SQUARE, W.

PREFACE.

Notwithstanding the great progress attained in our
knowledge of European Batracbians within the last
twenty years with regard to the distinction of species
and varieties, the study of their anatomy, life-histories,
and distribution, there exists at the present day no
work dealing with them as a whole from these different
aspects.

For twenty-five years a close student and collector
of these animals, which have always exercised an
extraordinary fascination on my mind, I have often
wished for an opportunity of supplying such a de-
sideratum, by utilising the enormous material which
had gradually accumulated in the literature, my own
notes, and the unrivalled collection in the British
Museum.

Thanks to the Council of the Ray Society, who, on
the special recommendation of Sir W. H. Flower,
acceded to my request, I have at last the satisfaction
of seeing my hopes realised, and, owing to the talent
of my artistic collaborators, Messrs. P. J. Smit and
J. Green, in a manner which fully satisfies my aspira-
tions. May those, few as they still are, who share
my fondness for this group of auimals endorse my
opinion of the beautiful and accurate illustrations
which are one of the principal features of this work,

11 PREFACE.

and derive from the perusal of these pages one-tenth
of the pleasure it has given me to write them ; I shall
feel amply rewarded for the trouble I have taken.

I would also express a hope that a little book of
this kind, embodying in a concise form the result of
much study, at present scattered in a multitude of
publications in some ten different languages, may
have the effect of stimulating interest to a subject
that has been too much neglected, and in the cultiva-
tion of which new workers will find much to repay
their efforts, especially if applied in other regions of
the globe, which, though much richer in Batrachians,
have as yet yielded little or nothing to our knowledge
of the life-histories.

This work has been planned with the view of
assisting the beginner as well as of affording new
information to the advanced student, and the synoptic
treatment has therefore been frequently resorted to in
the Introduction; whilst all descriptions in the sys-
tematic part are strictly comparative. Technical
terms have not been avoided, but the numerous figures
in the text should render them easily intelligible to
the beginner.

At the desire of the Council of the Ray Society this
volume is issued in two parts. The second part,
continuously paged and with bibliographical and alpha-
betical indexes, is to follow in a few months.

The plates, twenty-four in number, have been exe-
cuted by Mr. P. J. Smit and printed by Messrs.
Mintern Brothers. With two exceptions, all the
coloured figures have been taken from living speci-
mens. Five of the plates (I, II, III, XVI, XVII)
have already appeared in the ' Proceedings of the

PREFACE. Ill

Zoological Society' (1884, 18S5, 1891), to illustrate
papers of mine, and have been copied with the sanc-
tion of the Committee of Publication of that Society.

The figures in the text have been drawn by Mr. J.
Green, and reproduced by the Typographic-Etching
Company. Most of them are original ; if copied from
other works, the source has in every case been referred
to. A few cuts illustrating a paper by me on Tadpoles
in the c Proceedings of the Zoological Society ' for 1891,
and another by Dr. Ridewood on the development of
the hyoid, published in 1897, have been reproduced
by permission, for which I beg to tender my acknow-
ledgments to the Committee of Publication of the
Zoological Society.

1 also wish to publicly express my thanks to Sir
W. H. Flower, K.C.B., and to the Rev. Professor
Wiltshire for the interest they have shown in the
publication of this work; and to the various friends
and correspondents who have assisted me with notes
or material, especially Dr. W. G. Ridewood, to whom I
am indebted for many hints and preparations ; M. F.
Lataste, who has placed his private collection at my
disposal ; Dr. J. de Bedriaga, Count M. Peracca, Mr.
F. A. Moller, Dr. F. Werner, and Herr W. Wolter-
storff, who have supplied me with an abundance of
living specimens from France, Italy, Portugal, Austria,
and Germany ; and Messrs. Chevreux and Doumergue,
through whose kindness I have been able to make
observations on a large number of living specimens of
Discoglossus from Algeria.

G. A. B.

British Museum (Natural History) ;
October 28th, 1897.

THE

TAILLESS BATRACHIANS

OF

EUROPE.

INTRODUCTION

I. Classification ....
II. External Characters

III. Integument

IV. Dermal Secretion
V. Skeleton

VI. Viscera

VII. Habits

VIII. Voice

IX. Pairing and Oviposition
X. Spermatozoa ....

XL Eggs

XII. Development and Metamorphosis

XIII. Tadpoles

XIV. Hybrids

XV. Geographical Distribution .

page

2

9
21
30
33
49
57
61
64
75
78
85
99
112
115

A

INTRODUCTION.

I. Classification.

For many years the Tailless Batrachians (Ecaudata
s. Anuha) were classified in a very unsatisfactory
manner. The genera Bufo and Hyla were, it is true,
always regarded as the types of distinct groups: but
Pelodytes, Pelobates, Discoglossus, Bombinator, and
Alytes were placed with Bana, or in separate families
in most unnatural associations.

Dumeril and Bibeon, in the eighth volume of their
standard work, ' Erpetologie generale,' published 1841,
after dividing the order into two sub-orders, Phanero-
glosses and Phryna glosses (=■ Aglossa, the two exotic
genera Pipa and Xenopus), a primary division first
introduced by Wagler in 1830 and which has stood
the test of time, combined the Phaneroglossal genera
into three families — Raniformes, Hylaefovmes, Bufoni-
form.es ; the two latter containing each a single Euro-
pean genus, whilst under the former all others were
arranged in a series without further subdivision.

So unnatural an arrangement evoked criticism from
all who were acquainted with the life-histories of the
European forms ; and an excellent French observer of
these animals, A. Thomas, in a paper published in the
'Annales des Sciences naturelles,' 1854, dwelt upon
the correlation which exists between the shape of the
pupil and the mode of amplexation duriug parturition,
and proposed to group together on the one hand those
genera that have a horizontal pupil and an axillary
embrace, on the other those that combine a vertical
or triangular pupil with a lumbar embrace. These
divisions were later provided with names by Beuch in
1863 (Plagloglena, Orthoglena), and by de l'Isle in
1877 (Alamplexes, Tngidnamplexes) .

How exaggerated the importance attached to this
correlation, which, besides, holds good only for the
European forms, is now apparent to all. Yet the ar-
rangement proposed by Thomas was a decided advance

CLASSIFICATION. 6

on that of the ' Erpetologie generale,' and on the views
of J. Muller (1832), who overrated the taxonomic
importance of the structure of the ear, and inspired
the classification followed by Stannius in the ' Zoo-
tomie der Amphibien,' published in 1854.

The faults of the latter classification were further
developed by Gunther in 1858 (' Oat. Batr. Sal.'),
whose avowedly artificial scheme has been followed by
Fatio in 1872 and by De Betta in 1874.

Cope's epoch-making classification in 1865 (' Nat.
Hist. Review'), based on a correct appreciation of
the osteological characters, placed the matter on a
sound footing, and his groupings, slightly modified
in 1867, received confirmation, so far as European
genera are concerned, from Lataste's study of the
larval characters in 1878 and 1879. In revising the
classification in 1882 (' Cat. Batr. Ecaud.') I could
introduce but slight improvements to Cope's scheme,
as may be seen from the arrangement followed in this
work, in which the various groups are placed in
ascending order.

Principal Schemes of Classification.

1. Dcmeeil and Bibron, 1841,

Tongue absent : Phrynaglosses.

Tongue present : f toothed ;

Phaneroglosses.J digits

Upper jaw |

vtootkless

2. Stannius, 1854.

§ ("absent : Aglossa.
bo |

s i f absent

Mpv

without disks : Baniformes-

nth. disks : Hyleeformes

Bufoniformes Bufo.

fBana.

Discoglossus.

Pelodytes.

Alytes.

Pelobates.

Bombinator.

Hyla.

ch i present

P

GLOSSA

P«™°| present: (J±32£J-— * »*

Tympanum

\ pres<

[ die

_ ontna

sterni I p resent -.Banina
dilated at the end . Hyloidea

Pelobatoidea\^elolia-teSl

(Bombinator.

I Bufo.

(Alytes.
Bana
Hyla.

1NTE0DUCTI0N.

3. Gunthke, 1858.

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classification. 5

4. Bruch, 1863.

'vertical : Orthoglena . . . Pelobatides-

(Pelobates.
I Bombinator,

1 Pelodytes.
Pupil \Alytes.

(Hyloides . Hyla.
.round or horizontal : Plagioglena j Banoides . Bana.

\Bufonides . Bufo.

This arrangement is merely the application of the
views expressed by Thomas in 1854.

5. Cope, 1865.

Aglossa.

Bufoniformia . . Bufonidse . Bufo.

{Discoglossus.
Alytes.
Bombinator .

Scaphiopodidte [pf^ytes

Sylidse . . Hyla.
Raniformia ...... Banidse . . Bana.

The Bufoniformia correspond to the Bufoniformes
of Dumeril and Bibron ; the Arcifera are separated
from the Baniformia, which agree in the presence of
teeth, by the structure of the pectoral arch ; and the
Discoglossidae are defined for the first time by the
opisthoccelous vertebree provided with autogenous
ribs. Some further improvements were introduced in
1867, when the Bufoniformia were split into two
groups, based on the structure of the pectoral arch,
that group corresponding to the Baniformia among
the toothed forms, being named Firmisternia, a term
the signification of which I have extended to embrace
all non-arciferous forms, whether toothed or not — an
improvement since accepted by Cope himself.

6. Mivart, 1869, in one of his classical papers pub-
lished by the Zoological Society of London, endea-
voured to combine Giinther's and Cope's classifications,
but his arrangement does not differ materially from

6 INTRODUCTION.

the former author's so far as the grouping of the
European genera is concerned.

7. Lataste, 1879.

LiEVOGYRINID^E S. PrOCCELIPJE :

Vertebrae proccelous ; tadpole with"
sinistral spiraculum.

Mediogyrinid^ s. Opisthoccelid^

'Ranidse . Rana.

Hylidse . Hyla.

Bufonidse . Bufo.

Pelobatidie

(Pelobates.
\Pelodytes.

DM: ( j,. i •■, (Discoglossus.
Vertebrae opistboccelous ; tadpole^ " \Bombinator.

with medial spiraculum. \Alytidse . Alytes.

The Alytidm are separated from the DiscogJossidse
on account of the shape of the pupil and the aberrant
mode of parturition and development.

The classification adopted in this work is shown in
the following synopsis, which also includes a list of
all the species represented in Europe.

Order ECAUDATA.
Sub-order Phaneeoglossa.
Eustachian tubes separated ; tongue present.

Series A. — Aroifbea.

Coracoids and prsecoracoids connected by an arched
cartilage (the epicoracoid), that of the one side over-
lapping that of the other.

Family 1. — Discoglossid^e.

Vertebrae opistboccelous ; short autogenous ribs
attached to the anterior diapophyses ; diapophyses
of sacral vertebra dilated ; upper jaw toothed.

CLASSIFICATION. 7

1. Discoglossus, Otth, 1837.

1. pictus, Otth, 1837.

2. Bombinator, Merr., 1820.

2. igneus, Laur., 1768.

3. pachypus, Fitz., 1838.

3. 4Zyfe«, Wagl., 1830.

4. obstetricans, Laur., 1768.

5. cisternasii, Bosca, 1879.

Family 2. — Pelobatid^:.

Vertebrae proccelous ; * no ribs ; diapophyses of
sacral vertebra strongly dilated; upper jaw toothed.

4. Pelodytes, Fitz., 1838.

6. jmnctatus, Daud., 1802.

5. Pelobates, Wagl., 1830.

7. fuscus, Laur., 1768.

8. cultripes, Cuv., 1829.

Family 3. — Bufonid^:.

Vertebrae procoelous ; no ribs ; diapophyses of
sacral vertebra dilated ; jaws toothless.

6. Bnfo, Laur., 1768.

9. vulgaris, Laur., 1768.

10. viridis, Laur., 1768.

11. calamita, Laur., 1768.

Family 4. — Hylid^.

Vertebrae procoelous ; no ribs ; diapophyses of
sacral vertebra dilated ; upper jaw toothed ; terminal
phalanges claw-shaped.

7. Hyla, Laur., 1768.

12. arborea, L., 1766.

* In the European genera.

8 INTRODUCTION.

Series B. — Firmisternia.

Coracoids firmly united by a simple epicoracoid
cartilage ; prsecoracoids resting with their distal
extremity upon the coracoids, or connected with the
latter by the epicoracoid cartilage.

Family 5. — Ranid^;.

Vertebras proccelous ; no ribs; diapophyses of
sacral vertebra cylindrical ; upper jaw toothed.

8. Bana, L.. 1766.

13. esculenta, L., 1766.

14. arvalis, Nilss., 1842.

15. earner anis Blgr., 1886.

16. temporaria, L., 1766.

17. grxca, Blgr., 1891.

18. iberica, Blgr., 1879.

19. latastii, Blgr., 1879.

20. agilis, Thorn., 1855.

We are thus now acquainted with twenty well-
established species, as against twelve and eighteen
enumerated in the two preceding general works on
the Batrachians of Europe, viz. Schreiber's ' Herpe-
tologia Europasa,' 1875, and be Bedriaga's ' Frosch-
lurche Europas,' 1889.

Most of the additions made during the last twenty
years fall to the genus Rana, and are in fact dismem-
berments of the Linnean species R. temporaria. That
the forms here admitted as species are fully entitled
to that rank, no one acquainted with their histories
will contest for a moment. Nor, among recent acces-
sions to our knowledge, is the distinction of the two
species of Bombinator or the two of Alytes likely to
meet with any opposition. Bat the rank to be
assigned to the principal forms of Hyla arborea and
JRana esculenta, regarded by me as only varieties,
perhaps remains an open question in these days of
extreme multiplication of species.

EXTERNAL CHARACTERS. 9

II. External Characters.

As these Batrachians have no neck, the head passes
directly into the body, and, except in Pelobates, where
the skin is closely adherent to the rugose skull, the
demarcation is consequently difficult to trace. By
drawing a line connecting the articular extremities
of the mandible we obtain approximately the exact
length of the head, the greatest width being at the
commissures of the jaws. The head is more or less
flattened, with prominent eyeballs covered above by
the upper eyelid, which is continuous with the rest of
the cephalic integument. The upper surface of the
head may be limited on each side by an angular line
extending from the tip of the snout to the upper

Fig. 1.

Upper views of heads of (a) Rami temporaria and
(b) Bombinator igneus.

eyelid, the cantkus rostralis, continued behind as a
ridge or fold, the supra-temporal ridge (Fig. 1, a).
In some forms, such as Bombinator igneus (Fig. 1, b),
the snout is simply arched from one lip to the other,
and there is no trace of a canthus rostralis. The side
of the snout below the canthus rostralis is called the
loreal rep;ion. The lenp-th of the snout is taken bv
applying the points of the compasses to the anterior
border of the (bony) orbit and the very tip of the
snout, on the median line.

The nostrils are small valvular openings, situated
between the end of the snout and the eves, alter-

10 INTRODUCTION.

nately opening and closing as the throat is drawn in
and swollen out during respiration.

The eye is large, lateral (Fig. 1, a), or directed
upwards and outwards (Fig. 1, b), and protected by
three lids : the upper thick and coloured, the lower
rudimentary and immoveable, and the nictitating
membrane, transparent or partially pigmented, which
ascends over the eyeball to meet the upper lid. The
eye can be covered without the vision being much
impaired by elevation of the nictitating lid, as when
the frog is under water, or completely closed when
the eyeball is drawn in and the upper eyelid descends
to meet the lower. The iris is brilliantly coloured
with metallic pigment, golden, silvery, or bronzy,
with or without an admixture of red or black. The
pupil when fully distended is round and large, but
when contracted affects various shapes, which afford
important characters for the distinction of the genera.
Thus the contracted pupil of Bana, Hyla, and Bufo
is horizontal, oval with an angle in the lower border

Fig. 2.

® © 0

B C D

Different forms of pupils. A. Bombinator pacliypus. B. Alytes
obstetricans. c. Pelobatesfuscus. r>. Bana arvalis. E. Bufo
vulgaris.

in Bana (Fig. 2, d), and Bufo viridis and calamita, with
an upper and lower angle in Hi/la and Bufo vulgaris
(Fig. 2, e). That of Discoglossus and Bombinator may
be described as roundish or subtriangular according
to specimens, the lower angle being always distinct,
but the upper border sometimes convex, when the

EXTERNAL CHARACTERS. 11

pupil appears nearly round, or straight when the
snbtriangular form obtains (Fig. 2, a). In some
specimens of Bombinator the upper border is even
emarginate, so as to produce the shape of a heart ; and
when extremely contracted the pupil divides into
three branches, affecting the shape of a Y (Fig. 2, a)
This type of pupil approaches the vertical, and in
some specimens of Discoglossus the much-contracted
pupil is in fact a little deeper than broad. In Alytes,
Pelodytes, and Pelobates (Fig. 2, b, c) we find a regular
cat's pupil, vertically elliptical or club-shaped, linear
when fully contracted.

The tympanum, or drum of the ear, is absent in
Bombinator and Pelobates. When present it may be
concealed under the skin, as in some specimens of
Discoglossus, Pelodytes, and Bufo vulgaris, or appear
on the temple behind the eye as a round or oval disk
covered with thin skin (Fig. 1, a, p. 9).

The mouth is large, and cleft to beyond the eyes.
The jaws are edentulous in Bufo ; the upper is armed
with numerous closely-set teeth in the other genera —
ten to fifteen in each prasmaxillary, forty to fifty in
each maxillary. The teeth have sharply-pointed,
slightly-hooked crowns with long shafts applied to
the inner side of the prgemaxillary and maxillary
bones. The palate (Fig. 3, p. 12) is pierced by two
pairs of orifices, the choanse, or inner openings of the
nostrils anteriorly, and the openings of the Eusta-
chian or auditory tubes, situated near the commis-
sures of the jaws; the Eustachian tubes are extremely
fine, or sometimes even indistinct, in Bombinator. In
all the genera except Bufo there are two groups or
series of teeth, each implanted on an eminence of the
vomerine bones ; these vomerine teeth are situated
between or behind the choanse, as shown on p. 12,
Fig. 3, a, c, d.

The tongue, thick and papillose, flat, is attached in
front and in the middle, and the posterior portion, by
being bent over and rapidly thrust out of the mouth,

12

INTRODUCTION.

acts as an organ of prehension in most tailless Batra-
chians ; in the Disco gloss Idas, however, as in the
newts and salamanders, the tongue is entirely or
nearly entirely adherent to the floor of the mouth,
and the prey is seized by the jaws. The tongue
serves also as an organ of taste, for although frogs
seize almost any moving object, they will reject
before deglutition anything that is noxious to them,
as may be witnessed on offering a frog a brandling
or manure- worm (Alloloboplwra foetida), a ladybird
beetle, or a young Bombinator. Before being swal-
lowed, the food, if it be a small mollusc, crustacean,
or hard beetle, is crushed between the tongue, the
depressed eyeballs, and the vomerine teeth ; if the
latter be absent, as in Bufo, the sharp, sometimes

Fig. 3.

D

Open mouths of — A. Discoglossus pictus B. Bufo ealamita. c. Ilyla
arborea. D. Rana temporaria. Showing the shape of the tongue
and the disposition of the choanae, Eustachian tubes, and vomerine
teeth.

serrated edge of the palatine bones supplies their
function. The tongue is circular and entire in the

EXTERNAL CHARACTERS. 13

Discoglossidae (Fig. 3, a) ; circular and entire or
feebly nicked behind in the Pelobatidse and Hyla
(Fig. 3, c), resembling a mushroom when thrust out;
elliptical or pyriform and entire in Bufo (Fig. 3, b) ;
oval and forked behind in Bana (Fig. 3, d). In the
two latter genera the extensibility of the tongue is so
great that a large specimen is able to seize a prey two
inches distant from it.

Owing to the nature of the skin and the absence of
true ribs, the shape of the body varies enormously
according*- to the decree of moisture of the sub-
cutaneous sacs, the inflation of the lungs, or the
condition of the ovaries, and all measurements that
are not based on the bony framework are useless for
systematic purposes. The frogs of the genus Bana
appear hump-backed in a squatting posture, an angle
being produced at the articulation of the cylindrical
sacral diapophyses with the ilia; the other genera,
with flattened and dilated sacral diapophyses, have a
rounded back when at rest, as may be seen on com-
paring a true frog with a toad or tree-frog. The
vent, or cloacal opening, is a small rounded orifice at
the very posterior extremity of the body, above and
between the thighs.

The fore limb is divided into a brachium or arm, an
antebrachium or forearm, and a manus or hand with
four functional fingers, of which the third is the
longest. The length of the first finger as compared
to the second is often used as a specific character.
In order to preclude misunderstandings, it is well to
remark that when the first finger is stated to extend
as far as or beyond the second, the two are taken to
meet halfway. A rudimentary pollex is often indicated
externally by a tubercle at the base of the inner finger
(Fig. 4, a).

The hind limb is longer, usually very much longer,
than the fore limb, and divided into four distinct
segments — the femur or thigh, the crus or tibia, or
leg, the tarsus, and the pes or foot. Anatomically,

14

INTRODUCTION.

the tarsus is, of course, part of the foot proper; but
as it here forms a distinct segment, the term foot is
used in a restricted sense. The foot is measured from
the base of the metatarsal tubercle ; it has five elon-
gate toes, gradually increasing in length to the
fourth, the fifth being again shorter. There is in
addition a rudimentary sixth toe, the so-called prse-
halhtcV, conspicuous externally in the form of a
tubercle or spur at the base of the inner toe. In
Pelobates (Fig. 4, b) this prsehallux or inner metatarsal
tubercle acquires a very great development, is covered
by a horny sheath with sharp cutting edge, and serves
as a shovel for dio-o-ino; in the soil. There is often
another tubercle on the sole, at the base of the
fourth toe; this so-called outer metatarsal tubercle
(Fig. 4, o) is merely a thickening of the integument.

Fig. 4.

A. Hand of Alijtes obstetricans. b. Foot of Pelobates fuscus.
of Bufo calamita (lower views).

c. Foot

Other dermal tubercles are usually present, more or less
developed, single (Fig. 6, a, p. 16) or paired (Fig. 4, c),
under the digits at the articulations between the
phalanges; they are called subarticular tubercles.
The fingers are free except in Hyla arborea, in which
they are provided with a rudimentary web. The toes
are more or less webbed or bordered by membranes in
all our Batrachians, but the extent of the web varies
greatly according to the species, and is usually more
developed in males than in females, especially during
the breeding season. The web is usually smooth and
transparent; it is thicker, and often warty, especially
towards the margin, in Bafo.

EXTERNAL CHARACTERS.

15

In our single representative of the arboreal type, the
tree-frog — Hyla arbor ea — the extremities of the fingers
and toes are expanded into adhesive disks (Fig. 6, b),
which assist the animal in climbing on vertical smooth
surfaces. These disks do not act as suckers, as was
once believed, but adhere by rapid and intense pressure
of the distal phalanx and special muscles upon the
lower surface, which is at the same time provided
with numerous glands producing a viscous secretion.
The upward rotation of the terminal phalanx, the
swollen base of which is hinged upon an interarticular
cartilage situated under the extremity of the penulti-
mate phalaux, and can be raised or lowered like the
claw of a cat, is easily to be observed on a living-
specimen. When the disk does not adhere (Fig. 5), its
upper surface shows a short ridge produced by the ciaw-
like terminal phalanx, and the lower surface is convex
with some longitudinal grooves. During adhesion,
on the other hand, the claw-like phalanx is no longer
visible on the upper surface, but is indicated by a
groove, whilst the lower surface is flat and expanded.

Fig. 5.

Section through the extremity of the third toe of Ihjla arborea,
the bones being indicated by oblique bars, ph2, x>h3. Second
and third phalanges, ia. Interarticular cartilage, d. Ad-
hesive disk. st. Subarticular tubercle.

The adhesion of tree-frogs to smooth vertical sur-
faces is, however, not effected entirely through the
above-mentioned organs, their function being supple-
mented by the suctorial action of the belly, such as
is developed, though to a lesser degree, in the young
of most Batrachians which, devoid of digital expan-

16

INTRODUCTION.

sions, are nevertheless able to slowly ascend a pane of
glass.

In most of our genera the fourth and fifth toes are
cleft almost to the base, and the metatarsal portions of
them are separated by the web. In two genera, Bufo
and Hyla, this is not the case; the metatarsals of the
two outer toes are bound together by the common
integument, the angular divergence of the fifth toe
proceeding only from the basal phalanx, as may be
seen from the following figures (Fig. 6) representing
the foot of the common frog and that of the tree-
frog.

Fig. 6.

A B

Lower surface of foot of JRana temporaria (a) and Hyla arborea (b).

The tree-frop; differs from other Batrachians in
having the articulations of the limbs, especially the
wrist, marked above by a strong fold, reminding one
of the arms and legs of a jointed papier-mache doll.

The proportions of the hind limbs to the body are
best appreciated by bending the limb forwards against
the side, and ascertaining the position of the tibio-
tarsal or tarso-metatarsal articulation ; a certain allow-
ance must, however, be made in cases when the abdo-
men of a female is much distended with ripe ova. A
convenient method of appreciating the length of the
tibia as compared to the femur is to fold the limb so
as to place the former at a right angle to the axis of
the body, as shown in the accompanying figure (Fig. 7) ;
in which position the heels (tibio-tarsal articulation)
meet when the tibia is nearly equal to the thigh, fail
to meet when shorter, or overlap when longer.

EXTERNAL CHARACTERS. 17

Fig. 7.

c

Limbs of Hyla arborea, showing variations in the proportion
of the tibia. The dotted line indicates the middle line of
the body.

The great development of the hind limbs enables
these Batrachians to proceed by leaps ; there is, how-
ever, one exception among the European forms, viz.
Biifo calamita, in which the limbs are so short as to
render hopping impossible, and this toad progresses at
a running pace when pursued. The other extreme is
furnished by Bana agiiis, which owing to its extremely
lone; hind limbs is able to cover a distance of six feet
at one leap. The aquatic habits of a Batrachian can,
as a rule, be gauged by the extent of the web between
the toes : thus Bvfo calamita and Ah/tes, which have
only rudimentary webs, are very bad swimmers, and do
not venture far from the shore ; whilst Bana esculenta
and Bombinator, with their broadly- webbed toes, spend
nearly the whole of the active period in the water.
But there are exceptions : B debates, the toes of
which are completely webbed, sojourns but a very
short time in the water ; it is true that in this case
the web is probably of use for pushing aside the soil
in the process of digging.

The following synopsis of the species is based
entirely on external characters, and applies to both
sexes.

I. Tongue circular, entire, adherent, or slightly
free behind ; vomerine teeth behind the level of
the choanal ; first finger shorter than second
(Discoglossid^e).

18 INTRODUCTION.

A. Pupil round, subtriangular, or corcliform.

1. Vomerine teeth in long transverse series;

tympanum distinct or concealed under the
skin .... 1. Discoglossus pictus.

2. Vomerine teeth in two groups; no tym-

panum.
Tibia shorter than the foot

2. Bombinator igneus.
Tibia as lone: as or a little longer than the
foot .... 3. Bombinator pachypus.
p.. Pupil vertical ; vomerine teeth in short trans-
verse or slightly oblique series ; tympanum
distinct.
Three palmar tubercles ; fourth finger as long
as or slightly shorter than second

4. Alytes obstetricans.
Two palmar tubercles ; fourth, finger much
shorter than second 5. Alytes cisternasii.

II. Tongue circular, entire or slightly notched, and

free behind ; vomerine teeth between the

choanal ; pupil vertical (Pelobatid.e).

a. Tympanum distinct or concealed under the

skin ; toes webbed at the base and bordered ;

inner metatarsal tubercle small, soft

(I. Pelodytes punctatus.
p,. No tympanum ; toes nearly entirely webbed ;
inner metatarsal tubercle large, hard, com-
pressed, sharp-edged.
Interorbital space and occiput convex ; meta-
tarsal tubercle yellowish or pale brown

7. Pelobates fuscus.
Interorbital space and occiput flat; metatarsal
tubercle black . 8. Pelobates cultripes.

III. Tongue elliptical or pyriform, entire, free behind ;
no teeth ; pupil horizontal (Bufonid^e).
Toes at least nearly half webbed, with paired
subarticular tubercles ; no tarsal fold ; inter-

EXTERNAL CHARACTERS. 19

orbital space at least as broad as the upper
eyelid 9. Bufo vulgaris.

Toes at least nearly half webbed, with single
subarticular tubercles; a tarsal fold; fingers
very obtuse ; interorbital space narrower than
the upper eyelid . 10. Bufo viridis.

Toes webbed at the base, with paired subarticular
tubercles ; a tarsal fold usually present ; inter-
orbital space narrower than the upper eyelid ;
hind limb not, or but slightly, longer than head
and body . . . . 11. Bufo calamita.

IV. Tongue cordiforra, free behind; fingers and toes
dilated at the end into distinct disks ; fingers
webbed at the base; vomerine teeth between
the choana3 ; pupil horizontal (Hyltd^e).

12. Hyla arbor ea.

V. Tongue forked and free behind; pupil horizontal

(RANIDiE).

a. Vomerine teeth between or extending slightly

behind the level of the choanse; interorbital
space not more than half as broad as the
upper eyelid . .13. Bana esculenta.

b. Vomerine teeth extending behind the level of

the choana3 ; interorbital space at least one-
half the width of the upper eyelid.
1. Tibio-tarsal articulation rarely reaching the
tip of the snout, and never beyond ;
distance between the dorso-lateral folds
five to seven times in the length from
snout to vent.
Inner metatarsal tubercle large, hard, compressed,
measuring one-half to two-thirds its distance
from the tip of the inner toe ; first finger
extending beyond second ; tibia shorter than
the fore limb . . . 14. Bana arvalis.
Inner metatarsal tubercle small, soft, oval ; first
and second fingers equal, or first extending

20 INTRODUCTION.

slightly beyond second ; tibia as long as or
slightly shorter than the fore limb

15. Rana camerani.
Inner metatarsal tubercle small, soft, oval; first

finger extending beyond second ; tibia con-
siderably shorter than the fore limb

16. Rana temporaria.

2. Tibio-tarsal articulation reaching the tip of
the snout or beyond ; tibia nearly as long
as the fore limb; distance between the
dorso-lateral folds contained four to five
and a half times in the length from snout
to vent.

a. Tympanum not more than two-thirds the

diameter of the eye, distant from it.

Distance between the nostrils greater than the
interorbital width ; first and second fingers
equal, or first extending slightly beyond second;
inner metatarsal tubercle nearly half the length
of the inner toe, as long as the diameter of the
tympanum . . . 17. Rana grseca.

Distance between the nostrils greater than the
interorbital width ; first and second fingers
equal, or first extending slightly beyond second ;
inner metatarsal tubercle one-third the length
of the inner toe, shorter than the diameter of
the tympanum . . 18. Rana iberica.

Distance between the nostrils not greater than
the interorbital width ; first finger extending
beyond second; inner metatarsal tubercle one-
third the length of the inner toe, shorter than
the diameter of the tympanum

19. Rana, latastii.

b. Tympanum two-thirds to five-sixths the

diameter of the eye, and close to it ;
first finger extending beyond second ;
inner metatarsal tubercle very promi-
nent, but moderately large.

20. Rana agilis.

INTEGUMENT.

21

III. Integument.

The integument of the Bcaudata differs from that
of other Batrachians in the slight attachment to the

Fig. 8.

Rana temporaria, showing the dermal attachments and the dis-
position of the lymph- sacs, upper and lower views. The position
of the lymph-hearts is indicated by asterisks.

a. Abdominal sac.

b. Brachial ,,

c. Crural „

d. Dorsal ,,
/. Femoral „

g. Gular sac.

if. Interfe moral ..

I. Lateral ,,

p. Pectoral ,,

sb. Subbrachial sac.
sbp. Subplantar ,,
sf. Supra-femoral ,,
sp. Supra-plantar ,,

muscles, thus rendering the skinning of a frog a most
easy operation. But the body is not enclosed in a
loose sac, as is often stated in popular descriptions ;
there are definite lines of attachment (Fig. 8) by
means of connective tissue, which form the demarca-

22 INTRODUCTION.

tions between the lymph-sacs that exist on both the
dorsal and ventral surfaces immediately beneath the
integument.

The lymph, fed by the moisture absorbed through
the skin, is pumped into the veins by two pairs of
lymph-hearts, contractile muscular sacs, the anterior
of which are situated in the muscular tissue close to
the transverse processes of the third vertebra, the
posterior on the sides of the urostyle near its extre-
mity ; their pulsations may be observed on the back
of the living frog.

The number aud extent of the lymph-sacs varies in
the different genera. In Rana and Pelodytes there
are 22, 4 unpaired and 9 pairs. The dorsal sac
extends from the tip of the snout to above the vent,
and is limited by the line of attachment which corre-
sponds to the canthus rostralis, the supra-orbital
border, the supra-temporal fold, and the glandular
dorso-lateral fold. The lateral sac is limited below by
a band on each side of the belly, which may be marked
externally by a slight groove ; the abdominal surface of
the head and body is divided into three sacs, gular,
pectoral, and abdominal, with partitions across the
praacoracoid and coracoid regions. The further sacs
(brachial, subbrachial, femoral, supra-femoral, inter-
femoral, crural, supra- and sub-plantar) belong to the
limbs. In Hyla there is besides a close areolar attach-
ment of the abdominal integument. The Discoglos-
sidse differ in the absence of coracoid attachment,
or if such exists, as in Discoglossus, it is widely
interrupted on the middle line ; there is, therefore,
no distinct pectoral lymph-sac. In Pelohates the skin
of the belly is attached in its posterior half. In
Bufo the ventral integument is more or less broadly
attached, together with the whole of the dorsal
integument.

One of the most remarkable peculiarities connected
with the integument of Batrachians is the cutaneous
respiration, or the power which the surface of the

INTEGUMENT. 23

skin possesses of effecting those changes in the blood
which are usually performed by the lungs or gills,
and by which frogs may remain active for long
periods under water. Results of various experiments
have proved that pulmonary respiration alone is not
sufficient to support life without the aid of that of the
cutaneous surface ; whilst some of the tailed Batra-
chians have lately been shown to be deprived of both
lungs and gills, so that the contrary does not hold
good for them. But in order to carry out this im-
portant function it is of course necessary that the
surface be kept in a moist state, and this is effected
by a secretion of fluid from the skin itself.

Au immense number of close-set glandular caeca
open upon the surface of the skin, the slime-cells, in
addition to which a greater or lesser number of larger
poison-secreting glands are scattered or disposed in
very conspicuous prominent aggregations, such as the
dorso-lateral fold of liana, the large dorsal warts of
Bufo and Bombinator, and especially the so-called
parotoids, situated above the ear in Bufo and Ali/tes,
and a similar gland on the calf of Bufo calamita
(Fig. 9, p. 24). These glands are pierced with large
pores, visible to the naked eye or with the aid of a
low magnifier. A round gland, the frontal or pineal
gland, the homologue of the so-called parietal eye of
reptiles, is more or less discernible under the skin
of the forehead in front of and between the eyes ; this
gland, in the early stages, was connected with the
brain. The presence of lime concretions in the skin
of old specimens of Bufo vulgaris has been first
pointed out by Leydig. The same have been found
in B. calamita by 0. Seeck.

A character which does not appear to have been
noticed before is the presence of a filiform line or
raphe, on which the skin is much thinner, extending
along the middle line of the back from the snout to
the extremity of the coccygeal region in all specimens
of Bufo. This raphe, which may best be observed on

24

INTRODUCTION.

a stripped skin held against the light, gives rise to the
yellow vertebral line which is normal in Bufo calamita,
and only exceptional in B. viridis and vulgaris, inde-
pendently of another light vertebral streak which is
sometimes also present. The independence of the
two is most conspicuous in cases of deviation of the
former, as often happens in Bufo calamita, which
appears to be caused by the presence of large glands
in the course of the raphe necessitating a winding
(Fig. 9).

Fig. 9.

mm

: PS mm '

m

'•V ~> .- ■

Bvfo calamita, showing yellow raphe, independent of liglit
vertebral streak, and its deviation from the median line in
the middle of the back.

The epidermis is remarkable for the formation of
horny cells, as the spines on the warts of some toads
and Bombinator (Fig. 10, p. 29), or the sheaths on
the tips of the digits in Bufo and Pdobates, or on the
metatarsal tubercles when these are much developed.
There are also, in some Batrachians, special small,
pearl-like warts, mostly pigmentless, the "tact-spots"

INTEGUMENT.

-j:>

of Merkel, formed by the grouping together of
terminal ganglion-cells. These sense-organs are found
principally on the limbs and flanks of Bana esculenta,
more developed in males than in females; the small
warts on the fore limbs of Pelobates, which have by
some been compared with the nuptial horny excres-
cences, seem to be of the same nature. Very similar
productions appear during the breeding season in the
females of Bana temporaria and B. arvalis, but are of a
merely temporary character, and will be noticed in
the chapter' on Pairing and Oviposition, together with
the horny excrescences which arm the digits or other
parts in the males at the same period, and assist them
in maintaining their hold whilst pairing.

Pigment, usually confined to the cutis, sometimes
occurs in small quantity in the epidermis. As Leydig
has shown, five kinds of special cells may be pre-
sent in the cutis — black, yellow, red, white, and
metallic or iridescent, the last being the so-called
guanin-cells or iridocytes. The yellow and black pig-
ments combine to produce the bright green colour, as
in normal specimens of Bana esculenta tijpiea and
Eyla arborea : if the yellow pigment be absent, as in the
German specimens of Bana esculenta, var. ridibunda,
the black and gold produce the dull green or olive hne;
if both yellow and gold pigments are absent, the black
below the cloudy medium of connective tissue and
epidermis produces blue, as in the sky-blue specimens
of Bana esculenta and Ilyla arborea. In the play of
the special pigment-cells or chromatophores, which
contract or expand and radiate, we find the explana-
tion of the changes of colour which some specimens
undergo with so great rapidity. A tree-frog will turn
from yellow to green and black in less than an hour,
yellow when the black pigment contracts, black in the
opposite process.

That these rapid changes of colour harmonise,
within certain limits, with the surroundings is well
known. The researches of Dutartre on Bana escu-

26 INTRODUCTION.

lenta have shown that the changes are controlled
by the animal, the contraction or expansion of the
chromatophores not being due entirely to the direct
action of light and moisture upon the skin, but also to
a reflex action of the sympathetic nervous system
brought about by the visual impression. Comparative
experiments conducted on frogs possessed or deprived
of the sense of sight, and placed in different conditions
of light, show the blind individuals to change colour
much less rapidly.

The changes which we observe in Hyld, Rana, and
Bitfo are not limited to the ground colour, but also
extend to the markings. The tree-frog, normally
uniform, may become covered with lighter or darker
dots or spots, sometimes disposed with great regu-
larity and even assuming the form of cross-bars on
the limbs. An edible frog, of a fine grass-green with
few black spots above and with a pure white belly
when captured, may shortly after, when transferred to
an aquarium, appear largely marbled with black both
above and beneath.

F. Werner has proposed to group the European
Batrachians, with respect to this peculiarity, into
such as actually change colour {Alytes, Hyla, Bufo,
Ra i ia), and such as only lighten or darken their shade
(DisGoglossus, Pelodytes, Pelobates). He considers
Bombinator as representing a third category, no
changes of any kind taking place ; but I cannot agree
with him in this, as I have seen B. pachypus pass
from yellowish to a very dark brown, and also vary in
the presence, absence, or distinctness of the light and
dark markings.

In a recent work on East African Reptiles and
Batrachians G. Tornier endeavours to show that the
yellow and red pigments do not exist as chemical
entities, but are only degenerations of the brown
pigment or " melanin." And as a first " simple and
sure " proof of his contention he adduces the example
of Pelobates fuscus larvas, which according to him are

INTEGUMENT. 27

" quite black," and during metamorphosis assume the
light yellowish-grey ground colour through the greater
part of the pigment-cells turning pale. A more
glaring misrepresentation could not be imagined.
Far from being black, the tadpole of Pelobates fuscus
is normally pale brown, with darker spots and marb-
lings, as may be seen from the figure in this work.
And the tadpole of Pelobates ciiltrijpes, a species which
has the same dark spots as P. fuscus after transfor-
mation, is usually very pale reddish yellow. A proof
that the yellow and red pigments are chemically dis-
tinct from the brown or black is to be found in the
different action of alcohol on them. Thus yellow
pigment is rapidly destroyed ; therefore green frogs
turn olive or blue in spirit. Certain crimson pig-
ments, not unfrequent in frogs, resist the action of
alcohol and light better than the brown, which by dis-
coloration may turn to reddish brown or rufous, these
two reds strongly contrasting in specimens that have
long been in spirit, Tornier goes even so far as to doubt
the presence of any but the brown pigment in reptiles ;
surely the green of certain tree-snakes of the genera
Dryophis and Lachesis, which colour the spirit in which
they are preserved to the extent of resembling green
Chartreuse, is due to the presence of a chemically
distinct colouring matter, and has no relation whatever
to the green of frogs, which, produced, as we have seen,
by a combination of black and yellow, disappears in
spirit when the latter pigment is destroyed.

Another instance of the same author's carelessness
in the selection of examples is to be found a little
further in the same work (p. 130), where he declares
that Batrachian larva3 which develop in the light are,
with few exceptions, uniformly dark, or more usually
deep black, and retain that coloration until close to
the metamorphosis. I believe the exception is just
the other way. The embryos of Bombinator and
Alytes are yellowish or pale brown striped with
blackish, those of Hyla uniform yellowish, those of Bana

28 INTRODUCTION.

esculenta feebly pigmented; and of black tadpoles,
or of a dark brown approaching black, I only know
Bufo and liana temporarla; all tho other European
species are only exceptionally, not normally very dark.
That the light cannot have a great effect in producing
the pigment in Batrachian larvae is shown by Alytes
and Bombiuator, both of which develop the same
amount of pigmentation, the one under paternal care
never being exposed to daylight, the other reared in
pools or puddles exposed to the full action of the sun's
rays.

Cases of melanism, or better, absence of iridocytes
and all pigment but the brown or black, which may be
very abundantly and exclusively present, have been
observed by Heron-Royer in Alytes obstetricans, by
Alfred Duges, and more recently by Heron-Royer and
by Vaillant, in liana esculenta. I found a similar
young specimen of the latter species in a marsh near
the Belgian coast in 1884; the back was brown,
nearly black, the hinder side of the thighs and the
ventral surface pigmentless, transparent flesh-colour,
the iris black without any gold. A male specimen of
Bana grseca from Italy, showing the same abnormality
in the absence of all but the brown colour, was sent
to me last spring by Count Peracca, and is figured on
Plate XXII of this work.

Albinism, or absence of the brown pigment, on the
other hand, has been observed in a young Bombiuator
pachypus by Fatio, in larva3 of Alytes obstetricans by
Lataste and Heron-Royer, who succeeded in rearing
them through the metamorphosis, in a larva of Disco-
c/lossus p ictus by myself, in liana esculenta by Pavesi,
in larvae and young of Bufo viridis by Born, Camerano,
and myself, in larvae of Bana temporarla by Lessona,
Fischer-Sigwart, and Camerano ; an adult female of
the latter species found in England was exhibited
alive by Mr. Rowland Ward at the Linnean Society
in 1891, and another, from Wiltshire, presented by
Mr. W. Hannaford, is now preserved in the British

INTEGUMENT. 29

Museum. These albinos are of a uniform yellowish
flesh-colour, with silvery or golden iris and cherry-red
pupil ; the yellow ventral blotches are preserved in
Bombinator. A figure of a young Alytes albino reared
by Lataste is given on Plate VII. Albino specimens
of Alytes obstetricans obtained from tadpoles by Heron-
Royer paired in confinement, and produced offspring
that were likewise affected with complete albinism.
But for the difficulty in getting these Batrachians to
breed in captivity, a race of albinos could no doubt be
fixed by artificial selection, as in the case of the
axolotl, the numerous "white" examples of which
are said to be all descended from an albino male
which paired with normal females in the Jardin des
Plantes at Paris some thirty years ago.

Batrachians often "change skin," — about once a
month during the spring and summer. The thin, trans-
parent outer layer of the epidermis becomes detached
about the mouth, and, lubricated by an abundant secre-
tion of the slime glands, is pulled off by movements
of both fore and hind limbs, often in a single piece
turned inside out, especially when the operation is
performed in the water, and finally swallowed by its
owner. The casting of the epidermis on land is a
slower process, accompanied by ludicrous contortions ;
in the common toad it may last nearly an hour.

Fig. 10.

■ir'v .'."■ ■'V''v'-:i4.v<sf • ■'.: .

'<.''■+■

'•'>'

' ' "'V ■ L

$s4 '•'.•

■■■.-.-

-

Piece of dorsal integument of male Bombinator pachypus ( X 8),
showing warts and epidermal horny asperities.

30 INTRODUCTION.

IV. Dermal Secretion.

The skin of the dorsal surface of all Batrachians is,
as we have seen, studded with glands and follicles.
In various species, of which our Bufones are good
examples, some of the larger glands appear as very
prominent warts pierced with wide pores, especially a
large mass situated on each side of the back of the
head behind the eyes, which has received the mis-
nomer of parotid, a name now generally changed to
parotoid gland. These glands secrete, more or less
abundantly according to the species, a viscid or milky
more or less and variously odorous fluid, which is
exuded, when the animal is disturbed, by means of the
contraction of small muscles disposed in the skin
around the glands. If the larger glands be pressed
or subjected to an electric current the secretion is
squirted out with great force, and sometimes to a
considerable distance. Kobert recommends, as the
best means of obtaining an abundant supply of toad-
poison, the subcutaneous injection of a small dose of
chloride of barium ; after a few minutes the animal is
so covered with its secretion as to appear as if coated
with whitewash. The object of the secretion is to
guard Batrachians from the attacks of various enemies.
A clog will not often repeat the experiment of seizing
a toad; the signs of pain which it evinces after the
poisonous fluid has come into contact with its mouth
is familiar to most of those who have kept young
dogs in the country. That excellent Hungarian
naturalist, Prof. v. Mehely, relates the serious effects
of the poison of Bufo viridis on a small terrier which
accompanied him on a collecting excursion. A large
toad, found under a stone, was at once attacked by
the dog, but no sooner seized was let go again with
signs of great repulsion ; the toad had instantly become
covered with a thick white secretion. The dog ap-
proached it once more and then withdrew, sneezing and

DERMAL SECRETION. 31

howling and rubbing its foaming mouth on the grass.
After a few minutes the dog was seized with convul-
sions, and had to be carried home exhausted. On the
next day it had a swollen mouth and burning nose. It
did not completely recover until the following day.

That the toad's poison does not protect it against
snakes is well known, — our common Tropidonotus, for
instance, not feeling any more repulsion for it than for
a frog, although it will not eat Bombinator. It is the
experience of those who have handled freshly caught
specimens of the latter that the secretion of these
Batrachians acts as a sternutatory, and causes irritation
of the mucous membrane of the nose and conjunctiva,
the effect being comparable to the early stages of a
cold in the head. German violinists, it is said, when
suffering from moist hands, are accustomed to check
the perspiration by handling live toads. Many col-
lectors of Batrachians have learned, to their discom-
fiture, how the introduction of examples of certain
species in the bag containing the spoil of their excur-
sion may cause the death of the other prisoners, and
common tree-frogs are stated to have died from the
contact, in the same small terrarium, of examples of an
American species, Hyla versicolor, which, owing to its
warty skin, is often designated as a " tree-toad ; " for
although the poison has no effect on the skin of
individuals of the species which produces it, frogs of
different species, however closely allied, may poison
each other by mere contact. When ingurgitated or
inoculated the poison acts even on the individual by
which it has been secreted, as has been proved by
various experiments, but death is only produced by
using a stronger dose than is required for killing
individuals of another species.

The poison of Batrachians, unlike that of snakes,
is not a septic, but acts upon the heart and the central
nervous system. That of the common toad has beeu
compared, as regards its effects, to Digitalis and
Erythrophlaeum. For a long time authorities disagreed

32 INTRODUCTION.

as to the nature of the secretion ; some regarded it as
an acid, others as an alkaloid. Phisalix, in 1890,
claimed to have solved the question by reconciling
both views, he having found toads to be simultaneously
possessed of two different kinds of glands, different
both anatomically and physiologically. These are
mucous or slime glands spread over the greater part
of the body, the exudation of which is controlled by
the animal; and specific glands (the parotoids and the
larger dorsal glands), the product of which can only
be squirted out by foreign agency. The secretion of
the former glands is an alkaloid, and acts as a
narcotic ; that of the latter is an acid, and acts as a
convulsive. The anatomical differences of the two
kinds of glands have since been worked out by Paul
Schultz, who, however, regards the product of the
slime glands as innocuous.

Phisalix and Bertrand's recent investigations have
shown the active principle of toad-poison, the so-called
phrynin,to be also present in certain quantities in the
blood of these Batrachians.

Even the frogs of the genus Rana, so sensitive to the
poison of other Batrachians, are not free from toxic
properties. A goldfinch inoculated by Paul Bert with
the dermal secretion of Rana esculenta died in one
minute ; a frog of the same species in one hour and a
quarter.

The secretion of Alytes, Pelodytes, and Pel abates
smells like garlic, so strongly in the last that I have
observed the odour to remain for hours in a room
after I had purposely irritated a few specimens.
That of Bufo ealamita smells like boiled india-
rubber, like ignited gunpowder, or a Dutch clay pipe
smoked for the first time, according to Rosel's com-
parison; whilst that of its near ally, Bufo viriclis,
always reminds me of a linseed poultice.

A chemical comparative study of the dermal secre-
tion in the various types of European tailless Batra-
chians is still a desideratum.

SKELETON.

33

V. Skeleton.

The skull is more or less strongly depressed, and
remarkable for the extremely large orbits and the
reduction in the number of elements. We have to
distinguish between the chondroeranium., or primitive
skull — consisting mainly of cartilage with a few ossified
parts, with a large fontanelle or membranous space in
the interorbital region, and foramina for the exit of
the cranial nerves, — and the membrane hones, which
may be removed by maceration.

The ossified parts of the chondroeranium, or cartilage
bones (Fig. 11), are the ethmoid or sphenethmoid, form-
ing a ring round the anterior part of the brain- case, the

Fig. 11.

Chondroeranium
A. Upper view.

c. Brain-case.

e. Ethmoid.
eo. Exoccipital.
fe. Fontanelle.
fm. Foramen magnum.
fo. Optic foramen.
ft. Trigeminal foramen.

of Bana esculenta.
B. Lower view.
j. Jugal.

I. Lachrymal process.
n. Nasal capsule.
pr. Prootic.
pt. Pterygoid branch.
sp. Suspensorium.

(paired) prootic behind the orbit, the (paired) exoccipital
on each side of the foramen magnum, forming the
(paired) occipital condyle articulating with the first ver-
tebra, and a styliform jugal connecting the cartila-
ginous suspensorium of the mandible or the ossified
quadrate with the maxillary.

c

34

INTRODUCTION.

Skull, mandible, and hyoid of Discoglossus pietus.
A. Upper view of skull. B. Lower view of skull, c. Side view of

skull and mandible.

any. Angular.
ar. Articular.

c. Cornu of hyoid.
ca. Columella amis

d. Dentary.

e. Ethmoid.
eo. Exoccipital.
fp. Fronto-parietal.
hb. Body of hyoid.

j. Jugal.
m. Maxillary.
v. Nasal.

D. Lower view of mandible and hyoid.
p, Palatine.

pm. Pramiaxillary.

ppl. Postero-lateral process of

hyoid.

pro. Prootic.

ps. Parasphenoid.

pt. Pterygoid.

pth. Thyroid process of hyoid.

q. Quadrate.

sq. Squamosal.

v. Yomer.

vh. Ventral ossicle of hyoid.

SKELETON.

35

The membrane bones, all paired with the excep-
tion of the parasphenoid (and in Pelobates the fronto-
parietal), are the following : — Prsemaxillary, maxillary,
squamosal, pterygoid, palatine, vomer, parasphenoid,
nasal (often called prae frontal), and fronto-parietal.
A small ossification behind the narial opening is the
turbinal, regarded by some as the true nasal. The
cranial ossification may be feeble, and a considerable
portion of the cartilaginous primordial cranium remain
exposed, — as, for instance, in Bombinator, which,
together with Pelodytes, is remarkable for the lack of
palatine bones ; or it may be extraordinarily developed,
with superaddition of dermo-ossification, as in Pelo-
bates, one species of which (P. cultripes) is distin-
guished by the expansion of the fronto-parietal and

Fig. 13.

Upper view of skull of Pelobates cultripes.

squamosal, which join to form a complete roof over
the prootic (Fig. 13). The genus Pelobates is further
remarkable in having the fronto-parietal single,
whilst in all other European genera it is paired,
and in many species so separated along a portion
of the median line as to leave the endocranial or
fronto-parietal fontanelle exposed between the two.
The nasals may join in the middle line, or be more or
less widely separated from each other. The upper
lamina of the ethmoid is usually exposed between the
fronto-parietals and the nasals, and in Pelobates fuscus
it is so prolonged forwards as to reappear between
the nasals and the praBmaxillaries.

36 INTRODUCTION.

The squamosal is a hammer-shaped bone, the anterior
branch of which may be much reduced, as in Bafo,
long and free, as in Bana, or join the maxillary, as in
Discoglossus and Pelobates.

The vomers vary much in size according to the
genera ; they are largest in Discoglossus, where their
posterior toothed border covers over the palatines.
The latter bones, if present, are narrow and transverse,
and extend across the whole palate ; in Bufo their
lower surface forms a cutting, sometimes serrated edge,
which makes up for the absence of vomerine teeth.
The pterygoids are trifurcate, and the anterior branch
may or may not reach the palatines, or may be much
produced forwards when the latter bones are absent,
as in Bombinator. The parasphenoid is a large bone
shaped like a T, or like a dagger with very short
handle.

Most genera have a cartilaginous tympanic ring
applied to the outer surface of the hammer-shaped
squamosal, in the upper portion of which ends the
slender, ossified, stapedial rod (columella auris).
The tympanum is absent in Pelobates, and both the
tympanum and the stapedial rod in Bombinator,
which in this respect again resembles the tailed
Batrachians.

In Discoglossus three bones invest the Meckelian
cartilage which forms the ramus of the mandible : the
dentary in front, the angular or angulo-splenial behind,
and extending more to the front on the inner side,
and a small bone which articulates with the quadrate,
the articular. In Bana the articular remains cartila-
ginous, but the symphysial cartilages ossify, forming
the so-called mento-Meckelian or symphysial bones,
which in adult specimens fuse on the upper side
with the dentary (Fig. 14). These symphysial bones
are less distinct in Hyla and Bufo, less still or even
almost indistinguishable in the Pelobatidse and Disco-
glossidse.

The hyoid varies very considerably in the different

SKELETON. 37

genera. The body is broad with well-developed,
sometimes ossified posterolateral, but no anterior,
processes in the Discoglossidae (Fig. 12, d, p. 34) ;
with anterior processes and short postero-lateral
ones in the Pelobatidce and Bana (Fig. 14) ; elongate

Mandible and byoid of Bana esculenta, lower view.
ancj. Angular. pi. Lateral process of byoid.

c. Oornu of byoid. ppl. Postero-lateral process of byoid.

d. Dentary. I pth. Tbyroid process of byoid.

pa. Anterior process of byoid. | sij. Sympbysial(mento-Meckelian).

and without anterior processes in Bufo. In Pelodi/tes
and Pelobates the anterior processes are much expanded,
converge towards each other, and join the lateral pro-
cesses, enclosing a fenestra on each side of the hyoid
plate. The cornua are broad in the Discoglossidse,
moderately broad in Bufo, slender in Bana; they are
detached from the body of the hyoid in the Pelobatidce.
The thyroid processes are always ossified, and form a
pair of diverging styles, in front of which, on the
ventral side, a slender, single or paired, V- or H-shaped
ossification may be present, as in Discoglossus, Ahjtes,
and Pelodytes (Fig. 12, d, p. 34).

The vertebral column (Fig. 15) is formed normally
of ten segments, viz. eight dorsal vertebrae, one sacral,
and the urostyle or coccyx, — or of nine if the urostyle
be fused with the ninth vertebra, as is frequent in
Pelobates.

The vertebras are much depressed, especially the

38

INTRODUCTION.

so-called centra, the artioulary balls of which are
transversely elliptical, and the neural arch is either
closed above (vertebras imbricatim clauses, according to
the definition of Mertens), as in the Discoglossidas and
Pelobatidas, or notched between the zygapophyses so
as to expose the spinal cord between every two verte-
bras ; the latter type is most marked in Bana, in which,

Fig. 15.

c^*S>>

l

r?

B

Vertebral columns of (a) Discoglossus pictus and (b) Rana eseulenta,

upper and lower views.

the lateral openings for the exit of the spinal nerves
being also of large size, the vertebral column forms an
open work above and on the sides. Neural spines are
absent or represented by a low keel, which is much
prolonged posteriorly in Discoglossus and Pelobates.
The articulation is convexo-concave or opisthoccelous
in the Discoglossidas, concavo-convex or procoelous in
the other groups. In those forms in which the verte-
bras are procoelous the eighth is biconcave; the ninth
being invariably biconvex.

The first vertebra is devoid of transverse processes

SKELETON. 39

(diapophyses), but these are present on the succeeding
eight vertebras, and in the Discoglossidse alone the
first three are segmented off into short autogenous
ribs. The transverse processes of the ninth vertebra
give attachment to the ilia, and are therefore termed
sacral ; their form differs much according to the
genera, — cylindrical in Rana, flattened and subtri-
angular in Discoglossus, Alytes, Bufo, and Hyla, or
very strongly dilated in Bombinator, Pelodytes, and
Pelobates. In the latter genus the sacral processes are
often yielded by the urostyle. The diapophyses ter-
minate in cartilaginous epiphyses.

The urostyle, or coccyx, formed by coalescence of
at least two vertebrae, is a cylindrical solid rod with
more or less distinct dorsal crest, and articulates with
the sacral vertebra by two sockets, except in Bom-
binator, in which there is but one. In Pelobates the
urostyle is normally fused with the ninth vertebra,
and contributes to the sacrum ; in the Discoglossidse
a slender transverse process is present on each side
at the base of the urostyle.

Abnormalities in the vertebral column are not un-
frequent, and many remarkable cases have been
described by Goette, Lataste, Camerano, Bourne,
Adolphi, and Howes. Thus two vertebras may fuse,
or the number of segments may be increased to eleven,
as in a vertebral column of Alytes obstetricans described
by Lataste ; in this interesting specimen the ninth
vertebra bears on the left side the dilated sacral dia-
pophysis, whilst the diapophysis of the opposite side
is slender and does not differ from that of the vertebra
in front of it ; the tenth vertebra, on the other hand,
bears a dilated diapophysis on the right side and a
cylindrical one on the left ; the urostyle is quite
normal, with the basal processes developed as usual. I
have also examined a skeleton of Bombinator pachy pus,
with eleven segments, all, including the sacral ver-
tebra and the urostyle, appearing perfectly normal in
shape.

40

INTRODUCTION.

The pectoral arch falls into two distinct types, — the
arciferous (Fig. 16, a), in which the praecoracoid

Fig. 16.

Pectoral arch and fore limb of (a) Discoglossus pictus and (b) Bana

esculenta. Ventral view.

r. Radius.

c. Coracoid.
ec. Epicoracoid.

h. Humerus.
ost. Omosternum.
pc. Prsecoracoid.

s. Scapula.
ss. Supra-scapula.
st. Sternum.
Ulna.

u.

(praecoracoid + clavicle) and coracoid are widely
separated from each other distally, and connected by
an arched cartilage (the epicoracoid), the right usually
overlapping the left; and the firmisternal (Fig. 16, b),
in which both prascoracoid and coracoid nearly abut on
the middle line, and are only narrowly separated by
the fused epicoracoids. The latter type is exemplified
by the true frogs (Uanidse) in the adult state, although

SKELETON. 41

when quite young these Batrachians present a con-
dition similar to that which persists throughout life in
the other families.

A cartilage in the median line in front, supported
by a bony style in Bana, is the so-called omosternum ;
a larger one behind is the sternum. This is also
provided with a bony style in Ranas Pelodytes, and
Pelobates ; or a portion of it may be calcified, as in
Hyla and Bufo. In the Discoglossidse the sternum is
produced into two posteriorly diverging cartilaginous
or calcified styles.

The shoulder-blade is divided into two parts, the
ossified scapula proper and the supra-scapula, carti-
laginous or incompletely ossified, bent over at an angle
to the scapula.

The prascoracoid enters the glenoid cavity in the
Discoglossidx, Bufonidse, and Hylidse, but not in the
three other families, where the cavity is formed by
the scapula and coracoid.

The humerus is a feebly curved bone with a globular
distal condyle and a strong prasaxial crest, which is
more developed in males than in females, and which
in the former may be supplemented by a second crest
lower down. The head is replaced by an epiphysis,
which calcifies late, or never as in the Discoglossidse and
Pelobatidss. The radius and ulna are fused to a single
bone, which is shorter than the humerus. The ulnar
part projects beyond the radius at the articulation
with the humerus.

The carpal bones vary from six to eight. In the
more primitive carpus, such as is represented by the
Discoglossidse (Fig. 17, a), we find, following the
nomenclature of Howes and Ridewood, two elements
in contact with the radius-ulna, the radiale and ulnare ;
two, regarded as centralia, in the second row; and
four, plus a vestigial ligamentous fifth, in the third
row. The fourth distal carpale may fuse with the
outer centrale, as in Pelodytes (b), or also with the
third distal element, as in Bufo, Hyla, and Bana (c).

42

INTRODUCTION.

In the Pelobatidae and Bufonidas the inner central e is
iu contact with the radius.

Fig. 17.

Tm 1

n m iv v

A

Carpus of — A. Bombincdor pachypus. b. Pelodytes punctatus.
c. Ttcuia temporaria. (After Howes and Ridewood.)

R. Radius. U. Ulna. r. Radiale. u. Ulnare. c. Centrale.
1 — 1. Distal tai'salia. I — Y. Digits.

There are four functional fingers, with elongate
metacarpals, and 2, 2, 3, 3 phalanges. The distal
phalanx is pointed or slightly expanded at the end,
somewhat as in the human skeleton ; or bears a well-
marked transverse branch, giving it a T-shaped
appearance, as in R. grseca ; or it assumes a peculiar
claw-shaped conformation iu the Hylidse, which is
described and figured above, p. 15. In addition to
these four fingers there is a rudimentary thumb or
pollex, usually more developed in males than in
females, and which, besides its metacarpal, may show
one or two ossicles representing phalanges. The meta-
carpal of the pollex is very strongly developed in
the male Discoglossus, which is also remarkable for the
enlargement of the second metacarpal, bearing a crest
along the inner side.

The pelvis (Fig. 18) is much elongate, and shaped
like a pair of tongs, the free extremities of which are
attached to the diapophyses of the sacral vertebra.
The lateral branches are formed by the ilia, which,
with the ischia behind them, bound the acetabulum ;
the third element, the pubis, situated ventrally be-
tween the ilium and the ischium, is often cartilaginous
or calcified, and absent in Bombinator. In Pelobates

SKELETON.
Fig. IS.

43

il

Pelvis and hind limb of
Discoglossus pictus.

a. Astragalus.
c. Calcaneum.
/. Femur.
il. Ilium.

Ischium.

Pubis.

is.

P-

t. Tibia-fibula.

the pubis is represented by a
small ossified nodule, not enter-
ing the acetabulum. In the
Discoglossidae the acetabulum
remains open, as in the young
of other Batrachians.

The femur is more or less
strongly curved. The tibia
and fibula are fused to a

single nearly straight bone. The epiphyses of these

bones remain cartilaginous in the Discoglossidae and

Pelobatidm, calcify in the other families.

The tarsus is remarkable for the elongation of the

astragalus and calcaneum, which form an additional

44

INTBODUCTION.

segment to the limb, a sort of crus secondarium. This
is especially striking in Pelodytes, where the two
bones fuse, and resemble, but in length, the cms or
tibia-fibula.

In the Bufonidse, Hylidse, and Banidde a calcified
epiphysis unites the two bones at either extremity.
In liana and Bufo a small sesamoid bone is present on
the inner side of the proximal extremity of the tarsus.
A series of three or four small distal tarsals (Fig. 19)
intervenes between the astragalus and calcaneum and
the three inner metatarsals, whilst the two outer meta-
tarsals are directly in contact with the calcaneum.
The toes are five in number, with elongate meta-
tarsals and 2, 2, 3, 4, 3 phalanges. A rudimentary
inner digit, the so-called prsehallux, consists of a

Fig. 19.

A

Tarsus of — A. Bombinator pachypus. b. Pelodytes punctatus.
c. Rana temporaria.

metatarsal and one or two phalanges, the development
of which corresponds with that of the inner meta-
tarsal tubercle. In Bombinator, Alytes, and Pelodytes
the distal tarsals as well as the praehallux remain
cartilaginous throughout life ; in Discoglossus the
prsehallux ossifies, but the distal tarsals remain cartila-
ginous.

The following is a Synopsis of the Grenera and
Species, based on the osteological characters :

I. Prsecoracoid curved, connected with the coracoid
by an arched (epicoracoid) cartilage over-
lapping its fellow (Aroifeea).

SKELETON. 45

a. Vertebrae opistliocoelous, imbricate and com-
pletely covering the spinal cord above ;
autogenous ossified ribs attached to the
anterior diapophyses ; sacral vertebra with
dilated diapophyses overlapping the ilia;
urostyle with transverse processes at the
base ; omosternum cartilaginous or absent ;
sternum cartilaginous, with two posteriorly
diverging limbs ; eight elements in the
carpus, two in contact with radius-ulna ;
four distal tarsal elements ; long bones with
cartilaginous epiphyses (DiscoGLOSSiDiE).

1. Fronto-parietal fontanelle small, covered

over in very old specimens ; palatines
present; zygomatic branch of squamosal
forming a suture with maxillary ; second
rib with posterior process ; dorsal vertebras
with a strong postero-median neural pro-
cess ; sacral vertebra with moderately
dilated diapophyses and two condyles for
urostyle Uiscoglossus.

2. Fronto-parietal fontanelle very large; pala-

tines absent ; second rib with posterior
process ; postero-median neural process of
vertebra absent or very short ; sacral ver-
tebra with very strongly dilated diapo-
physes and a single condyle for urostyle.

Bombinator.

Tibia shorter than femur B. igneus.

Tibia as long as femur ... 7?. pachypus.

3. Fronto-parietal fontanelle large ; pala-

tines present ; a cartilaginous supra-
orbital ; second rib without process ; pos-
tero-median neural process of vertebra?
absent or very short ; sacral vertebra with
strongly dilated diapophyses and two
condyles for urostyle .... Alytes.
Fronto-parietal bones in contact only quite
posteriorly, embracing a very large sole-

46 INTRODUCTION.

shaped fontanelle ; metacarpal and phalanges
of fourth finsfer not more massive than the
others ....... A. obstetricans.

Fronto-parietal bones meeting iu the middle,
separating a large kite shaped f ontanelle in
front from a very small one behind ; meta-
carpal and phalanges of fourth finger very

massive A. cisternasii.

b. Vertebrae proccelous, without autogenous
ribs.
1 . Vertebrae with imbricate arches completely
covering the spinal cord above ; diapo-
physes of sacral vertebra very strongly
dilated, and sliding in a groove of the ilia ;
praecoracoid not entering glenoid cavity ;
omosternum cartilaginous ; sternum with
a bony style; long bones with cartilagi-
nous, non-calcified epiphyses ; three carpal
bones in contact with radius-ulna

(Pelobatidje),

a. A fronto-parietal fontanelle ; fronto-

parietals distinct ; palatines absent ;
postero-median neural process of ver-
tebrae very short or absent ; sacral ver-
tebra with two condyles for nrostyle ;
astragalus fused with calcaneum; seven
bones in carpus ; four cartilaginous
elements in distal tarsal row Pelodytes.

b. No fronto-parietal fontanelle; fronto-

parietal single, rugose ; squamosal ex-
panded, plate-like ; dorsal vertebrae with
a long postero-median neural process ;
sacrum usually fused with urostyle ;
astragalus and calcaneum distinct ; eight
bones in carpus, three in distal tarsal
row Velobates.

Squamosal not joining fronto-parietal ; ethmoid
reaching praemaxillaries . . P. fuscus.

Squamosal forming with fronto-parietal a broad

SKELETON. 47

bridge over the prootic ; ethmoid not
reaching praemaxillaries . P. cidtvipes.
2. Vertebrae notched between the zygapo-
physes ; diapophyses of sacral vertebra
moderately dilated ; sternum cartilagi-
nous ; praecoracoid entering the glenoid
cavity; six bones in carpus, three in distal
tarsal row ; long bones with calcified epi-
physes.

a. Distal phalanges obtuse ; three carpal

bones in contact with radius-ulna; omo-

sternum absent (Bufonid^e) . Bafo.

No fronto-parietal fontanelle ; urostyle longer

than the skull ...... B. vulgaris.

A small fronto-parietal fontanelle, disappearing
in old specimens ; urostyle slightly longer

than the skull B. viridis.

A large fronto-parietal fontanelle; urostyle as
long as the skull B. calamita.

b. Distal phalanges claw-shaped; two carpal

bones in contact with radius-ulna ; omo-
sternum cartilaginous (HvLiDiE) . Hyla.

II. Praecoracoid straight ; no arched epicoracoid
cartilage (Firmisternia).
Vertebras procoelous, notched between the zyga-
pophyses ; no autogenous ribs ; diapophyses
of sacral vertebra cylindrical ; omosternum
and sternum with a bony style ; six bones in
carpus, two in contact with radius-ulna ; three
distal tarsal bones ; long bones with calcified
epiphyses (Ranid2e) Raiia.

a. Zygomatic branch of squamosal veiw long,

nearly twice as long as the posterior ; fronto-
parietals very narrow, grooved ; second
diapophysis not once and a half as long as
the seventh R. esculenta.

b. Zygomatic branch of squamosal a little longer

or a little shorter than the posterior ; second

48 INTRODUCTION.

diapophysis usually about ouce and a half as
lous: as the seventh.

1. Fronto-parietals convex, narrow; nasals

with straight or slightly concave posterior
border.
Nasals widely separated from each other ; tibia
nearly twice as long as the tarsus.

B. a rv ah's.

Nasals narrowly separated ; tibia more than

twice as lono- as the tarsus . B. camerani.

2. Fronto-parietals flat, broad.

Nasals with concave posterior border; tibia
slightly longer than femur . B. temporaria.

Nasals with straight or convex posterior
border; tibia considerably longer than femur;
distal phalanges with very distinct trans-
verse expansion at the end* . . B. grseca.

Nasals with concave posterior border; tibia
considerably longer than femur . B. iberica.

Nasals with straight or feebly concave posterior
border; tibia considerably longer than femur.

B. latastii.

3. Fronto-parietals grooved, moderately broad ;

nasals with straight or slightly concave
posterior border ; tibia considerably longer
than femur B. agilis.

The above key, it must be observed, is drawn up
from skeletons of adult, or at least half-grown speci-
mens, and must necessarily often fail in its object if
applied to young ones.

* Quite as much developed as in the species on which the genus
Hylorana ov Limnodytes has been founded. I have pointed out that
certain Chinese and Burmese forms show a passage to exist from the
Rante temporarise to the Hyloranse, and this view is further confirmed
by a study of Rana grseca.

VISCERA.

49

VI. Viscera.

A general sketch of the internal soft anatomy is
not attempted here. We will merely indicate the prin-
cipal differences in the structure of the lungs and uro-
genital apparatus which are of special importance
from the point of view of the systematist. The
following figure will suffice to show the position and
relations of the various parts.

Fig. 20.
I ccc p vf

Longitudinal section through body of male Bana esculenta,
showing the viscera.

Large intestine.

Spleen.

Larynx.

Lung.

Pancreas.

Kidney.

a.

Auricle of heart.

i(J

ca.

Adipose bodies.

I

'/•

Bile-duct.

la

cl.

Cloaca.

P

h.

Liver.

pa

i.

Small intestine.

r

s. Stomach.

t. Testicle.

v. Ventricle of heart.
ve. Bladder.
vf. Gall-bladder.
vs. Vesicula seminis.

The lungs are a pair of sessile, or very shortly
pedunculate, oval sacs, often more or less pointed
posteriorly. Their walls are thick and spongy in
Bana, thin and diaphanous in the other genera.
Their periphery may be divided into numerous air-
cells, around which the ramifications of the pulmonary
artery are distributed, as in most genera; or the cells
may be few and the ramifications of the artery reduced

50

INTRODUCTION.

to five or six, as in Bombinator pachypus (Fig. 21, a).
Bomblnator igneus and Hyla are intermediate between
the two extremes. Owing to the filling up of the
abdominal cavity by the more voluminous genital
organs, the lungs are usually smaller in females
than in males. In the genus Ban a (Fig. 21, c)

Fig. 21.

4 mmmm

3 & MH'.[ ! '

13

Bombinator pachypus. B. Pelobates fuscus.
c. Rana temporaria.

the lungs, when fully distended, measure two-
thirds or more the length of the abdominal cavity
in males, one-third to one-half in females. In
Bombinator, Bufo, and Hyla they measure nearly
three-fourths that length in males, two-thirds in
females. In Pelobates they are equally developed in
both sexes, and larger than in any other genus, with
a seemingly superadded narrower terminal portion
bent inwards and forwards, tapering to a fine point
(Fig. 21, b) ; the length of the inflated lung consider-

VISCERA. 51

ably exceeds that of the abdominal cavity. The lungs
are smallest in Discoglossus, measuring only about half
the length of the abdominal cavitv in males.

The liver is divided into three lobes, one on the
right side, two on the left.

The kidneys are elongate, flattened, red bodies
attached to the dorsal body-wall, close to the middle
line, below the vertebral column ; from them the ducts
or ureters proceed to the cloaca, into which they open
by two small closely approximated apertures opposite
the opening of the urinary bladder.

In the males the testes lie at the inner side of and
ventrally to the kidneys, connected with them (except
in the Discoglossidss) by a number of delicate ducts,
vasa efferentia, which convey the spermatozoa into the
tubules of the kidneys, from which they escape by the
ureter. The male reproductive organs differ consider-
ably according to the species; and the testes, which
are not always symmetrical, vary accordiug to the
season and the individual, so that little importance
can be attached, from the systematic point of view, to
their size, shape, and pigmentation.*

The ovaries in the female correspond to the testes
in the male. The oviducts are lone1, much-convoluted
tubes, with thick gelatinous walls, in which the ova as
they descend become coated with their mucilaginous
envelope ; their open mouths are situated close to the
roots of the lungs, and their lower ends, much dilated,
have often been termed, improperly, uterus ; the ovi-
ducts open, together with the ureters, into the cloaca.

In both sexes two tufts of flattened yellow or orange
fatty appendages, the corpora adiposa, are attached
above the genital glands. They vary much in size,
according to the season, and are most developed just
before the breeding period. I have seen them of a
vermilion red in breeding specimens of Bufo calamita.

■ I have examined a breeding male, Bufo vulgaris, whose right testicle
was uniformly yellow, whilst the left was closely pigmented on about
two-thirds of its surface.

52

INTRODUCTION.

And finally, attention must be drawn to a pro-
blematic organ, peculiar to the males of the genus
Bufo, termed by Spengel " Bidder's organ," situated

Fig. 22.

r.<

ca

TS-

CCb

M/

B

D E

Right urogenital apparatus (ventral view, the testis shifted to
the side) of- — A. Discoglossus pictus. b. Bombinator pachypus.
c. Alytes obstetricans. D. Bnfa viridis. e. Rana temporaria.

b. Bidder's organ.
ca. Corpora adiposa.
m Miillenan duct.

r. Kidney.

t. Testis.

u. Ureter.
ve. Vas efferens.
vs. Vesicula seniinis.

VISCERA. 53

between the testicle and the adipose bodies. This
has been considered by some anatomists as a rudi-
mentary ovary, which it resembles in the ova-like
nature of its contents. There is, however, no ground
for regarding male toads as in any way normally
hermaphrodite, for Spengel has actually observed a
true hermaphrodite specimen of the common toad, in
which the problematic organ was developed in addition
to the ovary, aud that author therefore regards it as a
division of the testicle, endorsing the view of Bidder,
its discoverer.

Rudimentary oviducts, the Miillerian ducts, occur
in the males; most developed in Alytes, where, as
we have seen above, they are functional, viz. act as
seminal ducts, and in Bufo ; absent in the PelobatidsB
and Hylidas.

The following comparison is instituted on specimens
examined during the breeding period :

In Discoglossus the testes are remarkably large,
elongate, oval, ivory-white, longitudinally striated,
measuring two-thirds to five-sixths the length of the
kidneys. Contrary to what exist in all other Ecau-
data, there is but one vas efferens, of large calibre,
which does not open into the kidney, but is continued
uninterruptedly into the seminal duct, which extends
up to the front extremity of the kidney, terminating
in a short blind process. The vesicula seminis is very
large, and extends high up the side of the kidney.

In Bombinator pachypus the testes are yellow, un-
pigmented, twice to twice and a half as long as broad,
slightly flattened and rounded at both ends, nearly
half as long as the kidneys. The vasa efferentia open
into a canal along the inner edge of the kidney. This
canal communicates with the seminal duct at the
anterior extremity of the kidney, beyond which the
duct extends forwards as a blind process. A vesicula
seminis is absent. B. igneus differs in having the
testicles more or less pigmented, sometimes wholly
black. They measure two-fifths to one-half the length

54 INTRODUCTION.

of the kidney, and are nearly three times as long as
broad.

In Alytes obstetricans the testes are snbspherical or
shortly oval, yellow, usually more or less sprinkled
over with black pigment. There are two vasa effe-
rentia, which do not open into the kidney but pass into
the Mi'illerian duct, which functions as a seminal duct
distinct from the ureter, and bears, as a diverticulum,
a large strongly pigmented vesicula seminis.

In Pelobates fuscus the testes are oval, flattened,
their length one-third to three-fourths that of the
kidneys, the anterior extremity of which they nearly
reach; they are yellow, rarely feebly pigmented. A
vesicula seminis is absent.

Pelodytes punctatus agrees very closely with the
preceding. The testes are small, measuring only one-
third the length of the kidney, and much pigmented,
nearly black.

In Bufo vulgaris the testes are yellow, sometimes
more or less pigmented, elongate, cylindroid or
flattened, two or three times as long as broad, one-
half to three-fourths the length of the kidney, to the
anterior extremity of which they usually do not
extend. A vesicula seminis is absent. In B. viridis
the testes may be shorter, more oval, or cylindroid,
and more than half as long as the kidney; they are
more or less strongly pigmented, sometimes entirely
black, and do not extend to the anterior extremity
of the kidney. B. calamita has the testes elongate,
and, as in B. viridis, they are situated opposite the
middle of the kidney; they are so strongly pigmented
as to appear entirely or nearly entirely black.

In Ilyla arborea the testes are yellow, and resemble
those of Bufo vulgaris in size, form, and position. A
vesicula seminis is present.

In Bana esculenta the testes are yellow or orange,
short, oval, not half the length of the kidneys, to the
middle of which they are attached ; the vesicula
seminis is confounded with the ureter, which is of

VISCEKA. 55

large calibre and tapers gradually towards the cloaca.
R. temporaria has larger testes, measuring two-thirds
to three-fourths the length of the kidneys, nearer the
anterior than the posterior extremity of the latter, and
often more or less strongly pigmented ; the vesicula
seminis is large and distinct from the ureter, im-
mediately behind the kidney. In R. arvalis, R. grseca,
R. latastii, and R. agills the testes are, as a rule,
smaller, one-third to two-thirds the length of the
kidneys, and not or but feebly pigmented. In
R. temporaria, grseca, latastii, the veslcula seminis is
in contact with the kiduey, whilst in R. arvalis and
R. agilis it is a small sac, usually situated along the
ureter midway between its attachment to the kidney
and the cloaca. I have, however, dissected specimens
of Rana arvalis in which the vesicle was situated
very near the kidney.

The chief differences in the male urogenital appa-
ratus may be expressed in the following synopsis :

I. The spermatozoa are conveyed through the vasa

efferentia direct to the seminal duct; the latter
produced forwards beyond the kidney.

DlSCOGLOiSSIDiE.

a. The ureter functions as seminal duct.

A single vas efferens ; a vesicula seminis.

Discoglossm.

Several vasa efferentia opening into a canal along
the outer edge of the kidney ; no vesicula
seminis ..... Bombinator.

n. The Miillerian duct functions as semiual canal
distinct from the ureter ; two vasa efferentia
close together in front of the kidney ; a
vesicula seminis .... Alytes.

II. The spermatozoa are conveyed to the semiual

duct (ureter) through the kidney, in which the
anterior extremity of the duct is embedded.

56

INTRODUCTION.

a. No vesicula seminis.

No ovary-like body in front of the testis

Pelobatid^.

An ovary-like body (Bidder's organ) between the
testis and the corpora adiposa . Bueonimc.

d. A vesicula seminis formed by saccular en-
largement of the ureter. Hylidje, Ranid^;.

A table is added showing the proportions of some of
the principal viscera in different species, the measure-
ments, in millimetres, being taken from males during

the breeding season.

1.

2.

3.

4.

5.

6.

7.

8.

9.

10

Discoglossus piotus .

59

5

8

9

19

18

4

15

7

9

Bombinator igneus .

. 46

4

4

18

18

10

o
O

5

2

15

,, pachypus

44

4

4

JO

12

11

3

5 5

25

14

Alytes obstetricans .

37

4

5

]2

10

8

25

3

2

5

Pelobates fuscus

54

5

6

34

15

15

5

8

r

7

Peloclytes punctatus

38

3

4

14

9

9

6

35

9

Bufo vulgaris .

88

9

8

28

25

21

4

10

5

7

„ viridis .

70

9

i

25

25

17

5

10

4

18

,, calamita

66

9

7

26

23

16

5

i

3

19

Hijla arbor e a .

43

5

5

18

12

12

35

5

1-5

11

liana esculenta

72

8

7

26

25

15

6

6

5

16

,, arvalis .

61

6

6

20

19

17

6

5

o

o

8

„ temporaria .

75

6

6

35

24

19

6

12

6

17

,, greeca .

50

5-5

5-5

15

16

12

4

5

2-5

10

,, latastii .

51

4-5

4-5

15

13

12

35

4

25

8

,, agiiis

62

55

5-5

22

19

16

5

6-5

3

6

1. Length from snout to vent. 2. Length of ventricle of heart.
3. Breadth of ventricle of heart. 4. Length of inflated lung. 5. Length
of longest lohe of liver. 6. Length of kidney. 7. Breadth of kidney.
8. Length of testis (in case of difference between the two testes, the
longer is measured). 9. Breadth of testis. 10. Length of corpora
adiposa.

HABITS. 57

VII. Habits.

Frogs and toads were classified by the earlier authors
into terrestrial, aquatic, and arboreal. It would be
difficult now, nay impossible, to draw any sharp demar-
cation line between tbe three types, and all we can do
is to emphasise the differences which exist between the
extreme forms. Thus by opposing the extremes we
can show examples among the European representa-
tives of terrestrial and aquatic (Alytes obstetricans —
Eana esculenta) t fossorial and scansorial (Pelobates —
Hyla), saltatorial and cursorial (Rana agilis — Bufo
calamita), diurnal and nocturnal {Rana esculenta —
Alytes obstetricans) types, which are, however, con-
nected by insensible gradations. These adaj)tations
are revealed by certain structural characters, such as
the development of the web between the toes to facili-
tate natation, the great development of the inner
metatarsal tubercle to assist in burrowing, the adhesive
digital disks for climbing, the relative length of the
hind limbs for jumping, and the shape of the pupil,
whether round or horizontal or vertical, as indicative
of more or less diurnal or essentially nocturnal habits.
But here and there we meet with exceptions; such
are, for instance, the fully-webbed toes of Pelobates,
which spends but a short time of the year in the
water; the shovel-shaped metatarsal tubercle of liana
esculenta, var. lessonse, which does not burrow; or the
similarity in shape of the pupil in Rana and Bufo,
which yet differ so much in their habits. Still, on the
whole we may roughly incorporate the species outside
the breeding season into the following categories:

Diurnal and aquatic : Rana esculenta, Discoglossus,
Bomhinator.

Diurnal and terrestrial : Rana temporaria and allies.

Nocturnal and terrestrial : Bufo, Pelodytes, Alytes.

Nocturnal and fossorial : Pelobates.

Nocturnal and arboreal : Hyla.

That most, though not all, are aquatic during the

58 INTRODUCTION.

breeding season is mentioned in the chapter on
pairing and oviposition. All our species also spend a
certain period of their existence as gill-breathing,
fish-like larvas : there are exceptional forms which
entirely dispense with the larval life, and hop oat of
the egg in the perfect condition, but we have not to
deal with them here, for they are only found between
the tropics. Even our most aquatic species leave the
water after metamorphosis, and young specimens of
liana esculenta and Bombinator are terrestrial, or live
more on the border than actually in the water until
able to breed. Sexual maturity is not attained before
the third or fourth year in males, a year or two later
in females — of the larger species, at any rate. The
growth is slow, and so prolonged that of certain species,
such as liana and Bufo, one may say the older the
specimen the larger ; the gigantic size attained by
some in secluded localities appears to be mainly
due to the immunity they have enjoyed for a great
number of years. Individuals of the common toad
have been observed for very long periods, and it
is probably no exaggeration to assess the possible
duration of life in that species at a century. Among
the smaller forms a tree-frog has been kept in con-
finement for twenty-two years.

The food consists exclusively of live prey ; no
insect or worm will be taken that has not moved in
the presence of the consumer. But it is easy to deceive
a Batrachian by agitating any object ; the edible frog is
thus captured in France by means of a red rag put on
to a line. In fact, any moving object of commensu-
rate size will be taken, to be afterwards rejected if dis-
tasteful. Discoglossus and Boittliinatur alone among our
Batrachians are able to seize their prey under water.
The thoroughly nocturnal forms cannot be induced
to feed in the daytime. Frogs and toads occasionally
display cannibal instincts, swallowing young of their
own species. The prey is secured by throwing out
the tongue, except in the Disaxjlossidai, which, not

HABITS.

59

being provided with such a mechanism, simply seize
it with their jaws, as does a newt ; the hands often
assist in pushing it into the mouth. A certain
quantity of sand or fine gravel is usually found in
the stomach and the large intestine, together with
beetle elytra and other indigestible matter. As in
many birds, the faeces are enveloped in a muci-
laginous coating formed in the large intestine.

Although sensitive to cold and retiring at the
approach of winter, Batrachians do not fall into
complete lethargy. Hibernating specimens found in
holes, under heaps of manure or dead leaves, or even
buried in the mud under the w7ater, are only sluggish,
not dormant.

Salt is fatal to most Batrachians and to their eggs.
A frog soon perishes when thrown into sea water.
Yet Bufo calamita is in this respect exceptional,
often breeding in brackish pools or burrowing in sand
strongly impregnated with salt. The same toad is
also remarkable in being less partial to moisture than
any other of our Batrachians.

Among the species that remain concealed during
the daytime or in inauspicious weather, Bufo vulgaris
and Pelobates lead a solitary existence ; whilst Bufo
viridis and calamita and Alytes are of gregarious
habits, two individuals or more being usually found
in the same hole or under the same shelter.

Brief allusion should be made here to three popular
beliefs, which, although often refuted, still crop up
now and then, and, curiously, occasionally find cham-
pions in educated men.

The first, that toads squirt poison at their enemies,
is explained by the fact that these Batrachians, when
frightened and trying to escape, shoot out, to a con-
siderable distance, liquid from the vent. But this
liquid, tapped from the bladder, is as innocuous as
pure water, and has nothing to do with the poisonous
secretion of which toads are really possessed, but
unable to squirt out spontaneously.

60 INTRODUCTION.

The second is the supposed showers of frogs and
toads, believed to fall with the rain, and to have been
carried by a waterspout, some narrators of the
phenomenon even stating to have received them on
their hats or open umbrellas. But the fact that
these so-called showers consist of young frogs,
never of larva?, and only occur at the time of the
metamorphosis of the common species, when myriads
leave the water and conceal themselves in holes
and fissures in the soil, whence they suddenly emerge
when the rain falls, sufficiently accounts for the
phenomenon. Half blinded by the rain, people,
startled at the sudden appearance of these legions of
tiny Batrachians on the ground around them, actually
fancy they feel them falling, the delusion being further
enhanced in the case of baby tree-frogs, which, under
the circumstances, climb up their clothes.

The third belief, in live frogs and toads enclosed in
stones, hardly deserves refutation, when we know that
air, moisture, and food are indispensable to these
creatures. If every case could be properly investi-
gated, it would be found that the quarry workman,
from whom such tales invariably originate, has been
the victim of a delusion, and that the Batrachian he
fancied to have hopped out of the stone he was break-
ing lay concealed close to it unobserved, and jumped
from its retreat at the blow of the hammer.

And, finally, it remains to observe that our Batra-
chians do not bite, although some exotic forms do.
However, Pelobates, when irritated, assumes a very
aggressive attitude, screaming and jumping with open
mouth towards its tormentor, as if to snap at him.
No doubt these antics, accompanied by a repulsive
odour produced at the same time, must act as a
protection against snakes and carnivores. The Bom-
binator are also believed to startle pursuers by turning-
over and suddenly exhibiting the brilliant coloration
of their lower surfaces.

VOICE.

61

VIII. Voice.

All our male Batrachians are endowed with a voice,
which they produce at least during the pairing season.
Females are mute, or respond to the male by a mere
grunt. The larynx is provided with vocal cords,
which are set vibrating as the air is rapidly shifted
from the lungs into the buccal cavity. In many
species the sound is intensified by resonance in
special vocal sacs situated in the gular region, or at
the sides of the head behind the commissures of the
jaws. The vocal sacs are called internal when

Fig. 23.

K

>

B

A. Rana esculenta,^, with inflated external vocal sacs (lower view).

B. Rana temporaria, $, with inflated internal vocal sacs (lower view).

covered by the unmodified gular integument, how-
ever much this may be distended ; external when their
membrane, a diverticulum of the mylohyoid muscle,
projects through slits at the sides of the throat, as
in Rana esculenta, or when the skin is thinned and
converted into a bladder-like pouch, as in Hi/la
arborea. In some forms there are two distinct
sacs, in others but one. The air penetrates by one or
two openings in the floor of the mouth, small, rounded,
and situated near the commissures of the jaws, as

62

INTRODUCTION.

in Ran a, or slit-like, and situated at the side of the
tongue, as in most other genera. Pelodytes and Hyla
have two openings, whilst Bitfo has commonly but one,
either on the right side or on the left.

In Bombinator igneus internal vocal sacs are well
developed, but are formed by the skin of the floor of
the mouth, which is loose and plicate, and projects
through a slit dividing the submaxillary muscle into
an anterior and a posterior portion. Diseoglossus has
rudimentary vocal sacs of the same type, but they are
confined to the sides of the throat, near the mandible.
Bombinator pachypus, Alytes, Pelobates, Bufo vulgaris,
and Bana grseca, latastii, ibcriea, and agilis lack the
vocal sacs.

The mode of inflation of the vocal sac is best
observed in Hyla arborea, the bladder-like appendage
when blown being of enormous size, nearly as large
as the body, which, when the animal croaks, is much

Fig. 24.

Hyla arborea, $ , with the vocal sac in the collapsed and inflated

conditions.

thinned by the emptying of the lungs, as shown in
the accompanying figure (Fig. 24). This mechanism of
the shifting of the air from the lungs into the throat,
and vice versa, explains the fact, paradoxical in appear-
ance, of Batrachians being able to croak under water.
The voice varies very considerably according to the
species, and the names bombina, sonans, eampanisona,
ridibunda, cachinnans, &c, convey the impressions
made on the ears of the earlier observers. It affords
the trained collector one of the surest means of ascer-

voice. 63

taining the Batrachian fauna of a district during the
spring and early summer months; for although it
may not lead at once to the discovery of the performer,
owing to his most deceptive ventriloquial accomplish-
ments, the whistling argentine note of Alytes obstet-
ricans (often attributed by country people to the
salamander), the hoo-hoo of Bombinator pachypus, the
oonk-oonk of Bombinator igneus, the crech-creck-creck
of Pelodytes, the clock-clock-clock of Pelobatesfuscus, are
so characteristic as to be surely recognised by whom-
soever has heard them before. The sound uttered
by the breeding Bufo vulgaris may be compared to a
feeble oft-repeated bark ; that of Rana temporaria may
be rendered by grook, grook, and that of Rana agilis and
arvalis by co-co-co. But these are all comparatively
feeble monotonous sounds. If we turn to the loud
croakers, Rana esculenta, Hijla arborea, and Bufo
calamita, all three of which are provided with external
vocal sacs, we find a more varied performance, which
when produced by a large number of individuals, as is
usually the case, is simply deafening, and audible
miles away. The croaking of Rana esculenta has been
admirably rendered by Aristophanes in his chorus of
frogs, Bpe/cEK£/c££ /coct£ /coci£. Hi/la and Bufo calamita are
more difficult to imitate, their song consisting of an
extremely rapid succession of highly sonorous vibrat-
ing notes, bra or era.

These loud vocalists are in the habit of joining in
choruses, ceasing suddenly at the approach of the
listener. But they can easily be induced to resume
the concert by an imitation of their croak either by
the human voice or by strokes on a metal object. In
confinement they can also be enticed to croak b}r
beino- rubbed on the back.

Most frogs when seized by a snake or other enemy
utter a shrill cry. That of Pelobates fnscus is most
startling; and specimens of that species, when per-
sistently teased, can be aroused to a fit of anger,
accompanied by loud cries, lasting for several minutes.

64 INTRODUCTION.

IX. Pairing and Oviposition.

It is often stated in books that, in temperate
climates at least, tailless Batrachians have a fixed
annual period of reproduction, taking place in the end
of winter or in spring. This statement, based on a
generalisation of the familiar phenomenon presented
by our two common Northern species, liana tempo-
ral* ia and Bufo vulgaris, is erroneous, individuals of
some species — those of the family Discoglossidse, for
instance — breeding several times a year, at distant
intervals ; and between these two extreme types we
have almost every possible gradation.

In a first category, to which Bufo vulgaris, Bana
temporarta, and Bana arvalis belong, the pairing
season is of short duration, and although, with us,
dependent on atmospheric conditions and regulated
by the rise of the thermometer rather than by the
calendar, coincides almost to a day for all the in-
dividuals under equal climatic conditions. The males
are endowed with genesic fury to the extent of letting
themselves be mutilated or even immersed in spirit
without letting go their mate, to which they cling
tightly with their powerful fore limbs, awaiting the
expulsion of the ova to discharge on them their
seminal fluid. In their blind frenzy they will clasp
individuals of other kinds, even fishes, putrefied
corpses of females that have succumbed to their
embrace, or all sorts of floating objects. Common
toads may be fished by holding out to them a thick
stick, to which they cling with their arms. As males
of that species are always more numerous than
females, great fights ensue, in which the latter
sometimes perish under the pressure of several of
their would-be possessors.

PAIRING AND OVIPOSITION. 65

Even iu this category there are occasional excep-
tions. Such are the cases recorded by Martin and
Rollinat and W. Evans, of single couples of the
common toad found breeding near Argenton, in
France, on the 18th June, and near Edinburgh on
the 13th June, two or three months after the other
individuals in the same localities had deposited their
spawn and left the water. These exceptional cases
are probably due to some accident having prevented
the female from resorting to the breeding-place at
the proper time. Males, we know, may long retain
their genital ardour, so many of them, owing to their
excess in numbers, being unable to find a mate.

"We next find species which may also be said to
have a short breeding season, but which do not con-
gregate with the same ensemble, the condition of the
genital products not being the same at a given time
in all the individuals. In these species the female
does not, as a rule, enter the water until ready to
spawn, and the embrace is, in consequence, of short
duration, often taking place only at night. To this
category belong Pelobates, Pelodytes, Hyla, and Rana
agilis. The males are not, as a rule, animated with
the same frenzy as we observe in the first type, and
will often let go the females when disturbed or
handled.

A third category is represented by Bufo calamita,
Bufo viridis, and Rana esculenta, which spawn but
once a year, but the breeding season of which extends
over a longer period, viz. two or three months, within
the same district, with an ultimum mostly depending
on the weather.

Lastly, all the species of the family Discoglossidse,
and apparently also some individuals of Pelodytes, are
able to breed twice or more every year. Bombinator
spawns first in spring and again in summer, some-
times as late as August or the beginning of September ;
Discoglossus and Alytes as often as three or four times
at distant intervals during the warmer months, the

E

66 INTRODUCTION.

former (inhabiting the extreme south) from January
to October, the latter from the end of February or
beginning of March to the beginning of September.

The time at which each species commences to pair
varies, as we have seen, with the temperature, for the
early breeders at least. Thus in the south of Europe
these would start from January, whilst in the north or
at corresponding altitudes in the mountains represen-
tatives of the same species would be compelled to wait
until the thaw has set in. Again, exceptionally pro-
longed winters may cause the individuals of a species
to be retarded in this operation, so as to bring them
together with others which, under normal circum-
stances, do not breed until some weeks later. But,
on the whole, our Batrachians may be arranged in the
following order, beginning with the most precocious :

1. Discoglossus, liana teinporaria, B. arvalis.

2. Bana a gills, Alytes.

3. Bufo vulgaris, Belobates, Belodytes.

4. Bufo viridis, Bufo calamita, Hyla, Bombinator,

Bana esculent a.

Taking the three British species as an example,
Rana temporaria in normal years with us breeds in
the beginning of March, Bufo vulgaris in the beginning
of April, and Bufo calamita from the middle of May ;
their nuptial periods do not even overlap.

Leaving out the genus Alytes, which is altogether
exceptional in its mode of parturition, all our Batra-
chians are compelled to resort to the water for the
purpose of depositing their spawn. In some species,
such as our common frog, the males precede the
females, and may be found in winter in a semi-torpid
condition, but already attired in their full breeding
costume, at the bottom of pools thickly coated with
ice. The common toad travels long distances on land,
or following the course of brooks, in quest of suitable
breeding- places ; and if the sexes meet on the way the
male often secures immediate possession of the female,
accomplishing the rest of the journey on her back.

PAIRING AND OVIPOSITION. 67

The Natterjack toad, B. calamita, also often pairs on
land, and being a bad swimmer, and only spawning at
night, numbers in embrace may be found in the day-
time in boles on the banks, their presence being
revealed by their loud croak. The tree-frogs usually
pair only at night or in the evening. Thoroughly
aquatic species like Bombinator and Rana esculenta
resort, of course, to the water before mating.

Ahjtes, the most terrestrial of European Batrachians,
pairs and oviposits on land. The female never goes
into the water, and the male contents himself with a
hip-bath for the purpose of releasing his progeny from
the egg-capsules entrusted to his care.

Species differ greatly in the choice of a site for
their nursery. Alytes exercises the greatest judg-
ment, avoiding ponds or pools already largely stocked
with tadpoles of other species, or any water which is
not permanent. The common toad and the tree-
frog, also, are judicious in their choice, and their
offspring are never doomed through drying up of the
site selected, an eventuality which the common frog
and the Natterjack toad do not appear to be able to
foresee. The latter species especially often spawns
in roadside ditches and puddles of a most temporary
nature, although suitable places may be near by, and
we often find masses of their young tadpoles accumu-
lated in a small hole with scarcely any water left,
exposed to the sun -rays, where they must perish unless
saved by a timely rainfall. Frost often destroys the
brood of the common frog.

It has been observed by Heron-Royer, and I have
more than once been able to verify the fact, that
different species which breed in the same water some-
how manage to keep clear of each other. In a pond
near Paris the above-mentioned author found Rana
temporaria occupying the north and west, Rana agilis
and Bufo vulgaris the north-east, Peloch/tes punctatus
the south-east, and Hula arborea the south, east, and
west. In this country, where Rana temporaria and

68

INTRODUCTION,

Bufo vulgaris often breed in the same place, we always
find the tadpoles forming distinct colonies. A dip of
the net will secure specimens of one species only.
Courtship does not take place in any of the tailless

Fig. 25.

•w

.—

/ W

Pairs in embrace, a. Discoglossus pictus. B. Pelodytes punclatits.

Batrachians. The female is seized by the first comer,
and when he has once secured his hold he will remain
for a period varying between a few minutes and
several days in possession until the evacuation of the
ova, unless he be dislodged by a stronger competitor.

PAIRING AND OVIPOSITION.

69

Impregnation accompanies or immediately follows, in

one or more emissions, the discharge of the eggs.

The mode of embrace varies according to the species ;

it is axillary in all our genera with horizontal pupil,

and lumbar in the others, as shown in the following

synopsis :

Fig. 26.

- J||

•J <J

Pairs in embrace. A. Bufo vulgaris, b. Bana arvalis.

I. Male holding: the female round the waist.

a. The hands joining on the pubic region (Fig.
25, a).
Discoglossus, Bombinator, Ahjtes, Pelobates.

70 INTRODUCTION.

b. The forearms meeting on the pubic region
(Fig. 25, b) .... Pelodytes.

II. Male holding the female behind or above the
arms.

a. The hands dug into the axils or just above

them (Fig. 26, a).

Bufo vulgaris, B. calamlta, Hyla arborea.

b. The hands pressed against the breast (Fig.

26, b) Bufo viridis, Eana.

Owing to the mode of embrace the fore limbs of
the males are always stronger than those of the other
sex, and in some species acquire about the breeding-
time a great muscular development. The inner side
of the inner finger may develop a thick pad, supported
by a corresponding development of the bones of the
rudimentary pollex and first functional finger, which,
pressed against the breast or pubic region of the female,
assists in maintaining the hold, often leaving a deep
scar if the pairing lias been of prolonged duration.
There are developed in addition temporary horny
excrescences, in the form of small granules or spines,
which are present in most species, and afford useful
characters for their distinction during the breeding-
season. When this is over the excrescences are shed
in most species. They, however, often persist through-
out the year in Bufo, Discoglossus, and Bomblnator.
These nuptial asperities are absent in Ahjtes and
Pelobates, and scarcely distinguishable in our Hyla.
When present, their shape and coloration, as well as
their distribution on various parts of the limbs or even
of the body, vary according to the species. With
regard to distribution, wre find them located in fully
developed breeding individuals —

On the inner side of the two inner fingers, the ante-
brachium, the brachium, the breast at the base
of the arm, the chin, the belly, and along the
toes (Fig. 27, b) . . . Pelodytes.

On the inner and upper side of the three inner

PAIRING AND OVIPOSITION.

71

Fig. 27.

D

Nuptial excrescences of males, a. Discoglossus pidns. b. Pelodytes
punctatus. c. Bombinator pachypus (hand aud foot). D. Bufo
vulgaris (hand), e. Bana temporaria (hand).

i I INTRODUCTION.

fingers, on the chin, the belly, and on the free
edge of the web between the toes (Fig. 27, a).

Discoglossus.

On the inner side of the three inner fingers and

the antebrachium, and on the third, the second

and third, or the second, third, and fourth toes

(Fig. 27, c) . . Bombinator pachypus.

On the inner side of the two inner fingers and the

antebrachium . . Bombinator igneus.

On the inner side of the three inner fingers

(Fig. 27, d) Bufo.

On the inner and upper side of the inner finger
(Fig. 27, e) . . . . Rana.

At the base of the inner finger . . Hyla.

It sometimes happens that old females present at
least traces of these organs, the appanage of the males.
I have recorded such a case in a Rana temporaria with
eggs in the oviducts ; Bosca has found the same
anomaly in a Pelodytes punctatus, and Mehely in five
specimens of Bombinator pachypus. These horny
productions are usually dark brown or black ; they
are greyish in Rana esculenta and agilis, colourless
in Hyla arborea. Their aspect under the microscope,
and the differences they present according to the
species, have been studied by Lataste, who has shown
that they are low and obtuse in Rana esculenta and
agilis, erect and pointed in Rana temporaria and arvalis
and Discoglossus, pointed and oblique or hooked in
Bombinator and Pelodytes, pointed, erect, and crenu-
lated in Bufo vulgaris and viridis, and obtuse and
crenulated in Bufo calamita (Fig. 28).

As in all Batrachians in which no courtship takes
place, the males are not distinguished by any orna-
mental appendages or more vivid colours, but in Rana
temporaria and arvalis they acquire during the pairing-
season a peculiar swelling of the skin, by which the
back, and especially the gular region, take on a bluish
tinge and a flabby texture, strongly contrasting with
the aspect of the same parts later in the year. The

PAIRING AND OVIPOSITION.

73

females of the same species, on the other hand, develop
on the back and limbs, during the breeding season,
peculiar pearl-like structures of the epidermis, which
are regarded as sensory, being provided with a special
innervation, same as we find developed during the
spawning season in the males of many Cyprinoid and
Salmonoid fishes. These organs have been dealt with
at great length by Huber and Maurer.

Fig. 28.

MMMm?A

B

■ys.

HP P^

sf>MiP0n. f\:

D

1~«L

E

V*

■Sy*.^

W

,

Sections through the nuptial asperities of males, much en-
larged (after F. Lataste). A. Bana temporaria. b. Rana
esculenta. c. Bufo vulgaris. i>. Bufo calamita. E. Disco-
glossus pictus. F. Bombinator paclujpus.

The males of most of our Batrachians make them-
selves conspicuous during the pairing time by their
voice, whether they be provided with vocal sacs or
not. Although there are some, liana esculenta and
Hyla, in which the clamour is not restricted to the
breeding season, most others are comparatively mute
outside that period, and the vocal sacs undergo a sort
of regression, as is especially noticeable in Bana
temporaria and Bufo calamita.

74 INTRODUCTION.

Our larger species of toads and frogs do not
appear to be able to propagate until four or five
years old. But it has been ascertained on specimens
kept in confinement that Alytes, Bombinator, Pelobates,
and Hi/la — which are of proportionally larger size
immediately after transformation — attain much sooner
their full development, and may breed in the third
year of their existence.

A curious fact, for which we can at present find no
explanation, is the very considerable excess of males
over females in the genera Pelodijtes, Pelobates, and
Bufo, when adult at least, as has been observed not
only during the breeding season, but also at other
times, and especially during the winter, when numbers
congregate in holes. In other genera the proportion of
both sexes appears about equal ; Born and Yung have
even ascertained that in the young of the genus Bana
the females are in the proportion of 54 to 61 per cent.
It is believed that in some lower animals abundance
of food is an important factor determining sex. The
two above-mentioned physiologists have conducted a
series of experiments on tadpoles of frogs (Bana
temporar la), and by giving them a more abundant
or more artificial food have obtained a much hio-her
ratio of females, viz. 70 to 95 per cent. A. von
Griesheim, after examining the sexes of 440 specimens
of the common frog captured immediately after the
metamorphosis, found only 36 per cent, of males. It
would be highly interesting to make a similar investi-
gation on the young of some species, such as Bufo
vulgaris or Pelobates fuscus, in which the number of
adult males far exceeds that of females.

SPERMATOZOA. 75

X. Spermatozoa.

The seminal elements of tailless Batrachians Lave
received a great share of attention, not only on the
part of anatomists and physiologists, but even of
systematic workers. This is clue to the great amount
of difference in structure and size between these
elements in the various genera; and also to the fact
that otherwise closely allied species of the genus Rana
may differ in their spermatozoa more considerably
from each other than from others morphologically
more remote. The failure to obtain hybrids between
such near allies as the male Rana temjporaria and the
female R. arvalis has been attributed principally to
these differences ; for other species of the genus Bufo,
which stand very far apart, but which have almost
identical spermatozoa — as, for instance, B. vulgaris and
B. calamita, and B. vulgaris and B. viridis — cross with
comparative facility under artificial fecundation.

One great source of errors in describing and com-
paring the spermatozoa has been due to the changes
which these elements undergo in their development,
and care should be taken to ascertain whether or not
they are in a ripe condition. The safest way to avoid
mistakes is not to extract the spermatozoa from the
testicles, but to collect them in the seminal ducts
from specimens in embrace. This is, however, difficult
to do in species which are destitute of a seminal
bladder.

Thanks to the investigations of Leydig, La Valette
St. George, Spengel, and Pfliiger, which I have been
able to supplement for some of the less known species
of Rana, we are now well acquainted with the sperma-
tozoa of nearly all the European species.

Those of Discoglossus are remarkable for their great
size, measuring three millimetres, whilst those of other
species do not exceed one-tenth of a millimetre. The
so-called head is spirally wound like a corkscrew ; the

76

INTRODUCTION.

tail is about equally long, and provided with an ex-
tremely delicate, wavy, vibratile, hyaline membrane
or crest. As in the newts, they are collected together
in bundles, although devoid of a spermatophore.

Fig. 29.

a b c r> E F

Spermatozoa of Batrachians. A. Bombinator pachypus. B. Pelo-
bates fuscus. c. Bufo vulgaris, d. Hyla arborea. E. Rana
esculenta. F. Rana tem/poraria. (After La Valette St. George
and Leydig.)

In Bombinator a crest is present along the side of
the spindle-shaped head, and the tail is moderately
long, filiform, and devoid of a crest. The crest may
be partially detached from the head, and has been
taken to belong to the tail.

CD

In Alytes and Bufo the head is staff-shaped, pointed
in front ; the tail is about twice as long, with a deli-
cate crest.

In Pelobates the head is long and thin, with strong
spiral windings, corkscrew-shaped, passing gradually
into the extremely long, delicate, filiform tail.

The spermatozoa of Pelodytes bear much resem-
blance to the preceding, but the head is less twisted ;
the tail is also extremely long and filiform.

SPERMATOZOA. 77

In Ilj/la arbor ea the head is spindle-shaped, acutely
pointed in front, obtusely behind ; the tail filiform,
and about twice as long.

In Bana esculenta and arvalis the head is staff-
shaped and obtuse in front, and about half as long as
the filiform tail; aud in Bana temporaria and agilis
the head is very long and thin, not or but scarcely
longer than the tail, and attenuate into a fine point at
both ends.

The spermatozoa of B. grseca and latastii bear a
greater resemblance to those of B. arvalis, but have a
more elongate head, the diameter ten to twelve times
in the length ; the head is cylindrical, feebly attenuate,
and quite obtuse at both ends.

The spermatozoa of B. earner ani and B. iberica are
still unknown.

Attention has been drawn above to the importance
of securing for comparison the seminal elements in
their fully ripe condition. La Valette St. George has
shown that at certain stages of development these are
almost identical in B. esculenta and B. temporaria ;
and that accumulations of protoplasm at certain
points of the unripe element or spermatocyte, as he
calls it, produce a shape which may be very unlike
that of the true spermatozoon. So careful an observer
as Leydig has unfortunately fallen into the error of
figuring spermatocytes as spermatozoa in Bombinator,
Bufo, and Hyla.

Pfltiger has observed that spermatozoa wTith pointed
heads possess a greater facility for penetrating the
envelops of the ova of other species to reach the
nucleus, a fact which explains the different results
obtained in attempts at crossing in both directions,
to which highly interesting experiments allusion will
again be made in the chapter on hybrids.

78 INTRODUCTION.

XI. Eggs.

The eggs are spherical vitelline bodies, surrounded
by a thin, elastic, cortical membrane or chorion and
one or two gelatinous envelops, formed during their
passage down the oviducts : the outer capsule swells
out in the water after ovi position, and varies accord-
ing to the families or genera; the inner is absent in
the Pelobatidae. With the exception of Alytes, the
upper pole of the vitelline sphere is always pigmented,
varying from pale brown to black, and this coloration
may extend on the whole sphere, — as, for instance, in
Bufo vulgaris, or on the whole save a small whitish
spot at the lower pole, as in liana temporaria. The
inner mucilaginous envelop is formed in the upper
portion of the oviduct, and surrounds each ovum ; the
outer, which has been termed " adhesive envelop," is
formed lowrer down the oviduct, and either surrounds
each single ovum, whether free or agglutinated in
masses, or forms a common investment in which the
ova are disposed irregularly or in files.

All eggs, except those of Alytes, are deposited in
the water. If laid singly, each is fixed by its adhesive
envelop to submerged bodies ; if in strings or bands,
these are twined round plants. Egg-masses are usually
also fixed to plants, except those of Rana temporaria,
which are endowed with a peculiar buoyancy, and
simply float on the surface.

The eggs of some Batrachians have a strong odour
of fish, this being most noticeable, among European
forms, in Pelobates fuscus.

The amount of protection which the mucilaginous
envelops afford the embryo varies considerably. In
Discoglossus, Pelodytes, Pelobates, and Bufo they soon
partly dissolve, so as to release the embryos almost
before these are able to execute any spontaneous move-
ments ; the embryos, so to say, drop out and become
fixed by their adhesive subbuccal apparatus to the

EGGS. 79

outer surface of their remains. Iu Bombinator, Hi/la,
and Rana the embryo develops much further within
the egg-, and becomes liberated by its own action.

In Alytes, which is exceptional among European
Batrachians in the eggs being laid on land in strings,
taken charge of by the male parent, who carries them
about twisted round the legs, all the embryonic stages
are passed through within the egg, and the young is
born as a true tadpole. Contrary to the other Batra-
chians, the food-yolk of which is in insufficient quantity
to form an external appendage of the embryo, we find
here a large yolk-sac. These eggs, which measure
three and a half or four millimetres in diameter, are
of a uniform yellow colour, and are protected and
connected in rosary-like fashion by a comparatively
tough but highly elastic transparent investment,
which represents the adhesive envelop of the typical
Batrachian ovum, and contracts thread- or hair-like
between every two eggs.

The following synopsis shows the principal dif-
ferences between the eggs of the European species,
and will serve to their identification, the dimensions
given being those of recently deposited ones.

I. With large unpigmented vitelline sphere, 3^ — 4
millimetres in diameter, laid on land in rosary-
like strings, carried by the male twisted round
his legs, the larva leaving the envelops in an
advanced condition after the loss of the ex-
ternal gills and the formation of the spiraculum.

Alytes.

II. AVith small vitelline sphere (1 — 3 millimetres in

diameter), entirely or partially brown or black,

deposited in the water, the embryo leaving the

envelops with external gills or before their

appearance.

a. Deposited singly or in small groups of two to

twelve ; vitellus brown or black above, white

or yellowish beneath, the embryo leaving the

80 INTRODUCTION.

envelops with external gills or just before
their appearance, with the tail more or less
developed.
Vitellus 1 — 1| millimetres in diameter, black

above Discoglossus.

Vitellus 2 millimetres in diameter, brown above.

Bombinator.

b. Irregularly disposed iu a thick mucilaginous

band (formed by fusion in the cloaca of the
contents of the two oviducts) ; vitellus dark
brown or black, with the lower pole white,
1^ — 1\ millimetres in diameter, the embryo
leaving the envelops before the appearance
of the external gills and tailless.

Pelodytes, Pelobates.

c. In regular files in two long1 mucilaginous

strings ; vitellus entirely dark brown or
black, or with a whitish lower pole, the
embryo leaving the envelops before the
appearance of the external gills and with a

rudimentary tail Bnfo.

Vitellus 1^ — 2 millimetres in diameter; eggs, in
each string, in three or four files when floating,

in two when stretched B. vulgaris.

Vitellus 1 — \\ millimetres in diameter; eggs in
three or four files when floating, in two when

stretched B. viridis.

Vitellus 1 — 1^ millimetres in diameter, with
whitish lower pole ; eggs in two files when
floating, in one when stretched . B. calamita.

d. In large lumps ; upper pole brown or black,

lower whitish or yellowish ; the embryo

leaving the envelops with external gills and

a well-developed tail.

1. Envelop, when fully swollen out, measuring

3 — 4 millimetres in diameter; vitellus

yellowish white with brown upper pole, 1J

millimetres in diameter; embryo yellowish.

Ilij la.

Fio. 30.

@®

T,

D

• •

$

:^»:

9

m

h

9

«*'

G

(fti

m

■

G

Eggs of — a. Bambino tor pachypus. b. Discoglossus] pictus.
c. Alytes obstetrieann. r>. Pelodytes punctatus. E. Bufo
vulgaris. F. Hj/Za arborea. a. Rana temporaries.

F

82 INTRODUCTION.

i •

2. Envelop 7—10 millimetres in diameter
embryo brown or blackish . . . liana.

(a) Vitellus miparti, brown or blackish

above, yellowish or white below, or
the light spot covering at least the
lower third ; eggs submerged.

Diameter of vitellus 1^ — 2 millimetres.

B. esculenta, B. arvalis.

Diameter of vitellus 2 — 3 millimetres . B. agilis.

(b) Vitellus nearly entirely black, with a

small whitish lower pole.
Diameter of vitellus 2 — 3 millimetres; eggs float-
ing B,. temporaria.

Diameter of vitellus 1^ — If millimetres; eggs

submerged B. latastii.

It may be noticed that, so far as species are con-
cerned, the size of the egg stands in no relation to
that of the parent, a fact which is also conspicuous in
the tadpole. Tims the largest Batrachian, Bufo vul-
garis, has eggs no larger than the smallest, Belo-
dytes punctatus ; and in Bana temporaria, they are
considerably larger than in B. esculenta, which reaches
both in the larval and perfect states to a much greater
size. Heron-Royer has observed that young females
produce eggs of a somewhat smaller size than those
of full-grown specimens of the same species, and 1
have been able to verify his observation on Bana
temporaria.

The number of eggs also varies according to the
size of the female. Heron-Royer, who has counted
them in most of the European species, gives the
following numbers as the result of his computation :

Bufo viridis 10,000—12,000

Bana esculenta .... 10,000

Bufo vulgaris 4,972— 6,840

Bufo ealamita 3,000— 4,000

Bana temporaria .... 2,856 — 4,005
Pelobates fuscus .... 1,200— 2,236
Bana arvalis 1,000— 2,000

EGGS. 83

Pelodytes punctatus . . . 1,000 — 1,630

Bana agilis 669 — 1,200

Hyla arborea 800—1,000

These results, however, do not always agree with
my own, as will be seen further on under the various
specific headings.

In the Discoglossidx, which spawn two or three
times a vear, each brood consists of 300 to 1000
eggs in Discoglossus pictus, 80 to 100 in Bombinator
pachypus, and 20 to 90 in Alytes obstetricans. This
makes a probable maximum total of 3000 for the first
species, 300 for the second, and 270 for the third.
These numbers are, however, only approximative, for
no observer has yet been able to follow the spawning
of a given specimen throughout the year, and to record
the total number of eggs produced.

Manipulating frogs' eggs is not easy, on account
of the sticky mucilage which adheres to the fingers or
instruments, and the counting is a very trying opera-
tion. Approximate computations are more easily
arrived at with the egg-strings of toads, which can be
divided into small portions, in which the eggs are
counted and the result multiplied by measuring the
length of the whole strings.

Fig. 31.

Cleavage of egg of Bana tempora/ria (after Ecker).

After fecundation the vitelline sphere undergoes a
process of division, or segmentation, which can be
readily observed under the microscope or an ordinary
hand lens, and of which the above diagrammatic

84 INTRODUCTION.

figure from Ecker's ' Icories Physiologica3 ' will give
an idea.

This type of segmentation, in which the first furrow
completely cleaves the wrhole egg into two, each suc-
cessive one again subdividing that cell with which
it is related, the first two furrows vertical and the
third horizontal, is termed complete or holoblastic.
The first plane of division passes through the plane of
symmetry of the future embryo, the tissues of which
are formed from these cells. The egg of Alytes
contains a much greater quantity of nutritive matter,
and the segmentation is less regular and more limited ;
it belongs to the meroblastic type.

The time required for the eggs to hatch varies con-
siderably, according to the species. Thirty hours
may suffice for those of Discoglossus irictus, whilst
those of Alytes obstetricans take at least, under the
best conditions, nearly three weeks. The eggs of
other species fall between the two extremes, the
evolution being of course accelerated or retarded by
the variable temperature with which early breeders
have to contend, or even temporarily arrested when
the thermometer descends to freezing-point or below.

DEVELOPMENT AND METAMORPHOSIS. 85

XII. Development and Metamorphosis.

In the first condition, when the embryo has become
distinct from the vitelline sac, the head is large and
distinct from the elongate body, the tail absent or
rudimentary. The head is usually cleft below by a
median longitudinal groove, in the middle of which a
transverse or rhomboidal depression represents the
first rudiments of the mouth ; on each side, in front
of the mouth, a pit indicates the nostril ; behind it is
a grooved fold, the cephalic crescent, which develops
into a single or paired prominence, the holder —
"crochets "of Rusconi — acting as an adhesive apparatus
by means of which the helpless embryo fixes itself at
first to the outer surface of the mucilaginous envelop
of the egg, and later to weeds or submerged objects.
Eyes are absent. A small bud-like tubercle on each
side of the posterior border of the head is the rudi-
ment of the external gills, and vertical grooves in
front and behind the bud represent the visceral clefts,
the intervals between which will later become converted
into the four branchial arches.

The adhesive apparatus mentioned above varies con-
siderably, and its conformation affords the means of
distinguishing genera or even species at a period when
the tadpole characters are not yet developed. It is
mainly through the observations of Heron-Royer and
the more scientific researches of Thiele that we have
become acquainted with its modifications in most of
the European forms. Thiele has shown that the term
" sucker," which has been bestowed on this organ by
various authors, is a misnomer. There is no muscular
suctorial apparatus developed in connection with it ;
it is glandular and secretory, producing a sticky
mucus or slime, which serves to fasten the larva to its
resting-place.* The mode of its development and

* From this apparatus the ventral adhesive disk of certain tadpoles
living in mountain streams of the Himalayas, Burma, and the Malay
Archipelago is no doubt developed.

86

INTRODUCTION.

retrogression is shown in the following series of
figures of Bufo vulgaris (after Thiele).

Fig. 32.

Development and retrogression of the snbbuccal apparatus in

Bufo vulgaris.

It appears as a crescentic groove very early, before
any other organ, and disappears, after having under-
gone various changes, in the beginning of the tadpole
period. Its fullest development coincides with that of
the external gills. It is single and discoid, with a
crescentic groove, in Discoglossus; single and Y-shaped
in Pelodytes and Pelobates ; single and V-shaped or
crescentic in Bvfo ; and paired, forming two small
disks, in Bombinator, Bana, and Hyla. These disks
are close together in Bombinator, widely separated in
Ht/la and Bana temporaria and agilis, whilst they are
connected by a fine transverse ridge in Bana escu-
lenta, which thus affords a link between the single and
paired types of holders. Hyla differs from all others
in the disks shifting forwards so as to be, in the
later stages, situated on a level with the mouth.
The condition in Discoglossus appears to be the most
primitive, from which those shown by Bombinator,
Bana, and Hyla on the one hand, Pelobates and Bufo
on the other hand, may be derived. The following

DEVELOPMENT AND METAMORPHOSIS. 87

diagram, copied from Thiele's memoir, shows roughly
the shape of this organ, and its position with respect
to the mouth at the close of the first larval period.

Fig. 33.

p G H

A. Discoglossus pictus. b. Pelobates fuscus. c. Bufo vulgaris.
d. Bufo viridis. e. Rana esculenta. F. Rana temporaria.
G. Hyla arborea. H. Bombinator pachypus.

As the embryo grows the tail elongates and shows
a muscular portion with chevron-shaped myotomes,
bordered above and below by a membranous crest.
The gills become digitate or branched ; the olfactory
pits shift more forward, and become converted into
functional nostrils communicating with the mouth;
the eye may be detected at the side of the head,
appearing first as a pigmented ring under the trans-
parent epidermis; the mouth becomes bordered by
fleshy lips ; the anus is perforated ; and the larva
is able to feed, having thus far subsisted on the
vitellus contained in the abdomen.

The external gills, two or three in number on each
side, the second and third often rudimentary and con-
cealed under the first, are largest and persist longest
in Rana temporaria. In this species and in the toads
they are strongly pigmented, like the body ; in most
others they are not, or but feebly, pigmented. A little
smaller but likewise branched in Rana esculenta, Bufo
vulgaris, and Discoglossus, they remain very short and

88

INTRODUCTION.

Fig. 34.

-sp

unbranclied or bifid in Bufo viridis and calamita and in
Hyla arborea.

In Alytes, within the egg, the external gills are
extremely developed, but single; and the very long
and slender branches are confined to the ventral side,
as in other tailless Batrachians.

On entering the second period,
or true tadpole stage, the external
characters of which will be more
fully described in the following
chapter, an opercular fold covers
the external gills (the right some
time before the left in the forms
with lateral spiraculum), which
atrophy and are replaced by in-
ternal ones, small branched fila-
ments disposed along four cartila-
ginous arches. The anal tube is
developed ; the mouth acquires
horny mandibles and the lips
horny teeth ; the nostrils assume
a more dorsal position ; the sub-
buccal adhesive organ disappears ;
and the opercular fold having fused
with the skin above the gill-arches,
the head becomes confluent with
the globular swollen belly, in which
the extremely elongate intestine
shows through the transparent
abdominal membrane, coiled up
like a watch-spring (Fig. 34).
In the third period the hind limbs appear as buds
at the base of the tail, and gradually attain their full
development during the tadpole life. The fore limbs
grow simultaneously, and even more rapidly, but
remain concealed within a diverticulum of the bran-
chial chambers until fully formed, when they burst
through the skin (unless the left spiraculum be utilised
for the egress of the corresponding limb), leaving in

Young tadpole of Hyla
arborea, seen from
below, showing the
branchial arches {ba),
the heart (c), and the
coiled-up intestine
(i). X 3.

DEVELOPMENT AND METAMORPHOSIS.

89

front of them a small cleft, out of which the gill-
filaments of the branchial arches in process of dis-
appearance may often be seen projecting (Fig. 35).

Fig. 35

Alytes obstetricans, towards the end of the larval stage,
showing the gills (hr) protruding through the cleft in
front of the arm, and the median spiraculum (sjj).

Then only the caudal crests become reduced and the
tail gradually absorbed ; the gill-arches entirely dis-
appear ; the lungs, which had co-existed as accessory
respiratory and hydrostatic organs, assume alone (or,
rather, together with the skin) the respiratory func-

Fig. 36.

Development of Discoglossus pictus, x Ik.

tions; the horny armature of the mouth and lips is
shed in pieces ; the lips are absorbed and the buccal
cleft extends ; the eyes become free and acquire move-
able lids ; the lachrymal canal is shifted towards the

90

INTRODUCTION.

eye and perforates the lower eyelid ; the intestine
shortens; the anal tube ceases to function, and dis-
appears with the last vestiges of the caudal crest,
which had become detached from the vent ; and the
young frog, usually still bearing a stumpy tail, leaves
the water. The metamorphosis is completed.

The skull and its appendages also undergo very
important changes. The cranium in the tadpole is a
large undivided cartilage with narial openings and
large suborbital fenestras. The suspensorium of the
lower jaw is extremely elongate, and sends up a
strong process, connected with. the. cranium by a
bridge, in frout of the eye. The prasmaxillaries are

Fig. 37.

po rut

co-

pra

eo-

A. Skull of full-grown tadpole of Pelobates fnscus. B. The same
at the end of the metamorphosis, after the loss of the horny
beak. chy. Ceratohyal cartilage, eo. Exoccipital. fp. Fronto-
parietal. II. Lower labial cartilage, me. Mandibular (Mecke-
lian) cartilage, na. Nasal processes of chondrocranium. po.
Prseorbital process, pro. Prootic. su. Suspensorium (palato-
quadrate). ul. Upper labial cartilage.

represented by a single or paired cartilage, the upper
labial, loosely attached to the diverging anterior pro-
cesses of the* cranium, which supports the upper horny
beak; and to this corresponds a pair of cartilages,
the lower labials, ultimately the mento-Meckelian or
symphysial bones, supporting the lower beak and
attached to the short mandibular or Meckelian
cartilage. In the transformation of the mouth this
mandibular cartilage acquires a greater length, and
the suspensorium becomes reduced in proportion and
shifted backwards.

DEVELOPMENT AND METAMORPHOSIS. 91

The ceratohyal cartilage at first articulates by a
condyle with the suspensorium below its prseorbital
process, and extends across the gular region towards its
fellow, from which it is separated by a narrow space,
with one or two small cartilaginous pieces (basihyals,
Parker; copulse, Gaupp), behind which is a paired
plate (hi/obranchials, Parker; branchial plate, Gaupp).
To each branchial plate the four cartilaginous arches
bearing the tufted internal gills are attached by con-
nective tissue, and ultimately fuse with it (in the third
period). The homology of these branchial arches
has been the subject of much discussion. According
to Parker, they are to be regarded as subcutaneous
cartilages, and do not belong to the category of true
visceral arches ; they are called accordingly extra-
branchials — the true branchial arches of fishes and
tailed Batrachians being the ceratobranchials, repre-
sented by small styliform cartilages first attached to
the outer border of the hyobranchial cartilage, with
which they fuse, the fourth to persist throughout life,
and ossify as the thyrohyal. This view is strongly
refuted by the latest worker on the subject, Gaupp,
whose conclusions are as follows :

The hyobranchial apparatus of the larva consists
of two hyalia [= ceratohyals] (Fig. 38, ch), con-
nected mesially by a pars reuniens ; continuous with
it follows the copula [basihyal, bit'], which connects
the hyal with the branchial skeleton. The latter is
made up of two branchial plates [hyobranchials, hbr']
and four branchialia [ceratobranchials, cbr] on each
side, which pass into the branchial plates with their
inner extremities and into each other with their
dorso-lateral extremities. The whole hyobranchial
skeleton forms a cartilaginous continuum. For a
time a cartilaginous piece is developed in front of the
pars reuniens, which maybe regarded as & first copula.
During metamorphosis the larval apparatus is for the
greater part lost; the pars reuniens and the anterior
copula are absorbed ; the branchialia disappear with-

92 INTRODUCTION.

out a trace. The body of the hyoid cartilage of the
transformed animal is composed of the two united
branchial plates, the posterior copula, and of newly
formed cartilage, constituting the manubria of the
principal cornua, probably derived from the copula.

The alar and postero-lateral processes are new
formations which only appear comparatively late.
The so-called "processus thyroidei" [thyrohyals, Q
have nothing to do with the branchial processes (nor
with the ceratobranchials of Parker, which Gaupp
terms spiculci), but are remains of the posterior portion
of the larval branchial plates.

Thus, of the larval apparatus nothing is preserved
but the hyal arch, and a portion of the copula and of
the two branchial plates.

Dr. W. G. Ridewood has recently investigated the
development of the hyobranchial skeleton of Pelodytes,
and the careful figures he has drawn are reproduced
here with his kind permissiou. He agrees with Gaupp
in the main results, but shows the postero-lateral
process to be the persistent proximal portion of the
first ceratobranchial.

In Pelodytes as well as in Pelobates the anterior
processes are expanded and bent inwards, and the
cornua become broken, the anterior portion fusing
with the alar processes, whilst the posterior becomes
free from the hyoid plate, as may be seen on Fig. 38.

In order to complete this rapid sketch of the
evolution of the hyobranchial apparatus, which is of
great importance for the correct understanding of the
hyoid apparatus in the perfect form, it remains to
notice that in the Dlscoglossldx the first copula or
basihyal is of much larger size than in other frogs,
and persists longer ; whilst the second extends farther
back, and completely separates the hyobranchial
plates from each other.

The ossification of the thyroid processes takes place
only after transformation.

The only cranial ossifications to develop during the

DEVELOPMENT AND METAMORPHOSIS.
Fig. 38.

93

u: h.

if i

l. ctjrz

Transformation of the mandible and hyobranchial apparatus of
Pelodytes punctatus. (After Ridewood, ' P. Z. S.,' 1897.) Nine
different stages, No. 9 representing the adult. Figs. 1 — 7
dorsal view. Figs. 8 and 9 ventral view.

as. Angulo-splenial bone.
bh. Basihyal.
cbr. Ceratobranchial.
ch. Ceratohyal.
d. Dentary bone.
fl. Lateral foramen.
h. Hyoidean cornu.
ha. Convex surface by which
the ceratohyal articulates
with the palato-quadrate
cartilage.
hbr. Hyobranchial plate.
hgs. Hyoglossal sinus.
m. Internal or mesial part of
the mandibular cartilage.

m'. Lateral part of same.
mm. Mento-Meckelian (symphy-
sial) bone.
pa. Anterior process.
p>al. Antero-lateral process.
ppl. Postero-lateral process,
s. Space enclosed between the
hyobranchial plate and
the proximal ends of the
first and second cerato-
branchials.
sp. Spicules of branchial arches.
t. Thyrohyal.
if. Thyroid foramen.
v. Ventral splint-bone.

94 INTRODUCTION.

tadpole condition are the exoccipitals and prootics
(cartilage bones) and the fronto-parietals, or frontals
and parietals if these ossify from distinct centres, and
parasphenoid (membrane bones). The prsemaxillaries,
maxillaries, and squamosals, next in order, appear only
some time after the loss of the suctorial or larval
mouth.

Whilst the fore limbs are enclosed, each moiety of
the shoulder-girdle is widely separated from the other ;
the scapula is directed upwards, and the coracoid and
prEecoracoid form a loop turned downwards and
inwards. At the same period the ilium is per-
pendicular to the vertebral column.

The tail proper of the tadpole remains in a noto-
chordal condition, no cartilage being ever formed in
that region. But both dorsal and ventral cartilages
are developed in the basal portion of the caudal region,
and ossify as two dorsal elements or arches and a
single ventral strip behind the ninth or sacral verte"bra ;
they gradually fuse to a continuous cylinder, the
urostyle or coccyx, from which both chorda and spinal
cord ultimately disappear, to form in the adult a solid
bony rod.

The vertebras of the trunk are formed, as first
discovered by Dnges, on two different plans. In the
Discoglossidm, Pelobatidae, and Hylidse the chorda
remains for a long time exposed along the ventral
surface, and, owing to the absence of cartilaginous
formation around it, disappears without ever becoming
invested otherwise than by a thin elastic membrane ;
it can easily be stripped off below the vertebras in
specimens on the point of metamorphosing. This has
been termed by Gegenbaur the epichordal type. In
the Biifonidse and BanidsB, which represent the peri-
ehordal type, the greater share of the formation of the
whole vertebra falls to the (paired) dorsal cartilage,
but there is in addition a narrow ventral or hypo-
chordal cartilage which fuses with the dorsal or
becomes connected with it by calcified tissue ; the

DEVELOPMENT AND METAMORPHOSIS.

95

notocbord is thus completely surrounded by a thick
sheath in tadpoles with imperfectly developed limbs.
This mode of formation of both the arch aud the
greater part or whole of the so-called centrum from
the same cartilage explains why there is never a
neuro-central suture in these Batrachians.

Fig. 39.

Side view of cartilaginous skeleton of tadpole of Bombinator pachypus,
with outlines of tbe external parts. (After Goette.)

A.

Anus.

cor.

Coracoid and

ov.

Occipital vertebra

E.

Eye.

prajcoracoid.

p.

Pes.

M.

Mouth.

f-

Femur.

pe.

Pelvis.

N.

Nostril.

fi-

Fibula.

ru.

Radius-ulna.

8.

Spiraculum.

ll.

Humerus.

sc.

Scapula.

as.

Astragalus.

11.

Lower labial.

su.

Suspensorium.

ca.

Calcaneum.

VI.

Manus.

t.

Tibia.

ch.

Chorda.

me.

Meckelian.

ul.

Upper labial.

chy.

Ceratobyal.

V.

Vertebra.

During segmentation of the dorsal cartilages men-
tioned above, which send out the transverse pro-
cesses or diapophyses, there appears between each two
centra an intervertebral cartilage, out of which the
articulating knob of the centrum is formed, and be-
comes attached either to the vertebra anterior (pro-
ccelous type) or posterior (opisthocoelous type) to it,
if not remaining as an independent, intervertebral,
ossified sphere, as we sometimes find in specimens of
Pelobatidse. Ossification appears first in the neural
arch, next descends to the centrum, and sets in last
in the articulating condyle.

96

INTRODUCTION.

Fig. 40.

The annexed figure (40) of the vertebral column
(ventral aspect) of a recently transformed Pelobates
cidtripes shows well the independence of the inter-
vertebral spheres and the three elements which enter
into the formation of the urostyle.

A glance at this figure will also
show to what extenb the importance
of the mode of articulation of the
vertebras has been overrated by those
systematists who have proposed to
divide the tailless Batrachians pri-
marily into procoelian and opistho-
coelian types. As a matter of fact,
both occur in the family Pelobatirfae,
some of the exotic genera of which
are normally opisthoccelian.

During larval life the habits vary
considerably according to the species.
Thus Hyla is always on the move,
swimming about in every direction like a fish ; Bom-
binator, Alytes, Pelobates, and liana keep more to
the bottom, approaching the borders of the ponds
or pools during the hotter part of the day to sun
themselves in shallow water ; Discoglossus, Pelodytes,
and Bufo are in the habit of coming frequently to
take air on the surface, where they swim about with
upturned bellies.

The notion that tadpoles are exclusively herbivorous
has long been exploded ; they feed on both vegetable
and animal substances, and usually show a preference
for the latter, in addition to the microscopical organ-
isms contained in the ooze, which they swallow in large
quantities. Their carnivorous or even cannibal habits
are only too noticeable in confinement, and they have
been utilised for making small and delicate skeletons.
The tadpoles of Bombmator and Alytes are almost
exclusively carnivorous, the real difference between
them and the perfect animal being that the former
feed readily on dead or even decomposing matter,

DEVELOPMENT AND METAMOKPHOSIS. 97

whilst the latter takes only living prey. Nevertheless
the iutestiue in all tadpoles is exceedingly elongate,
being several times longer and of a greater calibre
than after the metamorphosis. Bataillon has ascer-
tained by direct experiment that the abbreviation of
the gut is not localised to any special region. Tad-
poles of Alijtes obstetricans having been anaesthetised
a few days before the egress of the fore limbs, the
abdominal cavity was cut opeu,and the intestine, after
having been slightly uncoiled, marked at regular dis-
tances by silk threads tied round it. The abdomen
was then sewn up, and the tadpoles, having recovered,
continued their evolution. At the close of the meta-
morphosis the gut was again examined, and it was
found that the threads, which were tied at intervals
of twenty millimetres, were then only seven or eight
millimetres apart.

I have measured the uncoiled digestive canal in
fully developed tadpoles and in recently transformed
young of the three following species. It will be seen
from the measurements, in millimetres, how consider-
able the range of variation between different forms
may be, both in the larval and perfect conditions.
Thus in Bana esculenta the length of the intestine of
the tadpole is nearly ten times that of the perfect
form, whilst in Ahjtes obstetricans its length is only
as 4 : 1 ; Pelodytes punctatus, with 6:1, is inter-
mediate. This difference is correlated with the
diet, Bana esculenta being the most herbivorous,
and Ahjtes obstetricans, as noticed above, almost ex-
clusively carnivorous in the tadpole condition.

Length of intestine in tadpole (i) and young (n) of —

i. ir.
Ahjtes obstetricans (25 mm. body-length) 235 60
Pelodi/tes punctatus (20 ,, ., ) 194 32

Bana esculenta (23 ,, ,, ) 472 45

Before closing this chapter on Development, the
highly interesting fact remains to be alluded to that

G

98

INTRODUCTION.

whereas the transformed tailless Batrachians, unlike
their tailed precursors, occupy too high a position in
the zoological scale to be able to regenerate ampu-
tated extremities, the hind limbs reproduce without
difficulty during the larval existence, a fact first
observed by Spallanzani in the last century, and
since verified by Grunther and Barfurth. In order
to be regenerated it is, however, necessary that the
limb be not completely removed. A basal fragment
of the femur must remain to give off the cartilage-
cells which, most curiously, will become differentiated
into the various segments, proceeding from the distal
to the proximal ; and the mutilation must have taken
place some time before the close of the third larval
period or true tadpole stage. It has been stated that
reproduction of the limbs can only be obtained in
young larva?, long before they have reached their full
size. This is, however, contrary to my own experience
on large, full-grown tadpoles of Alytes obstetricans, in
which the well-developed hind limb, after having been
amputated below the knee-joint, was regenerated with
the greatest facility.

Fig. 41.

Series of embryos of Bombinator paclnjpus. (Aiter Goette.)
Showing the formation of the larval month, together with the
atrophy of the adhesive disks; the gradual disappearance of
the external gills, and the formation of the median spiracnlum
by approximation of the two opercular slits ; and the develop-
ment of the gut (the abdominal membranes being removed).

.■SY.

.-- "

." c

7a.

la.

2.

3a.

2a.

-■

* 4. *

4a.

TADPOLES.

99

XIII. Tadpoles.

The term tadpole is restricted to the developmental
stages between the formation of the spiraculum and
the egress of the fore limbs, and corresponds to the
second and third periods of Duges. Descriptions are
preferably drawn up from specimens in the third
period, which begins with the budding of the hind
limbs.

Form. — The head and body are so fused that it is

w.„ j^h^MA

m

Open mouth of — A. Pelothjtes punctatus. B. Alytes obstetricans .
6.0. Buccal orifice, l.p. Labial papillae, m. Mandibles, t.
Series of labial teeth. (From ' P. Z. S.,' 1891.)

100

INTRODUCTION.

extremely difficult to discern the limit between the
two. The term " body " is, therefore, used in the
description as meaning both head and body. Its lon-
gitudinal measurement is taken to the origin of the
hind limbs. The tail consists of a fleshy muscular
portion bordered above and below by membranous
expansions, termed respectively the upper and lower
crest. By depth of the tail is meant its greatest
depth, crests included, and the length is measured
from the posterior extremity of the body.

Mouth (Fig. 42). — This term is used in its wider
sense, i. e. to include the much-developed lip surround-
ing, like a funnel directed downwards, the horny beak,
not unlike that of a cuttle-fish, which forms the
entrance to the mouth proper. The characters offered
by this circular lip are among the most important for
the distinction of species. The lip may be entirely
bordered by fleshy papillae (l.p.), or these may be re-
stricted to the sides, or to
the sides and the lower
border. Its inner surface
is furnished with ridges
armed with series of
minute bristle-like erect
horny teeth (t), each of
which, when strongly
magnified, is seen to be
formed of a column of
superposed cones, hol-
lowed out at the base
, and capping each other

Horny teeth of Bana acjilis (a) (Fig. 43) ; the Summit 01"
and Pelobatesfuscus (b). (After crown 0f each of these
Van Bambeke.) • -, -, ,

cones is expanded, spatu-
late, hooked backwards, and usually multicuspid.
The denticles are absent in Pelobates and Pelodytes,
and present in all the other genera. The number of
these columns is very great. F. E. Schulze has
counted as many as 1100 in the lip of Pelobates fuscus.

Fig. 43.

TADPOLES. 101

By drawing an imaginary line across between the
mandibles the lip may be divided into an upper and a
lower portion, the series of teeth above the upper
mandible being termed upper labial, those below the
lower mandible bein^ lower labial. These are described
as first, second, third, &c, proceeding from the outer
border towards the beak in both the upper and lower
sections of the lip, as shown in the accompanying
figures (p. 99). Each series is reckoned as one,
whether continuous or more or less interrupted in the
middle ; this method being far more simple, and at the
same time more correct, considering the great amount
of individual variation, than those used by some authors
who distinguish between "median" and "lateral"
series, according as to whether or not the series is
broken up in the middle. The first series, either in
the upper or lower division of the lip, may be marginal
(Fig. 42, a, t.l), or it may be within the border,
which is then occupied by fleshy papillae (Fig. 42,
b). The arrangement of the series is expressed
by a formula — § , for instance, indicating the number
in the upper and lower divisions, the figures being
separated by a transverse line corresponding to the
position of the horny beak. The labial teeth are
usually arranged in a single row on each ridge (Fig.
42, a) ; in the Disco glossidas, however, each ridge, or at
any rate the second, bears two or even three rows of
teeth (Fig. 42, b). The beak itself is made up of
horny elements like the labial teeth ; its edge, when
sufficiently magnified, is seen to be denticulate, each
denticle representing the cusp of a single tooth.

The inside of the mouth and the pharynx (Fig. 44)
are beset with long pointed mucous papillae, " taste-
organs " of F. E. Schulze, the longest of which border
the choanae (ch), one in front and one behind, and are
followed by a pair with a denticulate fold between
them ; a cushion-like, hemispherical swelling (t) repre-
sents the tongue, on the anterior border of which a
pair of papillae are inserted. A broad cleft with

102

INTRODUCTION.

denticulate anterior fold leads into the branchial
chambers (br), which contain the four branchial arches.

Fig. 44.

B

Transverse section through the head of larval Pelobates fuscus.
(After Schulze.) A. Upper part. b. Lower part. br. Bran-
chial chambers, ch. Choanse. 1. Lip. m. Mouth. ph. Pharynx.
t. Tongue.

These arches bear on the convex outer side the delicate
arborescent gills, and on the concave inner side de-
velop a membranous septum with vermicular perfora-
tions, a special sifting or filtering contrivance through
which the water absorbed by the mouth has to pass
before reaching the respiratory organs of the branchial
apparatus.

Spiraculum. — The water is expelled from the bran-
chial chambers by one or two tubes opening by one
orifice in all European Batrachians. This orifice is
the spiraculum. In the Discoglossidae two tubes are
present, which gradually converge towards each other
at the end of the first period of development, as si i own
on Fig. 41, p. 98, and in the tadpole proper discharge
through one transverse, slit-like or crescentic opening,
situated in the middle of the breast (Fig. 45, a, sp.).
In all other tailless Batrachians the tube is single, and

TADPOLES.

103

opens on the left side, straight backwards in Bufo,
backwards and upwards in the other genera (Fig. 45,
b, sp. ; Fig. 46, a, b).

Fig. 45.

sp-

an:

Lower view of tadpoles of — a. Alytes obstetricans.

(From 'P. Z. S.,* 1891.)

B. Hyla arborea.

The Lines of Muciferous Crypts or Sensory Canals. —
All tadpoles are provided with these organs, the homo-
logues of the lateral line in fishes ; they stand in imme-
diate relation to the nerve branches, and are regarded
as organs of a special sense possessed by aquatic
animals, feeling, in its broadest sense, having been
admitted as their possible use, and the function of
determining waves of vibration in the aqueous medium
having been suggested. The surface openings of these
tubules or canals form lines which may differ greatly

104

INTRODUCTION

as regards their degree of development in individuals
of the same species, and their arrangement also varies,
within certain limits, irrespective of the species. I

Fig. 46.

Tadpoles showing sensory canals. A. Rana agilis. b. Pelodytes
punctatus. c. Alytes obstetricans. (From ' P. Z. S.,' 1891.)

have found them usually most distinct in Pelodi/tes,
Pelobates caltripes, Rana agilis, R. latastii, and Bom-
binator igneus ; least in Bufo. I append figures (Fig.
46) of tadpoles of Rana agilis, Pelodytes punctatus,

m.n

■ .

..'■""■ "■■■■■ '"'•■'■...

*■

. . ... - .. " ■■

'■ M M—""

4.

': -■ -'-- •■

4<a .

j~^ ...

-

?*; MUUHHi -...""J"

»--' . ..'■sJ™'-'

H£>

( >..

...
- . .■•■■ ■*■<•> ■

■ ^ / /^r /■('// .,_ ■/'■f/,r, /( ussy .

TADPOLES.

105

and Alytes obstetricans, in which these organs are
represented by dotted lines. In addition to these
lines, all tadpoles show more or less distinctly a small
whitish gland in the middle of the head between the
eyes, the so-called frontal gland or pineal gland, which
in earty stages is connected with the brain. A glan-
dular streak extending from the nostril towards the
eye is the lachrymal canal.

The presence of a number of pulsating lymph-sacs
has lately been discovered by Weliky on each side of
the dorsal caudal crest.

Pigmentary Network. — In addition to the ordinary
pigment-cells, the tadpoles of afew genera (Discoglossus,
Bombinator, Pelodytes) present a system of fine black
lines, most apparent on the caudal crests and the more
feebly pigmented parts of the body, which afford
excellent characters for their identification. It is a
fact worthy of notice that when the end of the tail has
been injured these pigmentary lines are not reproduced
on the regenerated portion. Their aspect is shown in
the following figures (from 'P. Z. S.,' 1891), repre-
senting much-enlarged pieces of the upper caudal
crest of Pelodytes (a) and Discnglossus (b).

Fig. 47.

The following key has been drawn up to facilitate
the identification of the genera and species in the tad-
pole condition.

I. Spiraculum median ; anus median ; tail rounded
or obtuse at the end ; a papillose edge all round

106 INTRODUCTION.

the lip, sometimes narrowly interrupted above ;
labial teeth in § series, disposed in two or
three rows, at least in the second series.

a. Spiraculum equally distant from the anterior

and the posterior extremity of the body ; tail
at least once and a half the length of the body,
and three and a half or four times as long- as
deep ; caudal crests with a polygonal network
of fine black lines . . . Discoglossus.

b. Spiraculum nearer the posterior than the ante-

rior extremity of the body ; tail not more
than once and a half the length of the body,
twice to twice and a half as long as deep ;
caudal crests with fine black decussating lines .

BoMBINATOR.

Mouth trigonal . . . B. igneas.

Mouth elliptical . . . B. pachypus.

o. Spiraculum nearer the anterior than the poste-
rior extremity of the body ; tail at least once
and a half the length of the body, twice and
two-thirds to thrice and a half as long as
deep ; no black network . . Alytes.

II. Spiraculum sinistral ; labial teeth disposed in
a single row in each series or ridge.
a. Anus median.

1. Spiraculum directed upwards and back-
wards ; lower lip bordered with papillae ;
series of labial teeth f or f.

a. Tail obtusely pointed, with fine black de-

cussating lines ; an inverted fold on each
side of the lip, the upper edge of which
bears a long series of teeth ; beak white
with black ed^e . . Pelodytes.

O

b. Tail acutely pointed, without black lines ;

first series of upper labial teeth short ;

beak entirely black . . Pelobates.

Tail once and a half to twice as long as the

body .... P.fuscus.

TADPOLES. 107

Tail not more than once and a half as lono-
as the body . . .P. cultripes.

2. Spiraculum directed straight backwards,
tail rounded at the end ; both upper and
lower lip with toothed edge ; series of
labial teeth § . . . . Bdto.

Mouth at least as wide as the interocular
space, which is twice as great as the
distance between the nostrils ; second
upper series of labial teeth very narrowly
interrupted in the middle . B. vulgaris.

Mouth nearly as wide as the interocular space,
which measures once and a half the dis-
tance between the nostrils ; second upper
series of labial teeth more or less inter-
rupted in the middle . . B. viridis.

Mouth considerably narrower than the inter-
ocular space, which is nearly twice as great
as the distance between the nostrils ;
second upper series of labial teeth very
widely interrupted in the middle.

B. calamita.
b. Anus dextral ; spiraculum directed backwards
and upwards ; lower lip bordered with
papilla?.

1. Anus opening above the lower edge of the

tail ; upper caudal crest extending far
forwards on the back, almost to between
the eyes, which are lateral and visible
from the ventral as well as from the
dorsal aspect of the body ; series of labial
teeth § Hyla.

2. Anus opening close to the lower edge of the

tail ; upper caudal crest not extending
forwards beyond the vertical of the spira-
culum ; eyes on the upper surface. Rana.
a. Series of labial teeth —^-^-.
Interocular space at least twice the distance
between the nostrils, and much greater

108 INTRODUCTION.

than the width of the mouth ; tail acutely
pointed, at least nearly twice as long as
the body . . . . B. esculenta.

Interocular space but little wider than the

distance between the nostrils or the width

of the mouth ; tail once and two-thirds to

twice as long as the body . B. arvalis.

b. Series of labial teeth 3-^-A.

a. Tail obtusely pointed, once and a half to
twice as long as the body.

Second series of upper labial teeth widely in-
terrupted in the middle; first series of
lower labial teeth at least two-thirds the
length of the second ; width of mouth a
little less than the interocular width,
which equals about once and a half the dis-
tance between the nostrils . B. temporaria.

Second series of upper labial teeth continuous
or narrowly interrupted ; first series of
lower labial teeth at least two-thirds the
length of the second ; width of mouth
quite as great as the iuterocular space,
which equals about once and a half the
distance between the nostrils . B. grseca.

Second series of upper labial teeth widely in-
terrupted in the middle; first series of
lower labial teeth hardly half as long as
the second; width of mouth much less
than the interocular space, which equals
nearly twice the distance between the
nostrils . . . . . B. iberica.

(3. Tail acutely pointed or submucronate,
at least nearly twice as long as the
body.

Interocular space once and a half the width of
the mouth or the distance between the
nostrils ; no tubercle on the upper man-
dible . . . . . B. latastii.

Interocular space at least twice the width of

TADPOLES.

109

the mouth or the distance between the
nostrils ; usually a black tubercle on the
upper mandible . . . B. agilis.

With regard to size, the tadpoles rank as follows :

1.

o

3.

4.

5.

6.

7.

8.

9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.

Pel abates fuseus

,, cult rip es
Pana esculent a .
Alytes obstetricans

,, cisternasii
Pelodytes punctatus,
Bana agilis .
Bombinator igneus
Hi/la arborea
Bana ib erica .

,, grseca .

,, temporar i a
Bombinator pacing pus
Biifo viridis .
Bana latastii
,, arcalis
Discoglossus pictus
Bufo vulgaris
calamita,

5J

Maximum length
on record.
Millimetres.

175
120
111

90

69

65

59

50

49

49

48

46

45

44

44

43

33

32

30

But if we compare the maximum length of the tad-
pole with the maximum length of the adult, we

obtain the following order

1.

2.
3.
4.

5.
6.

Pelobates fuseus .
Alytes obstetricans
,, cisternasii .
Pelobates cultripes
Pelodytes punctatus
Bombinator iqneus

Maximum
length of adult from
snout to vent.
Millimetres.

80
54
40
88
45
50

110 INTRODUCTION.

Millimetres.

7. Ilijla arbor ea 50

8. liana iberlca 50

9. Bombinator pachypus 50

10. liana csculenta 125

11. ,, grseca 66

12. ,, agilis 90

13. ,, latastii 65

14. ,, arvalis 75

15. ,, temporaria 95

16. Bufo viridis 90

17. Dlscoglossus pictus 75

18. Bufo calamita 80

19. ,, vulgaris 180

It may be noticed that the five species which head
the list as having proportionally the largest tadpoles
are the only ones in which the pupil is vertical.

The structural differences which separate the genera
and species in their tadpole condition reflect, on the
whole, pretty accurately the system based upon the
perfect animals, although here and there the modifi-
cations are of unequal importance. We must bear in
mind, however, that such a correspondence, if existing
in the European Batrachians, is not universal. Larval
forms such as the tadpoles are outside the cycle of
recapitulation, the ontogeny being broken by the
intercalation of the larval phasis. The horny beak,
the circular lip with its horny armature, the spi-
raculum, the enclosure of the fore limbs in diverticula
of the branchial chambers, and such special adapta-
tions as the ventral disk or sucker of certain exotic
mountain forms, clearly point to tadpoles having
had a developmental history of their own. We need,
therefore, not be surprised at occasionally finding,
within the same genus, very different types of tad-
poles, or even a total suppression of the free larval
stages, as is actually the case in the large and widely
distributed genus Bana.

. '/>/

. in.

-

2 a .

iWiimfiS

■ >■ "

''■"

4a.

■•■■"■■ -•'-"-■•■.,»,

I '' n *.

..jit" jV .

"'■'•»* .

..r ■

7a

^'..'<r

7.

:

i

(wi******""'" »*, ""

■

L?S»

Ss "•:"'"" r

.

, *>> -

■" ■

' •, ■• jgSa^f^sss^^

— ^W3
5a.

•*^.5

1.

■ <:,'' ":■

«*?*?»■ jl»»"w. ■

TADPOLES. Ill

The tadpoles are represented, of the natural size,
on Pis. I, II, and III. The mouth (a) is enlarged
five diameters in figs. 1 and 2, PI. II ; seven diameters
in fig. 4, PL I ; fifteen diameters in figs. 3 and 5,
PI. II, and fig. 1, PI. I; ten diameters in the rest.

PLATE I.

Fig. 1. Discoglossus pictas. Montecristo.

2. Bomblnator igneus. Seeland, Deumark.

3. — pachypus. Mondorf, Luxem-
burg.

4. Alytes obstetrlcans. St. Germain, near Paris.

5. — — var. boscse. Sierra Es-

trella.

6. — cisternasii. Sierra Morena.

7. 8. Pelodijles punctatiis. Near St. Malo.

PLATE II.

Fig. 1. Pelobates fuscus. Prague.
,, 2. — cultripes. Near Bordeaux.
,, 3. Bufo vulgaris. Near London.
,, 4. — viridis. Breslau.
,, 5. — calamita. Near St. Malo.
„ 6, 7. Hyla arborea. Near St. Malo.
,,8. — — var. meridio nails. Near

Nice.

PLATE III.

Fig. 1 . Bana esculenta. Near St. Malo.

,, 2. — arvalis. Breslau.

,, 3. — temporaria. Near London.

,, 4. — grseca. Parnassos.

,, 5. — iberica. Coimbra.

,, 6. — latastli. Italy.

,, 7. — agilis. Near St. Malo.

112 INTRODUCTION,

XIV. Hybrids.

The fact, so often observed., tbat pairing takes place
between different species — as, for instance, between
Bana temporaria and Bufo vulgaris, Bana temporaria
and Pelobates fuscus, Pelodytes punctatus and Hyla,
arborea — has long ago given rise to the supposition
that hybrids are occasionally produced. At the end
of the last century, Spallanzani, after his successful
experiments on artificial fecundation, attempted to
raise hybrids between species of Bana, Bufo, and
Hyla, but without results.

Experiments were again instituted, from 1867 to
1872, by A. de l'lsle, who tried to hybridise the
French species of Bana and Bufo. The difficulty of
such experiments is very great, owing to the diversity
in the breeding season of the different species. Never-
theless de l'lsle succeeded in procuring suitable ex-
amples of three species of Bana, and artificially
impregnated ova of Bana esculenta and B. agilis with
the semen of B. temporaria, and ova of Bana tem-
poraria. with the semen of the two others. The ex-
periments, several times repeated, gave a negative
result, which the author attributes to the very con-
siderable difference between the genital organs of even
so closely allied species as B. temporaria and B. agilis.
But with Bufo vulgaris and B. calamita, which,
although morphologically more widely separated,
have almost identical genitalia, positive results were
obtained, although the development did not proceed
beyond the tadpole condition.

A few years later, in 1877 and 1878, the subject
was again taken up by Lataste, who succeeded in
fertilising eggs of Pelobates fuscus with the semen of
P. cultripes. But the few tadpoles obtained were
monstrous or dropsical, and not one survived to
develop the limbs.

HYBRIDS. 113

Experiments j conducted with the greatest care, were
instituted by Pfliiger in 1882. He was able to show
that different degrees exist in the action of the sperma-
tozoa upon the ova of different species. Thus the
spermatozoon of a newt (Molge alpestris, M. vulgaris)
acted upon the ovum of Bana temporaria to the extent
of producing a very irregular segmentation ; other
cross-fertilisations between various fiws and toads
extended to the formation of an embryo, of a tadpole,
or finally of a perfect Batrachian. These interesting
experiments were continued by Pfliiger and W. J.
Smith in 1883, and by Born in 1883 and 1884.

At first Pfliiger absolutely failed in crossing Bana
temporaria and It. ar calls, but ultimately both he and
Born succeeded in obtaining a few perfect hybrids (B.
I '<■ in pora via cT X B. arvalis ? ). Born also obtained
perfect hybrids of Bufo vulgaris and B. viridis crossed
in both directions; and, what is more extraordinary,
a few embryos, which did not develop beyond the
hatching period, from eggs of Bana esculenta fecun-
dated by Bufo viridis.

More extraordinary still is a case mentioned by
Heron-Royer. In March, 1881, he found a male Bana
temporaria pairing with a Belobates fuscus which had
begun to spawn. Having brought home the short
band of eggs, he was much surprised to see them
develop quite normally, and ultimately produce larva3
which were indistinguishable from those of Bana tern,-
poraria. ; and two that survived transformed into
normal specimens of B. temporaria. I cannot help
thinking that some confusion must have taken place :
may not, somehow or other, embryos of B. temporaria
have been substituted whilst Heron-Royer was observ-
ing their development ?

Hybrids between two closely allied species, Bom-
binator igneus and B. pachypus, were obtained by
Heron-Royer, and a second generation was produced
by crossing these hybrids with individuals of the
latter species. It also appears from remarks made

H

114 INTRODUCTION.

by v. Mehely that, in the few localities where the two
species co-exist, intermediate specimens occur which
are no doubt to be regarded as hybrids.

As to the supposed hybrids between Rana esculenta
and temporaria, noticed by Schlotthauber and others,
they are nothing but R. arvalis, a species which is in
many respects intermediate between the two supposed
parents, and with which the authors in question do
not seem to have been well acquainted.

Hybrid embryos between Rana esculenta and R.
arvalis appear to have been recently obtained by W.
Gebhardt.

So-called hybrids between Hyla arborea and II.
meridional is (bary tonus) , and Bufo viridis and B.
arabicus, obtained by Heron-Royer, are only mongrels,
as I do not consider the specific distinctions justified.

GEOGRAPHICAL DISTRIBUTION. 115

XV. Geographical Distribution.

The Western Palrearctic sub-region, to which Europe,
together with South-western Asia north of the 30th
parallel, and Africa north of the Sahara, belongs, is
characterised by the development of the Discor/lossidm
and Pelobatidae, the former family being represented
outside its limits by a single species in Manchuria,
Corea, and Northern China (Bombinator orientalis),
and another (Liopelma hochstetteri) in New Zealand,
of which it is the only Batrachian representative ;
whilst the latter, absent from the Eastern Palrearctic
sub-region, is again abundant in Southern China, the
Eastern Himalayas, Further India, the Malay Archi-
pelago and Papuasia, North America, and Mexico.

The Bufonidse, Hylidae, and Ranidae, of each of
which the type genus is represented in Europe, are
families of very extensive distribution. The first is
found all over the world except in Madagascar and
Australia; the second is absent from Africa south of
the Sahara, Madagascar, India, and the Malay Penin-
sula and Archipelago west of the Moluccas and
Sumba; and the third is absent from Australia (with
the exception of a single Papuan species found at
Cape York) and the greater part of South America.

Five species have a very wide range in the Palae-
arctic region. Three extend from Western Europe
and North Africa to China and Japan, viz. Bufo vul-
garis, Hi/la arborea, liana esculenta; Bana temporaria
ranges from Western Europe to the Pacific coast of
Northern Asia and Yesso, and Bufo viridis from the
Rhine and Alps, and North-west Africa to Mongolia,
Western China, and the Himalayas.

Of the species which are known to inhabit more
than one country, three may be regarded as strictly
Western, viz. Discoglossus pictus, Pelodytes punctatus,
and Pelobates cultripes. Bufo calamita and Alytes
obstetricans may also be described as Western forms,

116 INTRODUCTION.

generally distributed over France and the Spanish
Peninsula, becoming gradually more local and scarcer
east of the Rhine, and not extending into Russia nor
south of the Alps.

As Eastern forms we may regard Bombinator igneus,
Bufo viridis, and liana arvalis, which do not extend
to Belgium and France, although in North Africa
Bufo viridis reaches the Atlantic coast of Morocco,
Bana camerani (Caucasus, Armenia, North-west
Persia), and Bana grseca (Greece, Bosnia, Italy).
Pel abates fuscus is characteristic of Central Europe.

Bombinator pachypus and liana agilis have an exten-
sive but somewhat irregular distribution, being absent
from the north and north-east and the Pyrenean
Peninsula.

Three species have a very restricted range, viz.
Alytes cistemasii (Pyrenean Peninsula), Bana iberica
(Pyrenean Peninsula), and Bana. latastii (Italy).

Bana teniporaria may be regarded as a northern
species, reaching the extreme north and disappearing,
or confined to the mountains, in the south ; whilst
Bana esculenta and Hyla arborea extend very far
south, and are absent from the extreme north.

With regard to the range from north to south, the
twenty species may be classed in the following order :

Northern limit.

1. Bana temporaria 70°

2. Bufo vulgaris 65

3. Bana arvalis 60

* k [Bana esculenta 59

[Pel abates fuscus 59

r _ (Bufo viridis 58

' '[Bufo calamita 58

8. Hyla arborea, 57

9. Bombinator igneus 56

10. Bombinator pachypus 53

-. -. , 0 (Alytes obstetricans 52

[Bana agilis 52

13. Pelodytes punctatus 51

GEOGRAPHIC A I, DISTRIBUTION. 117

14. Pelobates cultripes 48°

15. liana latastii 46

16. liana, camerani 45

17. Rana grseca 44

,o ^q (Discoglossus pictus 43

e' {Rana iberica 43

20. Alijtes cistemasii 41

From south to north the following order obtains :

Southern limit.

(Bufo viridis 28°

1 — 3. -J Hyla arborea 28

[Rana esculenta 28

4. Discoglossus pictus 31

5. Bufo vulgaris 34

(Alytes obstetricans 36

Pelodytes punctaius 36

Pelobates cat tripes 36

Bufo calamita 36

1 ., \ Rana camerani 37

' ' \Rana agilis 37

12. Pelobates fuscus 38

(Bombinator pachypus ... 39

13 — 15.] Alytes cistemasii 39

[Rana gra&ca 39

16. Rana Iberica 40

-. „ ,o \ Rana temporaria 43

' ' [Rana latastii 43

iq 90 f Bombinator igneus 45

' [Rana arvalis 45

With regard to the vertical range, the species may
be arranged in the following order :

Maximum elevation

reached.

Feet.

1. Rana temporaria 10000

2. Rana camerani 8000

3. Bufo vulgaris 7000

118 INTRODUCTION.

Feet.

. f. fAlytes obstetricans .... 6500

4' °- \Bufo viridis 6500*

6. liana ijrseca 5800

7. Bombinator pachypus .... 5500

8. Junta iberica 4500

9. liana agilis 4200

10. Bufo cdlamita 4000

11. liana esculenta 3500

12. Hi/la arborea 3300

The other species do not ascend to 3000 feet.

The

following table

sh

OWS

; tl:

ie

distributio

n in t

-lie

different countries of Europe

I •

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

1.

Discc

ujlossus i^ictas

+

+

2

Bombinator igneus

+

+

+

+

o.

,, pachypus .

+

+

+

+

+

+

4.

Ahjtes obstetricans

+

+

+

+

+

5.

»j

cisternasii .

+

6.

Pelodytes punctatus

+

+

+

7.

Pdobates fuscus .

+

+

+

+

+

+

+

8.

*j

cultripes .

+

+

9.

Bufo

vulgaris

+

+

+

+

+

+

+

+

+

+

10.

))

viridis .

+

+

+

+

+

+

+

11.

calamita

+

+

+

+

+

+

+

12.

Hyla

arborea

+

+

4-

+

+

+

+

+

+

13.

Rana esculenta

+

+

+

+

+

+

+

+

+

+

14.

??

arvalis .

+

+

+

+

4-

15.

»»

camera hi

. . .

+

16.

)»

trmporaria .

+

+

+

+

+

+

+

-r-

+

+

17.

j »

grseca .

+

+

18.

5>

iberica .

• . .

+

19.

J)

latastii.

. • •

. . .

+

20.

Jl

agilis .

...

+'

+

+

+

+

1.

Great Britain ami Ireland.

2.

Scandinavia and D

en mark.

3.

Fiance.

4.

Belgium and Holh

md.

5

Germany and Swit

zerh

md.

6.

Spanish Peninsula

7.

Italy.

8.

Austria-Hungary <

md Roumania.

9.

Dalmatia, Bosnia,

and

countries

i south o

if the

Danube

10.

Russia.

* 15,000 feet in the Himalayas.

GEOGRAPHICAL DISTRIBUTION. 119

It may also not be without interest, in order to
bring out their associations, to enumerate the species
occurring in the environs (i. e. a radius of about fifteen
miles) of some of the principal towns the Batrachian
fauna of which lias been thoroughly searched by com-
petent workers. It will be seen, then, that no district
can boast of possessing more than ten out of the
twenty recognised species, and that, in spite of their
northern position, Paris and Bonn yield to no other
and are richer than many more southern localities.

1. London (three species).

Bufo vulgaris, B. calamita, Rana temporaria.

2. Copenhagen (nine species).

Bombinator igneus, Belobates fuscus, Bufo vul-
garis, B. viridis, B. calamita, Hi/la arborea,
Rana esculenta, B. arvalis, B. temporaria.

3. Paris (ten species).

Bombinator pachypus, Alytes obstetr leans, Belo-
dytes punctata*, Belobates fuscus, Bufo
vulgaris, B. calamita, Hula arborea, Rana
esculenta, R. temporaria, R. agilis.

4. Bordeaux (nine species).

Bombinator pachypus, Alytes obstetricans, Belo-
dytes punctatus, Belobates cultripes, Bufo
vulgaris, B. calamita, Hyla arborea, Rana
esculenta, R. agilis.

5. Brussels (five species).

Bufo vulgaris, B. calamita, Hyla arborea, Rana
esculenta, R. temporaria.

6. Berlin (nine species).

Bombinator igneus, Belobates fuscus, Bufo vul-
garis, l>. viridis, B. calamita, Hyla arborea,
Rana esculenta, R. arvalis, R. temporaria.

7. Bonn (ten species).

Bombinator pachypus, Alytes obstetricans, Belo-
bates fuscus, Bufo vulgaris, B. viridis, B.
calamita, Hyla arborea, Rana esculenta, R.
arvalis, R. temporaria.

120 INTRODUCTION.

8. Geneva (eight species).

Bombinator pachypus, Alytes obstetricans, Bufo
vulgaris, B. calamita, Hyla arborea, Rana
esculenta, B. temporaria-, B. agilis.

9. Coimbra (ten species).

Discoglossus pictns, Alytes obstetricans, A. cister-
nasii, Belodytes punctatus, Pelobates cut-
tripes, Bufo vulgaris, B. calamita, Hyla
arborea, Rana esculenta, R. iberica.

10. Turin (seven species).

Pel abates fuseus, Bufo vulgaris, B. vivid is, Hyla
arborea, Rana esculenta, R. latastii, R.
agilis.

11. Palermo (five species).

Discoglossus pictus, Bufo vulgaris, B. riridis,
Hyla arborea, Rana esculenta.

12. Vienna (ten species).

Bombinator igneus, B. pachypus, Pelobates
fuseus, Bufo nil gar is, B. viridis, Hyla
arborea, Rana esculenta, R. arvalis, R.
temporaria, R. agilis.

13. St. Petersburg (three species).

Bufo vulgaris, Rana arvalis, R. temporaria.

14. Moscow (eight species).

Bombinator igneus, Pelobates fuseus, Bufo vul-
garis, B. viridis, ILyla, arborea, Rana
esculenta, R. arvalis, R. temporaria.

And finally, in order to complete the sketch of the
Geographical Distribution, a list is appended of all
the tailless Batrachians known from other parts of
the Palasarctic region in its widest sense, viz. North
Africa and Asia north of the 30th parallel. Column A
stands for North-western Africa, B for North-eastern
Africa, C for South-western Asia, D for North-
western and Central Asia, E for Eastern Asia with
Tibet, and F for Japan. The names of species occur-
ring also in Europe are prefixed with an asterisk.

( i El (GRAPHICAL DISTRIBUTION.

121

Discoglossid.e.

*Discoylossus pictus, Otth .
Bombinator orlentalis, Blgr.

Pelobatid.e.

Pclodytes caucasicus, Blgr.
*Pelobates fuscus, Laur.

,, syriacus, Bttgr. .

BUFONID.E.

*Bufo vulgaris, Laur.
formosus, Blgr.
viridis, Laur.
raddii, Strauch
mauritanicus, Schleg
regularis, Reuss

Hylid.e.

*Hyla arborca, L.
,, stepheni, Blgr. .

RANID.E.

liana boulengeri, Gthr.
,, limnocharis, Boie
,, rugosa, Schleg.
* ,, esculenta, L.
,, piancyi, Lat.
arvalis, Nilss.

#

*

camcrani, Blgr.
,, macrocnernis, Blgr. .
* „ temporaria, L.
,, amurensis, Blgr.
,, martensii, Blgr.
,, japonica, Blgr.
¥ „ agilis, Tlioni. .

,, mascareniensis, D. & B.
., schmacheri, Bttgr. .
Rhacophorus buergeri, Schleg.
,, davidi, Sauv.

„ schlcgelii, Gthr.

A.

B.

C.

D

E.

F.

+

+
+
+

+

+

+

+

+

+
+

+

+

+

+

+
+

+

+

+

+

+

+

+
+

+
+

+

+
+

+

+

+

+

+

+

+

+
+

+

...

+

+
+
+
+

+

+
+

+

+

+
+

+
+

[At page 123.

DISCOGLOSSUS PICTUS.
BOMBINATOR IGNEUS.
rfl - PACHYPUS.

ALYTES OBSTETRICANS.

:P

CISTERNASII.

DISTRIBUTION OF EUROPEAN DISCOGLOSSID^E.

DISCOGLOSSIMl. 123

Order ECAUDATA.

Four limbs and no tail. Radius and ulna, and tibia
and fibula continent; tarsus (astragalus and calcaneum)
elongate, forming an additional segment in the biud
limb. Frontal bones confluent with parietals.

Sub-order PHANEROGLOSSA.
Eustachian tubes separated ; tongue present.

Series A.— ARCIFERA.

Pectoral arch vvith the opposite halves moveable, the
coracoids and praecoracoids connected by an arched
cartilage (the epicoracoid), that of the one side over-
lapping that of the other.

Family 1.— DISCOGLOSSULE.

Vertebrae opisthoccelous ; short ribs articulated to
the anterior diapophyses ; diapophyses of sacral
vertebra dilated. Upper jaw toothed.

The genera combined under this family constitute
a most interesting and perfectly natural group, as is
abundantly evidenced by the bony structure, the larval
characters, &c.

That they occupy the most lowly position among the
Ecaudata, and show the nearest approximation to the
Catidata, is another point on which there can be no
question. The opisthoccelous vertebrae with distinct
ribs, the increased number of carpal and tarsal ele-
ments, the non-extrusible tongue, the presence of the
azygos (posterior cardiual) vein discovered by Hoch-
stetter in Bombiuator, and since shown by Howes to be

124 DISOOGLOSSID.E.

fairly distinctive of the whole family, tlie structure of
the urogenital apparatus, together with other cha-
racters, give ample foundation to this proposition.

This family comprises only four genera. Three are

confined to the Palajarctic region and represented in

Europe; the fourth, Liopelma, Fitzinger, closely allied

to Ahjtes, is the only representative of the Batrachians

in New Zealand. The range of the species dealt with

in this work is shown on the accompanying map.

The European genera are distinguished as follows :

Tympanum present, distinct or hidden ; pupil

roundish or triangular; diapophyses of sacral

vertebra moderately dilated . 1. Discoglossus.

Tympanum absent; pupil roundish or triangular ;

diapophyses of sacral vertebra very strongly

dilated 2. Bombinator.

Tympanum distinct; pupil vertical; diapophyses
of sacral vertebra strongly dilated.

3. Ahjtes.

The relationships of these three genera cannot be
well expressed in a linear arrangement Discoglossus
is unquestionably the most generalised, and Bombinator
and Alytes are almost equally related to it, the latter
being, on the whole, more affine to Discoglosms than
to Bombinator, especially with regard to the osteological
characters.

1. Discoglossus.

Ottb, Neue Denksckr. allgeni. Sckweiz. Naturf. Ges., i, 1837, p. 6.

Pupil roundish or triangular. Vomerine teeth in
long transverse series behind the choana. Tongue
circular, entire, scarcely free behind. Tympanum
more or less distinct or concealed under the skin.
Fingers free, toes webbed ; outer metatarsals separated
by web. Diapophyses of sacral vertebra moderately
dilated. Urostyle articulated to two condyles.

A single species, confined to the western parts of
the Palasarctic region.

^

^

J

O1

DISCOGLOSSUS. 125

The pupil is described as roundish or triangular ;
in its contracted condition it varies considerably
according to individuals. It is never absolutely
round, for it forms an angle below, whence a subtri-
angular form results. I have seen specimens in which
the vertical diameter somewhat exceeds the horizontal,
so that the fully contracted pupil might be termed
vertically pear-shaped, or, the upper border forming
an open angle, kite-shaped.

1. DiSCOGLOSSUS PICTUS.

(Plate IV.)

Cetti, Anf. e Pose. Sard., iii, p. 38 (1777).

Rana tcnqioraria (non L.), Rozet, Voy. Alg., i, p. 230 (1833).

Discocflossus pictus, Ottb, N. Denbsebr. allgem. Schweiz. Nat.
Ges., i, 1837, p. C, figs. ; Tschndi, Class. Batr., p. SO (1838);
Bonaparte, Icon. Faun. Ital., Rett. Anf. (1838) ; Dumeiil &
Bibron, Erp. Gen., viii, p. 425 (1841); Giintber, Cat. Batr.
Sal., p. 35 (1858) ; Strancli, Erp. Alg., p. 77 (18G2) ; De Betta,
Atti 1st. Venet. (3), xiii, 1868, p. 77, and Faun. Ital., Rett.
Anf., p. G7 (1874); Scbreiber, Herp. Eur., p. 112 (1875);
Camerano, Atti Ace. Torin., xiv, 1879, p. 443, pi. — , figs.
6 — 8; Lataste, Act. Soc. Linn. Bord., xxxiii, 1879, p. 275,
pis. iii— v; Bedriaga, Bull. Soc. Nat. Mosc, 1881, p. 292;
Boulenger, Cat. Batr. Ecaud., p. 445 (1882); Camerano, Mem.
Ace. Torin. (2), xxxv, 1883, p. 204; Bedriaga, Arcb. f. Nat.,
1883, p. 254; Heron-Royer, Bull. Soc. Zool. France, 1885,
p. 565, pi. xiv ; F. E. Scbulze, Sitzb. Ges. Naturf. Fr. Berlin,
1886, pp. 5 and 31 ; Heron-Royer & Van Bambeke, Arcb.
Biol., ix, 1889, p. 280; Bedriaga, Bull. Soc. Nat. Mosc,
1889, p. 545, and Ampb. Rept, Portng., p. 22 (1889) ; Heron-
Royer, Bull. Soc. Et. Sc. Angers (2), xix, 1889, p. 160, pis. i
and ii; Boulenger, Trans. Zool. Soc, xiii, 1891, p. 160, and
Proc. Zool. Soc, 1891, p. 620, pi. xlvii, fig. 3; Anderson,
Proc. Zool. Soc, 1892, p. 24; Bedriaga, Suppl. Ampb. Port.,
p. 13 (1893) ; Mina-Palumbo, Nat. Sicib, xii, 1893, p. 262.

Discoglossus sarcitis, Tscbudi, in Ottb, 1. c, and Class. Batr.,
p. 80; Bonaparte, 1. c. ; Camerano, Atti Ace Torin., xiv,
1879, p. 437, pi. — , figs. 4 and 5.

Pseudis sardoa, Gene, Syn. Rept. Sard., p. 24, pi. v, figs. 1 — 3
& 6 (1839).

Rana picta, Sclilegel, in Wagner, Reisen Alg., iii, p. 131 (1841).

Colodactylus cxrulescens, Tscbudi, Faun. Per., Herp., p. 68, pi. xi,
fig. 2 (1845); Peters, Mon. Bed. Ac, 1873, p. 414, pi. iii,
fig. 1.

Discoglossus scovazzii, Camerano, Atti Ace. Torin., xiii, 1878,
p. 548, and xiv, 1879, p. 447, pi. — , figs. 1—3.

Discoglossus auritus, Heron-Royer, Bull. Soc Zool. France, 1888,
pp. 205 and 220, and Bull. Soc. Et. Sc. Angers (2), xix, 1889,
p. 177, pis. i and ii.

126

DISCOGLOSSID.E.

Vomerine teeth in a long, straight or slightly

curved series, narrowly interrupted in the middle,

behind the choanre, and extending outwards to the

Fig. 48. vertical of their outer borders or even

a little beyond. Tongue moderately

large, circular, thick, adherent, scarcely

free behind.

Head much depressed, a little broader
than long; snout rounded or obtusely
pointed, projecting considerably be-
yond the mouth, as long as or a little
longer than the diameter of the orbit ;
no canthus rostralis ; dorsal region slightly grooved ;
nostril a little nearer the tip of the snout than the
eye ; eye moderate ; interorbital space as broad as or
a little narrower than the upper eyelid, and equal to
the distance between the nostrils ; tympanum measur-
ing three-fifths to two-thirds the diameter of the eye,
sometimes feebly distinct, often completely hidden.

Open mouth.

Fig. 49.

A

Heads of males, upper views, a. Corunna.

r>. Sardinia.

Tlemsen.

DISCOGLOSSUS. 127

Fingers rather short, obtusely pointed, first shortest,
third longest, second and fourth equal ; no sub-
articular tubercles ; three palmar tubercles, the inner
(rudiment of pollex) largest and very prominent, the
two others flat and close together, at the base of the
third and fourth fingers.

Hind limb rather long; the tibio-tarsal articulation
reaches the tympanum, the eye, or the nostril ; tibia
longer than the femur, the heels overlapping when
the le^s are folded at rigdit angles to the rhachis.
Foot a little shorter than the tibia ; toes rather slender,
one-fourth or one-third webbed in females and young,
three-fourths or nearly entirely in adult males ; no
subarticular tubercles ; no tarsal fold ; a small rounded
inner metatarsal tubercle, the length of which equals
one-third to one-half the length of the inner toe.

Skin remarkably shiny in life, smooth or with small
warts or short glandular folds on the back ; a more or
less developed glandular lateral fold from the eye to
above the shoulder, often prolonged on the side of the
body to the inguinal region ; a fold of the skin may
be present across the occiput behind the eyes. Lower
parts smooth except the thighs, which are granulate
near the vent; throat and belly sometimes with
isolated small granules.

Coloration very variable. Pale brown, grey, greyish
olive, yellowish or red above, uniform or with dark,
often light-edged spots ; some specimens with a broad
yellow vertebral stripe, which may be bordered on
each side by one or two dark stripes ; the lateral
glandular fold light, often reddish or golden ; a dark
streak on each side from the tip of the snout to the
eye, and a dark temporal blotch or streak ; limbs with
transverse dark spots, sometimes forming regular
cross-bands ; a more or less distinct light triangular
or heart-shaped spot often present on the middle of
the back between the fore limbs, and another, more
in the form of a streak, on the coccygeal region.
The dorsal spots sometimes small and irregular, some-

128

DISCOGLOSSIDiE.

times forming handsome symmetrical markings; a large
triangular, heart-shaped, or chevron-shaped marking-
is usually present between the eyes. Lower parts
ivory-white, uniform or closely speckled with brown,
yellowish or more or less carneous under the limbs.
Iris golden in its upper portion, dark bronzy in its
lower two- thirds, the dark lateral streak of the head
passing through the ej^e ; sometimes entirely bronze-
brown, as in Bombinalor, with a fine golden border to
the pupil, interrupted at the lower angle.

Fig. 50.

Lower view of male with nuptial excrescences.

Male distinguished from the female by much stronger
muscular fore limbs, more developed inner carpal
tubercle, a great enlargement and flattening of the
inner finger, and more fully webbed toes, the web even
extending to the inner metatarsal tubercle. Vocal
sacs are in a rudimentary condition, and do not com-

DISCOGLOSSUS. 129

muuicate by any openings with the floor of the mouth.
Horny blackish excrescences are largely developed
and distributed as isolated minute spines over the
ventral surfaces and on the hind limbs ; they form
large groups on the inner palmar tubercle and the two
inner fingers, a band round the chin, and often border
the web of the foot. These excrescences usually
persist longer than in most other Batrachians, being
found, more or less developed, in adult specimens all
the year round.

Geographical Variations. — The polymorphism of
this species has given rise to the establishment of
various species and sub-species, which were believed
to be restricted to certain parts of the habitat of the
genus. Thus a Discoglossus sarcitis, Tschudi, was
stated to be peculiar to Sardinia and Corsica, and a
D. scovazzii, Camerano, was described from Morocco.

Lataste, in his monograph of 1879, had, it seemed,
disposed once for all of these supposed species by
showing the inconstancy of the characters adduced
for their separation ; and, for my part, after examin-
ing a very large material from almost every part of
the habitat, I fully agree with him. But, since
1879, the 1). sarcitis has been maintained, no longer
as a species, but as a sub-species, by Camerano,
and the D. scovazzii, or a form very closely agree-
ing with it, has been revived by Heron-Royer under
the name of D. auritus ; it may therefore be well
to explain in a few words why I take no notice of
them even as varieties. According to Camerano's
latest definition, I), sardus, from Sardinia, Corsica,
and neighbouring small islands, differs from the
typical D. pietus from Sicily, Malta, and the Spanish
Peninsula in having a less acuminate snout, more
robust limbs, and the length of the tibia contained
twice, or a little over twice, in the length of head and
body. The two former characters are really too slight
to be easily appreciable ; and besides, I find Maltese,
Algerian, and Portuguese specimens with the snout

I

130 DISCOGLOSSIMS.

quite as short and broadly rounded as in the Sardinian
form. As already observed by Lataste, this difference
is no greater than that between specimens of Bana
tem,poraria, and is certainly not more constant. With
regard to the third character, I need simply refer to my
tables of measurement and to Camerano's own table,
where Sardinian as well as Sicilian specimens are
shown to have the tibia less than half the length of
head and body.

Beclriaga also is inclined to retain D. sardus as a
distinct form, although admitting: that the existence
of annectant specimens renders a precise definition
impossible. On the whole, he finds the body shorter
in proportion to the head and limbs in Corsican and
Sardinian specimens than in those from Algeria and
Portugal. That there is nothing constant in that
supposed difference may be seen from the following
measurements (in millimetres) of four adult males, the
heads of which are figured above (p. 126), — a from
Corunna, b from Oran, c from Tlemsen, prov. Oran,
and d from Luras, Sardinia :

a.

b.

c.

d.

From snout to vent .

58 ..

. 62 ..

. 71 .

. 60

Length of head

18 ..

. 19 ..

. 20 ..

. 18

Width of head at angles of mouth .

19 ..

. 2L ..

. 23 .

. 21

,, „ below eyes

16 .

. 17-5..

. 20 ..

. 18

From end of snout to fore limb

25 .

. 25 ..

. 25 .

,. 25

Fore limb .

34 .

. 35 ..

. 35 .

35

Hind limb .

83 .

.. 95 ..

. 92 .

. 91

Tibia .

28 .

. 33 ..

. 32 .

.. 30

It will be noticed that the Spanish and Algerian
specimens a and b have the head a little longer in
proportion to the body than iu the Algerian c and the
Sardinian d ; that the width of the head below the eyes
is as great in proportion in the Algerian c as in the
Sardinian d; and that the hind limb is shorter in pro-
portion in the Algerian c than in the Sardinian d.

Heron-Koyer's D. auritus is founded on Algerian
specimens (which had been previously referred by
Camerano to D. scovazzii), supposed to differ from the
Europeans in having the tympanum distinct, the tern-

D1SC0GL0SSUS. 131

poral spot larger, and the inner metatarsal tubercle
smaller. But a male specimen from Algiers, collected
by Mr. Sclater, has the tympanum completely con-
cealed ; and, on the other hand, the organ is perfectly
distinct in a male from Corunna and in another col-
lected by M. Bosca on the Sierra Morena ; whilst a
number of specimens from Europe and Africa are in-
termediate between the two extremes. I have now
before me a large number of living specimens from
Oram In some the tympanum is very apparent, whilst
in others its presence can hardly be detected. The
shape and extent of the dark temporal band and the
size of the metatarsal tubercle are subject to variation
in specimens from the same locality, and I have failed
to find any constancy in the other very trivial distinc-
tive characters pointed out by Heron-Royer. Dr.
Anderson, who has collected specimens in Algeria,
some of which show no trace of a tympanum exter-
nally, concurs with me in rejecting I). auritus even as
a local form.

Specimens from Sardinia, Corsica, Montecristo, and
Giglio, it has been observed, never show the striped
form so frequent in Spain and Africa ; but neither do
the Maltese specimens, which are referred by Camerano
to J), j) ictus.

Measurements (in millimetres).

<? ?

1.

2.

3. 4.

5. 6.

7.

8.

From snout to vent

. 58 .

.55

...60 ... 73 .

..55. ..54..

.43 .

.76

Length of head

. 18 .

.16

...18 ... 21 .

..16. ..15..

.14 .

.19

Width of head

. 19 .

.18

...21 ... 25 .

..19....16..

.15 .

.23

Diameter of eye

. 5 .

. 4

... 5 ... 6 .

.. 5... 4..

. 35.

. 6

Interorbital width .

. 35.

.. 3-5... 4 ... 4-5.

.. 3... 3..

. 3 .

. 5

From eye to nostril

. 5 .

.. 4

... 4-5... 5-5.

.. 4... 4..

. 4 .

.. 5

„ ,, end of snout

. 8 .

P7.
.. t

5... 8-5... 10 .

.. 7... 7..

. 6 .

.. 9

Fore limb

. 34 .

..29

...35 ... 40 .

..27. ..26.

.24 .

..35

Hind limb .

. 83 .

..80

...91 ...105 .

..84.. .77..

.64 .

..95

Tibia

. 28 .

..26

...30 ... 35

..26. ..25.

.20 .

..33

Foot .

. 26 .

..25

...26 ... 32

..24. ..22.

.18 .

..30

1, 5. Corunna : Seoaue.

4. Oran : Cbevreux.

2, 6. Coimbra : Gadow.

7. Giglio :

Florence

Mus.

3. Luras, Sardinia : Camerano.

8. Algiers

: H^ron-!

itoyer

132 DISCOGLOSSID^.

Skeleton. — Ethmoid short, not extending poste-
riorly beyond the anterior third or two-fifths of the
parasphenoid, embracing above a large fontanelle
which extends to the line of the anterior borders of
the orbits, and which remains exposed between the
diverging anterior borders of the fronto-parietals ex-
cept in very old specimens ; its upper lamina obtusely
pointed and partly covered by the nasals, which are
large and form a long suture in the median line.
Fronto-parietals large, in contact along their posterior
two-thirds or more. Zygomatic branch of the squa-
mosal joining by suture an ascending process of the
maxillary. Vomers large, their posterior toothed
border nearly straight, and covering the feeble pala-
tines, narrowly separated from each other in the
median line ; pterygoids regularly trifurcate, with a
small round outer wing-like expansion at the meeting-
point of the three branches, joining a corresponding
process of the angular bone of the mandible ; the
inner branch of the pterygoid in contact with the
parasphenoid, which is _L-shaped, obtuse or truncated,
or indentated anteriorly, and not quite reaching the
vomers. Mento-Meckelians quite indistinct.

Hyoid a large, broad cartilaginous plate with
rounded lateral wings, small postero-lateral processes,
and proximally slender, mesially broad and lamellar
ceratohyal cornua without forward processes ; in addi-
tion to the rather large thyrohyals, which meet with
their inner angles, there is a V-shaped, slender, ventral
ossification, which is in contact with the latter at its
apex, and reaches forward to the anterior notch of the
cartilaginous plate ; this ossification often paired, the
two branches being disconnected at the apex.

Vertebral column twice to twice and one-third as
long as the skull. Vertebrae imbricate and completely
covering the spinal cord above, with a long postero-
median neural process. Second, third, and fourth
vertebras with short diapophyses, to which short ribs
are attached. The first rib is the shortest, horizontal

DISCOGLOSSUS.
Fig. 51.

133

Wliole skeleton of female and fore limb of male.

134 DISCOGLOSSIDiE.

or directed a little forwards ; the second is the longest,
nearly horizontal or directed a little backwards, and
sends off from the middle of its upper border a short,
slender process, directed outwards and backwards ; the
third rib, which may be somewhat curved, is directed
slightly backwards, and in length is intermediate be-
tween the first and second. The following vertebras
have no distinct ribs, but the diapophyses are longer
and more slender, the last two or three beiug turned
more or less distinctly forwards. The diapophyses
of the sacral vertebra, which cover the anterior extre-
mities of the ilia, on which they may shift forwards
and backwards to some extent, are moderately dilated,
their diameter at the free extremity being somewhat
less than their length. Two condyles for articulation
with the urostyle. The latter is a little longer than
the rest of the vertebral column, and bears on each
side at its base a short, slender, transverse process
directed obliquely backwards.

Preecoracoids slender, curved, entering the glenoid
cavity ; coracoids stronger, feebly curved ; supra-
scapula ossified ; omosternum cartilaginous ; sternum
produced into two long, slender, diverging processes.
Humerus once and a half to once and two-thirds as
long as radius-ulna. Carpus with eight bones, two of
which are in contact with radius-ulna ; a single bone in
the pollex, very large in the male ; metacarpal of inner
digit enlarged in the male, with strong inner crest.

Pelvis nearly two-thirds the length of the vertebral
column ; pubis cartilaginous ; acetabulum open. Femur
and tibia with the epiphyses cartilaginous, the former
strongly sinuous and shorter than the latter ; astra-
galus a little longer than calcaneum, from which it is
perfectly distinct throughout, owing to the absence of
calcified epiphyses, aucl one-half or rather more than
half the length of tibia-fibula; three tarsalia in the
distal row, one or two of which may ossify ; and two
bones to the praahallux. Distal phalanges obtuse,
slightly expanded at the apex.

DISCOGLOSSUS.

135

Measurements op Skeleton (in millimetres).

Length of skull

Width of skull

Least interorbital width

Dorsal vertebral column

Urostyle

Humerus

Radius-ulna

Manus

Pelvis

Femur

Tibia

Tarsus

20
24
4
18
19
17
11
17
27
26
31
17

?
15
17

3
17
19
12

8
11
23
21
24
13

Pes .... 25 ... 21

Habits. — Discoglossus pictus resembles the true
frogs in the quickness of its movements. It is
active by day as well as by night, and usually found
in or about water, objecting neither to brackish pools
nor to cold running mountain streams. Like its
relative Bombinator, and unlike all other European
Ecaudates, it is able to seize its prey under water, in
the same manner as newts do, and is therefore easily
fed in confinement with earthworms dropped into
the tank with which the terrarium must be provided.
Although particularly shy when handled, it does well
in captivity, and will usually take food immediately on
arriving after a long journey in a small box. When
seized it becomes covered with a slimy secretion,
which renders it very difficult to handle, but which
has no special odour nor any irritating action on our
mucous membranes, so far as I can judge from experi-
ments upon myself. Like Lataste, who was the first
to study the habits in captivity, and who later had
the good fortune to observe this Batrachian in a wild
state in Algeria, I have never heard the rutting male
produce more than a feeble note, sounding to my ear
as ha-a, ha-a-a, or wa, wa, wa-iva-wa, issued in rapid
succession, although specimens have repeatedly paired
and bred in my terrarium. But according to Heron-
Royer, specimens under his observation uttered at
night a constantly repeated ra-a, ra-a, loud enough
to disturb him when produced in a yard outside his

136

DISCOGLOSSIDJ].

window. Discoglossus cannot be described as mute,
but its voice is certainly the feeblest among all our
members of the tailless tribe.

The pairing season extends from January, in
Algeria, to September and October, and the female
is able to spawu three times in the year. The

embrace, which never lasts

Fig. 52.

J**0*ik^

long,

is lumbar, the

Male and female pairing.

male's hands joining on the pubic region. The re-
markably small eggs are produced slowly, one at a
time, and feebly adhere to the bottom of the pool or
tank, where they form a single layer, more or less
closely set. Only exceptionally have I found eggs
attached to the weeds with which I had furnished the
small tank in the terrarium, the pair keeping to the
bottom during oviposition. The spermatozoa, which
from their large size can be detected with the naked
eye, are ejaculated in several bundles.

The larvae, as a rule, live only from one to two
months before transforming, and keep small, being in
this respect comparable to those of the common toad ;
but tadpoles born late in the summer may remain for
nine or ten months in that condition.

Specimens which I received from Algeria in January
and February, 1897, although in apparently excellent
health and readily feeding on worms and insects, did

DISCOGLOSSUS. 137

not begin to pair, so far as I could observe, until the
beginning of June. The first eggs, about 400 in
number, I found on the bottom of the tank on the
morning of June 7th, the embryos emerging on the
10th; whilst on the 14th, or exactly a week later, the
young had lost their external gills and entered the
tadpole stage, measuring 11 mm. total length, buds
of the hind limbs beinsr discernible on the 17th.
On the 23rd the largest specimens measured 21 mm.,
and on the 3rd July some had reached their full
length, viz. 25 to 30 mm. On the 5th July some
had put on the striped livery. The fore limbs first
appeared on the 6th, and the metamorphosis was
completed by the 8th, when the first specimens left
the water with a reduced caudal appendage, and
measuring 8 to 10 mm. from snout to vent, the last
metamorphosing as late as August 6th. Striped speci-
mens were in the proportion of one to four spotted or
immaculate.

On the morning of June 14th the tank again con-
tained eggs, 895 in number, which, from their con-
dition, must have been deposited in the night of the
12th — 13th (I was absent on Sunday, 13th), and the
embryos of which were liberated in the afternoon of
the 14th, thus less than forty-eight hours after the
deposition of the eggs — a process of development
more rapid than has been observed in any other
species of Batrachians. The young from this brood
began to leave the water on July 20th, whilst others
retained the larval condition until October 5th. None
were of the striped form. A third lot of 936 eggs
were deposited during the night of July 6th — 7th ;
they hatched in the night of 8th — 9th, and the young
completed their metamorphosis between August 16th
and October, a few being still in the tadpole condi-
tion whilst these lines are passing through the press.
About two-thirds of them belonged to the striped
form. In all these cases I was struck by the ex-
tremely small proportion of eggs that did not develop,

138 DISCOGLOSSIU^J.

and the very slight amount of mortality that took
place among the embryos ; whilst of the tadpoles all
except such as I killed for study went through the
metamorphosis. In fact, the success in rearing these
broods of Discoglossus has been greater than I have
experienced in dealing with any other kind of tailless
Batrachians under similar conditions. I had no diffi-
culty in feeding the tiny young on aphides.

On the 17th July eggs had again been produced
during the night, but they numbered only 306. They
hatched on the 18th ; most of them were destroyed
shortly after by a fungus. A further oviposition took
place under my eyes in the afternoon of July 31st,
341 eggs being produced. The male was spotted, the
female striped. The same pair remained in the tank,
which, on the morning of August 2nd, contained a
further lot of eggs, 543 in number — laid, I have every
reason to believe, by the same female. Both these
broods hatched in less than thirty-six hours. The
first went through their evolution without mishap,
metamorphosis beginning on September 11th; whilst
the second perished wholesale as embryos through a
fungus. Among the former, contrary to my expecta-
tion, not one developed into the striped form, but I
had the pleasure of obtaining an albino tadpole, flesh-
colour with large pale golden spots; the so-called
pigmentary network was distinguishable, of an
opaque white. The albinism was, however, not per-
fect, for the pupil of the eye was black instead of red,
and later in life a small quantity of brown pigment
appeared on the back. Further ovipositions took place
at night on August 18—19, 20—21, 23—24, and 26—
27, the number of eggs being 314, 830, 598, 360. All
these broods did well, and at the time this account of
them is written I have a large number of tadpoles,
which will probably not transform until the coming
spring. They differ greatly in their degree of deve-
lopment, irrespective of age, measuring from 11 to 30
millimetres total length.

DISCOGLOSSUS. 139

Eggs. — Small, 1 to 1^ mm. in diameter, dark brown
or black, with the lower third or fourth white or grey,
in isolated capsules measuring
not more than 3 or 4 mm. in ^ @5%«V£)^ ._.
diameter. The number of eggs
does not seem ever to exceed FlG- 53-

1000, and may be as few as 300. Eclosion takes place
on the second to sixth day. The embryo, which then
measures 3 or 4 mm., is blackish-brown ; it has a very
short tail, but the external gills have not yet appeared.

Tadpole (PI. I, fig. 1). — Length of body once and
two-thirds its width, one-half to two-thirds the length
of the tail. Eyes on the upper surface of the body,
the distance between them about once and a half the
distance between the nostrils, equal to or slightly less
than the width of the mouth. Spiraculum in the
mid- ventral line, equally distant from either extremity
of the body. Anal opening median, larger than the
spiraculum. Tail three to four times as long as deep,
broadly rounded at the end, both upper and lower
crests but very feebly convex, the former not extending
upon the back ; the depth of the muscular part at its
base one-half to two-thirds the total depth.

Mouth elliptical. Beak white, edged with black.
Lips bordered by a single series of papillae, which are
usually narrowly interrupted in the middle of the
upper lip ; a well-marked chink on each side of the
lower lip. Series of labial teeth •§-, occupying the
whole width of the lips, the third lower interrupted
in the middle; the first upper and the first lower series
formed of one or two rows of teeth, the others con-
stantly of two.

The ordinary lines of crypts on the head are dis-
tinguishable, and often also one running along each
side of the back.

Brown or olive above, grey or whitish below,
speckled with gold ; caudal crests whitish, uniform or
with small brown dots. Colour-dimorphism is dis-
cernible some time before the egress of the fore limbs ;

140 DISCOGLOSSIDxE.

advanced tadpoles may be marked with a light, dark-
edged vertebral stripe, whilst this is absent in other
specimens ont of the same brood. The whole body and
tail with a network of fine brown lines forming polygonal
meshes ; this network most easily traceableon the tail.

Total length, 33 mm.; body, 12 mm.; width of body,
7 mm. ; tail, 21 mm. ; depth of tail, 6 mm.

Habitat. — Discoglossus pictus inhabits South-western
Europe and North-west Africa. It is found nearly all
over Central, Western, and Southern Spain and
Portugal, but appears to be absent from the east coast
as well as from the Balearic Islands, a fact which is the
more surprising since the species is again abundant in
Corsica, Giglio, Montecristo, Sardinia, Sicily and small
neighbouring islands, Malta and Gozo. In Africa its
habitat extends all over Morocco, Algeria, and Tunisia
north of the Sahara; also on Gralita Island, oft' the
coast of Tunisia. It inhabits the coasts and plains as
well as the hills, having been found by Bedriaga in
Corsica from sea level to an altitude of 2450 feet.

The occurrence of Discoglossus in Santa Maura,
Ionian Islands, whence a single specimen is stated by
De Betta to have been obtained by the late A. Ninni
in 1863, has not been confirmed bv more recent ex-
plorers of those islands, and Bedriaga informs us that
the specimens put up under that name in the Athens
Museum belong; to Bana esculenta.

A few years ago the late Heron-Rover introduced
the species into France near Amboise, in Touraine,
where it appears to have become perfectly acclimatised.
Several thousands of young specimens have also been
turned loose in the neighbourhood of Argenton, dep.
Indre, by M. Rollinat in 1892 and 1893.

PI. IV shows specimens from Oran, for which I am
indebted to the kindness of Messrs. E. Chevreux and
Doumergue. The right-hand figure represents a full-
grown male in nuptial costume ; the two others repre-
sent females of the spotted and striped forms.

5

N

■J

1

N5

■

Ss

<s

BOMBINATOB. 141

2. BOMBINATOR.
Merrein, Tent. Syst. Amph., p. 178 (1820), partim.

Pupil roundish, triangular, or cordiform. Vomerine
teeth in two short transverse groups behind the
choanjfi. Tongue circular, entire, adherent. No
tympanum. Fingers free, toes webbed; outer meta-
tarsals separated by web. No palatine bones. Dia-
pophyses of sacral vertebra very strongly dilated.
Urostyle articulated to a single condyle.

This genus includes three closely allied species.
Two are European ; the third, Bombinator orientaUs,
Blgr., inhabits North-eastern Asia, from Manchuria to
Northern China.

Apart from the coloration, the two European
species are easily distinguished by the length of the
tibia or crus, which is shorter than the foot, measured
from the base of the inner metatarsal tubercle, in
B. igneus, and as long as the foot or even a little
longer in B. pachypus. Males are besides distinguished
by the presence in the former and the absence in the
latter of internal vocal sacs ; during the breeding
season the third finger and one or several of the toes
are provided with plates of black horny asperities in
B. pachypus, which are never present on those parts in
B. igneus.

Although nearly allied, and actually known to pro-
duce fertile hybrids,* the right to specific distinction
of the two European forms, so long confounded
under the name of B. igneus, is now established beyond
contest.

* That this affords no absolute criterion in mattei-s of specific dis-
tinctions is shown, among Batrachians, by the two newts, Molge cristata
and M. marmorata, producing the hybrid known as M. biasii, which
proves to be fertile for one or two generations at least.

142 DISCOGLOSSID/E.

2. BOMBIXATOR 1GNEUS.

(Plates V and VI.)

Linnasus, Faun. Suec, p. 94 (1746).

? Bana variegata, Linnaeus. Syst. Nat., ed. 10, p. 211 (1757).

Bana bombina, Linnaeus. Faun. Suec, ed. 2, p. 101 (1761), and
Syst. Nat, ed. 12, i, p. 355 (1766).

Bufo igneus, Laurenti, Syn. Rept., pp. 29 and 129 (1768);
Schneider, Hist. Ainph., i, p. 187 (1799).

Bufo ignicolor, Lacepede, Quadr. Ov., i, Syn. Meth., and p. 595
(1788).

Bana rubeta. Lindaker, Abb. Bohm. Ges. Wiss., i, 1791, p. 112.

Bombinator igneus, Merrem, Tent. Syst. Amph., p. 179 (1820);
Graven burst, Delic. Mus. Vratisl., p. 67 (1829); Schulz,
Faun. March, p. 470 (1845) ; Nilsson, Skand. Faun., Anrf.,
p. 94 (1842); Collin, Naturh. Tidsskr. (3), vi, 1869, p. 307;
Boulenger, Proc. Zool. Soc, 1886, p. 500, pi. 1, fig. 2 ; Heron-
Royer, Bull. Soc. Zool. France, 1887, p. 640, pi. xii; Bou-
lenger, Bull. Soc. Zool. France, 1888, p. 174; Wolterstorff.
Zeitschr. f. ges. Naturw., lvi, 1889, p. 29, and Jabrb. Ver.
Magdeb., 1S90, p. 318; Heron-Royer, Bull. Soc. Et. Sc.
Anders (2), xx, 1891, p. 201; Mehely, Zool. Anz., 1891,
p. 269, and Math. Term. Kozl. (Budapest), xxiv, 1S91, p. 558,
pi. i ; Erwin Schulze, Schr. Nat. Ver. Harz, vi, 1891, p 40;
Boulenger, Proc. Zool. Soc, 1891, p. 621. pi. xlvii, fig. 4;
Mehely, Math. Nat. Ber. Ungarn, x, 1892, p. 61, pi. iv ;
Werner, Rept. Amph. Oesterr.-Ung., p. 109 (1897).

Bombinator igneus, part., Bonaparte, Icon. Faun. Ital., Rett.
Anf. (1838); Schreiber, Herp. Eur., p. 95 (1875).

Bombinator bombinus, Gluckselig. Lotos, i, 1851, p. 224 ; Bedriaga,
Bull. Soc. Nat. Mosc, 1889, p. 581; Sasserno, Boll. Mus.
Torin., iv, 1889, No. 68; Diirigen, Deutschl. Ampb., p. 552,
pi. ii, figs. 3 and 4 (1S97).

Vomerine teeth in two roundish or transversely oval
groups, close together just behind the level of the

Fig. 54. Fig. 55.

Open mouth. Upper view of head.

choanaa. Openings of the Eustachian tubes minute.
Tongue moderately large, circular, comparatively
thin, entirely adherent.

BOMBINATOR. 143

Head depressed, as long as broad, or slightly broader
than long; snout rounded, scarcely projecting, as long
as the diameter of the orbit ; no canthus rostralis ; no
loreal groove ; nostril a little nearer the eye than the
tip of the snout ; eye rather small, very prominent,
directed upwards and outwards ; interorbital space
narrower than the upper eyelid and nearly equal to
the distance between the nostrils.

Fingers short, obtuse, first shortest, third longest,
fourth a little longer than second ; no subarticular
tubercles ; two or three round palmar tubercles, inner
usually largest and more prominent.

Hind limb stout, little longer than head and body ;
the tibio-tarsal articulation reaches the axil or the
shoulder, the tarso-metatarsal the commissure of the
jaws or the eye ; tibia a little shorter than the femur,
the heels not meeting: when the leo;s are folded at
right angles to the rhachis. Foot longer than the
tibia ; toes short, obtuse, flattened, at least two-thirds
webbed in the adult, sometimes nearly entirely webbed,
but the membrane always with deep crescentic emargi-
nation ; in young specimens the toes may be barely half
webbed ; no subarticular tubercles ; a very small and
feebly prominent round inner metatarsal tubercle.

Upper parts covered with more or less prominent,
round or oval, distinctly porous warts ; each may be
tipped with closely set black asperities, none of which,
as a rule, are developed into true spines, although
there is sometimes a conical central eminence. Some
of the warts may run together and form short chains,
particularly an oblique, posteriorly converging pair
between the shoulders, and a parotoid-like one on
each side behind the eyes; a distinct fold usually
extends below the latter, from the eye to the base of
the arm. Lower parts smooth, or with small scattered
granules, each of which is tipped with black horny
matter as on the dorsal warts ; lower surface of thio-hs
with laro-er flat granules ; a more or less strono- o-ular
fold separating the head from the body.

144 discoglossidj:.

Grey, olive, or rarely bright grass-green above, with
distant symmetrical blackish or dark bottle-green
spots disposed on the larger warts. Usually, in German
specimens, a pair of pale green or bright green roundish
spots between the shoulders. Austrian specimens
often lack the green colour, and Mehely found it in
two specimens only, out of fifty collected in Hungary.
A dark streak from the end of the snout to the eye ;
upper lip with dark vertical bars; limbs, including
the digits, with more or less regular dark ci'oss-bars.
Lower parts bluish-black, usually with white dots,
and more or less numerous, insuliform or vermicular,
bright orange to vermilion markings, the most con-
stant of which are a pair of pectoral spots and a pair
of lumbar cross-bars; the latter sometimes nearly
continuous, like a belt across the praspubic region ; the
spots sometimes absent on the throat and belly, which
then are entirely black, dotted Avith white ; a small
white spot sometimes present on the chin. A large
bright spot on the inner side of the palmar, and
another on the inner side of the plantar surface;
these spots are not confluent with those on the
forearm and tarsus, and do not extend to the tip of
the inner digit ; one specimen, from Vienna, is excep-
tional in having the plantar and tarsal spots confluent
on the right side. The tips of all the digits black or
yellowish-white, never bright orange or red.

Iris golden, much sprinkled over with brown, or
bronzy brown with a golden circle round the pupil ;
the o'olden border broader above than below and at
the sides ; the pupil round with a lower angle, or sub-
triangular, rarely notched above as in the following
species, — in fact, perfectly similar to that of Discoglossus.

Male distinguished from the female by a usually
somewhat shorter body, a rather broader head,
stronger fore limbs, and especially by the presence of
a pair of internal vocal sacs formed by the skin of the
floor of the mouth, which is loose and plicate, and
projects through a slit dividing the submaxillary

BOMBINATOE. 145

muscle into an anterior and a posterior portion ; when
these sacs are fully inflated the throat is globular and
larger than the rest of the head. During the breeding
season black horny excrescences form a band along
the inner side of the forearm, and cover the inner
carpal tubercle as well as the inner side of the first
and second fingers.

Measurements (in millimetres).

3 ?

1.

2.

->
3.

i —
4.

5.

— \
6.

From snout to vent .

. 49 ..

. 46 ..

. 42 .

.. 50 ..

. 46 ..

. 41

Length of bead

. 15 ..

. 14 .

.. 13 .

.. 15 .

. 13 ..

. 12

Width of head

. 16 ..

. 15 ..

. 14 ..

. 16 ..

. 14 ..

. 12

Diameter of eye

o

Q

o
O

o

o

. 3

Interorbital width

o

o

. 1-5..

2

2

1-5

From eye to nostril .

." 2-5 '.'.

o

. 2 ..

.' 2 '.',

','. 2 5.'.'

o

,, ,, to end of snout

. 6 ..

'. 5 "

. 5 ..

. 5 ..

.. 5'5..

. 5

Fore limb

. 22 ..

. 21 ..

. 19 ..

. 21 ..

. 20 ..

. 19

Hind limb

. 57 ..

. 52 .

. 48 ..

. 55 .,

. 52 ..

. 48

Tibia .

. 16 .,

.. 14 .

. 13 .

. 15 ..

. 14 ..

13

Foot .

. 19 ..

. 16 .

.. 16 .

. 17

17 .

15

1,4. Kullen, Sweden : Lilljeborg.

2. 5. Berlin : Boulenger.

3, 6. Holies, Hungary: Mebely.

Skeleton. — Skull very flat and feebly ossified, the
enormous fontanelle being but narrowly bordered at
the sides, or at the sides and behind, by the much-
reduced fronto-parietal bones ; ethmoid short and
nearly entirely exposed, its upper lamina with convex
anterior border, and in contact with or narrowly
separated from the nasals; the latter bones have a
concave anterior and a convex posterior border, and
are in contact with each other in the median line.
Squamosal feeble, with short z}rgomatic, and no pos-
terior branch. Vomers thin ossifications narrowly
separated in the middle line ; no palatine bones ; the
anterior branch of the trifurcate pterygoid much
elongate, extending to the anterior border of the orbit,
where it curves inwards to make up for the missing
palatine, the inner branch short and widely separated
from the parasphenoid ; latter shaped like a dagger
with very short handle, obtuse or truncated anteriorly,

K

146 DISC0GL0SSIDJ5.

not reaching the vomers, leaving the ethmoid un-
covered, or covering but a small portion of it. Stapes
absent. No well-defined mento-Meckelian bones.

Hyoid a large broad plate with two pairs of ossifica-
tions ; ceratohyal cornua broad and without forward
processes; postero-lateral process large, ossified at the
base, this ossification extending to the body of the
hyoid and sometimes nearly meeting its fellow in front
of the likewise ossified but slender thyrohyals.

Vertebral column twice and one- third to twice and
a half as long as the skull. Vertebras imbricate, the
column closed above. Second, third, and fourth ver-
tebrae with short diapophyses and small distinct ribs,
the second of which is the longest, and bears a small
posterior process as in Diseoglossus. The first diapo-
physis is directed forwards, the second, third, and
fourth are nearly horizontal, and the three following
are again directed forwards. The diapophysis of the
sacral vertebra are very strongly dilated, and cover
the anterior third of the pelvis ; their longitudinal
diameter, without the cartilaginous epiphysis, equals
about twice the transverse diameter ; this vertebra
has a single condyle for articulation with the urostyle.
As in Diseoglossus, the latter bone has a posteriorly
directed, slender transverse process on each side at
its base, but usually more developed, its length often
exceeding that of any of the processes of the presacral
region ; it is sometimes followed by a second similar
process ; the length of the urostyle equals that of
the skull, or of the six or seven anterior vertebrae.

Proecoracoids moderately slender, strongly curved,
entering the glenoid cavity ; coracoids little stronger,
feebly curved ; supra-scapula cartilaginous, or partially
ossified ; sternum cartilaginous, produced into two
long, slender, diverging processes. Humerus once
and one-third as long as radius-ulna. Carpus with
eight elements, as in Diseoglossus, two of which are
in contact with radius-ulna ; two bones in the pollex.

Pelvis measuring three-fifths to two-thirds the

BOM RI NATO It.

147

length of the vertebral column ; no pubis, ilium and
ischium meeting in the acetabulum. Femur slightly
curved, a little longer than the tibia; calcaneum
slightly longer than the astragalus, two-thirds to three-

Fig. 56.

Skeleton of male.

fourths the length of the tibia ; three cartilaginous
tarsalia, and one or two minute cartilages in the prae-
hallux. Distal phalanges obtuse, slightly expanded at
the apex. The epiphyses of all the longbonesuncalcified.

148 DISCOGLOSSID^].

Measurements of Skeleton (in millimetres),

(J

?

Length of skull

13

11

Width of skull

14

12

Least interorbital width .

2-5 ..

.

Dorsal vertebral column .

It]

15

Urostyle

13

12

Humerus

9

8

Radius-ulna

rt

/

6

Manns

10

9

Pelvis

20

17

Femur

14

13

Tibia

13

12

Tarsus

0

8

Pes ...

18

16

Habits. — This species is diurnal and thoroughly
aquatic during its period of activity, being found in
small ponds or pools or among the grass on the
borders. It is very agile in the water, and proceeds
by small leaps on land. It likes to rest near the
surface of the water with spread-out limbs, nothing
but the eyes and nose emerging, and dives to con-
ceal itself in holes or under stones or bury in the
mud when disturbed. I found it pairing near Berlin
in the latter half of May. The male embraces the
female round the waist, and, swelling out its throat,
utters at short intervals a somewhat powerful and
melancholy note, lioolc, It oak, or hoonl; hoonlc, whence
the name " Unke " by which it is generally known in
Germany. The pairing season extends over two or
three months, and Heron-Rover has ascertained that the
female spawns two or three times in the year at distant
intervals. The eggs are produced singly or a few at
a time, and become fixed by their gelatinous capsules
to submerged grass or other weeds ; this operation is
performed slowly, and may last from one to three
days, with several hours' interruption.

Males appear to reach sexual maturity much sooner
than females. I have found pairing male specimens
not more than 28 mm. long from snout to vent,
although the youug that have just completed the
metamorphosis measure as much as 15 to 20.

BOMBINATOK. 149

The skin of the upper parts produces an acrid
secretion, very similar to that of the crested newt,
fresh-caught specimens becoming covered with white
froth when handled ; and this exudation causes great
irritation to our mucous membranes, fits of sneezing
and running of the eyes being the usual consequence
of simply looking into the bag in which the captives
are being brought home. When surprised on land,
and unable to escape in the water, this Batrachian,
in common with its congener B. pachypus, makes
ridiculous contortions, ultimately feigns death, and
concavely bending its spine turns up the head and
hind part of the body, as well as the limbs which
are folded over, thus exposing the brilliantly coloured
lower surfaces ; at the same time covering its eyes
with its hands as if not to see the danger.

Eggs. — The vitellus measures about 2 mm. in
diameter ; the upper hemisphere is dark brown, the
lower yellowish-white. There are two gelatinous
capsules, the outer of which measures 7 or 8 mm.
in diameter. The eggs being laid singly or in small
groups of seldom more than ten at intervals of several
hours, the exact number produced by one female has
not yet been ascertained with absolute certainty, but
appears to be from 80 to 100 for each brood. The
embryo escapes from its envelop after a week in a
more advanced condition than Discoglossus, provided
with small external gills and a well-developed tail, and
more or less distinctly striped as in that of Alytes.

Tadpole (PI. I, fig. 2). — Length of body once and
one-fourth to once and one-third its width, two- thirds
to four-fifths the length of the tail. Eyes on the
upper surface of the body, the distance between them
twice and a half to three times as great as that
between the nostrils, equal to or slightly less than the
width of the mouth. Spiraculum in the mid-ventral
line, nearer the posterior than the anterior extremity
of the body. Anal opening median, much larger than
the spiraculum. Tail twice to twice and a half as long

150 DISCOGLOSSIM;.

as deep, ending in an obtuse point ; the upper crest
convex, not or but slightly deeper than the lower,
and extending upon the back ; the muscular part at
its base two-fifths to one-half the total depth.

Mouth triangular. Beak white, edged with black.
Lips bordered by a series of papilla? ; a well-marked
chink on each side of the lower lip ; series of labial
teeth § , occupying the whole width of the lips, all
uninterrupted, or the third lower broken up in the
middle ; the first upper and the first lower series
formed of two or three rows of teeth, the others of
two, three, or four.

Well-marked series of muciferous crypts ; one on
each side of the head, from above the upper lip,
passing above the nostril and bordering the eye, then
descending towards the upper lip, where it curves and
ascends to below the eye; two on each side of the
back, beginning at some distance behind the eye, the
upper extending to the dorsal portion of the muscular
part of the tail, the lower very short and parallel to
the upper ; and finally a short curved one on each
side of the belly.

Brown above, greyish-white below ; the series of
crypts whitish ; tail greyish, with or without small
brown spots. A network of fine blackish lines
crossing each other at right angles is spread over
the whole tadpole.

Total length 50 mm. ; body, 20 ; width of body, 16 ;
tail, 30; depth of tail, 15.

Habitat. — Bombinator hjnem must be regarded as
an Eastern species, essentially bound to the plain.
It inhabits Russia, west of the Volga, between lat. 47°
and 56°, Southern Sweden (Scania), Denmark, North
Germany (as far west as the Weser and Oldenburg),
Austria-Hungary, and Moldavia. As pointed out by
Wolterstorff and confirmed by v. Mehely, it avoids
hilly districts, where it is replaced by its congener
B.pachypus; only in a few places in Thuringia, Lower
Austria, and Transylvania are the two species to be

. y /

VI

/. 2. ■ '>/■'//,/>//,<"/<■>/ /,//,,/,. . ". - 4v//////^A/ Afr// //////. y

BOMBINATOR. 151

found together. The highest altitude at whicli it has
yet been found is about 800 feet in Transylvania.

The two specimens represented on PI. V, left hand,
are from Magdeburg, kindly sent me by Mr. W. Wol-
terstorff. The male shows the ventral aspect and the
inflated gular sac ; the other specimen is a female.

Two more specimens, both males, are figured, lower
aspects, on PI. VI ; fig. 1 from Vienna, fig. 2 from
Holies, Hungary.

3. BOMBINATOR PAGHYPUS.

(Plates V and VI.)

Roesel, Hist. Ran., p. 97, pis. xxii and xxiii (1758).

Bufo salsus, Schrank, Naturhistor. Briefe, i, p. 308 (1785) ;

Schneider, Hist. Ampb., i, p. 213 (1799).
Bana salsa, Gnielin, Syst. Nat., i, p. 1049 (1789).
Bana sonans, Lacepede, Quadr. Ov., i, Syn. Meth., and p. 535,

pi. xxxvii (1788).
Bana bombina, Sturm, Deutschl. Faun., iii (1797).
Bana ignea, Shaw, Gen. Zool., iii, p. 116, pi. xxxv (1802).
Bufo bombinus, Daudin, Hist. Rain. Gren. Crap., p. 75 (1803), and

Hist. Rept., viii, p. 146 (1803).
Bufo pluvialis, Daudin, Hist. Rain., Sec., pi. lxxxvi.
Bombina ignea, Oken, Lehrb. Naturg., iii, p. 207 (1816) ; Koch,

in Sturm, Deutschl. Faun., iii (1828) ; Reider & Hahn,

Faun. Boic, iii (1832j.
Bombinator igneus, part., Bonaparte, Icon. Faun. Ital., Rett.

Anf. (1838); Schreiber, Herp. Eur., p. 95 (1875) ; Boulenger,

Cat. Batr. Ecaud., p. 447 (1882).
Bombinator pabhypus, Fitzinger, in Bonaparte. 1. c. ; Boulenger,

Bull. Sue. Zool. France, 1888, p. 175; Heron-Royer & Van

Bambeke, Arch. Biol., ix, 1889, p. 282; Bedriaga, Bull. Soc.

Nat. Mosc, 1889, p. 566; Sasserno, Boll. Mus. Turin., iv,

1889, No. 68; Heron-Royer, Bull. Soc. Et. So. Angers (2),

xx, 1891, p. 211; Mehely, Zool. Anz., 1891, p. 269, and

Math. Term. Kozl. (Budapest), xxiv, 1891, p. 563, pi. ii ;

Boulenger, Proc. Zool. Sue, 1891, p. 621, pi. xlvii, fig. 5;

Mehely, Math. Nat. Ber. Ungarn, x, 1892, p. 68, pi. v;

Martin & Rollinat, Vert. Dep. Indre, p. 352 (1894);

Buuleuger, Boll. Mus. Turin., xi, 1896, No. 261 ; Werner,

Rept. Ampb. Oesterr.-Ung., p. 106 (1897) ; Diirigen,

Deutschl. Amph., p. 543, pi. ii, figs. 1 and 2 (1897).
Bombinator brevipes, Blasius, Isis, 1839, p. 667, and Ber. 19. Vers.

Deutsch. Naturf. Brunswick, p. 81 (1841); Erwin Scbulze,

Schr. Nat. Ver. Harz, vi, 1891, p. 40.
Bombinator igneus, Duvernoy, Regne Anim., Rept., pi. xxxix,

fig. 1 (1836) ; Dumeril & Bibron, Erp. Gen., viii, p. 487

(1841); Bruch,Wiirzb. Nat, Zeitschr ,iv, 1863, p. 96; Fatio,

152 niseoGLossiD.K

Vert. Suisse, iii, p. 368 (1872) ; Koch, Ber. Seuck. Ges., 1872,
p. 162; De Betta, Faun. Ital., Rett. Anf., p. 70 (1874);
Lataste, Herp. Gir., p. 275 (1876); Lejdig, An. Batr., p. 50
(1877); Bedriaga, Zool. Anz., 1879, p. 664; Camerano, Atti
Ace. Tor. (2), xxxv, 1883, p. 211, figs.

Bombinator variegatus, Bedriaga, Bull. Soc. Nat. Muse, 1881,
p. 291.

Bombinator bombinus, Boulenger, Proc. Zool. Soc., 1886, p. 499,
pi. 1, fig. 1; Hcron-Royer, Bull. Soc. Zool. France, 1887,
p. 610 ; Wolterstorff, Zeitscbr. f. ges. Naturw., lvi, 1888, p. 28 ;
Erwin Schulze, Zool. Anz., 1891, p. 161.

Although closely allied to the preceding, with which
it has long been confounded, this species is easily
distinguished by several important characters.

The general habitus is stouter still. Head con-
stantly broader than long ; snout more broadly
rounded, as long as or slightly shorter than the
diameter of the orbit ; eye rather larger ; nostril
equally distant from the end of the snout and the
eye, or slightly nearer the latter. In some males the
tarso-metatarsal articulation reaches a little beyond
the eye. Tibia as long as the femur, and as long as
or slightly longer than the foot ; the heels usually
meet when the legs are folded against the thighs.
The foot is usually rather swollen, the digits shorter,
and the web fuller, often with rectilinear border in
the males. Warts on the upper surfaces usually more
prominent and more crowded, not forming sym-
metrical chains ; parotoid gland very rarely distinct.

Fig. 57.

■^*.->-.'V, A ■' '-I'-'S '■■'■'■ '■'■'; ■■"'* ' ' '■■■ ■'■■

•< .; ijt - ■/,.-?"■ ■- , -v.3- '■■"■* •■' ■* -.v,-.-: •

''■.-- 1 ■ ■ '■■• .'-■ -, . . - ■ 'irl
■a- " ■' r/* ' ■ ■■$ >■•.-.•' * :;•:"-*': ** '

- &m?m m i m ■!.--a-'\- mm

Piece of dorsal skin of male ( X 8).

The black horny excrescences on the warts con-
sisting in the males of regular spines, distinguishable
with the aid of an ordinary hand lens, surrounded
with very small points (Fig. 57); in female specimens,

B0MB1NAT0R. 153

however, these excrescences are often not different
from those of B. igneus, which in the males may
exceptionally assume the form of spines. Gular fold
usually absent or rather indistinct; sometimes, how-
ever, as well marked as in B. igneus.

Upper parts yellowish, greyish-brown, or olive, as if
powdered with metallic bronzy dust, uniform or with
dark spots disposed as in B. igneus; usually a pair of
more or less distinct, light yellowish or whitish, round
spots between the shoulders, and another pair further
back in the middle of the body. Black has been as-
signed as the ground colour of the lower surfaces in B.
igneus ; in this species in most cases these parts should
be described as yellow, varying from pale straw-colour
to orange, with greyish-blue or black spots or mar-
blings, which may be lighter in the centre. In some
specimens, however, the black is more diffused, and
might be regarded as the ground colour on which
small yellow blotches are irregularly distributed, in
addition to white dots, as in B. igneus ; such is the
specimen from Basle figured on PI. VI, fig. 4, and
I have seen similar ones from Belgium and Normandy.

The yellow of the lower surface of the thigh sends
up a process which is well visible from behind when
the animal swims, as a patch halfway between the
vent and the leg, and the bright colour of the lower
surface of the arm often extends uninterrupted across
the breast, which it never does in B. igneus. The
fingers and toes are tipped with bright yellow, and
the yellow palmar and plantar spots involve the whole
of the inner digit, unless interrupted by a small black
spot, sometimes the whole of the second, and may
even extend on their upper surface; the plantar spot
is often, but by no means always, confluent with the
tarsal. In specimens from the peninsula of Italy
(the typical B. pachyjpus of Fitzinger), such as the
one from Calabria figured on PL VI, fig. 3, the
yellow tarsal spot is absent or reduced to a few small
marblings, and the lower surface of the tibia may also

154 DISOOGLOSSIDiE.

be uniform black, or grey spotted with black. In
these peninsular specimens, which strongly contrast
with those from north of the Po, the yellow of the
under surface of the arm does not extend across the
breast as usual in the individuals from North Italy,
Austria-Hungary, and Germany (var. brevipes), and
as shown by the example from Schemnitz, Hungary,
figured on PI. VI, fig. 5; but the breast is either
uniform blue, grey, or blackish, or bears a pair of
isolated spots as in B. igneus. In France, Belgium,
and Switzerland we find individuals bridging over the
two forms. Specimens, eight in number, obtained by
M. Lataste at Boulay-les-Trous, near Paris, are very
interesting in this respect. In all the plantar spot is
completely separated from the tarsal, which may be
large or broken up and reduced to a few marblings ;
in some the spots on the breast are completely
detached from those of the fore limbs, and the inner
digits of both hand and foot are not completely
involved in the palmar or plantar spot. Detached
spots on the breast aud tarsus also occur in Belgian
examples. In 25 from near Dinant I count 16 with
the plantar spot separated from the tarsal on both
sides, 3 with the two confluent, and 6 in which the
right side belongs to the one, and the left to the other
of the two categories. In 25 from Mondorf, Luxem-
burg, 7 belong to the first category, 12 to the second,
and 6 to either. A specimen from Goslar, Harz, has
also the plantar and tarsal spots disconnected.

Some of the principal variations observed in the
markings of the lower parts are represented on PL VI.

Iris more uniformly bronzy than in the preceding
species, with the golden edge to the pupil finer, less
distinct ; pupil roundish, with lower angle triangular,
or, more frequently, cordiform, Y-shaped when fully
contracted.

Very young specimens bluish-white or greyish
beneath, with black spots and large yellow blotches
on the limbs (Bufo salsus, Schrank). In the second

BOMBINATOR. 155

year the yellow pigment extends to the other parts of
the under surface. The bright colours are never fully
developed in specimens under 20 mm. body-length.

The male of this species differs from that of the
preceding in the total absence of vocal sacs, the sub-
maxillary muscle being undivided ; and in the presence,
during the breeding season, of a band of black horny
excrescences on the inner side of the third finger as
well as on the first and second, and of small round or
oval groups of similar excrescences under the penulti-
mate phalanx of the third, the second and third, or
the second, third, and fourth toes, as first pointed out

Fig. 58.

Lower view of fore limb and foot of male, showing nuptial

excrescences.

by Bruch in 1863, although their discovery is usually
attributed to Ley dig, who first gave a figure of them.
The spiny horny excrescences of the upper parts are
more developed in the males than in the females, the
difference being most marked during the breeding
season, the development of these excrescences being-
correlated with that of the so-called copulatory or
clasping plates which arm the fingers and toes.

Geographical Variations. — As we have seen above
in dealing with the markings of the lower parts, it is
possible to correlate some of the variations with the
habitat. Thus I think I can in every case recognise
specimens from the mountains of the Italian Peninsula,
which represent the typical B. pachypus, by the pre-
dominance of orange or yellow on the belly and the
lower surface of the thighs, its absence or its reduc-
tion to spots on the breast, and its absence on the
tarsi; the habitus is particularly massive, and the size
larger. Specimens from other parts of Europe may be

156 DlSCOGLOSSlD^l.

grouped together as a var. brevipes, Blasius, in which
we notice a gradual predominance of yellow together
with an attenuation in its intensity as we proceed from
west to east. There is yet a third form, from Monte-
negro, with which I am only acquainted through Schrei-
ber's description, and which perhaps deserves to rank
as a variety ; the lower parts are black, without or
with small and isolated yellow spots. The skin is
described as comparatively smooth, with widely sepa-
rated warts covered with a black horny layer.

Measurements (in millimetres).

c? 2

1. 2. 3. 4. 5. 6. 7. 8.

From snout to vent .45 ...44 ...48 ...46 ...44 ...45 ...50 ...42

Length of head . . 13 ...13 ..'.14 ...13 ...13 ...13 ...14 ...12

Width of head . .15 ...15 ...18 ...15 ...14 ...14 ...17 ...13

Diameter of eye . 4 ... 35... 4 ... 4 ... 3'5... 35... 4 ... 3

Interorbital width . 2 ... 2 ... 25... 25... 2 ... 2 ... 3 ... 2

From eye to nostril . 2'5... 2'5... 25... 3 ... 2'5... 25... 25... 25

,, „ endofsnout 6 ... 5 ... 6 ... b' ... 5 ... 6 ... 6 ... 5

Fore limb . .23 ...22 ...25 ...23 ...19 ...20 ...23 ...19

Hind limb . .51 ...50 ...57 ...52 ...46 ...51 ...53 ...46

Tibia . . .16 ...15 ...17 ...16 ...14 ...15 ...17 ...14

Foot . . .16 ...15 ...17 ...16 ...14 ...15 ...17 ...14

1, 5. Mondorf, Luxemburg : Bouleiiger.

2, 6. Marcellise, Prov. Verona : De Betta.
3. S. San Bruno, Calabria : Giglioli.

4, 8. Parnassos : Kriiper.

7. Arsoli, Rome : Vinciguerra.

Skeleton. — The osteological differences between
this and the preceding species are very slight indeed.
The most constant is found in the length of the tibia,
which equals that of the femur. The sacral diapo-
physes are usually not quite so strongly dilated, and
the bones are nearly white instead of yellow as usual
in the allied species.

As has been shown by Goette, whose great work
1 Entwickelungsgeschichte der Unke ' deals with this

species, by Camerano (' Atti Ace. Torin.,' xv, 1880,
p. 445) and by Sasserno (op. cit., xxiv; 1889, p. 703),
there often occur curious individual anomalies in the
posterior vertebras, which may result in asymmetry of

BOMBINATOR.

157

the sacrum or in the presence of two sacral diapo-
physes, the second of which is produced by enlarge-
ment of the process at the base of the nrostyle, and
connected distally with the one in front of it by a
common cartilaginous epiphysis. Both Camerano
(loc. cit.) and Howes (' Proc. Anat. Soc. Gr. Brit.,'
1890, p. xvi) have recorded a curious anomaly similar
to that observed by Lataste in an AJytes obstetricans :
the sacral process on the right side is normal, whilst
that on the left side is formed by the tenth vertebra,
or base of the urostyle. The latter bone is sometimes
provided with two processes on each side instead of
one. I have examined a specimen from Amboise,
preserved in M. Lataste's collection, which is re-
markable for having nine prsesacral vertebra?, and
perfectly normal sacral vertebra and urostyle.

Measurements of Skeleton (in millimetres).

s

?

Length of skull

12

11

Width of skull

14

12

Least interorbital width .

2-5 ..

Dorsal vertebral column .

15

14

Urostyle

11

12

Humerus

,

10

8

Radius-ulna

,

65 ..

6

Manus

,

8

7

Pelvis

, ,

17

17

Femur

.

14

13

Tibia

14

13

Tarsus

t .

9

8

Pes .

, ,

14

13

Habits. — B. pachypus has much the same habits as
its congener, but is less particular in the choice of the
localities. Any sort of pools, warm or cold, clear or
dirty, with or without vegetation, even the merest
puddles, appear a satisfactory resort to this livery
little Batrachian, which may be heard in spring and
summer uttering its low and mournful note, hoo, hoo.
This sound is often produced by large numbers in a
small pool exposed to the full rays of the sun. Pairing
takes place two or three times in the year, and spawn
is mostly found from the end of May to the middle

158 DISCOGLOSSID^E.

of July, sometimes as late as September, as observed
by Pfli'iger near Bonn. Specimens from Belgium kept
by me in confinement were still pairing on August
18th, and Mr. A. Pam informs me that some brought
home by him this summer from Switzerland were ob-
served in embrace as late as October 13th.

Mehely found spawn as early as the 2nd of May in
Transylvania. Spallanzani, in the last century, ob-
served it pairing in May in the mountains near
Modena, and he tells us that during his journey to
Switzerland in 1779 the same species was pairing in
July and August. I have myself observed it in
embrace in the end of May near Salzburg and near
Bordeaux, and spawning in Belgium in June and
July. Two years ago, around Freiburg in Baden, the
breeding*1 seemed to be over in the beginning; of
August. On the whole, as observed by Werner, this
species appears to breed a little later than its con-
gener ; near Vienna, where the two occur, the season
has been observed by that authority to begin in the
first half of May for B. igneus, in the second half of
the same month for B, pachypus. The eggs are
usually attached to weeds as in the other species ;
but as they are sometimes laid in puddles without any
trace of vegetation, they then simply drop to the
bottom. The lack of vegetation is no hindrance to
the development of the larvas, which are mainly car-
nivorous. Recently transformed young were found
by Schrank in the last century in the salt water of
the saline caves of Berchtesgaden, Austria.

These Batrachians hibernate on land in holes or
under stones. In early spring, in Belgium, I have
found several specimens together in their winter
quarters — deep recesses between stones, which they
shared with Salamandra maculosa. They do not
appear to emerge before the end of April, when they
travel considerable distances in search of suitable
breeding localities, breaking the journey in any small
temporary puddle they come across. I much doubt

B0MBINAT0R.

159

their ever spending the winter in the mud at the
bottom of pools, as they are believed by Fatio occa-
sionally to do.

Eggs. — Do not differ materially from those of the pre-
ceding species. Upper hemisphere pale brown, lower
yellowish- white. The figure here given is a copy of
Heron- K oyer's.

Fig. 59.

Tadpole (PI. I, fig. 3). — One very striking character
distinguishes this tadpole from the preceding, viz. the
shape of the mouth, which is elliptical as in other
genera of Discoglossidse. The tail is, as a rule, rather
shorter, and the muciferous crypts are hardly distin-
guishable in the tadpoles obtained by me ; however,
it is very probable that the latter difference would not
prove constant if tested on more extensive material.

Body 17 mm. ; width of body 14; tail 20; depth of
tail 10.

Habitat. — Bombinator pachypus has a very exten-
sive range in Europe. It is found nearly all over
France with the exception of the north coast of

160 DISCOGLOSSIDJK.

Brittany, in the plains as well as in the hills, but
very local in many parts, and does not ascend the
Pyrenees ; South-eastern Holland ; Belgium south of
the Sambre and the Meuse, and in the calcareous
localities near Tournay ; Germany all along the Rhine,
but not higher up than 2000 feet in Baden ; South
Germany, Switzerland, in the valleys as well as on
the mountains up to 3900 feet. East of the Weser in
Germany this species enters into competition with
B. igneus, and, as first pointed out by Wolterstorff,
becomes restricted to the hills ; this being the case
wherever the two species co-exist, as in Austria-
Hungary and Moldavia. Its highest recorded oc-
currence in Transylvania is at 3900 feet, as in
Switzerland, whilst in the Tyrol it reaches to 4850
feet, and in Bosnia to 5500 feet. In Italy the
species is found in Lombardy and Venetia,* in the
Apuan Alps up to 4500 feet, and along the chain of
the Apennines from Emilia to Calabria, between COO
and 3000 feet, as I am informed by Prof. Giglioli. It
is on record from the Etna, but not from other parts of
Sicily. East of the Adriatic it occurs in Dalmatia,
Bosnia, Herzegovina, Montenegro, Turkey (Adria-
nople), and Greece (Parnassos, 3000 feet).

Mehely has recently expressed the opinion that
B. pachypus is to be regarded as a typical hill or
mountain form, which only descends to the plain in
the cooler northern regions, like Bona temporaria.
This is certainly not the case. In the south-west of
France the species is quite abundant at sea level ; I
have myself collected it on the banks of the Garonne,
in the immediate vicinity of Bordeaux, where it is
plentiful in small ditches bordering the vineyards ;
and E. de Betta found it numerous in the plains
of Venetia.

* The species is absent from Piedmont. Some hundred specimens
from Venetia were turned loose in the neighbourhood of Turin by
Count Peracca about ten years ago.

BOMBINATOR. 161

Male (ventral view) and female (dorsal view)
specimens from near Dinant, Belgium, are figured on
the right side of PI. V. Ventral views of specimens
from the Serra San Bruno, Calabria (fig. 3), Basle,
Switzerland (fig. 4), and Schemnitz, Hungary (fig. 5),
are given on PI. VI.

Hybrids. — Mehely has pointed out that in the few
places in Hungary where the two species occur
together, intermediate specimens are fouud such as
are not to be procured from other localities. These
specimens are no doubt hybrids. Heron-Royer suc-
ceeded in crossing the two species in both directions,
and four nearly adult hybrids are described and
figured by him in 'Mem. Soc. Zool. France,' 1891,
p. 81. These hybrids, which proved fertile, and pro-
duced a second generation, were nearly intermediate
between the parent forms, but somewhat nearer
B. igneus, whether the latter was the father or the
mother : the ventral spots were lemon-yellow on a
black ground ; the digits were not tipped with bright
yellow ; the inner finger and the inner toe were yellow
in one specimen, in the others the inner finger was
black, but the inner toe yellow.

162 DISC0GL0SS1D/E.

3. Alytes.

Wagler, Syst. Amph., p. 206 (1830).

Pupil vertical. Vomerine teeth in transverse or
slightly oblique series behind the choanse. Tongue
circular, entire, slightly free behind. Tympanum
distinct. Fingers free, toes webbed ; outer meta-
tarsals separated by web. Diapophyses of sacral
vertebra strongly dilated. Urostyle articulated to
two condyles.

Two species, both of which inhabit Europe. The
presence of three palmar tubercles in A. obstetricans,
and of two in A. cisternasii, and the short swollen
outer finger in the latter easily distinguishes them, in
addition to several other characters, external and
osteological.

The characters in which the latter species differs
from the former are adaptations to more burrowing
habits, and of too slight importance, in my opinion, to
justify the genus Ammoryctis, proposed by Lataste in
1879 for the then newly discovered Alytes cisternasii,
which has since proved to be endowed with the same
extraordinary nursing habits as its long known and
famous congener. It may be added, as a further
argument against generic distinction, that the tad-
poles of the two species resemble each other so closely
as to be almost indistinguishable before the limbs
have made their appearance ; this being the only
instance that I know among Batrachians of well-
marked species not being differentiated in their larval
condition. The contrary would, however, in my
opinion, be no valid objection to generic association,
for reasons explained in the Introduction, p. 110.

3

Mt

:*W

\

I

\-

\

V

1

Si

ALYTES. 163

4. Alytes ojjstetricans.
(Piute VII.)

Deniours, Hist. Acad. Sc., 1741, p. 28, and Mem. Ac. Sc, 1778,
p. 13.

Bufo obstetricans, Lam-en ti, Syn. Rept., pp. 28 and 128 (1768) ;
Brongniart, Bull. Soc. Philom., ii, 1801, p. 91, pi. vi, fig. 4;
Daudin, Hist. Rain. Gren. Crap., p. 87, pi. xxxii, fig. 1
(1803), and Hist. Rept., viii, p. 176 (1803) ; Zavvadski, Faun.
Gal.-Bukow. Wirbelth, p. 156 (1840).

Rana campanisona, Laurenti, 1. c, p. 30.

liana obstetricans, Wolf, in Sturm, Deutscnl. Faun., iii, Heft 4
(1805).

Bombinator obstetricans, Merrem, Tent. Syst. Amph., p. 179
(1820) ; Gravenhorst, Delic. Mus. Vratisl., p. 68 (1829).

Bufo campcuusonus, Goldfuss, Handb. d. Zool., p. 484 (1820).

Alytes obstetricans, Wagler, Icon. Amph., pi. xxii, figs. 3 — 5 (1830) ;
Schinz, Faun. Helv., p. 145 (1837) ; Tschudi, Isis, 1837, p. 702 ;
Bonaparte, Icon. Faun. Ital., Rett. Anf (1838); Dumeril &
Bibron, Erp. Gen., viii, p. 467 (1841); Giintber, Cat. Batr.
Sal., p. 38 (1858); Brucb, Wi'uzb. Nat. Zeitscbr., iv,
1863, p. 92, and Ber. Offenb. Ver., v, 1864, p. 51, fig.;
Fatio, Vert. Suisse, iii, p. 358 (1872); Kocb, Ber. Senck.
Ges., 1872, p. 155; Schreiber, Herp. Eur., p. 102 (1875);
De l'lsle, Ann. Sc. Nat. (3), xx, 1876, No. 7; Lataste,
Herp. Gir., p. 249 (1876); Leydig, An. Batr., p. 64 (1877);
Lataste, Bull. Soc. Zool. France, 1877, p. 281; F. Miiller,
Verh. Nat. Ges. Basel, vi, 1877, p. 420; Boulenger, Cat.
Batr. Ecaud., p. 448 (1882); Heron-Royer, Bull. Soc. Zool.
France, 1883, p. 417, pi. xiii, and 1886, p. 671 ; Nebring,
Sitzb. Ges. Nat. Fr. Berlin, 1887, p. 48; Heller, Zool. Gart.,
xxix, 1888, p. 180; Heron-Royer &. Van Bambeke, Arch.
Biol., ix, 1889, p. 285; Bedriaga, Bull. Soc. Nat. Mosc.,1889,
p. 596; Fatio, Vert. Suisse, v, App., p. 7 (1890); Wolter-
storlf, Zool. Anz., 1891, p. 65 ; Erwin Scbulze, Schr. Nat. Ver.
Harz, vi, 1891, p. 42; Boulenger, Proc. Zool. Soc, 1891,
p. 622, pi. xlvii, figs. 6 & 7; Wolterstorff, Zool. Anz., 1893,
p. 151 ; Oudetuans, Tijdschr. Dierk. Ver. (2), iv, 1893, Vers).,
p. xlii; Martin & Rollinat, Vert. Dep. Indre, p. 356 (1894);
Werner, Rept. Amph. Oesterr.-Ung., p. Ill (18L»7) ; Diirigen,
Deutschl. Amph., p. 558, pi. i, fig. 4, and pi. ii, fig. 6 (1897).

Obstetricans vulgaris, Duges, Rech. Ostcol. Batr., p. 7 (1834).

Alytes obstetricans delislei, Lataste, Rev. Int. Sc, iv, 1879, p. 543.

Alytes obstetricans boscai, Lataste, 1. c, and Act. Soc Linn. Bord.,
xxxiv, 1880, p. 181, pi. xi; Bedriaga, Amph. Rept. Portug.,
p. 25 (1889), and Larves Batr. Portug., p. 12 (1891).

Alytes boscai, Heron-Royer & Van Bambeke, 1. c, p. 289.

Vomerine teeth in a transverse series, interrupted
in the middle, behind the choanse; the series extends

164 DlSCOGLOSSID^.

outwards to the vertical of tbe inner or of the outer
border of the choanse. Tongue large, thick, circular,
slightly free behind.

Fig. 60. Head large, moderately depressed,

broader than long; snout rounded,
slightly projecting, as long as the
diameter of the orbit ; can thus ros-
tralis rounded ; loreal region grooved,
nostril a little nearer the eye than
the tip of the snout ; eye large, very
prominent; interorbital space as broad
as the upper eyelid, and as broad as

Open mouth. Qr & Httle broader tban tbe clisfcance

between the nostrils ; tympanum circular, three-fifths
to four-fifths the diameter of the eye.

Fingers rather short, depressed, obtuse, first shorter
than second, as long as fourth or slightly shorter,
third longest ; no subarticular tubercles ; F 61
three prominent palmar tubercles, corre-
sponding in position to the base of the first,
third, and fourth fingers, inner oval, outer
round and largest, median round and
smallest.

Hind limb short; the tibio-tarsal articula- Lower
tion reaches the shoulder in the female, the view of
tympanum in the male ; the tarso-metatarsal e b '
articulation reaches the eye in the female, the tip of
the snout or between the eye and the tip of the snout
in the male; tibia as long as the femur, the heels
meeting when the legs are folded at right angles to
the rhachis. Foot as long as or a little shorter than
the tibia ; toes short, depressed, obtusely pointed, one-
third to one-half webbed, and bordered; no subarticular
tubercles ; a small, rounded, inner metatarsal tubercle.

Upper parts covered with small smooth warts,
limbs nearly smooth, with a glandular thickening on the
forearm and another on the calf; a small, more or less
distinct parotoid gland behind the eye above the tym-
panum, usually followed by a series of smaller glands

ALYTES. 165

along each side of the body ; a more or less distinct
round gland behind the angle of the mouth. Throat,
belly, and lower surface of thighs, or belly and pubic
region granulate ; a strong gular fold separating the
throat from the breast.

Greyish or pale brown above, speckled with darker
or with small greyish-olive or greenish spots, some-
times with red or reddish centres ; an ill-defined
dark canthal streak sometimes present ; a light
band across the anterior half of the interocular
space, and a triangular, cordiform, or A-shaped
light marking between the shoulders are often dis-
tinguishable, and between these two light markings a
dark triangle or X ; the glands of the lateral series
often orange or red. Lower parts dirty white or
greyish, with the granules of a pure white; limbs
carneous ; throat and breast often speckled with grey,
especially on the sides. Iris pale golden, rarely
silvery, veined with black.

Male difficult to distinguish from the female. The
body is, however, somewhat shorter, and the fore
limbs are a little stronger.

Geographical Variations. — This species varies very
little in France, Germany, and Switzerland. But the
specimens from the Spanish Peninsula show less
uniformity; the more divergent may be distinguished
at a glance, and have been named var. hoscae,
Lataste. The skin is usually smoother, the parotoid
glands are smaller, and the dorsal spots are generally
larger and better defined, sometimes forming mar-
blings ; these characters combine to produce a physi-
ognomy somewhat suggestive of a young Pelobates
cultripes. I am, however, unable to confirm the abso-
lute constancy of the various structural differences by
which this variety is to be distinguished from the
typical form ; measurements of the skull and vertebral
column of some Rhenish and Spanish specimens show
me that the proportions may, exceptionally, be the
same.

166

DISCOGLOSSID^.

Measurements (in millimetres),

3

1.

2.

3.

4.

5.

6.

From snout to vent

. 38 ..

. 46 ..

. 41 ..

. 43 .

. 51

... 43

Length of head

. 14 ..

. 16 ..

. 15 .

. 15 ..

. 15

.. 15

Width of head

. 16 ..

. 18 ..

. 17 ..

. 17 ..

. 19

. 17

Diameter of eye

. 4-5 ..

. 5 ..

. 4 5 .

. 4 5.

. 5

.. 45

Interorbital width .

. 3 ..

. 4 ..

. 35 .

. 3-5 .

. 4

.. 4

From eye to nostril

. 4 .

. 4 5..

. 4 .

. 4 .

. 45

... 4

„ ,, end of snout

. 6-5 .

. 8 ..

4

7

. 8

i

Diameter of tympanum

o
O

. 35 .

. 3 .

. 3 .

. 4.

... 3

Fore limb

. 24 ..

. 26 ..

. 25 .

. 24 .

. 25

.. 23

Hind limb .

. 52 .

. 59 .

. 54 .

. 52 .

. 5s

.. 52

Tibia

. 15 .

. 17 ..

. 16 .

. 16 .

. 17

... 16

Foot

. 15 .

. 10 ..

. 15 .

. 15 .

. 16

.. 14

1. Paris : Lataste.

2. French Jura : Ling Roth.

3. Hameln, Weser ; Wolterstorff.

4, 6. Valencia, Spain : Bosca
(var. boscai).
5. Paris: Heron-Royer.

According to Fatio the female reaches a length of 54 mm.

Skeleton. — Ethmoid short, embracing above a large
fontanelle which extends nearly to the line of the
anterior borders of the orbits ; fronto-parietals very
narrow, in contact with each other only quite poste-
riori}*, with an inward process in the middle which
gives the fontanelle the shape of the sole of a shoe ;
ethmoid rounded or somewhat produced anteriorly,
exposed between the posteriorly diverging inner
borders of the nasals, which are large and in contact
with each other in front or along the greater part of
their length ; a cartilaginous supra-orbital plate is
articulated to the nasal and the anterior third of the
fronto-parietal. Zygomatic branch of the squamosal
very short. Vomers large, narrowly separated from
each other on the median line, their posterior toothed
borders partly covering the feeble palatines ; pterygoid
regularly triradiate, the anterior branch joining the
maxillary and the slender palatine ; parasphenoid
X-shaped, obtuse or truncated in front, not quite
reaching the vomers, and separated from the ptery-
goids. Mento-Meckelians very indistinct.

Hyoid a large, broad, cartilaginous plate with
rounded lateral wings, small postero-lateral processes,
and long cornua ; a pair of diverging, ossified stylo-

ALYTES.

107

hyals, apart at the base, in front of which a slender V-
or Y-shaped ossicle is developed on the ventral side.

Vertebral column twice or a little more or a little
less than twice as long as the skull. Spine closed

Fig. 62.

Skeleton of male.

above, the neural arch without or with a very short
postero-median process. Second, third, and fourth
vertebrae with short diapophyses, with short auto-
genous ribs. The first rib the shortest and directed

168

DISCOGLOSSIDJE.

a little forwards ; the second the longest, nearly hori-
zontal ; the third intermediate in length between the
other two, and horizontal or directed slightly back-
wards ; the fourth or fourth and fifth diapophyses
nearly horizontal, the sixth and seventh, sometimes
also the fifth, directed forwards. Diapophyses of the
sacral vertebra triangular, their distal diameter equal-
ling or slightly exceeding their length ; two condyles
for articulation with the urostyle. Latter nearly as
long as or a little shorter than the eight anterior
vertebra? taken together, with a rather strong, curved
or posteriorly hooked transverse process at the base.

Prascoracoids slender, curved, meeting at an acute
angle, entering the glenoid cavity ; coracoids stronger,
nearly straight; supra-scapula ossified; omosternum
absent; cartilaginous sternum with two feebly calci-
fied posteriorly diverging styles. Carpus with eight
elements, two of which are in contact with radius-
ulna ; a single bone in the pollex.

Pelvis three-fifths to two-thirds the length of the
vertebral column; pubis cartilaginous; acetabulum
open. Femur and tibia equal in length, the former
strongly curved; astragalus nearly two-thirds the
length of the tibia ; three small cartilaginous tarsalia ;
one cartilaginous element to the prsehallux. Distal
phalanges obtuse, slightly expanded at the apex.

Measurements of Skkleton (in mil

F. typica.

<?

¥

Length of sk

all

13 .

. 14

Width of sku

11

10 .

. 17

Least interoi

bital w

dth

4 .

4

Dorsal vertebral column

13 .

. 15

Urostyle

12 .

. 12

Humerus

.

10 .

. 10

Radius-ulna

7 .

rr
(

Manus

12 .

. 11

Pelvis

.

10 .

. IS

Femur

15 .

. 15

Tibia

15 .

. 15

Tarsus

.

10 .

. 10

Pes

.

17 .

. 10

i met res).

Var.

boscte.

T~

~?

12

... 13

14

... 10

35

... 35

11

... 12

11

... 13

9

... 10

0

t

10

... 11

16

... 18

13

... 15

13

... 15

8

... 10

13

... 15

ALTTES. 169

Habits. — The life-history of this Batrachian is one
of great interest, but difficult to observe owing to its
nocturnal habits and shy disposition. First dis-
covered in the act of parturition by Demours, in the
middle of the last century, on the border of a small
pond in the Jardin des Plantes, we have bad to wait
until the year 1870 for a truly scientific account,
based on exact observations, of the mode of oviposition
of the so-called " crapaud accoucheur de sa femelle."

This we owe to that most excellent and patient
naturalist, Arthur de l'lsle du Dreneuf, who, residing
near Nantes, in a locality where the species is excep-
tionally abundant, spent in three consecutive years
more than fifty nights in the open in order to unravel
the mystery, and whose notes taken on the spot cover
250 foolscap pages of small handwriting. The account
given bereafter of the pairing and oviposition is com-
piled from that source. Although I have stayed at
places where Ahjtcs is not uncommon, and have made
nocturnal excursions with a lantern in order to
witness the operation, I have, in common with most
herpetologists, hitherto failed in my object.

Ahjtes is nocturnal and slow in its movements ; it
progresses mostly crawling, but sometimes by short
leaps, even when embarrassed with the eggs. It is
able to burrow chiefly by means of the fore limbs, but
usually selects for its retreat holes made by small
mammals, or interstices between stones. Turning
over large stones in the vicinity of the water wdiere
its tadpole attracts notice, is the surest means of
securing specimens in the daytime. Towards evening
it reveals its presence by a clear whistling note, which
has often been compared to the sound of a little bell,
or to a chime when produced by numerous individuals.

The breeding season lasts throughout spring and
summer, and the female is able to spawn two, three,
or even four times in the year. The season seems,
however, at its height in May and June in the plain ;
later, of course, in the mountains.

170 . DISCOGLOSSIDJl.

Pairing and oviposition take place on land. The
male seizes the female round the waist; but during
impregnation, or when he proceeds to lade himself"
with the ova, he clasps her round the head in front of
the fore limbs. Males are more numerous, and often
dispute about the females. When he has secured un-
disturbed possession the male strongly bends the body
so as to bring his heels close to the cloacal region of
the female, which he proceeds to lubrify by rapid move-
ments of the inner toes, at times introduced into the
cloaca, the two feet working alternately. This sort of
raking consists of 1100 to 1300 strokes for the two
feet, divided into 15 to 21 periods of 45 to 103 strokes,
with intervals of two or three minutes' rest, and lasts
about twenty-five minutes. Then the movements sud-
denly stop, the female extends her hind limbs, tightly
embracing those of the male, which are bent at angles
at the knees, the tarsi erect and pressed close together ;
the eggs are suddenly expelled, with noise, as if by ex-
plosion, and fall into the rhomboidal receptacle formed
between the pelvic limbs and bounded behind and
beneath by the tarsi and feet of the male. The yellow
eggs, as if threaded together by elastic filaments, form
a large mass, two to four layers of about ten, in this
receptacle.

The very moment the eggs are emitted the male
unclasps the waist of the female and shifts his hold to
the base of the head ; the body then stretched out,
but the legs remaining in the same position as before,
fecundation commences; it takes place in two or three
emissions at short intervals. The seminal fluid is
much diluted with the contents of the urinary bladder,
which soaks the envelops of the eggs. An interval
of ten to fifteen minutes' rest follows before the male
proceeds to attach the eggs to his legs. This is done
in the following manner : — Still holding his mate
round the head, he draws out the legs so as to stretch
the elastic connections between the egg's sticking to
his tarsi ; then, folding one of the limbs, bringing the

MATES.

171

knee to the level of the sacrum, and stretching it out
again, he passes it, toes first, into the egg-mass; the
other limb follows in like manner. These movements
are repeated several times, and accompanied by one
or two seminal ejaculations. Thus the strings of eggs
come to be fastened round the legs, where they will
remain until eclosion. All the time this operation has
lasted, about ten minutes, the female has remained
motionless, still connected with the eggs by the last
elastic filaments from the oviducts. These two threads
stretch to a length of 4 to 12 inches before breaking
when the pair separates.

Thus laden, and yet so little impeded in his
movements as to occasionally resort again to hymen
during the nursing period, and successfully add
on a second burden, the male retires to his usual
retreat, Q-o'mg- about at night in order to feed him-
self and to keep up the moisture of the eggs, even
resorting* to a short immersion in the water during
exceptionally dry nights. The development within
the egcr takes about three weeks, sometimes a little
less, often a little more. The male enters the water
with his burden ; the larva?, in the full tadpole con-
dition, measuring 14 to 1 7 mm., bite their way through
the tough envelop, which is not abandoned by the
father until all the young are liberated.

The tadpole is found in spring and summer, at all
stages of development, in small reservoirs, cow-ponds,
flooded quarries, pits in brick-fields, &c. The tadpoles
of the late broods hibernate under the ice, concealed
in recesses but not torpid. Some specimens, at least
in confinement, remain nearly two years before trans-
forming, but, as a rule, larva? born early in the spring
accomplish their metamorphosis within three to five
months. The duration of the larval life varies, how-
ever, in individuals of the same brood; and as such as
have not sufficiently developed their limbs to take to
the land in the autumn have necessarily to post-
pone the metamorphosis until the following spring, it

172 DISCOGLOSSlDtf!.

follows that whereas some may have spent only three
or four months in the water, their brothers will
sojourn for a whole year or more in that element.
Young reared by me measured, immediately after meta-
morphosis, from 18 to 22 mm. from snout to vent.

Eggs. — Large, the vitellus measuring 3J to 5 mm. in

diameter, varying in colour from pale straw-yellow to

bright yellow. They are coated with

Fig. 63. two transparent gelatinous envelops,

the outer of which is tough and elastic,

and forms the threads by which the eggs

are connected in two rosary-like series

as they issue from the cloaca. These

connecting threads measure, without

great tension, from 4 to 7 mm., and

according to the number of eggs, which

varies between 18 and 54, the whole

rosary has a length of 70 cm. to 2 m. Males observed

by Geisenheyner and Melsheimer with 126 and 150

eggs were no doubt nursing double or treble broods.

When fresh laid the eggs are nearly spherical,
but they soon acquire a more transversely oval
shape. Through the transparent capsules the
whole development can be easily followed. An
enormously large vitelline sac is present, and the em-
bryo develops uncommonly long, unpigmented gills,
one only on each side, with a large number of slender
branches along the ventral side of the trunk. These
gills are absorbed and replaced by internal ones, and
the transformations which accompany the passage from
the first or embryonic to the second or tadpole period
are all effected within the egg-capsules. The embryos
are at first uniform yellowish-white ; the pigment,
when it appears, forms two brown stripes, but before
hatching the little tadpole has put on his grey,
more or less spotted coloration.

Tadpole (PI. I, figs. 4 and 5). — Length of body once
and one-third to once and a half its width, two-fifths
to one-half the length of the tail. Nostrils nearly

ALYTfiS. 173

halfway between the end of the snout and the eyes.
Eyes on the upper surface of the body, the distance
between them about twice as great as that between
the nostrils, and equal to or slightly greater than the
width of the mouth. Spiraculum in the mid-ventral
line, a little nearer the anterior than the posterior
extremity of the body. Anal opening median, very
much larger than the spiraculum. Tail twice and
two-thirds to thrice as long as deep, ending in an
obtuse point ; the upper crest convex, usually a little
deeper than the lower, and extending but very slightly
upon the back; the depth of the muscular portion, at
its base, about half the total depth.

Beak white, with a broad black margin. Lip
entirely surrounded by a series of papillae. Labial
teeth in § series, occupying nearly the whole width of
the inner surface of the lip, all continuous, or the third
lower narrowly broken up in the middle ; the first
upper and the first lower series composed of one or
two rows of teeth, the others of two or three.

Lines of crypts usually very indistinct; all that can
be distinguished being the usual lines from the end of
the snout between the nostrils, bordering the eyes
above, behind and below, and forming a loop on each
side of the upper lip, a line beginning at a considerable
distance behind the eye along each side of the back to
the upper border of the muscular part of the tail, and
another very short line close to and parallel with the
anterior extremity of the latter. But in a fine
specimen from Bellaigues, Switzerland, I find the
lines much more distinct and blackish ; in addition to
the series described above, it shows the second dorsal
line prolonged to the base of the tail, which also
bears two lines, the upper being on the supra-caudal
crest ; a short series descends vertically from below
the centre of the eye, another, curved, from below the
anterior extremity of the dorsal lines, a third on each
side of the mouth, and a fourth extends on each side
of the belly, from the level of the spiraculum nearly

174 DISCOGLOSSID,*:.

to the origin of the hind limbs, its anterior extremity
bent downwards and forwards. This specimen is
figured above (p. 104, Fig. 46, c).

Lead-grey to blackish above, uniform or with round
blackish spots ; sides with large silvery or pale golden
spots ; tail with numerous dark brown dots or round
black spots, which are very apparent on the greyish-
white crests. Nearly black tadpoles as well as albinos
are on record.

Total length 80 mm.; body, 28; width of body,
21; tail, 52 ; depth of tail, 19. Exceptionally grows
to 90 mm.

Habitat. — The midwife toad is common in France
nearly everywhere ; it is also found in Belgium (pro-
vinces Namur, Liege, and Luxemburg), South-eastern
Holland (Limburg), Luxemburg, Switzerland, Vorarl-
berg, in Germany along the Rhine, and locally in hilly
districts as far east as Brunswick and Thuriugia
(Hameln on the Weser being the northernmost, Nord-
hausen and Eisenach the easternmost points of its
ascertained distribution), and all over Spain and
Portugal. It ascends to 5000 feet in the Alps of
Switzerland, and to 6500 feet in the Pyrenees (Lac
Bleu, Hautes-Pyrenees), where the snow is not
absent for more than three months.

Specimens have been introduced by Knauthe near
Schlaupitz, in Silesia, some years ago, and appear
to have established themselves there. The species is
stated to occur in Bukowina, whence it was described
by Zawadski in 1840. It would be highly interesting
to ascertain whether it does really extend so far to the
east ; for the present the statement can only be
accepted with caution.

In France the midwife toad is found in the plains
as well as in the mountains ; common at Biarritz, close
to the sea, it extends high up the Pyrenees. I have
also found it on the coast of Normandy, and, as Prof.
Bavay informs me, it occurs on the coast of Finistere.
East of the Rhine, however, its habitat, like that of

ALYTES.

175

Bombinator pachypas, becomes restricted to the hills.
It by no means avoids the dwellings of man, and may
be found in great numbers in old walls of villages as
well as in gardens and waste grounds of towns. It is
still to be met with in the Jardin des Plantes, where it
was first discovered in the middle of the last century.

Four specimens are figured on PI. VII, viz. a male
from Paris, laden with eggs ; a female from Liege, on
the left hand ; a young albino from Geuevilliers near
Paris, reared by M. Lataste ; and on the right hand a
specimen from Valencia, Spain (var. hoscx).

5. Alytes oisternasii.
(Plate VIII.)

Alytes cisternasi, Boscji, An. Sue. Esp., viii, 1879, p. 217; Bou-
lenger, Cat. Batr. Ecaud., p. 449 (1882); Heron-Royer &
Van Bambeke, Arch. Biol., ix, 1889, p. 289; Bedriaga, Bull.
Soc. Nat. Mosc, 1889, p. 617, Ampb. Rept. Portug., p. 27
(1889), and Larves Batr. Portug., p. 1-41 (1891); Boulenger,
Proc. Zool. Soc , 1891, p. 624, pi. xlvii, fig. 8.

Ammoryctis cisternasi, Lataste, C. R. Ac. Sc, lxxxviii, 1879, p.
983; Bosca, Bull. Soc. Zool. France, 1880, p. 252, and An.
Soc. Esp., x, 1881, pi. ii, figs. 1 — 6.

Vomerine teeth in two transverse series, widely sepa-
rated from each other in the middle, behind the choanse ;

these series often slightly oblique,
forming together an angle pointing
forwards. Tongue large, circular,
slightly free behind.

Head large, moderately depressed,

Fig. 64.

much broader than long ; snout round-
ed, projecting, shorter than the dia-
meter of the orbit; canthus rostralis
rounded, loreal region grooved ; nos-
tril nearer the tip of the snout than
the eye ; eye large, very prominent ;
interorbital space broader than the upper eyelid, as

Open mouth.

176 discoglossidj;.

broad as the distance between the nostrils ; tympanum
three-fifths to two-thirds the diameter of the eye.

Fingers rather short, depressed, obtuse, first shorter
than second, third a little longer than second, fourth
shortest ; fourth finger very thick, stump-
like, brown and horny at the end ; no sub-
articular tubercles : two prominent palmar
tubercles, inner small and oval, outer very
large, oval, oblique. Brachiununearly en-
tirely embedded in the skin of the body.

Hind limb short ; the tibio-tarsal articula-
tion reaches the axil or the shoulder ; the Lower view
tarso-metatarsal articulation reaches the tym- tand
panum or the eye ; tibia as long as the femur,
the heels meetiug when the limbs are folded. Foot
as long as or a little shorter than the tibia ; toes
short, depressed, obtusely pointed, one-third webbed ;
no subarticular tubercles ; a small, oval inner meta-
tarsal tubercle.

Upper parts covered with very small smooth warts ;
snout smooth ; a series of enlarged warts along the
upper eyelid ; a very small parotoid gland. Lower
parts coarsely granulate; a strong gular fold.

Greyish or brownish above, with small dark spots ;
a light cross-bar between the eyes, and three light
spots on a line between the shoulders ; the enlarged
warts on the eyelids whitish or orange ; some of the
warts on the sides also whitish or orange. Some
specimens have the markings better defined, and a
light triangular or cordiform spot between the
shoulders is accompanied by an oblique, curved
light band on each side of the back, beginning
from the supra-temporal fold. These markings,
which may also be observed, though less defined, in
the preceding species, are highly suggestive of those
of certain specimens of Discoglossus pictus, and afford
further evidence of the close relationship, so long
overlooked, which exists between the two genera.
Lower parts whitish. Iris golden.

ALYTES.

177

Measurements (in millimetres).

1.

2.

3.

&

?

?

From snout to vent .

38

35

.. 40

Length of head

12

11

.. 12

Width of head

16

14

.. 17

Diameter of eye

4

4

.. 4-5

Interorbital width

4

4

.. 4-5

From eye to nostril .

3-5 ..

3

.. 35

„ ,, end of snout

6

5-5 .

.. 6

Diameter of tympanum

2-5 ..

2-5 .

.. 3

Fore limb

20

19

.. 19

Hind limb

45

42

.. 45

Tibia .

13

12

.. 14

Foot .

13

125 .

.. 12-5

1, 2, Merida, Spain : Bosca. 3. Badajos, Spain : Bosca.

Skeleton. — Apart from the stouter and shorter
limb-bones, the skeleton of this species differs from
that of the preceding in the larger nasals, which over-
lap the ethmoid and the inner borders of which meet
along: their entire length, in the constricted fronto-
parietals, and in the shape of the frontoparietal
fontanelle, which is divided into two portions between
which the fronto-parietals form a bridge ; the anterior
fontanelle is large and kite-shaped, the posterior very
small, a mere foramen between the fronto-parietals.

Fig. 66.

Upper view of skull and lower view of left forearm and hand.

The coracoids and prgecoracoids are more flattened
and more robust, and the metacarpal and phalanges
of the outer finger are shorter and more massive ; the
terminal one in particular being extremely short,
nearly hemispherical. The femur is slightly longer
than the tibia.

The more strongly ossified cranium is correlated

M

178

DISCOGLOSSID.E.

with the burrowing habits of this
differences noticed between this
obstetricans are merely of degree.

species,
species

and the
and A.

Measurements op Skeleton (in millimetres).

Length of skull .

12

Width of skull .

16

Least interorbital width

3

Dorsal vertebral column

12

Urostyle .

10

Humerus .

9

Radius-ulna

6-5

Manus

8-5

Pelvis

16

Femur

12

Tibia

11

Tarsus

7-5

Pes

12

Habits. — This species is a burrower in sandy locali-
ties, digging by means of the outer edge of the hands.
In accordance with these habits its lungs are more
developed than in the allied species. Bosca having
found the male carrying the eggs, in April, there is no
doubt the breeding habits are much the same as in
A. obstetricans, although the pairing has not yet
been actually observed. The tadpole is to be found
all the year round.

Tadpole (PL I, fig. 6). — I am not able to detect any
character by which this tadpole may be surely distin-
guished from that of A. obstetricans. The spots on the
tail are, however, smaller, more crowded, and forming
vermiculations on the muscular portion, the space oc-
cupied by the lateral groove being usually free from
spots.

Length of body, 20 mm.; width of body, 15 mm.;
tail, 42 mm. ; depth of tail, 14.

Habitat. — Spain (Aragon, New Castille, Estrema-
dura) and Portugal (Douro, Beira, and Alemtejo).

The specimen figured on PI. VIII is a female, one of
the types from Spain, preserved in M. Lataste's col-
lection.

[At j>aye 179.

PELOBATES FUSCUS.

„ CULTRIPES.

DISTRIBUTION OF EUROPEAN PELOBA.TIDJ3.

PELODYTES. 179

Family 2.— PELOBATIILE.

Vertebras procoelous (in the European genera), with-
out autogenous ribs ; diapophyses of sacral vertebra
very strongly dilated. Upper jaw toothed.

The PelobafidcB occupy an intermediate position
between the Discoglossidce and the Bufonidse. Seven
genera are known, inhabiting Europe and South-
western Asia, South-eastern Asia and New Guinea, and
Xorth America and Mexico.*

The two European genera are easily distinguished : —
Pelodytes, slender and frog-like, with deeply cleft toes,
and a more or less distinct tympanum ; Pelobates, stout
and toad-like, with broadly webbed toes, no tympanum,
the skin of the head adherent to the skull, and a large,
sharp-edged, shovel-shaped tarsal tubercle.

Both genera must be looked upon as highly special-
ised forms of a group which has, in all probability,
sprung up from the Discoglossidse, with which some
of the exotic o-enera are nearer connected through

«... O

a character to which great importance has been
attached, viz. the opisthoccelous vertebras.

A map is here given showing the range of the three
European species of this family.

4. Pelodytes.

Fitzinger, in Bonaparte, Icon. Faun. Ital., Rett. Ant'. (1838).

Pupil vertical. Vomerine teeth in two small groups
between the choanas. Tongue subcircular, entire or
slightly nicked, and free behind. Tympanum present,

* Exotic genera :

Scaphiopus, Holbr. E. and S. United States, Mexico.

Batrachopsis, Blgr. New Guinea.

Leptobrachium, Tsch. S. China, Further India and Malay

Archipelago.
Megalophrys, Kuhl. Malay Peninsula and Archipelago.
Asteroplnjs, Tsch. New Guinea.
The last two have opisthoccelous vertebrae.

180 PELOBATIDJL

more or less distinct or bidden under the skin.
Fingers free, toes webbed at tbe base and more or
less distinctly bordered ; outer metatarsals separated
by web. Proximal tarsal bones fused. Vertebras pro-
ccelous ; diapophyses of sacral vertebra very strongly
dilated ; two condyles for articulation with urostyle.
Omosternum cartilaginous; sternum with a bony style.

This genus includes two species : — P. punctatus,
Fitz., of Western Continental Europe, and P. caucasi-
cus, Blgr., recently discovered on the Asiatic side of
the Caucasus at an altitude of 7000 feet. The latter
has been made the type of a distinct genus, Pelody-
topsis, by Nikolski, on grounds too trivial to deserve
recognition.

Perhaps this Pelodytes caucasicus may yet be found
in Cis-Caucasia, and vield a further addition to the
European fauna. It is distinguished from its con-
gener in the longer hind limbs, the tibio-tarsal articu-
lation reaching the tip of the snout, and the somewhat
more posterior position of the groups of vomerine
teeth ; the toes are not so strongly fringed in the
male, the fore limbs are more robust, and the black
horny nuptial excrescences are much more strongly
developed on the warts and ridges of the upper sur-
faces.

6. Pelodytes punctatus.
(Plate VIII.)

Rana punctata, Daudin, Hist. Rain. Gren. Crap., p. 51, pi. xvi,
fig. 1 (1803), and Hist. Rept., viii, p. 100 (1803).

Rana plicata, Daudin, 11. cc, pp. 53, 102.

Rana daudinii, Merrem, Tent. Syst. Ainph., p. 177 (1820).

Bombinator plicatus, Fitzinger, N. Class. Rept., p. 65 (1826).

Obstetricans punctatus, Duges, Rech. Osteol. Batr.. p. 7 (1834).

Alytes punctatus, Tscliudi, Class. Batr., p. 84 (1838).

Pelodytes punctatus (Fitzinger), Bonaparte, Faun. Ital., Rett.
Anf. (1838) ; Dumeril & Bibron, Erp. Gen., viii, p. 463 (1841) ;
Thomas, Ann. Sc. Nat. (4), i, 1854, p. 290; Giinther, Cat.
Batr. Sal., p. 35 (1858) ; Fatio, Vert. Suisse, iii, p. 353 (1872) ;
De Betta, Faun. Ital., Rett. Anf., p. 68 (1874); Scbreiber,
Herp. Eur., p. 99 (1875) ; Lataste, Herp. Gironde, p. 242

J

V. mi.

"*?

m >-.

jf/r-

"■♦

w®

■ /////,-. j ,-/■)/,///?/■>/?.

'y;c ^

A

// // C /ff//f./

PELODYTES.

181

Fig. 67.

(1876); Heron-Royer, Bull. Soc. Zool. France, 1878, pp. 128
& 299, pi. iii, and 1879, p. 229, pis. x and xi; Boulenger,
Bull. Soc. Zool. France, 1880, p. 225, and Cat. Batr. Ecaud.,
p. 438 (1882); Peracca, Boll. Mus. Torin., i, 1886, No. 1;
Ht'ron-Royer, Bull. Soc. Et. Sc. Angers, xv, 1886, p. 91 ;
Heron-Royer & Van Bambeke, Arch. Biol., ix, 1889, p. 277,
pi. xx, figs. 5— 12; Bedriaga, Bull. Soc. Nat. Mosc, 1889,
p. 533. and Aniph. Rept. Portug., p. 20 (1890) ; Boulenger,
Proc. Zool. Soc, 1891, p. 617, pi. xlvii, figs. 1 & 2; Bedriaga,
Ampb. Rept. Portug., Suppl., p. 11 (1893); Martin &
Rollinat, Vert. Dep. Indre, p. 349 (1894).
Pelodytes daudini, Bosca, Bull. Soc. Zool. France, 1880, p. 255.

Vomerine teetli in two short transverse or slightly
oblique series on a line with the anterior borders of
the choauae ; these series either close
top-ether or nearer the choanaa than
each other. Tongue large, circular,
entire or feebly nicked behind.

Head, much depressed, as long as
broad or slightly broader than long ;
snout rounded, projecting beyond the
mouth, as long as the diameter of the
orbit; canthus rostralis rounded; loreal
region grooved; nostril midway between
the eye and the tip of the snout or a little
nearer the latter ; eye large, prominent ; interorbital
space as broad as or a little narrower than the upper
eyelid, and equal to the distance between the nostrils;
tympanum usually moderately distinct, sometimes
hidden, one-half to three-fifths the diameter of the eye.

Fingers rather elongate, somewhat swollen at the
end, third much the lougest, first a little shorter than
second, which is a little shorter than fourth ; basal
subarticular tubercles distinct; three palmar tubercles,
median smallest and circular.

Hind limb rather slender ; the tibio-tarsal articu-
lation reaches the eye or between the eye and the
nostril ; tibia a little longer than femur, the heels
overlapping when the legs are folded at right angles
to the rhachis. Foot as long as or a little longer
than the tibia; toes slender, webbed at the base, the
web extending as a fringe along each side to the

Open mouth.

182 PELOBATIDiE.

tips, which are slightly swollen; subarticular tubercles
feeble or indistinct; no tarsal fold; a small, rounded
inner metatarsal tubercle.

Upper surfaces covered with more or less prominent
porous warts of unequal size; the larger warts on the
back sometimes disposed in wavy longitudinal series ;
a strong glandular fold from the eye to above the
shoulder, passing above the tympanum, and sometimes
followed along each side of the body by a chain of
large warts. Lower surfaces smooth except the pos-
terior part of the belly and the thighs, which are
coarsely granulate.

Greyish or pale greyish -olive above, with small irre-
gular dark olive, bottle-green , or bright green spots
(whence the name "Persille" by which this Batrachian
is known in some parts of France), which may be
confluent into more or less regular cross-bars on the
limbs; between the eyes two oblique streaks of the
dark colour, converging backwards, are usually dis-
tinct ; a dark streak extends from the tip of the snout
to the eye, involving the nostril, and often reappears
behind the eye along the supra-temporal fold ; two or
three dark vertical bars, sometimes broken up into
spots, on the upper lip ; the larger warts on the sides
sometimes orange or rusty red ; a light X-shaped
marking on the ground colour is usually more or less
distinction the anterior part of the body, the anterior
branches extending to the upper eyelids, the posterior
to the sacral region. Lower parts white, often
yellowish on the limbs and rosy about the inguinal
region. Iris golden, more or less pigmented with
brown on the lower half, or bronzy-brown with a fine
pale golden border to the pupil.

Male distinguished from the female by a shorter
body, longer and more robust fore limbs, a greater
development of the dermal borders of the toes, and
the presence of an internal vocal sac which communi-
cates with the mouth through a long slit on each side
of the tongue ; owing to the pigmentation of the vocal

PELODYTES.

183

Fig. 68.

sac the throat appears pale purplish-grey. During
the breeding season horny excrescences are much de-
veloped on the lower sur-
faces, and similar pro-
ductions of small size are
sometimes also scattered
on the upper surfaces, the
largest crowning the prin-
cipal warts ; no trace of
them being found nor-
mally in the females. These
excrescences, which are of
a dark purplish-brown or
black colour, are disposed
as follows on the lower sur-
faces : — A rounded group
on each side of the breast,
at the base of the arm ;
another on the arm; a band
along1 the inner side of
the fore- arm ; and a band
along the inner side of the
two inner fingers ; there
are besides, very often, a
points on the abdominal

Lower

band on
granules,

the chin, isolated
and two linear

series along the toes, interrupted at the articulations.

Measurements (in millimetres).

From snout to vent .
Length of head
Width of head ■
Diameter of eye
Interorbital width .
From eye to nostril .

„ „ end of snout
Tympanum .
Fore limb
Hind limb
Tibia
Foot

1.

38

12

12
35
2-5
3-5
6
2

24
59
18
20

2.

41
13
15

4

3

3-5

/

2
29
68
20

3.

40

14

14
4

3-5
35

4.

40
13
13

4

o

o

5.

43
15

16
4
3
35

/

. b .

/

2-5..

2

. 25

26 ..

22

29

63 ..

'. 60 '.

. 71

19 ..

. 18 .

. 21

20 ..

. 18 .

. 23

1, 4. Paris : Heron-Royer.

2, 5. Nice : Bedriaga.

3,6. Valencia: Lilford

6.

44
14
14

4

3

3-5

7
o

25
61
18
1°

184 PELOBATID.E.

Skeleton. — The skull, so far as the shape and the
extent of ossification are concerned, is intermediate
between that of Discoglossus and that of Bombinator,
agreeing with the latter in the absence of palatine
bones. Ethmoid short, not extending posteriorly
beyond the anterior two fifths of the basisphenoid ; the
lower anterior lamina rounded or truncate, and extend-
ing nearly to the posterior border of the vomers;
the upper obtusely pointed and extending to between
the nasals. The large chondrocranial fontanelle is
only partially covered by the fronto-parietals, these
bones being in contact with each other to but a small
extent either behind or in the middle. Nasals mode-
rately large, tear-shaped, not meeting on the median
line. Zygomatic branch of the squamosal short.
Vomers moderately large, narrowly separated from
each other on the median line ; {pterygoids rather
slender, the anterior branch longest and joining
the maxillary; parasphenoid dagger-shaped, sharply
pointed in front, and extending forwards to the line
of the anterior borders of the orbits, separated from
the pterygoids. Mento-Meckelian bones distinguish-
able only from the inner side of the mandible.

Hyoid a large cartilaginous plate with slender
postero-lateral processes ; anterior processes much
expanded, turned inwards and closely approximating
in front, and confluent with the lateral wings, enclos-
ing a small fenestra ; thyrohyals slender, diverging,
narrowly separated from each other at the base ; a
pair of slender ossifications in front of the thyroidals
on the ventral side, distinct and L-shaped or united
and H-shaped. Ceratohyal cornua with the posterior
portion detached from the hyoid plate, the anterior
portion being included in the hyoid plate.

Vertebral column twice or a little less than twice as
long as the skull. Spine closed above ; neural arch
without or with a very short postero-median pro-
cess. First diapophysis obliquely directed forwards ;
second, third, and fourth horizontal ; second longest.

l'KLODYTES.

1 s:>

fourth small; fifth, sixth, and seventh small, slender
and directed forwards.

Fig. 69.

Skeleton of male.

Sacral vertebra with very strongly dilated diapo-
physis, the transverse diameter of which is nearly

186

PELOBATTD.E.

three times in the longitudinal, but little shorter than
the urostyle, which articulates by two sockets, and
measures three-fifths to two-thirds the length of the
remainder of the vertebral column.

Preecoracoids slender, strongly curved ; coracoids
nearly straight, dilated at both eDds ; a cartilaginous
omosternum ; sternum with a bony style dilated proxi-
mally, and a large discoid cartilage. Humerus con-
siderably longer than radius-ulna. Carpus with seven
elements, three of which are in contact with radius-
ulna ; a single bone in the pollex.

Pelvis three-quarters the length of the vertebral
column ; ilia with an inner groove, embracing the
sacral processes; the pubis does not ossify. Femur
and tibia with the epiphysis cartilaginous ; tibia a
little longer than the femur, nearly as long as the
pelvis ; calcaneum and astragalus completely fused
into a sino-le slender bone with a minute foramen in
the middle, resembling the tibia, of which it measures
three-fifths the length ; the elements of the distal row
of the tarsus are not ossified, and the prashallux
contains two cartilages. Distal phalanx of all digits
slightly expanded at the apex.

Measurements of Skeleton (in millimetres).

3

?

Length of skull

11

14

Width of skull

11

15

Least interorbital width .

o

o

Dorsal vertebral column .

12

17

Urostyle

7

11

Humerus

8-5 ..

11

Radius-ulna

6

8

Manus

9

12

Pelvis

15

20

Femur

14

17

Tibia

16

18

Tarsus

9

11

Pes

17

20

Habits.— This small and graceful Batrachian is essen-
tially nocturnal ; only during the breeding season is it
met with in the daytime. Its long limbs enable it

PELODYTES. 187

to jump in the manner of the true frogs, covering
1-4 or 1 5 inches at one leap, and it is besides a good
climber, ascending smooth vertical surfaces by adhesion
of the abdominal surface. Nor, in spite of its nearly
free toes, is it a bad swimmer, often frequenting deep
ponds during the pairing time. Its dermal secretion
bears a strong smell of garlic As often heard in
the evening, its voice is feeble, a sort of crecJc-creck-
crech, which cannot be better compared than to the
creaking of a leather shoe. But during the pairing
the male produces, under water, a more sonorous and
very different note — co-ack, to which the female re-
sponds by a feeble coo.

As a general rule, Pelodytes punctatus breeds, in
France, between the end of February and the begin-
ning of April ; but instances of its breeding in May,
July, August, September, October, and November
have been placed on record by A. Thomas, Lataste,
Bedriaga, and myself. Thomas, who first observed
the breeding habits of this Batrachian, near Nantes,
stated that it spawns twice in the year, in the
spring and in the autumn. According to Heron-
Royer, another French observer who devoted special
attention to the habits of the same animal, Pelo-
dytes breeds only once a year, in the spring.
Bedriaga has observed the spawning to take place in
the south of France in the spring and again in the
autumn. On the 30th of October, 1896, he was so kind
as to send me from Nice a large number of breeding
specimens, — among them, however, only two females,
one of which spawned on the way and the other
reached me with eggs in the oviducts. Having,
besides, myself found Pelodytes pairing on the north
coast of Brittany at the end of August, 1 have no
doubt Heron-Royer was wrong, and that the autumnal
hymen will also be found to take place near Paris,
where the species is so plentiful.

The mode of embrace, which falls in the lumbar
type, is peculiar. Owing to the thin waist of the

188

PELOBATIOE.

female the elbows of the male join on the pubic region,
and the forearms are carried forwards, pressed against
the middle line of the belly, the hands meeting without
intercrossing the fingers. Sometimes, however, the
embrace is less close, the elbows fitting into the groins,
and the forearms forming an angle on the belly. The
pairing is usually of short duration, a few hours only,

Fig. 70.

Lower view of male and female pairing.

and the male will, as a rule, let go if handled. Speci-
mens kept in confinement by me last winter paired in
January, and remained together for nearly three weeks.

Stagnant water of all sorts is resorted to for the
purpose of spawning, provided it be furnished with
vegetation, preferably Graminacea, round the stems
of which the bands of ova are twined.

The larvas of the spring broods transform in July
and August ; but late offspring of course hibernate
in the larval condition, and tadpoles may therefore be
found throughout the year.

Males are more frequent than females; out of 150
specimens collected by Lataste in the Gironde during
the spring of 1877, only 20 were female.

PELODTTES. 189

Eggs. — Small, measuring 1| to 2 mm. in diameter ;
blackish, with the lower pole white. Laid in two

strings which coalesce in the cloaca, FlG 71
thus forming: a single lengthened

band, often broken into several ■ L

pieces, twined round the stalks or ^&'ik'>>&&

small reeds or grass below the /^iJiSJ^^

surface of the water. The muci- $*!*>££*$*•
laginous capsule surrounding each <fj^*>jp*
egg measures 2\ to 3 mm. ; the
outer envelop, which in the toads
forms a sort of tube protecting the
eggs, is here absent; and when, as
sometimes happens, the band is
deposited in fragments, the egg-
masses embracing a stalk may
have more the appearance of little
balls, resembling, but for the darker
colour, those of Hyla arbor ea.

Heron-Royer has ascertained the number of ova in
a brood to vary between 1000 and 1600, and the
embryo to leave the mucilaginous envelop on the
fifth day, at a very low stage of development, tailless,
and having not yet developed the gills.

Tadpole (PI. I, figs. 7 and 8). — Length of body
rather more than once and a half its width, and not
quite two-thirds the length of the tail. Nostrils half-
way between the end of the snout and the eyes, or a
little nearer the latter. Eyes on the upper surface of
the body, equidistant from the end of the suout and
the spiraculum, the distance between them about twice
as great as that between the nostrils, and equal to the
width of the mouth. Spiraculum on the left side,
directed upwards and backwards, nearly equidistant
from either extremity of the body, visible from above
and from below. Anal opening median, much larger
than the opening of the spiraculum. Tail twice and a
half to three times as long as deep, ending in an
obtuse point ; the upper crest very convex, deeper

190 TELOBATlDii.

than the lower, and rarely extending forwards as far
as the level of the spiraculum ; the depth of the mus-
cular portion at its base one-third to two-fifths of the
greatest total depth.

Beak white, with a black margin. An inverted
fold at the side of the lip ; this is furnished with a
single row of papillae except on the upper border,
which is toothed. Labial teeth in \ or \ series, the
second and third, both above and below, the longest;
the first and second series in both divisions of the lip
uninterrupted, or the second upper with very slight
median interruption, the others separated in the
middle and gradually decreasing in length to the fifth,
which if present is short. According to Bedriaga,
there may be as many as six series of teeth on the
lower lip, the first three of which are uninterrupted.

Lines of crypts usually very apparent, but some-
times very indistinct. On the head they approach
each other between the nostrils and completely border
the eye posteriorly, the anterior extremities of this
naso-orbital hoop approaching each other above the
upper lip. Of the two dorsal lines, which diverge
posteriorly, the upper, extending to the dorsal edge of
the muscular portion of the tail, is interrupted at a
short distance behind the eye ; its anterior portion
may even descend to join the lower line, which thus
appears bifurcated in front; the lower line extends,
usually uninterrupted, from behind the eye to the
middle of the muscular portion of the tail, where it is
lost ; both lines, however, may stop short of the tail.
A sinuous line, on the flanks, curved above the spira-
culum, not bent upwards posteriorly, sometimes ex-
tending nearlv to the orio-in of the hind limbs. In
addition to these lines a short horizontal branch
originates above the upper lip, and bifurcating below
the vertical of the anterior border of the eye, forms a
hoop which descends to the sides of the throat.

Coloration usually varying from pale grey to olive-
brown above, the sides with pale metallic spots ; the

PELODYTES. 191

lines of crypts whitish ; caudal crests greyish, with
blackish spots and white dots, and pale metallic spots.
Some specimens, however, have the tail almost spot-
less; in others, on the contrary, it is very closely
spotted, but always less abundantly on the lower crest
than on the upper. Lower parts pale grey with silvery
spots. Tail and the greater part of the body with fine
black decussating lines. I have observed at St. Eno-
gat, in Brittany, a colony of abnormal tadpoles, not
exceeding 40 mm. in length when fully developed,
which were nearly black on the back, the tail dark
brown with the black decussating lines very crowded,
the belly of a beautiful steel-blue, and the lines of
crypts quite indistinct (see PI. I, fig. 8).

The largest tadpole obtained by me in Brittany
measures 57 mm., body 21, width of body 15, tail 36,
depth of tail 14. One from Nice measures 65 mm.

Habitat. — Pelodytes punctatus inhabits nearly the
whole of France, from the coasts of Brittany, Nor-
mandy, and the Pas-de-Calais to the foot of the
Pyrenees and the Mediterranean littoral. It has not
been found in the North-eastern Departments Nord,
Ardennes, and Lorraine, which seem to be outside the
eastern limit of its distribution. The only localities
east of Prance whence this species has as yet been
recorded are Castino in Southern Piedmont, and Loano
in Eastern Liguria. In the Pyrenean peninsula it is
on record from the Portuguese provinces Minho,
Douro, Alemtejo, and Algarve, and the Spanish pro-
vinces New Castille, Valencia, and Andalusia. In no
part of its habitat does this species extend to any
considerable altitude ; in France it avoids the central
plateau, and has not been found on the Pyrenees,
although inhabiting Spain. The highest recorded
altitude is 1450 feet at Castino, Piedmont.

The specimens, male and female, figured on PI. VIII,
are from Nice, received from Dr. J. de Bedriaga.

192 PELOBATIDiE.

5. Pelobates.

Wagler, Sjst. Amph., p. 206 (1830).

Pupil vertical. Vomerine teeth in short transverse
series between the choanse. Tongue circular, entire
or slightly nicked, and free behind. Tympanum
absent. Fingers free, toes webbed ; outer metatarsals
separated by web. Inner metatarsal tubercle shovel-
shaped. Vertebrae proccelous ; diapophyses of sacral
vertebra very strongly dilated ; urostyle usually fused
with sacrum. Omosternum cartilaginous ; sternum
with a bony style.

Three species of this essentially fossorial genus
are known : two from Europe, and one, P. syriacus,
Boettger, from Asia Minor and Syria.

The European species are easily distinguished from
each other. In P. fuscus the occiput is humped, the
orbit is not completely closed by bone behind, and the
metatarsal spur or shovel is yellowish or pale brown.
In P. cultrvpes the occiput is plane or gently arched,
the cranial roof completely surrounds the orbit, and
the metatarsal spur is black.

P. syriacus agrees with P. cultrvpes in the form of
the head and the structure of the skull, with P. fuscus
in the colour of the spur.

The genus Didocus, Cope (type Bana calcarata,
Michahelles), is founded on a young Pelobates cultripes,
and is therefore a synonym of Pelobates, as pointed
out further on in a foot-note. The adult Pelobates
cultripes has also been raised to generic rank by the
same author, but quite unnecessarily in my opinion,
the only distinctive feature being the greater develop-
ment of the cranial roof — a mere difference of degree,
the importance of which is outweighed by the close
resemblance which this species bears in all other re-
spects to its ally P. fuscus.

H

jm

X

PELOBATES. 193

7. Pelobates f use us.
(Plate IX.)

Rosel, Hist. Ran., p. 69, pis. xvii — six (1758).

Bufo fuscus, Laurenti, Syn. Rept., pp.28 & 122 (1768); Schneider,
Hist. Amph., i, p. 196 (1799); Daudin, Hist. Rain. Gren.
Crap., p. 81, pi. lxxx, fig. 1 (1803), and Hist. Rept., viii, p. 161
(1803) ; Duvernoy, Regne Aniin., Rept., pi. xxxviii, fig. 1
(1836).

Rana vespertina, Pallas, Reise Russ. R., i, p. 458 (1771).

Rana fusca, Meyer, Syn. Rept., p. 10 (1795) ; Sturm, Deutschl.
Faun., iii (1797); Gravenhorst, Delic. Mus. Vratisl., p. 32
(1829).

Bufo vespertinus, Schneider, Hist. Amph., i, p. 225 (1799).

Rana alliacea, Shaw, Gen. Zool., iii, p. 1-46, pis. xli & xlii (1802).

Bombina fusca, Koch, in Sturm, Deutschl. Faun., iii (1828).

Bombina marmorata, Koch, 1. c. ; Hahn, Faun. Boic, pi. xxi,
fig. b (1832).

Bombinator fuscus, Fitzinger, N. Class. Rept., p. 65 (1826).

Pelobates fuscus, Wagler, Syst. Amph., p. 342 (1830) ; Tschudi,
Class. Batr., p. 83 (1838); Duineril & Bibron, Erp. Gen.,
viii, p. 477 (1841); Steenstrup, Isis, 1841, p. 900; Nilsson,
Skand. Faun., Amf., p. 96 (1842) ; Giinther, Cat. Batr. Sal.,
p. 40 (1858) ; Bruch, Wiirzb. Naturh. Zeitschr., ii, 1861, p.
178. and iii, 1862, p. 181; Collin, Naturh. Tidsskr. (3), vi,
1869, p. 316 ; Fatio, Vert. Suisse, iii, p. 376 (1872) ; C. Koch,
Ber. Senck. Ges., 1872, p. 151 ; Cornalia, Atti Soc. Ital., xvi,
1873, p. 96, pis. ii & iii ; De Betta, Faun. Ital., Rett. Anf.,
p. 71 (1874) ; Schreiber, Herp. Eur., p. 90, fig. (1875) ; Ley-
dig, An. Batr., p. 77 (1877) ; Lessona, Atti Ac. Lincei,
Mem. Sc. Fis., i, 1877, p. 1077, pi. iii; Boulenger, Cat.
Batr. Ecaud., p. 437 (1882) ; Fatio, Vert. Suisse, iv, App.,
p. v (1882) ; Camerano, Mem. Ace. Torin. (2), xxxv, 1883, p.
217 ; Heron-Royer, Bull. Soc. Et. Sc. Angers, xv, 1886, p. 61 ;
Camerano, Boll. Mus. Torin., i, 1886, No. 9; Boulenger,
Bull. Soc. Zool. France, 1888, pp. 115 & 162; Peracca,
Boll. Mus. Torin., iii, 1888, No. 46; Wolterstorff, Zool. Anz.,

1888, p. 672; Heron-Royer & Van Bambeke, Arch. Biol., ix,

1889, p. 269 ; Bedriaga, Bull. Soc. Nat. Mosc, 1889, p. 493 ;
Boulenger, Proc. Zool. Soc, 1891, p. 614, pi. xlvi, fig. 7;
Zander, Corr.-Bl. Nat. Ver. Riga, No. 38, 1895, p. 62;
"Werner, Rept. Amph. Oesterr.-Ung., p. 104 (1897) ; Diirigen,
Deutschl. Amph., p. 519, pi. ii, fig. 7, and pi. iii, fig. 1 (1897).

Cultripes minor, Miiller, Isis, 1832, p. 538, and Zeitschr. f. Phy-
siol., iv, 1832, p. 212.

Pelobates insubricus, Cornalia, Atti Soc. Ven. -Trent., ii, 1873,
p. 44.

Pelobates latifrons, Heron-Royer, Bull. Soc. Zool. France, 1888,
pp. 85, 108, 117, figs. ; Heron-Royer & Van Bambeke, 1. c,
p. 272.

N

194

PELOBATID^E.

Fig. 72.

Open mouth.

Vomerine teeth in two strong, transverse, slightly
oblique or slightly curved series between the choanaB,

the inner borders of which they touch,
and narrowly separated from each
other in the middle. Eustachian tubes
very small. Tongue large, thick, cir-
cular, sometimes entire, usually feebly
nicked behind.

Head very convex, broader than
long, the occiput gibbose; the skin
adherent to the skull; snout rounded,
projecting beyond the mouth, as long-
as or a little longer than the diameter
of the orbit ; no canthus rostralis ; nostril midway
between the eye and the tip of the snout ; eye large,
prominent ; interorbital space convex, much broader
than the upper eyelid, and a little greater than the
distance between the nostrils ; a sort of low knob
behind the eye, formed by the squamosal bone.

Fingers moderately elongate, pointed, third much
the longest, first and fourth equal and slightly longer
than second ; subarticular tubercles very indistinct ;
two rather indistinct carpal tubercles.

Hind limbs robust and short, with swollen calves ;
the tibio-tarsal articulation reaches the shoulder or
the commissure of the mouth, and the tarso-metatarsal
the eye, or between the eye and the nostril ; tibia
shorter than the femur, the heels being widely sepa-
rated from each other when the legs are folded.
Foot much longer than the tibia ; toes short, pointed,

broadly webbed, at least two-
thirds webbed in summer, the
web reaching the tips of all the
toes in the spring ; subarticular
tubercles indistinct ; a very large,
compressed, sharp-edged inner
metatarsal tubercle, placed ob-
liquely to the axis of the foot
like the inner toe, which it equals or slightly exceeds
in length.

Fig. 73.

Lowei' view of foot.

• PELOBATES. 195

Skin smooth on the head and limbs ; sometimes
smooth on the back, usually covered with small flat
warts of unequal size ; lower parts smooth, with a few
granules on the pubic region.

Coloration very variable. Ground colour olive-grey,
pale brown, yellow, yellowish-white, or brownish-
white, with dark-edged chestnut-brown, olive-brown,
or reddish -brown spots, which often simulate larger
and smaller islands with numerous indentations, or
may be confluent into a pair of longitudinal bands,
bordering a pale area of the ground colour along the
spine ; sometimes the spots are very small and nume-
rous, giving the animal a speckled appearance ; the
spots often form irregular bands across the hind
limbs; a dark streak usually extends on each side of
the snout to the eye, involving the nostril, and a dark,
continuous or interrupted band sometimes extends
from one eyelid to the other across the crown ; a light
streak of the ground colour is often present on the
coccygeal region. In addition to these markings there
are often brick-red or vermilion spots or dots, which
may be confined to the sides, or so profusely scattered
on the whole of the upper surfaces as to almost obscure
the ground colour. Lower parts dirty white, uniform,
or spotted or speckled with greyish- F 74

brown, the soles often greyish-
brown ; metatarsal spur yellowish,
pale horn-colour, or reddish-brown.
Iris golden or copper-colour, some-
times fire-colour.

Male distinguished from the fe-
male by a large, oval, smooth gland
on the upper surface of the bra-
chium. Vocal sacs and copulatory
excrescences are absent; during Upper view of foi-e limb
the breeding season, however, small of hreed™s mal*-
pearl-like, uncolourecl excrescences are scattered on
the upper surface of the antebrachium and the
fingers.

196 PELOBATID^.

Measurements (in millimetres).

1. 2. 3. 4. 5. 6.

From snout to vent . . 50 ... 55 ... 65 ... 69 ... 72 ... 61

Length of head . . 17 ... 19 ... 20 ... 20 ... 20 ... 19

Width of head . . 22 ... 23 ... 24 ...25 ... 24 ... 23

Diameter of eye . . 6 ... 6 ... 7 ... 6 ... 7 ... 6

Interorbital width . . 6... 6 ... 7 ... 7 ... 7... 7

From eye to nostril . . 4 ... 4 ... 4-5... 4-5... 5 ... 5

„ „ end of snout . S ... 8 ... 9 ... 10 ... 10 ... 10

Fore limb . . .33 ... 33 ... 38 ... 38 ... 38 ... 37

Hind limb . . . 70 ... 71 ... 81 ...81 ... 85 ... 80

Tibia . . • 19 ... 19 ... 22 ...23 ... 24 ... 22

Metatarsal tubercle . . 5 ... 5 ... 5 ... 5-5... 5 ... 5-5

Inner toe (from tubercle) . 4 ... 4 ... 5 ... 45... 5 . . . 4-5

1. Drancy (Seine-et-Oise): Paratre. I 4. Paris: Heron-Royer.
2, 6. Turin : Peracca. 5. Halle/S. : Wolterstorff.

3. Szamos-Ujvar (Transylvania) : Mehely.

Skeleton. — Skull very strongly ossified, the maxil-
laries, squamosals, nasals, ethmoid, and fronto-parietal
rugose, granular, or pitted. Ethmoid entirely ossified,
confluent with the likewise ossified nasal capsule, and
produced forwards beyoud the nasals to between the
asceuding processes of the praemaxillaries ; only a
small triangular portion of it is exposed between the
nasals and the fronto-parietal. Nasals large, in con-
tact with each other between the nares. Fronto-
parietal single, widening behind into a more or less
developed angular process on each side ; this process
is more or less widely separated from a corresponding
one of the squamosal, with which it is connected by
ligament only, the prootic being entirely exposed
above. Transverse branch of squamosal expanded,
plate-like, suturally united in front with the maxillary.

The degree of roughness of the f ronto-parietals and
the development of the post-orbital processes vary
according to individuals, the bone being generally less
developed in this respect in specimens from France
than in those from Germany, Austria, and Italy. In
quite young specimens the fronto-parietal is nearly

PELOBATES.
Fig. 75.

197

Skeleton o£ male from Transylvania.

198 PELOBATIDJ).

smooth, and the dermo-ossification, as it develops,
produces granular asperities which may become con-
fluent into alveolar ridges, producing a pitted appear-
ance. In very old specimens the sutures between the
nasals, the ethmoid, and the fronto-parietal may become
obliterated.

Vomers rather large, variable in shape, extending
or not to the palatines, which are strongly ossified.
On the palatal side the ethmoid extends posteriorly to
about the middle of the length of the parasphenoid.
Latter large, _L-shaped, its anterior pointed extremity
produced to between the palatines. Pterygoids not
reaching the palatines, extensively in contact with
the parasphenoid. Stapes absent.

Mento-Meckelian or symphysial bones distinct on
the inner side only.

Hyoidalarge broad cartilaginous plate, with slender
postero-lateral processes ; anterior processes much
expanded and confluent with the lateral wings, en-
closing a small fenestra ; ceratohyal cornua with
short posterior portion detached from the body of
the hyoid ; thyroh}rals large, massive, in contact at
the base, diverging and more or less expanded poste-
riorly.

Vertebral column once and a half to twice as long
as the skull. Spine closed above, the neural arch
produced posteriorly into a strong median process
between the zygapophyses. Three anterior diapo-
physes strong and long, especially the second, which
bears a more or less distinct dorsal knob or process
as on the corresponding rib of Discoglossus, — the first
directed obliquely forwards, the second and third
nearly horizontal; the following short and slender,
directed forwards. Sacral vertebra with very strongly
dilated diapophyses, the transverse diameter of which
is twice and a half to nearly three times in the axial ;
urostyle short, not longer than the sacral wings, and
fused with the sacrum, to the diapophyses of which
its anterior portion contributes if these processes be

PKLOBATES.

199

not entirely furnished by it, as is sometimes the
case.*

Praacoracoids slender, strongly curved, not entering
the glenoid cavity ; coracoids nearly straight, strongly
expanded distally ; a cartilaginous omosternum ; ster-
num with a bony style dilated proximally and a large
discoid cartilage. Humerus considerably longer than
radius-ulna. Eight bones in carpus, three of which are
in contact with radius-ulna ; a single bone in the pollex.

Pelvis five-sevenths to five-sixths the length of the
vertebral column ; ilia with an inner upper groove
into which the sacral processes slide; pubis absent
or reduced to a small bony nodule, not entering the
acetabulum. Femur and tibia with cartilaginous epi-
physes ; tibia shorter than the femur, which is shorter
than the pelvis. Calcaneum slightly shorter than
astragalus, half as long as tibia; three distal tarsal
bones ; prsehallux with a single very large, compressed,
curved phalanx. Terminal phalanges pointed.

Measurements op Skeleton (in millimetres).

Length of skull

Width of skull

Least interorbital width .

Dorsal vertebral column .

Urostyle (with sacral vertebra)

Humerus

Radius-ulna

Maim s

Pelvis

Femur

Tibia

Tarsus

Pes

* I have long ago pointed out that the vertebral column figured by
Gene in 1839 as of Bombinator belongs to a Pelobates. I should not
allude to this again were it not for the fact that Gene's error has crept
into Bateson's ' Study of Variation,' p. 127, in a paragraph marked
with an asterisk as a sign of special importance to the example quoted,
and with the addition, by the compiler, of a further error, viz. that the
specimen came from Sardinia. The sacrum in the Gene specimen, as
well as in others that have since been described and figured by Adolphi
(' Morphol. Jahrb.,' xx, 1895, p. 449, pi. xix), is formed entirely by the
processes at the base of the urostyle, and there are thus nine instead of
eight presacral vertebra?.

<?

?

14

17

18

24-

5

6

14

18

10-5 ..

13

12

15

8

10

12

16

20

22

17

22

15

19

7

9

23

28

200 pelobattd,e.

Habits. — Pelobates fuscus is a thoroughly burrowing
Batrachian, spending the greater part of its existence
several feet below the surface of the ground, the
capacious lungs filled with a good provision of air ; it
buries itself in a nearly erect squatting posture by
means of the large sharp-edged horny shovels with
which its feet are armed, the digging being effected
by alternate lateral movements of the heels, inter-
rupted now and then for a short rest; on sand or a
very loose soil less than a minute may suffice for the
animal to disappear from sight, the soil falling over
and around it so as not to leave a trace of a burrow.
The English name " spade-foot," which is applied to
its North American representative, Seaphiopus, would
be a most appropriate one. Owing to these habits it
is only found in localities where the soil is light or
sandy; in cultivated districts the extensive growth of
asparagus is an almost infallible indication of its pre-
sence. In the summer months it emerges an hour or
two after sunset, and hopping along frog-like, sets
forth in search of its food, consisting of worms, slugs,
and all kinds of insects, especially beetles. When
suddenly seized it produces a dermal exudation, smell-
ing like garlic, and usually utters a startling shrill cry
much like that of a kitten, at the same time opening-
its mouth in a defiant attitude. Some specimens
when repeatedly teased — pinches in the leg and light
pokes in the sides being the best method to employ
to witness the sight — can be roused to what appears
to be a fit of anger lasting several minutes, the animal
screaming in a most extraordinary manner, and jump-
ing madly with open mouth as if to snap at its perse-
cutor. The prolonged screams produced on those occa-
sions can only be compared, in a diminutive way, to
those of an infant. At the same time the luno-s are
inflated to their utmost, and the body, swollen like a
ball, is raised above ground. A specimen is figured
in this attitude on PI. IX, right hand. These antics
are very similar to those performed by Ceratophrys

PELOBATES. 201

omata, a South American Cystignathoid, of likewise
nocturnal and burrowing habits, specimens of which
have often been kept in confinement in this country;
an important difference being, however, that a large
Ceratophrys is able to inflict a very painful bite on the
finger of the unwary.

This Batrachian appears in the daytime only during
the breeding season, which takes place in March and
even as late as May* in Italy, between the end of
March and the beginning of May in Central and
Northern Europe, stray pairs occurring exceptionally
as late as the 21st of July, according to an observation
made by Prof. Van Bambeke at Ghent, Belgium, in
1875. At the time of pairing, at which pools or deep
ditches are resorted to, the males, much more nume-
rous than the females, utter under water a mono-
tonous, constantly repeated note — clock- clock, clock-
clock-clock, — produced by alternately shifting the air
backwards and forwards from the capacious lungs
into the buccal cavity. The female, which answers
by a sort of grunt or a deep tock-tock-tock, is seized
round the waist, and the eggs are expelled, more or
less rapidly, either immediately or within a few days.

The length of the larval life varies considerably; as
a ride, the final transformation takes place from the
beginning to the end of summer, but several cases of
larval hibernation have been observed by Pfliiger and
Kollmann.

Eggs. — The contents of the two oviducts fuse in the
cloaca, and are expelled in a thick band 15 to 20 mm.
in diameter when swollen up, in which the ova are
irregularly disposed at small interspaces. These
bands, which have a strong smell of fish, are twisted
round weeds by the female as they are laid. The ova

* According to Spallanzani, who described the oviposition in a pair
brought to him in May, 1780, by a fisherman at Pavia, and who
received afterwards other pairs of the same species (CEuvres de
Spallanzani, III, ' Experiences pour servir a l'histoire de la Genera-
tion,' Pavie, 1787, p. 137). As Crivelli and Camerano have shown,
Spallanzani was the first after Posel to observe this curious Batra-
chian, which was not rediscovered in Italy until ninety years later.

202

PELOBATID.E,

are small, 2 to 2-J mm. in diameter, grey or brown
with white lower pole, and develop rapidly ; the em-
bryo, gill-less and tailless, breaks through about the
fifth or sixth dav, and attaches itself, through its
adhesive subcephalic apparatus, to the outer surface

of the mucilage.

Fig. 76.

rTl

Tadpole (PI. II, fig. 1). — Length of body once and
a half to twice its width, oue-half to two-thirds the
length of the tail. Nostrils a little nearer the eyes
than the end of the snout. Eyes on the upper sur-
face, equidistant from the end of the snout and the
spiraculum, the distance between them at least twice,
sometimes nearty three times as great as that between
the nostrils, and greater than the width of the mouth.
Spiraculum on the left side, directed upwards aud
backwards, equidistant from either extremity of the
body or a little nearer the anterior extremity, visible
from above and from below. Anal opening median,
a little larger than the spiraculum, and close to the
body. Tail twice and a half to thrice and one-fifth as
long as deep, acutely pointed ; upper crest convex,
slightly deeper than the lower, not extending far
upon the back ; the depth of the muscular portion
at its base about half the greatest total depth.

Beak black. Lip bordered with papilla?, which form
two or more rows on the sides ; the papillose border
interrupted mesially by a narrow toothed descending
lobe, which appears at first sight as continuous with
the second upper series of teeth. This anterior series
is followed by three or four other series of teeth,
which are all widely interrupted in the middle and

PELOBATES. 203

gradually decrease in length, — the fourth, if at all
present, being extremely short. On the lower lip we
see likewise a short outer series, followed by three or
four much longer ones, all of which, with the occa-
sional exception of the first, are interrupted in the
middle, and may be more or less broken up on the
sides. The series of labial teeth may therefore be
formulated as \ or -f. Small isolated teeth may also
be scattered on the papilla? at the angles of the
mouth.

I have not been able to distinguish lines of crypts
beyond the two series which run from the end of the
snout to the upper border of the eyes, passing be-
tween the nostrils. I am therefore unable to iudge
whether the dorsal lines have been correctly figured
by Lessona, who rightly regards the figure given by
Corn alia as fanciful.

The advanced tadpole is brown or olive-brown above,
with or without small darker spots, greyish-white or
bluish beneath ; sides with roundish whitish or pale
golden spots; tail pale brown, with small grey and
whitish spots.

The body usually reaches at least the size of a
pigeon's egg, but not unfrequently exceeds that size.
The largest specimen in the British Museum, from
Prague, measures 125 mm., body 38, width of body
25, tail 87, depth of tail 27. The largest specimen on
record is one preserved in the Berlin Museum, obtained
in December, 1867, on the Jungfernheide, near Berlin,
and which measures 175 mm.

Habitat. — The range of this species is an extensive
but broken one, its occurrence being obviously depen-
dent on the nature of the soil, and restricted to the
plain, the highest elevation at which it has }7et been
recorded being 1400 feet (Val di Susa, Piedmont). P.
fuscus also appears to avoid competition with its con-
gener P. cultrijjes ; the limit of the range of the two
closely meet without anywhere overlapping. It in-
habits the north-east and east of France, from Flanders

20-k PELOBATID^l.

to the Jura, westwards to Paris, the Dep. Sarthe, and
along the Loire; Belgium (Prov. Limburg, Antwerp,
and East Flanders), South Holland, Luxemburg, Swit-
zerland near Basle, Germany (except Wiirtemberg),
Austria-Hungary, Carniola, North Italy (Piedmont,
Lombardy, Venetia, Emilia), Denmark, Southern
Sweden and Gotland, and Russia, northwards to the
Gulf of Riga and eastwards to the Ural River and
the Kirghiz Steppes, the Emba River being its eastern-
most limit.

The species has quite recently been recorded from
Lenkoran, south-west coast of the Caspian Sea, by
Zander.*

The reported occurrence of Pelobates fuscus in the
Cyclades is probably to be ascribed to a confusion
with Bufo viridis, some specimens of which closely
resemble this species in their markings.

The specimens represented on PI. IX are from the
environs of Turin, presented by my friend Count
Peracca, to whom I am indebted for so much material
utilised in the preparation of this work. The left-
hand figure is that of a female; the others represent
males, one in the act of disappearing under the sand,
another in the irritated attitude described above.

* Dr. Zander kindly informs me that the specimens Lave been pre-
sented by him to the Senckenberg Museum, Frankfort (M.), and that
bis determination bas been confirmed by Prof. Boettger.

"

Is

>

pelobates. 205

8. Pelobates cultripes.
(Plate X.)

Buna cultripes, Ouvier, Regne Anim., 2nd ed., ii, p. 105 (1829).

Rana calcarata, Michahelles, Isis, 1830, p. 807, pi. — .

Cultripes provincialis, Muller, Isis, 1S32, p. 538, and Zeitscbr. f.

Physiol, iv, 1832, p. 212.
Bufo calcaratus, Scbinz, Nat. Rept., p. 233, pi. xcvi, fig. 2 (1833).
Bombinator fuscus, Duges, Recb. Osteol. Batr., p. 7, pi. ii (1834).
Belobates cultripes, Tschudi, Class. Batr., p. 83 (1S3S) ; Duuieril

& Bibron, Erp. Gen., viii, p. 483 (1841) ; Gunther. Cat.

Batr. Sal., p. 41 (1858) ; Schreiber, Herp. Ear., p. 92 (1875) ;

Lataste, Herp. Gir., p. 263 (1876); Boulenger, Cat. Batr.

Ecaud., p. 438 (1882); Heron-Royer, Bull. Soc. Et. Sc.

Angers, xv, 1886, p. 77; Heron-Royer <fc Van Bambeke,

Arch. Biol., ix, 1889, p. 275; Bedriaga, Bull. Soc. Nat.

Mosc, 1889, p. 519, and Anmh. Rept. Portug., p. 19 (1889) ;

Boulenger, Proc. ZooL Soc, 1890, p. 664, and 1891, p. 616, pi.

xlvi, fig. 8.
Belobates fuscus, part., Bonaparte, Icon. Faun. Ital., Rett. Anf.,

(1838).
Didocus* calcaratus, Cope, Journ. Ac. Philad. (2), vi, 1866, p. 81.

In the tongue and vomerine teeth this species
entirely agrees with its congener, but it differs in the
occasional presence of small, grain-like teeth on the
pterygoids and parasphenoid ; these teeth are quite
rudimentary, and the mucous membrane has to be
removed to ascertain their presence. I have found
them in three specimens from France (Nantes, Bor-
deaux, south of France). In one specimen there
are about ten teeth on the parasphenoid, at the base
of the longitudinal branch of the J_-shaped bone, and
two pterygoid teeth close together on the left side ;
another has the same number on the parasphenoid and
a series of four on the pterygoids; and in a third
there are eight teeth on the right pterygoid, none
being present on the parasphenoid nor on the right
pterygoid.

* The genus Didocus was founded upon a young specimen, on the
erroneous assumption that in Belobates cultripes the temporal roof is
developed before the tail of the larva has disappeared. I have examined
the skeleton of a young but fully transformed specimen from Beziers,
Herault, which shows both a large fronto-parietal fontanelle, bordered
by smooth fronto-parietals, and an uncovered temple, thus answering
in every respect the definition of the genus so rashly established by
Cope more than twenty years ago, and still maintained by him in his
latest writings.

206 PELOBATID.E.

Head as in P. fuscus, but rather larger in propor-
tion, and not humped on the occiput nor behind the
eyes, the skull forming a complete, evenly curved
rugose casque entirely surrounding the orbits ; inter-
orbital space nearly flat, usually much broader than
the distance between the nostrils.

Shape and proportions of the limbs much as in
P. fuscus, but digits more obtuse, and metatarsal
shovel sharper and larger, always exceeding the
leno'th of the inner toe.

Skin smooth or densely covered above with very
small round warts.

Yellowish, whitish, or greenish-yellow above,
speckled, spotted or marbled with brown, olive, or
greyish-olive, the spots small or large, and but rarely
confluent into longitudinal bands. A small male
specimen, collected by M. Lataste at Greilhan,
Gironde, has four perfectly regular dark dorsal
stripes, the median pair extending forwards to the
interorbital region, where they meet a dark cross-bar.
White beneath, uniform or speckled with greyish-
olive ; metatarsal spur black ; toes often tipped with
a black horny layer. Iris silvery or greenish-grey,
vermiculate with black.

External male sexual characters as in P. fuscus.

In describing Pelobates syriacus, Boettger (' Zool.
Anz.,' 1889, No. 302) mentions breeding specimens of
Pelobates cultripes to be possessed of nuptial asperities
on the inner side of the three inner fingers and on the
tips of the toes. This extraordinary statement is based
on a confusion with the American Scaphiopus liam-
monclii, Baircl (dugesii, Brocchi), breeding males of
which were erroneously labelled ' Pelobates cultripes,
Spain,' in the Senckenberg Museum. The so-called
nuptial asperities of the tips of the toes are nothing
but the blackish horny sheaths which cap them, as
well as the tarsal spur, at all seasons. I am indebted
to the kindness of Prof. Boettger for an opportunity
of examining these specimens.

PELOKATES.

207

Measurements (in millimetres).

S

From snout to vent
Length of bead
Width of head
Diameter of eye
Interorbital width
From eye to nostril

„ ,, end of snout
Fore limb
Hind limb
Tibia .

Metatarsal tubercle .
Inner toe (from tubercle)

1.
66
1'2
26

8

6-,

5

10
40
S8
24

6-

5

2.
77

■16

29

9

8

7

13

45

100

28

3.

4.

5.

57 ..

51 .

.. 88 ...

20 ..

17 .

.. 27

25 ..

21 .

.. 36

8 ..

7 .

.. 9

7

6-5.

.. 7

5-5..

5

.. 8

10 ..

9 .

.. 14

35 ..

27 .

.. 47

so ..

67 .

.. 118

22

19 .

.. 32

6 '.'.

5 .

.. 8

5 ..

35.

.. 7

6.

62

20

25

8

6

5

10

34

79

22

6

4

1. Nantes :

2. Seville:
3, 4. Merida :

Paris Museum.
Calderon.
Bosca.

5. Badajos : Bosca.

6. Abrantes : Gadow.

Skeleton. — The trunk and limbs agree with those of
its congener, but the skull is very different, owing to the

Fig. 77.

Upper view of skull.

more extensive dermo-ossification of the post-orbital
region, which forms a complete roof over the prootic,
only the posterior portion of the latter being visible
from above. This roof, which completely closes the
orbit behind, is formed by the fronto-parietal and the
squamosal ; the latter bone has a convex posterior
border, separated from the likewise curved posterior
border of the fronto-parietal plate by a notch, the
angle of which corresponds to the suture between the
two bones.

208

PELOBATIL\E.

All the bones of the upper surface are closely
studded with granular or subcorneal asperities. A
further important difference in the skull of this species
compared to P. fuscus is the absence of the extra-
ordinary forward prolongation of the ethmoid ; this
bone does not extend beyond the nasals.

Duges, basing his observations on this species, has
denied the fusion of the sacral vertebra with the
coccyx described by Mertens in Pelobates fuscus, with
which P. cultrvpes was then confounded ; he observes,
however, that the articulation, by means of one con-
dyle, is an almost immoveable one.

In a specimen from Bordeaux, from which I prepared
the first skeleton, I found matters as stated by Duges,
whilst in two other skeletons, from Bordeaux and
Avignon, the two bones are as completely fused as in
P. fuscus. As the ankylosis of the sacrum and coccyx
has often been given as a generic character of Pelo-
bates, it is important to note the inconstancy of the
character in this species at least.

Measurements op Skeleton (in millimetres).

<?

9

Length of skull

23

25

Width of skull

29

31

Least interorbital width .

7

75

Dorsal vertebral column .

24

26

Urostyle (with sacr

al vertebra)

17

18

Humerus

21

20

Radius-ulna

.

14

13

Manus

,

20

19

Pelvis

, ,

34

38

Femur

. ,

28

30

Tibia

25

26

Tarsus

.

13

13

Pes

.

34

36

Habits. — Much the same as those of its congener.
P. cultripes seems, however, more partial to the coast,
being found in abundance, in France, on the sandy
dunes of the Atlantic littoral. It breeds in the end
of March or April in France, in March in Spain. Its
note, which I have only heard in specimens captured

PELOBATES. 209

in summer, may bo well rendered by a guttural co,
co, co, co, co ; its nuptial call is said to be the same.

Eggs. — According to Heron-Rover, they differ but
slightly from those of P. fuscus ; the mucilaginous
band is a little narrower and flatter, thus much re-
sembling that of Pelodijtes punctatus.

Tadpole (PL II, fig. 2). — The differences from P.
fuscus are but slight, and most of those that have
hitherto been relied upon prove not to be absolutely
constant. Thus the nostrils, as I now find from
Portuguese specimens received from Dr. de Bedriaga
since the publication of my ' Synopsis of the Tadpoles,'
may be as wide apart as in P. fuscus. The series of
labial teeth are more broken up, and their arrange-
ment is therefore less easily expressed by a formula.
The tail is shorter, not more than once and a half the
length of the body. The lines of crypts are usually
more distinct, owing to the black colour of the tubules
and the lighter colour of the body. Their disposition
is well shown in the example figured. The following
description is taken from five specimens obtained by
M. Lataste near Bordeaux :

Length of body once and two-thirds to once and
three-fourths its width, once and oue-fourth to once
and two-fifths in the length of the tail. Nostrils half-
way between the eyes and the end of the snout, or a
little nearer the latter. Eyes on the upper surface, a
little nearer the spiraculum than the end of the snout,
the distance between them nearly twice as great as
that between the nostrils, which equals the width of
the mouth. Spiraculum equidistant from either ex-
tremity of the body. Tail once and two-thirds to
once and five-sixths as long as deep, pointed, the
muscular portion not half the total depth.

Specimens preserved in spirit, of which I have a
good number before me at the present moment, are
pale greyish or brownish above and on the sides, the
belly dirty white with round white spots. Muscular
part of tail pale reddish-brown, with some darker

o

210 PELOBATID^.

brown spots ; caudal crests greyish, more or less
distinctly spotted or freckled with brown.

The coloration in life is described by Lataste as
reddish-yellow above, greyish or bluish -white beneath ;
tail with small brown spots. Some twenty years ago
I saw a number of large specimens in a small tank in
the Botanic Gardens at Bordeaux. My recollection
is that they were of a very pale brown, without distinct
markings.

Total length 100 mm.; body 39; tail 61 ; depth of
tail 25.

According to Duges the size of the body may equal
a hen's egg.

Habitat. — France, Spain, and Portugal. In France
this species has been found along the west coast
to a little north of the mouth of the Loire, and in
the southern departments Haute-Garonne, Pyrenees-
Orien tales, Aude, Herault, Gard, Bouches-du-llhone,
and Vaucluse ; in Spain in the provinces Galicia,
Estremadura, New Castille, Valencia, Andalusia ; and
in Portugal in the provinces Douro and Algarve. It
probably occurs, as stated by Bosca, all round the
coasts of the Pyrenean Peninsula.

The record of the occurrence of Pelobates cultrvpesm
Syria is due to a confusion with an allied species,
Pelobates si/riacns, to which allusion is made above,
p. 192.

The two specimens figured on PI. X were obtained
near Bordeaux by M. F. Lataste.

ntlNTED BY AViLkUD AND SON,
BARTHOLOMEW CLOSE, E.C., AND 20 UANOVEE SQUARE, W.