Chapter IV: ECAUDATA (22).--Frogs and toads. Four limbs and no tail. Radius (1)

confluent with ulna, and tibia with fibula; tarsus (astragalus and
calcaneum) elongate, forming an additional segment in the hind limb.
Caudal vertebrae fused into a urostyle or coccyx. Frontal bones
confluent with parietals.

This order embraces about 1300 species, of which some 40 are fossil,
divided into two sub-orders and sixteen families:--

A. Aglossa,--Eustachian tubes united into a single ostium pharyngeum;
no tongue. DACTYLETHRIDAE, PIPIDAE.

B. Phaneroglossa,--Eustachian tubes separated; tongue present.
DISCOGLOSSIDAE, PELOBATIDAE, HEMIPHRACTIDAE, AMPHIGNATHODONTIDAE,
HYLIDAE, BUFONIDAE, DENDROPHRYNISCIDAE, CYSTIGNATHIDAE, DYSCOPHIDAE,
GENYOPHRYNIDAE, ENGYSTOMATIDAE, CERATOBATRACHIDAE, RANIDAE,
DENDROBATIDAE.

The Phaneroglossa are divided into two groups; _Arcifera_ and
_Firmisternia_, representing two stages of evolution. The family
characters are mainly derived from the dilatation or non-dilatation of
the sacral diapophyses, and the presence of teeth in one or both jaws,
or their absence. The _Discoglossidae_ are noteworthy for the presence
of short ribs to some of the vertebrae, and in some other points also
they approach the tailed batrachians; they may be safely regarded as,
on the whole, the most generalized of known Ecaudata. Distinct ribs
are present at an early age in the Aglossa, as discovered by W.G.
Ridewood (23). The recent addition of a third genus of Aglossa,
_Hymenochirus_ (24) from tropical Africa, combining characters of
_Pipa_ and _Xenopus_, has removed every doubt as to the real affinity
which connects these genera. _Hymenochirus_ is further remarkable for
the presence of only six distinct pieces in the vertebral column,
which is thus the most abbreviated among all the vertebrata.

Frogs and toads occur wherever insect food is procurable, and their
distribution is a world-wide one, with the exception of many islands.
Thus New Caledonia, which has a rich and quite special lizard-fauna,
has no batrachians of its own, although the Australian _Hyla aurea_
has been introduced with success. New Zealand possesses only one
species (_Liopelma hochstetteri_), which appears to be rare and
restricted to the North Island. The forest regions of southern Asia,
Africa and South America are particularly rich in species.

According to our present knowledge, the Ecaudata can be traced about
as far back in time as the Caudata. An unmistakable batrachian of this
order, referred by its describer to _Palaeobatrachus_, a determination
which is only provisional, has been discovered in the Kimmeridgian of
the Sierra del Montsech, Catalonia (25), in a therefore somewhat older
formation than the Wealden Caudata _Hylaeobatrachus_.

Apart from a few unsatisfactory remains from the Eocene of Wyoming,
fossil tailless batrachians are otherwise only known from the
Oligocene, Miocene and Pliocene of Europe and India. These forms
differ very little from those that live at the present day in the same
part of the world, and some of the genera (_Discoglossus, Bufo,
Oxyglossus, Rana_) are even identical. _Palaeobatrachus_ (26), of
which a number of species represented by skeletons of the perfect form
and of the tadpole have been described from Miocene beds in Germany,
Bohemia and France, seems to be referable to the _Pelobatidae_; this
genus has been considered as possibly one of the Aglossa, but the
absence of ribs in the larvae speaks against such an association.

Numerous additions have been made to our knowledge of the development
and nursing habits, which are extremely varied, some forms dispensing
with or hurrying through the metamorphoses and hopping out of the egg
in the perfect condition (27).

_Skeleton._--In the earliest forms of this order, the Stegocephalia,
we meet with considerable variety in the constitution of the
vertebrae, and these modifications have been used for their
classification. All agree, however, in having each vertebra formed of
at least two pieces, the suture between which persists throughout
life. In this they differ from the three orders which have living
representatives. Even the inferior arches or chevrons of the tail of
salamanders are continuously ossified with the centra. As a matter of
fact, these vertebrae have no centra proper, that part which should
correspond with the centrum being formed, as a study of the
development has shown (H. Gadow, 14), by the meeting and subsequent
complete co-ossification of the two chief dorsal and ventral pairs of
elements (tail-vertebrae of Caudata), or entirely by the pair of
dorsal elements. In the Ecaudata, the vertebrae of the trunk are
formed on two different plans. In some the notochord 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 be easily
stripped off below the vertebrae in larval specimens on the point of
metamorphosing. This has been termed the _epichordal_ type. In others,
which represent the _perichordal_ 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 hypochordal
cartilage which fuses with the dorsal or becomes connected with it by
calcified tissue; the notochord is thus completely surrounded by a
thick sheath in tadpoles with imperfectly developed limbs. This mode
of formation of both the arch and 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.

During segmentation of the dorsal cartilages mentioned above, which
send out the transverse processes of diapophyses, there appears
between each two centra an intervertebral cartilage, out of which the
articulating condyle of the centrum is formed, and becomes attached
either to the vertebra anterior (precoelous type) or posterior
(opisthocoelous type) to it, if not remaining as an independent,
intervertebral, ossified sphere, as we sometimes find in specimens of
_Pelobatidae_.

In the Caudata and Apoda, cartilage often persists between the
vertebrae; this cartilage may become imperfectly separated into a
cup-and-ball portion, the cup belonging to the posterior end of the
vertebra. In such cases the distinction between amphicoelous and
opisthocoelous vertebrae rests merely on a question of ossification,
and has occasionally given rise to misunderstandings in the use of
these terms.

Amphicoelous (bi-concave) vertebrae are found in the Apoda and in some
of the Caudata; opisthocoelous (convexo-concave) vertebrae in the
higher Caudata and in the lower Ecaudata; whilst the great majority of
the Ecaudata have procoelous (concavo-convex) vertebrae.

rp, The rhinal process.
pnl, The praenasal processes.
an, The alinasal processes, shown by the removal of part of the
floor of the left nasal chamber.
AO., The antorbital process.
pd, The pedicle of the suspensorium continued into cv, the
ventral crus of the suspensorium.
cd, Its dorsal crus.
tt, The tegmen tympani.
SE, The sphen-ethmoid.
EO., The exoccipitals.
Qu.J., The quadratojugal.
II. V. VI. Foramina by which the optic, trigeminal and abortio dura,
and abducens nerves leave the skull.]

All living batrachians, and some of the Stegocephalia, have transverse
processes on the vertebrae that succeed the atlas (fig. 4), some of
which, in the Caudata, are divided into a dorsal and a ventral
portion. Ribs are present in the lower Ecaudata (_Discoglossidae_ and
larval Aglossa), but they are never connected with a sternum. It is in
fact doubtful whether the so-called sternum of batrachians, in most
cases a mere plate of cartilage, has been correctly identified as
such. When limbs are present, one vertebra, rarely two (fig. 5) or
three, are distinguished as sacral, giving attachment to the ilia. In
the Ecaudata, the form of the transverse processes of the sacral
vertebra varies very considerably, and has afforded important
characters to the systematist. In accordance with the saltatorial
habits of the members of this order, the vertebrae, which number from
40 to 60 in the Caudata, to upwards of 200 in the Apoda, have become
reduced to 10 as the normal number, viz., eight praecaudal, one sacral
and an elongate coccyx or urostyle, formed by coalescence of at least
two vertebrae. In some genera this coccyx is fused with the ninth
vertebra, and contributes to the sacrum, whilst in a few others the
number of segments is still further reduced by the co-ossification of
one or two vertebrae preceding that corresponding to the normal sacral
and by the fusion of the two first vertebrae, the extreme of reduction
being found in the genus _Hymenochirus_, the vertebral column of which
is figured here (fig 6.)

As stated above in the definition of the order, the Stegocephalia have
retained most of the cranial bones which are to be found in the
Crossopterygian fishes, and it is worthy of note that the bones termed
post-temporals may give attachment to a further bone so prolonged
backwards as to suggest the probability of the skull being connected
with the shoulder-girdle, as in most teleostome fishes. This
supposition is supported by a specimen from the Lower Permian of
Autun, determined as _Actinodon frossardi_, acquired in 1902 by the
British Museum, which shows a bone, similar to the so-called "epiotic
cornu" of the microsaurians, _Ceraterpeton_ and _Scincosaurus_, to
have the relations of the supra-cleithrum of fishes, thus confirming a
suggestion made by C.W. Andrews (28). As in fishes also, the sensory
canal system must have been highly developed on the skulls of many
labyrinthodonts, and the impressions left by these canals have been
utilized by morphologists for homologizing the various elements of the
cranial roof with those of Crossopterygians. The pineal foramen, in
the parietal bones, is as constantly present as it is absent in the
other orders. Although not strictly forming part of the skull,
allusion should be made here to the ring of sclerotic plates which has
been found in many of the Stegocephalia, and which is only found
elsewhere in a few Crossopterygian fishes as well as in many reptiles
and birds.

In the orders which are still represented at the present day, the
bones of the skull are reduced in number and the "primordial skull,"
or chondrocranium (fig. 7), remains to a greater or less extent
unossified, even in the adult. Huxley's figures of the skull of a
caccilian (_Ichthyophis glutinosus_), fig. 8, of a perennibranchiate
urodele (_Necturus maculosus = Menobranchus lateralis_), fig. 9, and
of a frog (_Rana esculenta_), fig. 10, are here given for comparison.

The skull, in the _Apoda_, is remarkably solid and compact, and it
possesses a postorbital or postfrontal bone (marked 1 in the figure)
which does not exist in any of the other living batrachians. The
squamosal bone is large and either in contact with the frontals and
parietals or separated from them by a vacuity; the orbit is sometimes
roofed over by bone. The presence, in some genera, of a second row of
mandibular teeth seems to indicate the former existence of a splenial
element, such as exists in _Siren_ among the Caudata and apparently in
the labyrinthodonts.

In the Caudata, the frontals remain likewise distinct from the
parietals, whilst in the Ecaudata the two elements are fused into one,
and in a few forms (Aglossa, some _Pelobalidae_) the paired condition
of these bones has disappeared in the adult. Prefrontal bones are
present in the _Salamandridae_ and _Amphiumidae_, but absent (or fused
with the nasals) in the other Caudata and in the Ecaudata. In most of
the former the palatines fuse with the vomers, whilst they remain
distinct, unless entirely lost, in the latter. The vomer is single, or
absent, in the Aglossa. In the lower jaw of most of the Ecaudata the
symphysial cartilages ossify separately from the dentary bones,
forming the so-called mento-meckelian bones; but these symphysial
bones, so distinct in the frog, are less so in the _Hylidae_ and
_Bufonidae_, almost indistinguishable in the _Pelobatidae_ and
_Discoglossidae_, whilst in the Aglossa they do not exist any more
than in the other orders of batrachians.

No batrachian is known to possess an ossified azygous supra-occipital.

VII.p, Posterior division of the seventh nerve.
VII. Chorda tympani
V^1, V^2, V^3, First, second and third divisions of the trigeminal.
s.s.l, Stapedio-suspensorial ligament.
h.s.l, Hyo-suspensorial ligament.
m.h.l, Mandibulo-hyoid ligament.
a, Ascending process of the suspensorium.
p, Pterygo-palatine process.
q, Quadrate process.
o, Otic process.
Na, Posterior nares.
Mck, Meckel's cartilage.
Gl (fig. 10), The position of the glottis.
Bb^1, Bb^2, Basilbranchials.]

Although there are four branchial arches in all the larval forms of
the three orders, and throughout life in the _Sirenidae_, the
perennibranchiate _Proteidae_ have only three (see fig. 11). In the
adult Apoda these arches and the hyoid fuse into three transverse,
curved or angular bones (see fig. 13), the two posterior disconnected
from the hyoid. In the Ecaudata, as shown by F. Gaupp (29) and by W.G.
Ridewood (30), the whole hyobranchial apparatus forms a cartilaginous
continuum, and during metamorphosis the branchialia disappear without
a trace. The hyoid of the adult frog (fig. 12) consists of a plate of
cartilage with two slender cornua, three processes on each side, and
two long bony rods behind, termed the thyro-hyals, which embrace the
larynx. In the Aglossa, which are remarkable for the large size and
complexity of the larynx, the thyro-hyal bones are incorporated into
the laryngeal apparatus, whilst the recently discovered _Hymenochirus_
is further remarkable for the large size and ossification of the
hyoidean cornua (ceratohyals), a feature which, though not uncommon
among the salamanders, is unique among the Ecaudata (31).

Pmx, Premaxilla.
Mx, Maxilla.
Vo, Vomer.
Na, Nasal.
S.e, Sphen-ethmoid.
Fr, Frontal.
Pa, Parietal.
E.O, Exoccipital.
Ep, Epiotic process.
Pr.O, Pro-otic.
t.t, Tegmentympani.
Sq, Squamosal.
Q.J, Quadrato-jugal.
Pt1, Pterygoid, anterior process.
Pt2, Internal process.
Pt3, Posterior or external process.
Ca, Columella auris.
St, Stapes.
Hy, Hyoidean cornu.
P.S, Parasphenoid.
An, Angulate.
D, Dentale.
V, Foramen of exit of the trigeminal.
H, Of the optic.
X, Of the pneumogastric and glosso-pharyngeal nerves.
V1. Foramen by which the orbito-nasal or first division of the fifth
passes to the nasal cavity.]

The pectoral girdle of the Stegocephalia is, of course, only known
from the ossified elements, the identification of which has given rise
to some diversity of opinion. But C. Gegenbaur's (32) interpretation
may be regarded as final. He has shown that, as in the Crossopterygian
and Chondrostean ganoid fishes, there are two clavicular elements on
each side; the lower corresponds to the clavicle of reptiles and
higher vertebrates, whilst the upper corresponds to the clavicle of
teleostean fishes, and has been named by him "cleithrum." As stated
above, there is strong evidence in favour of the view that some forms
at least possessed in addition a "supracleithrum," corresponding to
the supra-clavicle of bony fishes. The element often termed "coracoid"
in these fossils would be the scapula. The clavicles rest on a large
discoidal, rhomboidal, or T-shaped median bone, which clearly
corresponds to the interclavicle of reptiles.

The pectoral girdle of the living types of batrachians is
distinguishable into a scapular, a coracoidal, and a praecoracoidal
region. In most of the Caudata the scapular region alone ossifies, but
in the Ecaudata the coracoid is bony and a clavicle is frequently
developed over the praecoracoid cartilage. In these batrachians the
pectoral arch falls into two distinct types--the _arciferous_, in
which the precoracoid (+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_, in which both precoracoid and coracoid nearly abut on
the median line, and are only narrowly separated by the more or less
fused epicoracoids. The former type is exemplified by the toads and
the lower Ecaudata, whilst the latter is characteristic of the true
frogs (_Ranidae_), although when quite young these batrachians present
a condition similar to that which persists throughout life in their
lower relatives. A cartilage in the median line in front of the
precoracoids, sometimes supported by a bony style, is the so-called
Omosternum; a large one behind the cora-coids, also sometimes provided
with a bony style, has been called the sternum. But these names will
probably have to be changed when the homologies of these parts are
better understood.

Hh, Hypo-hyal.
Ch, Cerato-hyal.
Bb^1, First basibranchial.
Bb^2, Ossified second basibranchial.
Ep.b^1, Ep.b^2, Ep.b^3, First, second and third epibranchials.
Gl, Glottis.]

The pelvic arch of some of the Stegocephalia contained a well ossified
pubic element, whilst in all other batrachians only the ilium, or the
ilium and the ischium are ossified. In the Ecaudata the ilium is
greatly elongated and the pubis and ischium are flattened, discoidal,
and closely applied to their fellows by their inner surfaces; the
pelvic girdle looks like a pair of tongs.

The long bones of the limbs consist of an axis of cartilage; the
extremities of the cartilages frequently undergo calcification and are
thus converted into epiphyses. In the Ecaudata the radius and ulna
coalesce into one bone. The carpus, which remains cartilaginous in
many of the Stegocephalia and Caudata, contains six to eight elements
when the manus is fully developed, whilst the number is reduced in
those forms which have only two or three digits. Except in some of the
Stegocephalia, there are only four functional digits in the manus, but
the Ecaudata have a more or less distinct rudiment of pollex; in the
Caudata it seems to be the outer digit which has been suppressed, as
atavistic reappearance of a fifth digit takes place on the outer side
of the manus, as it does on the pes in those forms in which the toes
are reduced to four. The usual number of phalanges is 2, 2, 3, 2 in
the Stegocephalia and Caudata, 2, 2, 3, 3 in the Ecaudata. In the foot
the digits usually number five, and the phalanges 2, 2, 3, 3, 2 in the
Caudata, 2, 2, 3, 4, 3 in the Stegocephalia and Ecaudata. There are
occasionally intercalary ossifications between the two distal
phalanges (33). There are usually nine tarsal elements in the Caudata;
this number is reduced in the Ecaudata, in which the two bones of the
proximal row (sometimes coalesced) are much elongated and form an
additional segment to the greatly lengthened hind-limb, a sort of
_crus secundarium_. In the Ecaudata also, the tibia and fibula
coalesce into one bone, and two or three small bones on the inner side
of the tarsus form what has been regarded as a rudimentary digit or
"prehallux."

_Integument._--In all recent batrachians, the skin is naked, or if
small scales are present, as in many of the Apoda, they are concealed
in the skin. The extinct Stegocephalia, on the other hand, were mostly
protected, on the ventral surface at least, by an armour of
overlapping round, oval, or rhomboidal scales, often very similar to
those of Crossopterygian or ganoid fishes, and likewise disposed in
transverse oblique lines converging forwards on the middle line of the
belly. Sometimes these scales assumed the importance of scutes and
formed a carapace, as in the "batrachian armadillo" discovered by E.D.
Cope. A few frogs have the skin of the back studded with stellate bony
deposits (_Phyllomedusa, Nototrema_), whilst two genera are remarkable
for possessing a bony dorsal shield, free from the vertebrae
(_Ceratorphrys_) or ankylosed to them (_Brachycephalus_). None of the
Stegocephalia appears to have been provided with claws, but some
living batrachians (_Onychodactylus, Xenopus, Hymenochirus_) have the
tips of some or all of the digits protected by a claw-like horny
sheath.

The integument of tailed and tailless batrachians is remarkable for
the great abundance of follicular glands, of which there may be two
kinds, each having a special secretion, which is always more or less
acrid and irritating, and affords a means of defence against the
attacks of many carnivorous animals. A great deal has been published
on the poisonous secretion of batrachians (34), which is utilized by
the Indians of South America for poisoning their arrows. Some of the
poison-secreting glands attain a greater complication of structure and
are remarkable for their large size, such as the so-called "parotoid"
glands on the back of the head in toads and salamanders.

Mn, Mandible.
Hy, Hyoid.
Br^1 Br^2, Br^3, Branchial arches.
Gl, Glottis.
Tr, Trachea.
Ivc, Inferior vena cava.
V, Ventricle.
Au, Auricles.
Rsvc, Lsvc, right and left superior cavae.
Ta, Truncus arteriosus.
Ao, Left aortic arch.
P.A. Right pulmonary artery. The pericardium (lightly shaded)
extends as far as the bifurcation of the synangium.]

In all larval forms, in the Caudata, and in a few of the Ecaudata
(_Xenopus_, for instance), the epidermis becomes modified in relation
with the termination of sensory nerves, and gives rise to organs of
the same nature as those of the lateral line of fishes. In addition to
diffuse pigment (mostly in the epidermis), the skin contains granular
pigment stored up in cells, the chromatophores, restricted to the
cutis, which are highly mobile and send out branches which, by
contraction and expansion, may rapidly alter the coloration, most
batrachians being in this respect quite comparable to the famous
chameleons. Besides white (guanine) cells, the pigment includes black,
brown, yellow and red. The green and blue, so frequent in frogs and
newts, are merely subjective colours, due to interference. On the
mechanism of the change of colour, cf. W. Biedermann (35).

One of the interesting recent discoveries is that of the "hairy" frog
(_Trichobatrachus_), in which the sides of the body and limbs are
covered with long villosities, the function of which is still unknown
(36).

The nuptial horny asperities with which the males of many batrachians
are provided, for the purpose of clinging to the females, will be
noticed below, under the heading _Pairing and Oviposition_.

_Dentition._--In the Microsauria and Branchiosauria among the
Stegocephalia, as in the other orders, the hollow, conical or slightly
curved teeth exhibit simple or only slightly folded walls. But in the
Labyrinthodonta, grooves are more or less marked along the teeth and
give rise to folds of the wall which, extending inwards and ramifying,
produce the complicated structure, exhibited by transverse sections,
whence these batrachians derive their name; a somewhat similar
complexity of structure is known in some holoptychian (dendrodont)
Crossopterygian fishes. In the remarkable salamander _Autodax_, the
teeth in the jaws are compressed, sharp-edged, lancet shaped. The
teeth are not implanted in sockets, but become ankylosed with the
bones that bear them, and are replaced by others developed at their
bases. Teeth are present in the jaws of all known Stegocephalia and
Apoda and of nearly all Caudata, _Siren_ alone presenting plates of
horn upon the gingival surfaces of the premaxillae and of the dentary
elements of the mandible. But they are nearly always absent in the
lower jaw of the Ecaudata (exceptions in _Hemiphractus,
Amphignathodon, Amphodus, Ceratobatrachus_, the male of
_Dimorphognathus_), many of which (toads, for instance) are entirely
edentulous.

There is great variety in the distribution of the teeth on the palate.
They may occur simultaneously on the vomers, the palatines, the
pterygoids and the parasphenoid in some of the Stegocephalia
(_Dawsonia, Seeleya, Acanthostoma_), on the vomers, palatines and
parasphenoid in many salamandrids (_Plethodontinae_ and
_Desmognathinae_), on the vomers, pterygoids and parasphenoid (some
_Pelobates_), on the vomers and parasphenoid (_Triprion, Amphodus_),
whilst in the majority or other batrachians they are confined to the
vomers and palatines or to the vomers alone (37).

As regards the alimentary organs, it will suffice to state, in this
very brief sketch, that all batrachians being carnivorous in their
perfect condition, the intestine is never very long and its
convolutions are few and simple. But the larvae of the Ecaudata are
mainly herbivorous and the digestive tract is accordingly extremely
elongate and coiled up like the spring of a watch. The gullet is
short, except in the Apoda. The tongue is rudimentary in the
perennibranchiatea Caudata, well developed, and often protrusile, in
the _Salamandridae_ and most of the Ecaudata, totally absent in the
Aglossa.

The organs of circulation cannot be dealt with here; the most
important addition made to our knowledge in recent years being found
in the contributions of F. Hochstetter (38) and of G.B. Howes (39),
dealing with the azygous (posterior) cardinal veins in salamanders and
some of the Ecaudata. The heart is situated quite forward, in the
gular or pectoral region, even in those tailed batrachians which have
a serpentiform body, whilst in the Apoda (fig. 13) it is moved back to
a distance which is comparable to that it occupies in most of the
snakes.

_The Respiratory Organs._--The larynx, which is rudimentary in most of
the Caudata and in the Apoda, is highly developed in the Ecaudata, and
becomes the instrument of the powerful voice with which many of the
frogs and toads are provided. The lungs are long simple tubes in some
of the perennibranchiate Caudata; they generally shorten or become
cellular in the salamandrids, and attain their highest development in
the Ecaudata, especially in such forms as the burrowing _Pelobates_.
Although the lungs are present in such forms as preserve the gills
throughout life, it is highly remarkable that quite a number of
abranchiate salamanders, belonging mostly to the subfamilies
_Desmognathinae_ and _Plethodontinae_, are devoid of lungs and breathe
entirely by the skin and by the bucco-pharyngeal mucose membrane (20).
Some of the _Salamandrinae_ show the intermediate conditions which
have led to the suppression of the trachea and lungs. In the Apoda, as
in many serpentiform reptiles, one of the lungs, either the right or
the left, is much less developed than the other, often very short.

_Urino-genital Organs._--The genital glands, ovaries and testes, are
attached to the dorsal wall of the body-cavity, in the immediate
vicinity of the kidneys, with which the male glands are intimately
connected. The oviducts are long, usually more or less convoluted
tubes which open posteriorly into the cloaca, while their anterior
aperture is situated far forward, sometimes close to the root of the
lung; their walls secrete a gelatinous substance which invests the ova
as they descend. In most male batrachians the testes are drained by
transverse canals which open into a longitudinal duct, which also
receives the canals of the kidneys, so that this common duct conveys
both sperma and urine. In some of the discogloesid frogs, however, the
seminal duct is quite independent of the kidney, which has its own
canal, or true ureter. Many of the Ecaudata have remnants of oviducts,
or Mullerian ducts, most developed in _Bufo_, which genus is also
remarkable as possessing a problematic organ, Bidder's organ, situated
between the testis and the adipose or fat-bodies that surmount it.
This has been regarded by some anatomists as a rudimentary ovary.
Female salamandrids are provided with a _receptaculum seminis_.
Copulatory organs are absent, except in the Apoda, in which a portion
of the cloaca can be everted and acts as a penis. The urinary bladder
is always large.

The spermatozoa have received a great share of attention, on the part
not only of anatomists and physiologists, but even of systematic
workers (40). This is due 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 may differ very
considerably in this respect. The failure to obtain hybrids between
certain species of _Rana_ has been attributed principally to these
differences. The spermatozoa of _Discoglossus_ are remarkable for
their great size, measuring three millimetres in length.

_Pairing and Oviposition_--Batrachians may be divided into four
categories under this head:--(1) no amplexation; (2) amplexation
without internal fecundation; (3) amplexation with internal
fecundation; (4) copulation proper. The first category embraces many
aquatic newts, the second nearly all the Ecaudata, the third the rest
of the Caudata, and the fourth the Apoda.

In the typical newts (_Molge_) of Europe, the males are adorned during
the breeding season with bright colours and crests or other ornamental
dermal appendages, and, resorting to the water, they engage in a
lengthy courtship accompanied by lively evolutions around the females,
near which they deposit their spermatozoa in bundles on a gelatinous
mass, the spermatophore, probably secreted by the cloacal gland. This
arrangement facilitates the internal fecundation of the female without
copulation, the female absorbs the spermatozoa by squeezing them out
of the spermatophore between the cloacal lips. Other newts, and many
salamanders, whether terrestrial or aquatic, pair, the male embracing
the female about the fore limbs or in the pelvic region, and the males
of such forms are invariably devoid of ornamental secondary sexual
characters; but in spite of this amplexation the same mode of
fecundation by means of a spermatophore is resorted to, although it
may happen that the contents of the spermatophore are absorbed direct
from the cloaca of the male. The spermatozoa thus reach the eggs in
the oviducts, where they may develop entirely, some of the salamanders
being viviparous.

In all the tailless batrachians (with the exception of a single known
viviparous toad), the male clings to the female round the breast, at
the arm-pits, or round the waist, and awaits, often for hours or days,
the deposition of the ova, which are immediately fecundated by several
seminal emissions.

The fourth category is represented by the Apoda or Caecilians in
which, as we have stated above, the male is provided with an
intromittent organ. Some of these batrachians are viviparous.

In those species in which the embrace is of long duration the limbs
of the male, usually the fore limbs (pleurodele newt, Ecaudata),
rarely the hind limbs (a few American and European newts), according
to the mode of amplexation, acquire a greater development, and are
often armed with temporary horny excrescences which drop off after the
pairing season. These asperities usually form brush-like patches on
the inner side of one or more of the digits, but may extend over the
inner surface of the limbs and on the breast and chin; the use of them
on these parts is sufficiently obvious, but they are sometimes also
present, without apparent function, on various parts of the foot, as
in _Discoglossus, Bombinator_, and _Pelodytes_. In some species of the
South American frogs of the genus _Leptodactylus_ the breast and hands
are armed with very large spines, which inflict deep wounds on the
female held in embrace.

In most of the Caudata, the eggs are deposited singly in the axils of
water plants or on leaves which the female folds over the egg with her
hind limbs. The eggs are also deposited singly in some of the lower
Ecaudata. In many of the Ecaudata, and in a few of the Caudata and
Apoda, the eggs are laid in strings or bands which are twined round
aquatic plants or carried by the parent; whilst in other Ecaudata they
form large masses which either float on the surface of the water or
sink to the bottom.

A few batrachians retain the ova within the oviducts until the young
have undergone part or the whole of the metamorphosis. Viviparous
parturition is known among the Caudata (_Salamandra, Spelerpes
fuscus_), and the Apoda (_Dermophis thomensis, Typhlonectes
compressicauda_); also in a little toad (_Pseudophryne vivipara_)
recently discovered in German East Africa (41).

_Development and Metamorphosis._--In a great number of batrachians,
including most of the European species, the egg is small and the
food-yolk is in insufficient quantity to form an external appendage of
the embryo. But in a few European and North American species, and in a
great many inhabitants of the tropics, the egg is large and a
considerable portion of it persists for a long time as a yolk-sac.
Although the segmentation is always complete, it is very irregular in
these types, some of which make a distinct approach to the meroblastic
egg.

With the exception of a number of forms in which the whole development
takes place within the egg or in the body of the mother, batrachians
undergo metamorphoses, the young passing through a free-swimming,
gill-breathing period of considerable duration, during which their
appearance, structure, and often their _regime_, are essentially
different from those of the mature form. Even the fossil Stegocephalia
underwent metamorphosis, as we know from various larval remains first
described as _Branchiosaurus_. They are less marked or more gradual in
the Apoda and Caudata than in Ecaudata, in which the stage known as
tadpole is very unlike the frog or toad into which it rather suddenly
passes (see TADPOLE). In the Caudata, external gills (three on each
side) persist until the close of the metamorphosis, whilst in the
Apoda and Ecaudata they exist only during the earlier periods, being
afterwards replaced by internal gills.

Many cases are known in which the young batrachian enters the world in
the perfect condition, as in the black salamander of the Alps
(_Salamandra atra_), the cave salamander (_Spelerpes fuscus_), the
caecinan _Typhlonectes_, and a number of frogs, such as _Pipa,
Rhinoderma, Hylodes_, some _Nototrema, Rana opisthodon_, &c. A fairly
complete bibliographical index to these cases and the most remarkable
instances of parental care in tailless batrachians will be found in
the interesting articles by Lilian V. Sampson (42), and by G. Brandes
and W. Schoenichen (43). It will suffice to indicate here in a
synoptic form, as was done by the present writer many years ago, when
our knowledge of these wonders of batrachian life was far less
advanced than it is now, the principal modes of protection which are
resorted to:--

1. Protection by means of nests or nurseries.

A. In enclosures in the water.--_Hylafaber_.
B. In nests in holes near the water.--_Rhacophorus, Leptodactylus_.
C. In nests overhanging the water.--_Rhacophorus, Chiromantis,
Phyllpmedusa_.
D. On trees or in moss away from the water.--_Rana opisthodon,
Hylodes, Hylelia platycephala_.
E. In a gelatinous bag in the water.--_Phrynixalus, Salamandrella_.

2. Direct nursing by the parents.

A. Tadpoles transported from one place to another.--_Dendrebates,
Phyllobates, Sooglossus_.
B. Eggs protected by the parents who coil themselves round or "sit"
on them.--_Mantophryne, Desmognathus, Autodax, Plethodon,
Cryptobranchus, Amphiuma, Ichthyophis, Hypogeophis, Siphonops_.
C. Eggs carried by the parents.
(a) Round the legs, by the male.--_Alytes_.
(b) On the back, by the female.
(1) Exposed.--_Hyla goeldii, H. evansii, Ceratohyla_.
(2) In cell-like pouches.--_Pipa_.
(3) In a common pouch.--_Nototrema, Amphignathodon_.
(c) On the belly.
(1) Exposed, by the female.--_Rhacophorus reticulatus_.
(2) In a pouch (the produced vocal sac), by the
male.--_Rhinoderma_.
(d) In the mouth, by the female.--_Hylambates brevirostris_.

_Geographical Distribution._--If a division of the world according to
its batrachian faunae were to be attempted, it would differ very
considerably from that which would answer for the principal groups of
reptiles, the lizards especially. We should have four great
realms:--(1) Europe and Northern and Temperate Asia, Africa north of
the Sahara (palaearctic region) and North and Central America
(nearctic region); (2) Africa and South-Eastern Asia (Ethiopian and
Indian region); (3) South America (neotropical region); and (4)
Australia (Australian region). The first would be characterized by the
Caudata, which are almost confined to it (although a few species
penetrate into the Indian and neotropical regions), the
_Discoglossidae_, mostly Europaeo-Asiatic, but one genus in
California, and the numerous _Pelobatidae_; the second by the presence
of Apoda, the prevalence of firmisternal Ecaudata and the absence of
_Hylidae_; the third by the presence of Apoda, the prevalence of
arciferous Ecaudata and the scarcity of _Ranidae_, the fourth by the
prevalence of arciferous Ecaudata and the absence of _Ranidae_, as
well as by the absence of either Caudata or Apoda. Madagascar might
almost stand as a fifth division of the world, characterized by the
total absence of Caudata, Apoda, and arciferous Ecaudata. But the
close relation of its very rich frog-fauna to that of the Ethiopian
and Indian regions speaks against attaching too great importance to
these negative features. It may be noted here that no two parts of the
world differ so considerably in their Ecaudata as do Madagascar and
Australia, the former having only Firmisternia, the latter only
Arcifera. Although there is much similarity between the Apoda of
Africa and of South America, one genus being even common to both parts
of the world, the frogs are extremely different, apart from the
numerous representatives of the widely distributed genus _Bufo_. It
may be said that, on the whole, the distribution of the batrachians
agrees to some extent with that of fresh-water fishes, except for the
much less marked affinity between South America and Africa, although
even among the former we have the striking example of the distribution
of the very natural group of the aglossal batrachians, represented by
_Pipa_ in South America and by _Xenopus_ and _Hymenochirus_ in Africa.

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BATRACHOMYOMACHIA (Gr. [Greek: Batrachos], "frog," [Greek: mus], "mouse," and [Greek: machae], "battle"), the "Battle of Frogs and Mice," a comic epic or parody on the _Iliad_, definitely attributed to Homer by the Romans, but according to Plutarch (_De Herodoti Malignitate_, 43) the work of Pigres of Halicarnassus, the brother (or son) of Artemisia, queen of Caria and ally of Xerxes. Some modern scholars, however, assign it to an anonymous poet of the time of Alexander the Great.

Edition by A. Ludwich (1896).

BATTA, an Anglo-Indian military term, probably derived from the Canarese _bhatta_ (rice in the husk), meaning a special allowance made to officers, soldiers, or other public servants in the field.

BATTAGLIA, a town of Venetia, Italy, in the province of Padua, 11 m. S.S.W. by rail from Padua. Pop. (1901) 4456. It lies at the edge of the volcanic Euganean Hills, and is noted for its warm saline springs and natural vapour grotto. A fine palace was erected in the Palladian style in the 17th century by Marchese Benedetto Selvatico-Estense, then owner of the springs.

BATTAKHIN, African "Arabs" of Semitic stock. They occupy the banks of the Blue Nile near Khartum, and it was against them that General Gordon fought most of his battles near the town. Their sheikh, El Obeid, routed Gordon's troops on the 4th of September 1884, a defeat which led to the close investment of Khartum. In the 18th century James Bruce described them as "a thieving, pilfering lot."

BATTALION, a unit of military organization consisting of four or more companies of infantry. The term is used in nearly every army, and is derived through Fr. from It. _battaglione_, Med. Lat. _battalia_ (see BATTLE). "Battalion" in the 16th and 17th centuries implied a unit of infantry forming part of the line of battle, but at first meant an unusually large _battalia_ or a single large body of men formed of several _battalias_. In the British regular service the infantry battalion is commanded by a lieut.-colonel, who is assisted by an adjutant, and consists at war strength of about 1000 bayonets in eight companies. Engineers, train, certain kinds of artillery, and more rarely cavalry are also organized in battalions in some countries.

BATTAMBANG, or BATTAMBONG (locally _Phralabong_), the chief town of the north-western division of Cambodia, formerly capital of Monton Kmer, i.e. "The Cambodian Division," one of the eastern provinces of Siam, now included in the French protectorate of Cambodia. It is situated in 103 deg. 6' E., 13 deg. 6' N., in the midst of a fertile plain and on the river Sang Ke, which flows eastwards and falls into the Tonle or Tale Sap, the great lake of Cambodia. The town is a collection of bamboo houses of no importance, but there is a walled enceinte of some historical interest. Trade is small and is carried on by Chinese settlers, chiefly overland with Bangkok, but to a small extent also by water with Saigon. The population is about 5000, two-thirds Cambodian and the remainder Chinese and Siamese. The language is Cambodian.

Battambang was taken by the Siamese when they overran the kingdom of Cambodia towards the end of the 18th century, and was recognized by the French as belonging to Siam when the frontier of Cambodia was adjusted by treaty in 1867-1872. In another treaty in 1893, Siam bound herself to maintain no armed forces there other than police, but this arrangement was annulled by the treaty of 1904, by which Battambang was definitely admitted to lie within the French sphere of influence. Under a further treaty in March 1907 (see SIAM), the district of Battambang was finally ceded to the French.

BATTANNI, or BHITANI, a small tribe on the Waziri border of the North-West Frontier Province of India. The Battannis hold the hills on the borders of Tank and Bannu in the Dera Ismail Khan district, from the Gabar mountain on the north to the Gomal valley on the south. They are only 3000 fighting men strong, and are generally regarded as the jackals of the Waziris. Their chief importance arises from the fact that no raids can be carried into British districts by the Mahsud Waziris without passing through Battanni territory. A small British expedition against the Battannis was led by Lt.-Col. Rynd in 1880. Under the excitement caused by the preaching of a fanatical mullah the Mahsud Waziris had attacked the town of Gomal. The Battannis failed to supply information as to their movements, and gave them a passage through their lands. The British troops accordingly stormed the Hinis Tangi defile in face of opposition, and burned the village of Jandola.

BATTAS (Dutch _Battaks_), the inhabitants of the formerly independent Batta country, in the central highlands of Sumatra, now for the most part subjugated to the Dutch government. The still independent area extends from 98 deg.-99 deg. 35' E., and 2 deg.-3 deg. 25' S. North-east of Toba Lake dwell the Timor Battas, and west of it the Pakpak, but on its north (in the mountains which border on the east coast residency) the Karo Battas form a special group, which, by its dialects and ethnological character, appears to be allied to the Gajus and Allas occupying the interior of Achin. The origin of the Battas is doubtful. It is not known whether they were settled in Sumatra before the Hindu period. Their language contains words of Sanskrit origin and others referable to Javanese, Malay and Tagal influence. Their domain has been doubtless much curtailed, and their absorption into the Achin and Malay population seems to have been long going on. The Battas are undoubtedly of Malayan stock, and by most authorities are affiliated to that Indonesian pre-Malayan race which peopled the Indian Archipelago, expelling the aboriginal negritos, and in turn themselves submitting to the civilized Malays. In many points the Battas are physically quite different from the Malay type. The average height of the men is 5 ft. 4 in.; of the women 4 ft. 8 in. In general build they are rather thickset, with broad shoulders and fairly muscular limbs. The colour of the skin ranges from dark brown to a yellowish tint, the darkness apparently quite independent of climatic influences or distinction of race. The skull is rather oval than round. In marked contrast to the Malay type are the large, black, long-shaped eyes, beneath heavy, black or dark brown eyebrows. The cheek-bones are somewhat prominent, but less so than among the Malays. The Battas are dirty in their dress and dwellings and eat any kind of food, though they live chiefly on rice. They are remarkable as a people who in many ways are cultured and possess a written language of their own, and yet are cannibals. The more civilized of them around Lake Toba are good agriculturists and stock-breeders, and understand iron-smelting. They weave and dye cotton, make jewellery and krisses which are often of exquisite workmanship, bake pottery, and build picturesque chalet-like houses of two storeys. They have an organized government, hereditary chiefs, popular assemblies, and a written civil and penal code. There is even an antiquated postal system, the letter-boxes being the hollow tree trunks at crossroads. Yet in spite of this comparative culture the Battas have long been notorious for the most revolting forms of cannibalism. (See _Memoirs of the Life, &c., of Sir T.S. Raffles_, 1830.)

The Battas are the only lettered people of the Indian Archipelago who are not Mahommedans. Their religion is mainly confined to a belief in evil spirits; but they recognize three gods, a Creator, a Preserver and a Destroyer, a trinity suggestive of Hindu influence.

Up to the publication of Dr H.N. van der Tuuk's essay, _Over schrift en uitspraak der Tobasche taal_ (1855), our knowledge of the Batta language was confined to lists of words more or less complete, chiefly to be found in W. Marsden's _Miscellaneous Works_, in F.W. Junghuhn's _Battalander_, and in the _Tijdschrift van het Bataviaasch Genootschap, _vol. iii. (1855). By his exhaustive works (_Bataksch Leesboek_, in 4 vols., 1861-1862; _Bataksch-nederduitsch Woordenboek_, 1861; _Tobasche Spraakkunst_, 1864-1867) van der Tuuk made the Batta language the most accessible of the various tongues spoken in Sumatra. According to him, it is nearest akin to the old Javanese and Tagal, but A. Schreiber (_Die Battas in ihrem Verhaltnis zu den Malaien von Sumatra_, 1874) endeavoured to prove its closer affinity with the Malay proper. Like most languages spoken by less civilized tribes, Batta is poor in general terms, but abounds in terms for special objects. The number of dialects is three, viz. the Toba, the Mandailing and the Dairi dialects; the first and second have again two subdivisions each. The Battas further possess six peculiar or recondite modes of speech, such as the _hata andung_, or language of the wakes, and the _hata poda_ or the soothsayer's language. A fair acquaintance with reading and writing is very general among them. Their alphabet is said, with the Rejang and Lampong alphabets, to be of Indian origin. The language is written on bark or bamboo staves from bottom to top, the lines being arranged from left to right. The literature consists chiefly in books on witchcraft, in stories, riddles, incantations, &c., and is mostly in prose, occasionally varied by verse.[1]

See also "Reisen nach dem Toba See," _Petermanns Mitteil_. (1883);
Modigliani, _Fra i Batacchi indipendenti_ (Rome, 1892); Neumann, "Het
Pane- en Bilastroomgebied," _Tydschr. Aardr. Gen._, 1885-1887; Van
Dijk in the same periodical (1890-1895); Wing Easton in the _Jaarboek
voor het Mynwezen_, 1894; Niemann in the _Encyclopaedia van
Nederlandsch-Indie_, under the heading _Bataks_, with very detailed
bibliography; Baron J. v. Brenner, _Besuch bei den Kannibalen
Sumatras_ (Wurzburg, 1893); H. Breitenstein, _21 Jahre in Indien,
Java, Sumatra_ (Leipzig, 1899-1900); G.P. Rouffaer, _Die Batik-Kunst
in niederlandisch-Indien und ihre Geschichte_ (Haarlem, 1899).

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