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527
BATRACHIA

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.

1911 Britannica - Batrachia - Ichthyophis glutinosus1.png
Fig. 13.—Ventral view of
the head and trunk of
Ichthyophis glutinosus.
Mn, Mandible.

Hy, Hyoid.
Br1 Br2, Br3, 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 Müllerian 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