The quadrato-jugal elements are reduced to ligaments. In many Salamandrinae the large orbito-temporal space is divided into an orbital and a temporal fossa by an arch which is formed by the meeting of two corresponding processes from the squamosal and frontal bones respectively. This bridge is rarely bony (Salamandrina, Triton), mostly ligamentous;–apparently a reminiscence of the Stegocephalous condition. The two premaxillary bones are liable to fuse into one, for instance in Cryptobranchus, generally in adult Tritons. They are most reduced, and are toothless, in Siren.
The two maxillary bones are absent only in Necturus, Proteus, Typhlomolge, and Siren. Their posterior end is frequently free, loosely connected by ligaments with the pterygoid in Cryptobranchus; or with the distal portion of the quadrate, and in this case either just touching it (Tylototriton), or forming a broad junction (Pachytriton).
Each half of the lower jaw consists of a dentary, articular and angulo-splenial. The splenial remains as a separate element in Siren; in others only during the larval period. There are no mento-Meckelian elements.
Skeleton of the Anura
The vertebral column.–The distinctive peculiarities of the vertebrae of the Anura are that they are notocentrous, and that about a dozen of them are modified and fused into an os coccygeum. The whole column is the most specialised found in the Vertebrata; and various stages are rapidly hurried through and obscured caenogenetically during the embryonic development. Paired cartilages appear on the dorsal side of the thin chordal sheath, and whilst tending to enclose the spinal cord in a canal, their bases grow head- and tail-wards into what will ultimately become the intervertebral region. This extension of cartilage leads to a fusion with that of the next following pair of arches, so that the axial column at this early stage consists of a right and left longitudinal ridge of cartilage which sends off dorsal processes, neural arches, in metameric succession. Next, the intervertebral cartilage increases in such a way as to constrict the chorda either laterally (Rana) or obliquely from above downwards and inwards (Bufo, Hyla). We recognise in this cartilage the interdorsalia. Ventral arcualia are late and much obscured. There is scarcely any cartilage which could represent the interventralia, the intervertebral cartilage being almost entirely made up of the interdorsalia. These fuse together and form a disc or nodule, which later fuses either with the vertebra in front, and in this case fits into a cup carried by the vertebra next behind (procoelous vertebrae), or the knob is added to the front end of the vertebra, fitting into a cup formed by the tail end of the vertebra next in front (opisthocoelous vertebrae). Much later than the two longitudinal dorsal bands there appears on the ventral side an unpaired band in which appear metamerically repeated swellings of cartilage, likewise unpaired. These swellings become confluent, in a way similar to that which produced the dorsal bands, and form the unpaired ventral band of cartilage, the hypochordal cartilage of some authors. The swellings in this band, equivalent to the basiventralia, become semilunar in a transverse view, their horns tending upwards towards the basidorsal cartilages, but there is no actual meeting. Both dorsal and ventral elements are, however, joined together and form the chief portion of the vertebrae, owing to the rapidly proceeding calcification and later ossification of the all-surrounding "membrana reuniens" or skeletogenous layer so far as that is not cartilaginous.
Procoelous vertebrae exist in the overwhelming majority of Anura; opisthocoelous are those of the Aglossa, the Discoglossidae, and of some Pelobatidae. The systematic value of this pro- or opistho-coelous character has been much exaggerated. We have seen that the centra of the vertebrae of the Anura are formed entirely by the interdorsal elements, hence the term "notocentrous," and these centra sometimes remain in adult specimens of Pelobates as separately ossified and calcified pieces, not fused with the rest of the vertebrae. This important discovery has been made by Boulenger, but Stannius had previously mentioned a specimen of Pelobates in which the second and fourth vertebrae are biconvex, the third, sixth, and eighth biconcave. Moreover, since the sacral vertebra, generally the ninth, in all the Anura is invariably biconvex, the eighth being biconcave in the procoelous families, opisthocoelous like the remaining seven vertebrae in the other families, it is not difficult to imagine that in the Anura the production of pro- or opistho-coelous vertebrae depends simply upon the centra or articulating knobs happening to fuse either with the hind or the front end of the vertebrae. This must of course ultimately be determined by a mechanical problem of motion.
A second type of the vertebrae amongst the Anura is the epichordal type, an exaggeration in degree of the notocentrous tendencies of the more usual perichordal arrangement. It shows, namely, the almost complete suppression of all the ventral cartilaginous elements, so that the chorda remains for a long time on the ventral surface of the axial column in the shape of a flattened longitudinal band. These two types are not unconnected. The suppression of the ventral elements applies most typically to the trunk region, while hypochordal cartilage exists in the anterior cervical vertebrae, and above all in the coccyx. Typically epichordal are the vertebrae of Pipa, Xenopus, Bombinator, Pelobates, Discoglossus and Alytes. It is significant that the epichordal often coincide with opisthocoelous vertebrae, and still more suggestive is the fact that Bombinator is eminently aquatic, Pipa and Xenopus entirely so, having lost the tympanum, at least externally. The epichordal feature is not necessarily indicative of relationship. It has probably been developed independently in various groups, in correlation with a resumption of aquatic life. Various genera of Pelobatidae and most likely some Cystignathidae, e.g. Pseudis, will not improbably connect the two types and their several correlated features, for instance, the frequent reduction of the tympanic cavity.
The os coccygeum has retained rather primitive features in so far as much dorsal and ventral cartilage is developed; but this has almost entirely lost its metameric arrangement, and the posterior half of the coccyx is formed chiefly by the ventral mass of cartilage, while the dorsal elements are more or less reduced. Only two vertebrae, generally the tenth and eleventh of the whole column, are clearly visible, each being composed of a pair of dorsal and a pair of ventral cartilaginous blocks. The sacral vertebra articulates with the coccyx by one or two convexities, but in the Aglossa, in some Pelobatidae, and a few others, the coccyx is fused with the sacral vertebra. Beyond the first and second component vertebrae of the embryonic coccyx, the cartilage is continued in the shape of two dorsal, and one ventral, bands, which soon fuse with each other. Dorsally this cartilage surrounds the spinal cord; the latter degenerates towards the end of the tadpole-stage, leaving, however, the empty spinal canal. The chorda, completely surrounded by cartilage, persists into the post-larval stage, but is destroyed long before the creature attains maturity. Ultimately the whole coccyx ossifies.
The tail proper, namely that portion which is absorbed during the metamorphosis, remains throughout its existence in an apparently primitive condition. The chorda dorsalis and the spinal cord extend through its whole length, surrounded by continuous connective tissue without any cartilage; in fact it represents a piece of typical vertebral column before the appearance of cartilage. The reduction of this swimming organ begins at the hind end.
The vertebral column of the adult.–The first vertebra (we will call it the atlas since it carries the skull) is not, as in the Urodela, provided with an odontoid process. It articulates by two cups with the condyles of the occiput. In some Anura it co-ossifies, rather incompletely, with the second vertebra, regularly in the fossil Palaeobatrachus, often in Ceratophrys, Breviceps, and occasionally in Pelobates, Bufo, Rana, and Xenopus. This is, however, no justification for looking upon the first vertebra as a complex of two vertebrae, although the atlas is frequently very thick and broad, and even carries, in the Aglossa, considerable lateral wings or diapophyses. Those of the trunk-vertebrae are often very long, acting thereby as substitutes for ribs which are absent, except on the second, third, and fourth vertebrae of the Discoglossidae, and on the second and third of the Aglossa. In the adult Aglossa these ribs fuse with the processes which carry them.