put. It consists of (1) the basihyal variously called copula, or corpus linguae, or unpaired middle portion. (2) The urohyal likewise unpaired, rested ventrally on the larynx. (3) The os entoglossum originally paired, but coalescing into an arrow-headed piece, attached to the anterior end of the basihyal and lodged in the tongue proper. It is homologous with the distal ends of the ceratohyals or ventral elements of the hyoidean or second visceral arch. The dorsal or hyomandibular portion of this same arch is transformed into the auditory chain, ending in the fenestra ovalis. (4) A pair of thyrohyals, homologous with the posterior hyoid horns of mammals, i.e. third visceral or first branchial arch. As the most developed pair in birds they are commonly, although wrongly, called the hyoid horns. They articulate upon facets of the hinder outer corners of the basihyal.
The vertebrae are stereospondylous, the centrum or body and the arch being completely fused into one mass, leaving not even a neuro-central suture. The arch alone sends out processes, viz. the spinous process, the anterior and posterior oblique (commonly called pre- and post-zygapophyses), and the transverse processes. The latter articulate with the tuberculum of the corresponding rib, while the capitulum articulates by a knob on the side of the anterior end of the centrum. In the cervical region the ribs are much reduced, fused with their vertebrae and enclosing the transverse canal or foramen. When the vertebrae are free their centra articulate with each other by complicated joints, exhibiting four types. (1) Amphicoelous; each end of the centrum is concave; this, the lowest condition, is embryonic, but was retained in Archaeopteryx and in the thoracic vertebrae of Ichthyornis. (2) Procoelous, concave in front; only in the atlas, for the reception of the occipital condyle. (3) Opisthocoelous, or concave behind, only occasionally found in the thoracic region, e.g. Sphenisci. (4) Heterocoelous (fig. 8) or saddle-shaped; the anterior surface is concave in a transverse, but convex in a vertical direction, which on posterior surface shows the conditions reversed. This is the most perfect arrangement attained by the vertebral column, and is typical of, and restricted to, birds. The intervertebral joints are further complicated by the interposition of a cartilaginous or fibrous pad or ring. This pad varies much; it is morphologically the homologue of the pair of basiventral elements which by their lateral extension give origin to the corresponding ribs. Later those pads fuse with the anterior end of the centrum of the vertebra to which they belong; where the vertebral column is rendered inflexible, the disks are ossified with the centra and all trace of them is lost. Sometimes the pad is reduced to a ventral semi-ring or meniscus; it retains its largest almost original shape and size in the second vertebra, the axis or epistropheus, where it forms a separately ossifying piece which connects, and coössifies with, the odontoid process (the centrum of the atlas) and the centrum of the second vertebra. Sometimes the ventral portions of these pads form paired or unpaired little ossifications, then generally described as intercentra; such are not uncommon on the tail. The atlas is composed of three pieces; a pair of lateral elements (the right and left dorsal arch pieces) joining above the spinal cord, and a ventral piece equivalent to the first basiventral elements, i.e. serially homologous with the intervertebral pads. In the adults the atlas forms a more or less solid ring. A remnant of the chorda dorsalis and its sheath persists as the ligamentum suspensorium between the central portions of the successive vertebrae.
|Fig. 8.—A cervical vertebra from the middle of the neck of a|
Fowl; natural size. a, Side view; b, upper view; c, lower view;
pr.z, pre-zygapophyses; pt.z, post-zygapophyses.
In birds we distinguish between the following regions of the axial skeleton. (1) Cervical vertebrae, or those between the skull and the first vertebra which is connected with the sternum by a pair of complete ribs. The last 1 to 5 of these vertebrae have movable ribs which do not reach the sternum, and are called cervico-dorsals. (2) Dorsals, those which begin with the first thoracic rib, and end at the last that is not fused with the ilium. The term “lumbar” vertebrae is inapplicable to birds. (3) Pelvic, all those which are fused with the iliac portion of the pelvis, generally a considerable number. (4) Caudal, those which are not connected with the pelvis. It is to be noted that often no absolute line of demarcation can be drawn in regard to these regions, their definitions being rather convenient than morphological.
|Fig. 9.—The “sacrum” of a young|
Fowl; natural size, seen from below.
d.l, Dorso-lumbar, s, sacral, c, caudal
In comparison with all other vertebrates the number of neck-vertebrae of the birds is considerably increased; the lowest number, 14 to 15, is that of most Passeres and many other Coraciomorphae; the largest numbers, 20 or 21, are found in the ostrich, 23 in Cygnus olor and 25 in the black swan. Dorsal vertebrae frequently have a ventral outgrowth of the centrum; these hypapophyses may be simple vertical blades, ⊥-shaped, or paired knobs; they serve for the attachment of the thoracic origin of the longus collianticus muscle, reaching their greatest development in Sphenisci and Colymbidae. In many birds some of the thoracic vertebrae are more or less coössified, in most pigeons for instance the 15th to 17th; in most Galli the last cervical and the next three or four thoracics are coalesced, &c. The pelvic vertebrae include of course the sacrum. There are only two or three vertebrae which are equivalent to those of the reptiles; these true sacrals are situated in a level just behind the acetabulum; as a rule between these two primary sacral vertebrae issues the last of the spinal nerves which contributes to the composition of the sciadic plexus. These true sacrals alone are connected with the ilium by processes which are really equivalent to modified ribs; but the pelvis of birds extends considerably farther forwards and backwards, gradually coming into contact with other vertebrae, which in various ways send out connecting transverse processes or buttresses, and thus become pre- and post-sacral vertebrae (fig. 9). The most anterior part of the ilium often overlaps one or more short lumbar ribs and fuses with them, or even a long, complete thoracic rib. Similarly during the growth of the bird the posterior end of the ilium connects itself with the transverse processes of vertebrae which were originally free, thus transforming them from caudals into secondary post-sacrals. Individual, specific and generic variations are frequent.
|Fig. 10.—A side view of the|
The last six or seven caudal vertebrae coalesce into the pygostyle, an upright blade which carries the rectrices. Such a pygostyle is absent in Archaeopteryx, Hesperornis, Tinami and Ratitae, but it occurs individually in old specimens of the ostrich and the kiwi. In Ichthyornis it is very small. In all the Neornithes the total number of caudal vertebrae, inclusive of those which coalesce, is reduced to at least 13.
Sternum (figs. 10 and 11).—Characteristic features of the sternum are the following. There is a well-marked processus lateralis anterior (the right and left together equivalent to the mammalian manubrium), which is the product of two or three ribs, the dorsal parts of which reduced ribs remain as cervico-dorsal ribs. Then follows the rib-bearing portion and then the processus lateralis posterior; this also is the product of ribs, consequently the right and left processes together are equivalent to the xiphoid process or xiphisternum of the mammals. The lateral process in most birds sends out an outgrowth, directed out and upwards, overlapping some of the ribs, the processus obliquus. The median and posterior extension of the body of the sternum is a direct outgrowth of the latter, therefore