The arrangement of the proximal and middle segments of the limb is fairly constant in all reptiles with limbs adapted for walking, and the tibia and fibula are always separate. The pes is however subject to a considerable amount of variation, especially as regards the tarsus. In some Chelonia the tarsus like the carpus has an extremely simple arrangement, consisting of a proximal row of three bones, the tibiale, intermedium and fibulare, a centrale, and a distal row of five tarsalia. In most living reptiles, however, the tibiale and intermedium are as in mammals united, forming the astragalus. In Crocodiles (fig. 48, B, 15) the centrale is also united with the tibiale while the distal tarsalia are very slightly developed. The calcaneum in Crocodiles is drawn out into a long process forming a heel in a manner almost unique among Sauropsida. In Sphenodon and Lacertilia the tibia and fibula articulate with a single large bone representing the whole proximal row of tarsalia.
The pes is generally pentedactylate, but in some Crocodiles the fifth digit is vestigial (fig. 48, B), and in some Dinosauria (fig. 35) there are only three digits. The North American Dinosaurs present a continuous series ranging from a pentedactylate plantigrade form like Morosaurus, to such a form as Hallopus with a highly digitigrade and specialised pes reduced to three functional digits, and a vestigial fifth metatarsal. The second, third and fourth metatarsals in this form are nearly two-thirds as long as the femur, and the calcaneum is drawn out into a heel much as it is in most mammals.
In Lacertilia, Orthopoda and many Chelonia, the ankle joint comes to lie between the proximal and distal row of tarsals as in birds.
CHAPTER XVII.
CLASS. AVES[94].
Birds form a large and extremely homogeneous class of the vertebrata, and are readily distinguished from all other animals by the possession of an epidermal exoskeleton having the form of feathers. Feathers differ from hairs in the fact that they grow from papillae formed of both the horny and the Malpighian layer of the epidermis, which papillae at first project from the surface, and only subsequently become imbedded in pits of the dermis. A dermal exoskeleton does not occur in birds.
The endoskeleton is characterised by its lightness, the large bones being generally hollow; but the pneumaticity does not vary in proportion to the power of flight. The cervical part of the vertebral column is very long and flexible, while the post-cervical portion is generally very rigid, owing to the fusion of many of the vertebrae, especially in the lumbar and sacral regions. The vertebrae are generally without epiphyses to their centra. The cervical vertebrae in living forms have saddle-shaped articulating surfaces, and many of them bear ribs. The thoracic ribs in almost all birds have large uncinate processes. The sternum is very large, and the ribs are always attached to its sides, not as in many reptiles to any backwardly-projecting process or processes. The sternum ossifies from two or more centres.
The skull is extremely light, and its component bones show a great tendency to fuse together completely. The facial part of the skull is prolonged into a beak, chiefly formed of the premaxillae; this beak is in all modern birds devoid of teeth, and is coated externally with a horny epidermal sheath. The quadrate is large and freely movable. The supratemporal arcade[1] is imperfect, while the infratemporal arcade[95] is complete. There are no postorbital or postfrontal bones. Neither parotic processes nor an interparietal foramen occur. There are commonly large pre-orbital vacuities. The palatines and pterygoids never form a secondary bony palate as in Crocodiles. Part of the floor of the skull is formed by a wide basitemporal (paired in the embryo) which is continued in front as a long slender rostrum; these structures have replaced the parasphenoid of Ichthyopsids. Cartilage or bone is always developed in the sclerotic. The first branchial arch is well developed, the hyoid arch but slightly. The coracoids are large, and the clavicles are nearly always united forming the furcula. There is no separate interclavicle and hardly any trace of a precoracoid.
The anterior limbs form wings, and the manus is in the adult always much modified, never having more than three digits. The three bones of the pelvis are, except in Archaeornithes, always ankylosed together in the adult, and the ilium is greatly prolonged in front of the acetabulum, which is perforated. The ilia are not connected with the sacrum by ossified sacral ribs. The pubes and ischia are directed backwards parallel to one another, and except in a very few forms never meet their fellows in ventral symphyses. The fibula is generally much reduced. The proximal tarsal bones are always ankylosed to the tibia, and the distal tarsals to the metatarsals, so that the ankle joint is intertarsal. The first metatarsal is nearly always free. The pes never has more than four digits in the adult.