CHAPTER VIII.
GENERAL ACCOUNT OF THE SKELETON IN FISHES[39].
EXOSKELETON.
The most primitive type of exoskeleton is that found in Elasmobranchs and formed of placoid scales; these are tooth-like structures consisting of dentine and bone capped with enamel, and have been already described (p. 4). In most Elasmobranchs they are small and their distribution is fairly uniform, but in the Thornback skate, Raia clavata, they have the form of larger, more scattered spines. In adult Holocephali and in Polyodon and Torpedo there is no exoskeleton, in young Holocephali, however, there are a few small dorsal ossifications.
The plates or scales of many Ganoids may have been formed by the gradual fusion of elements similar to these placoid scales, and often bear a number of little tooth-like processes. In Lepidosteus, Polypterus, and many extinct species, these ganoid scales, which are rhomboidal in form and united to one another by a peg and socket articulation, enclose the body in a complete armour. In Trissolepis part of the tail is covered by rhomboidal scales, while rounded scales cover the trunk and remainder of the tail. Acipenser and Scaphirhynchus have large dermal bony plates which are not rhomboidal in shape and do not cover the whole body. In Acipenser a single row extends along the middle of the back and two along each side.
The majority of Teleosteans have thin flattened scales which differ from those of Ganoids in being entirely mesodermal in origin, containing no enamel. There are two principal types of Teleostean scales, the cycloid and ctenoid. A cycloid scale is a flat thin scale with concentric markings and an entire posterior margin. A ctenoid scale differs in having its posterior margin pectinate. The Dipnoi have overlapping cycloid scales. The rounded scales of Amia and of many fossil ganoids such as Holoptychius are shaped like cycloid scales, but differ from them in being more or less coated with enamel. In Eels and some other Teleosteans the scales are completely degenerate and have almost disappeared. Some Teleosteans, like Diodon hystrix, have scales with triradiate roots from which arise long sharp spines directed backwards. These scales, which resemble teeth, contain no enamel; they become erect when the fish inflates its body into a globular form. Many Siluroids have dermal armour in the form of large bony plates which are confined to the anterior part of the body. In Ostracion the whole body is covered by hexagonal plates, closely united together.
The fin-rays are structures of dermal origin which entirely or partially support the unpaired fins, and assist the bony or cartilaginous endoskeleton in the support of the paired fins.
In Elasmobranchs, Dipnoi, and Chondrosteous ganoids the skeletons of the fins are, as a rule, about half of exoskeletal, half of endoskeletal origin, the proximal and inner portion being cartilaginous and endoskeletal, the distal and outer portion being exoskeletal, and consisting of horny or of more or less calcified fin-rays. In bony Ganoids and Teleosteans the endoskeletal parts are greatly reduced and the fins come to consist mainly of the fin-rays, which are ossified and frequently become flattened at their distal ends.
The fin-rays of the ventral part of the caudal fin are carried by the haemal arches; those of the dorsal and anal fins and of the dorsal part of the caudal fin generally by interspinous bones, which in adult Teleosteans alternate with the neural and haemal spines. In Dipnoi these interspinous bones articulate with the neural and haemal spines. In many Siluroids the anterior rays of the dorsal and pectoral fins are developed into large spines which often articulate with the endoskeleton, or are sometimes fused with the dermal armour plates. Similar spines may occur in Ganoids in front of both the dorsal and anal fins. Polypterus has a small spine or fulcrum in front of each segment of the dorsal fin. Such spines are often found fossilised, and are known as ichthyodorulites.
Similar spines are found in many Elasmobranchs, but they are simply inserted in the flesh, not articulated to the endoskeleton. They also differ from the spines of Teleosteans and Ganoids in the fact that they are covered with enamel, and often have their edges serrated like teeth. In the extinct Acanthodii they generally occur in front of all the fins, paired and unpaired.
In Trygon, the Sting-ray, the tail bears a serrated spine which is used for purposes of offence and defence. Many ichthyodorulites may have been spines of this nature fixed to the tail, rather than spines situated in front of the fins. The spines, which are always found in front of the dorsal fin in Holocephali, agree with those of Elasmobranchs in containing enamel, and with those of Teleosteans in being articulated to the endoskeleton.