Page:EB1911 - Volume 14.djvu/272

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260
[ANATOMY
ICHTHYOLOGY

In such an archaic Selachian as Pleuracanthus the fin is clearly of the biserial archipterygial type, but the lateral rays are reduced (pectoral) or absent (pelvic) (fig. 17, a) on one side of the axis. In a typical adult Selachian the pectoral fin skeleton has little apparent resemblance to the biserial archipterygium—the numerous outwardly directed rays springing from a series of large basal cartilages (pro-, meso- and metapterygium). The condition in the young (e.g. fig. 17, b, Acanthias) hints strongly, however, at the possibility of the fin skeleton being really a modified biserial archipterygium, and that the basal cartilages represent the greatly enlarged axis which has become fixed back along the side of the body. In Crossopterygians (Polypterus) the highly peculiar fin skeleton (fig. 18) while still in the embryonic cartilaginous stage is clearly referable to a similar condition. In the Actinopterygians—with the increased development of dermal fin rays—there comes about reduction of the primitive limb skeleton. The axis becomes particularly reduced, and the fin comes to be attached directly to the pectoral girdle by a number of basal pieces (Teleosts) probably representing vestigial rays (cf. fig. 19).

EB1911 Ichthyology - Skeleton of Pectoral Limb of Polypterus.jpg

From Budgett, Trans. Zool. Soc. London, xvi, part vii. From Wiedersheim’s Verg. Anat. der Wirbeltiere, by permission of Gustav Fischer.

Fig. 18.—Skeleton of Pectoral Limb of Polypterus. a, 30 mm. larva. b, Adult.

EB1911 Ichthyology - Skeleton of Pectoral Limb of Amia.jpg

From Wiedersheim’s Verg. Anat. der Wirbeltiere, by permission of Gustav Fischer.

Fig. 19.—Skeleton of Pectoral Fin of Amia.

Views on the general morphology of the fin skeleton are strongly affected by the view held as to the mode of evolution of the fins. By upholders of the lateral fold hypothesis the type of fin skeleton described for Cladoselache[1] is regarded as particularly primitive. It is, however, by no means clear that the obscure basal structures figured (Fig. 20) in this fin do not really represent the pressed back axis as in Pleuracanthus.

EB1911 Ichthyology - Skeleton of Pectoral Fin of Cladoselache.jpg

From Bashford Dean, Mem. N.Y. Acad. of Science.

Fig. 20.—Skeleton of Pectoral Fin of Cladoselache.

EB1911 Ichthyology - Placoid elements of Raia clavata (male).jpg

Fig. 21.—Placoid elements of a male Thorn-back, Raia clavata.

The pelvic fin skeleton, while built obviously on the same plan as the pectoral, is liable to much modification and frequently degeneration.

Osseous or Bony Skeleton.—The most ancient type of bony skeleton appears to be represented in the placoid elements such as are seen in the skin of the Selachian (fig. 21). Each placoid element consists of a spine with a broadly expanded base embedded in the dermis. The base is composed of bone: the spine of the somewhat modified bone known as dentine. Ensheathing the tip of the spine is a layer of extremely hard enamel formed by the inner surface of the ectoderm which originally covered it. Such typical placoid scales are well seen on any ordinary skate. In the groups of fishes above the Selachians, the coating of placoid elements shows various modifications. The spines disappear, though they may be present for a time in early development. The bony basal plates tend to undergo fusion—in certain cases they form a continuous bony cuirass (various Siluroids, trunk-fishes) formed of large plates jointed together at their edges. More usually the plates are small and regular in size. In Crossopterygians and Lepidosteus and in many extinct forms the scales are of the ganoid type, being rhomboidal and having their outer layer composed of hard glistening ganoine. In other Teleostomes the scales are as a rule thin, rounded and overlapping—the so-called cycloid type (fig. 22, A); where the posterior edge shows toothlike projections the scale is termed ctenoid (fig. 22, B). In various Teleosts the scales are vestigial (eel); in others (as in most electric fishes) they have completely disappeared.

EB1911 Ichthyology - Cycloid Scale of Scopelus resplendens and Ctenoid Scale of Lethrinus.jpg

Fig. 22.—A, Cycloid Scale of Scopelus resplendens (magn.). B, Ctenoid Scale of Lethrinus (magn.).

Teeth.—Certain of the placoid elements belonging to that part of the skin which gives rise to the lining of the stomodaeum have their spines enlarged or otherwise modified to form teeth. In the majority of fishes these remain simple, conical structures: in some of the larger sharks (Carcharodon) they become flattened into trenchant blades with serrated edges: in certain rays (Myliobatis) they form a pavement of flattened plates suited for crushing molluscan shells. In the young Neoceratodus[2] there are numerous small conical teeth, the bases of which become connected by a kind of spongework of bony trabeculae. As development goes on a large basal mass is formed which becomes the functional tooth plate of the adult, the original separate denticles disappearing completely. In the other two surviving Dipnoans, similar large teeth exist, though here there is no longer trace in ontogeny of their formation by the basal fusion of originally separate denticles. In the Selachians the bony skeleton is restricted to the placoid elements. In the Teleostomes and the Dipnoans the original cartilaginous skeleton becomes to a great extent unsheathed or replaced by bony tissue. It seems highly probable that the more deeply seated osseous elements occurring in these as in the higher groups arose in the course of evolution by the spreading inwards of bony trabeculae from the bases of the placoid elements. Such a method has been demonstrated as occurring in individual development in the case of certain of the more superficially placed bones.[3]

The placoid element with its cap of enamel secreted by the ectoderm is probably originally derived from a local thickening of the basement membrane which with the external cuticle may be looked on as the most ancient skeletal structure in the Metazoa. The basal plate appears to have been a later development than the spine; in the palaeozoic Coelolepidae[4] the basal plate is apparently not yet developed.

Only a brief summary can be given here of the leading features in the osteology of fishes. Care must be taken not to assume that bony elements bearing the same name in fishes and in other groups, or even in the various sub-divisions of the fishes, are necessarily strictly homologous. In all probability bony elements occupying similar positions and described by the same anatomical

  1. Bashford Dean, Journ. Morph. ix. (1894) 87, and Trans. New York Acad. Sci. xiii. (1894) 115.
  2. R. Semon, Zool. Forschungsreisen, Band i. § 115.
  3. O. Hertwig, Arch. mikr. Anat. xi. (1874).
  4. R. H. Traquair, Trans. Roy. Soc. Edin. xxxix. (1899).