treating of the subjects dealt with, and in particular I should like to mention that the section devoted to the skeleton of mammals is, as it could hardly fail to be, to a considerable extent based on Sir W.H. Flower's Osteology of the Mammalia.
SIDNEY H. REYNOLDS.
March 10, 1897.
CHAPTER I.
INTRODUCTORY ACCOUNT OF THE SKELETON IN GENERAL.
By the term skeleton is meant the hard structures whose function is to support or to protect the softer tissues of the animal body.
The skeleton is divisible into
A. The Exoskeleton, which is external;
B. The Endoskeleton, which is as a rule internal; though in some cases, e.g. the antlers of deer, endoskeletal structures become, as development proceeds, external.
In Invertebrates the hard, supporting structures of the body are mainly exoskeletal, in Vertebrates they are mainly endoskeletal; but the endoskeleton includes, especially in the skull, a number of elements, the dermal or membrane bones, which are shown by development to have been originally of external origin. These membrane bones are so intimately related to the true endoskeleton that they will be described with it. The simplest and lowest types of both vertebrate and invertebrate animals have unsegmented skeletons; with the need for flexibility however segmentation arose both in the case of the invertebrate exoskeleton and the vertebrate endoskeleton. The exoskeleton in vertebrates is phylogenetically older than the endoskeleton, as is indicated by both palaeontology and embryology. Palaeontological evidence is afforded by the fact that all the lower groups of vertebrates—Fish, Amphibia, and Reptiles—had in former geological periods a greater proportion of species protected by well-developed dermal armour than is the case at present. Embryological evidence tends the same way, inasmuch as dermal ossifications appear much earlier in the developing animal than do the ossifications in the endoskeleton.
Skeletal structures may be derived from each of the three germinal layers. Thus hairs and feathers are epiblastic in origin, bones are mesoblastic, and the notochord is hypoblastic.
The different types of skeletal structures may now be considered and classified more fully.
A. Exoskeletal structures.
I. Epiblastic (epidermal).
Exoskeletal structures of epiblastic origin may be developed on both the inner and outer surfaces of the Malpighian layer of the epidermis[1] . Those developed on the outer surface include hairs, feathers, scales, nails, beaks and tortoiseshell; and are specially found in vertebrates higher than fishes. Those developed on the inner surface of the Malpighian layer include only the enamel of teeth and some kinds of scales. With the exception of feathers, which are partly formed from the horny layer, all these parts are mainly derived from the Malpighian layer of the epidermis.
Hairs are slender, elongated structures which arise by the proliferation of cells from the Malpighian layer of the epidermis. These cells in the case of each hair form a short papilla, which sinks inwards and becomes imbedded at the bottom of a follicle in the dermis. Each hair is normally composed of an inner cellular pithy portion containing much air, and an outer denser cortical portion of a horny nature. Sometimes, as in Deer, the hair is mainly formed of the pithy portion, and is then easily broken. Sometimes the horny part predominates, as in the bristles of Pigs. A highly vascular dermal papilla projects into the base of the hair.
Feathers, like hairs, arise from epidermal papillae which become imbedded in pits in the dermis. But the feather germ differs from the hair germ, in the fact that it first grows out like a cone on the surface of the epidermis, and that the horny as well as the Malpighian layer takes part in its formation.
Nails, claws, hoofs, and the horns of Oxen are also epidermal, as are such structures as the scales of reptiles, of birds' feet, and of Manis among mammals, the rattle of the rattlesnake, the nasal horns of Rhinoceros, and the baleen of whales. All these structures will be described later.
Nails arise in the interior of the epidermis by the thickening and cornification of the stratum lucidum. The outer border of the nail soon becomes free, and growth takes place by additions to the inner surface and attached end.
When a nail tapers to a sharp point it is called a claw. In many cases the nails more or less surround the ends of the digits by which they are borne.
Horny beaks of epidermal origin occur casing the jaw-bones in several widely distinct groups of animals. Thus among reptiles they are found in Chelonia (tortoises and turtles) as well as in some extinct forms; they occur in all living birds, in Ornithorhynchus among mammals, and in the larvae of many Amphibia.
In a few animals, such as Lampreys and Ornithorhynchus, the jaws bear horny tooth-like structures of epidermal origin.
The enamel of teeth and of placoid scales is also epiblastic in origin[2] , and it may be well at this point to give some account of the structure of teeth, though they are partly mesoblastic in origin. The simplest teeth are those met with in sharks and dogfish, where they are merely the slightly modified scales developed in the integument of the mouth. They pass by quite insensible gradations into normal placoid scales, such as cover the general surface of the body. A placoid scale[3] is developed on a papilla of the dermis which projects outwards and backwards, and is covered by the columnar Malpighian layer of the epidermis. The outer layer of the dermal papilla then gradually becomes converted into dentine and bone, while enamel is developed on the inner side of the Malpighian layer, forming a cap to the scale. The Malpighian and horny layers of the epidermis get rubbed off the enamel cap, so that it comes to project freely on the surface of the body.
As regards their attachment teeth may be (1) attached to the fibrous integument of the mouth, or (2) fixed to the jaws or other bones of the mouth, or (3) planted in grooves, or (4) in definite sockets in the jaw-bones (see p. 107).
Teeth in general consist of three tissues, enamel, dentine and cement, enclosing a central pulp-cavity containing blood-vessels and nerves. Enamel is, however, often absent, as in all living Edentates.
Enamel generally forms the outermost layer of the crown or visible part of the tooth; it is the hardest tissue occurring in the animal body and consists of prismatic fibres arranged at right angles to the surface of the tooth. It is characterised by its bluish-white translucent appearance.
II. Mesoblastic (mesodermal).
Dentine or ivory generally forms the main mass of a tooth. It is a hard, white substance allied to bone. When examined microscopically dentine is seen to be traversed by great numbers of nearly parallel branching tubules which radiate outwards from the pulp-cavity. In fishes as a rule, and sometimes in other animals, a variety of dentine containing blood-vessels occurs, this is called vasodentine.
Cement or crusta petrosa forms the outermost layer of the root of the tooth. In composition and structure it is practically identical with bone. In the more complicated mammalian teeth, besides enveloping the root, it fills up the spaces between the folds of the enamel.
The hard parts of a tooth commonly enclose a central pulp-cavity into which projects the pulp, a papilla of the dermis including blood-vessels and nerves. As long as growth continues