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PROBOSCIDEA


in one extinct species of true elephant (Elephas planifrons) as regards some of these teeth.

(Flower's Osteology of Mammalia.)

Fig. 2.—Section of the Skull of the African Elephant (Elephas africanus) taken to the left of the middle line, and including the vomer (Vo) and the mesethmoid (ME).

an, Anterior, pn, Posterior nasal aperture.

Most proboscideans are animals of large dimensions, and some are the most colossal of land mammals. The head is of great proportionate Physical Characteristics. size; and, as the brain-case increases but little in bulk during growth, while the exterior wall of the skull is required to be of great superficial extent to support the trunk and the ponderous tusks, and to afford space for the attachment of muscles of sufficient size and strength to wield the skull thus heavily weighted, an extraordinary development of air-cells takes place in the cancellous tissue of nearly all the bones of the cranium. These cells are not only formed in the walls of the cranium proper, but are also largely developed in the nasal bones and upper part of the premaxillae and maxillae, the bones forming the palate and the basi-cranial axis, and even extend into the interior of the ossified mesethmoid and vomer. Where two originally distinct bones come into contact, the cells pass freely from one to the other, and almost all the sutures become obliterated in old animals. The intercellular lamellae in the great mass which surrounds the brain-cavity superiorly and laterally mostly radiate from the inner to the outer table, but in the other bones their direction is more irregular. Like the similar but less developed air-cells in the skulls of many other mammals, they all communicate with the nasal passages, and they are entirely secondary to the original growth of the bones, their development having scarcely commenced in the new-born animal, and gradually enlarge as the growth of the creature proceeds. The nasal bones are very short, and the anterior nasal aperture situated high in the face. The zygomatic arch is slender and straight, the jugal bone being small, and forming only the middle part of the arch, the anterior part of which (unlike that of true Ungulates) is formed only by the maxilla. The maxillo-turbinals are rudimentary, the elongated proboscis supplying their place functionally in warming and clearing from dust the inspired air.

The neck is very short. The limbs, as already mentioned, are long and stout, and remarkable for the great length of the upper segment (especially the femur) as compared with the lower segment, as represented by the foot. It is owing to this and the vertical position of the femur that the knee-joint in the hind-leg is placed much lower, and is more conspicuous externally than in most quadrupedal mammals; and this having been erroneously compared with the hock-joint or ankle of the more ordinary ungulates, gave rise to the popular fallacy that the joints of the elephant's leg bend in a contrary direction to that of other mammals. There is no round ligament in the hip-joint, or third trochanter to the femur. The radius and ulna are distinct, though fixed in a crossed or prone position; and the fibula also is quite separated from the tibia. The feet are short and broad, the carpal and tarsal bones being very square, with flattened surfaces for articulation; the astragalus especially differs from that of the more typical ungulates in its flatness, in the absence of distinct pulley-like articular surface at either extremity, and in having no articular facet for the cuboid. The fibula articulates with the calcaneum, as in the artiodactyle sub-order of Ungulata. Of the five toes present on each foot, the middle one is somewhat the largest, while the lateral ones are the smallest, and generally lack (especially in the hind-foot) the complete number of phalanges. The terminal phalanges are all small, irregular in form, and late in ossification. The whole are encased in a common integument, with a flat, subcircular, truncated sole, the only external indication of the toes being the broad oval nails or hoofs arranged in a Semicircle around the front edge of the sole. The hind foot is smaller and narrower than the front. The liver is small and simple, and there is no gall-bladder. In form the brain resembles that of the lower orders of mammals in that the cerebellum is entirely behind and uncovered by the cerebrum, but the hemispheres of the latter are richly convoluted.

Elephants are exclusively vegetable-feeders, living, chiefly on leaves and young branches of forest trees and various kinds of herbage, or roots, which they gather and convey to their mouth by a very mobile proboscis, an organ which combines in a marvellous manner strength with dexterity of application, and is a necessary compensation for the shortness and inflexibility of the neck, as it is by this that many of the functions of the lips of other animals are performed. By its means elephants are enabled to drink without bending the head or limbs. The end of the trunk being dipped, for instance, into a stream or pool, a forcible inspiration fills the two capacious air-passages in its interior with water, which, on the tip of the trunk being turned upwards and inserted into the mouth, is ejected by a blowing action, and swallowed. Or if the animal wishes to refresh and cool its skin, it can throw the water in a copious stream over any part of its surface. Elephants can also throw dust and sand over their bodies by the same means and for the same purpose, and they have frequently been observed fanning themselves with boughs held in the trunk.

The following are the distinctive features of the genus Elephas, the type of the family Elephantidae: Dentition: i. 1/0, c. 0/0, m. 6/6 = 26. The incisors variable, but usually of very large size, especially in the male sex, directed somewhat outwards, and curved upwards, without enamel except on the apex before it is worn; preceded by small milk-incisors. The molars succeed each other by horizontal replacement from before backwards, never more than one or part of two being in use on each side of each jaw at the same time; each composed of numerous flattened enamel-covered plates or ridges of dentine, projecting from a common many-rooted base, surrounded and united together by cement. The number of plates increases from the anterior to the posterior molar in regular succession, varying in the different species, but the third and fourth (or the last milk-molar and the first true molar), and these only, have the same number of ridges, which always exceeds five. Skull of adult very high and globular. Lower jaw ending in front in a deflected, spout-like symphysis. Vertebrae: C. 7, D. 19-21, L. 3-4, S. 4, C. 26-33.

The two existing species of elephant are the Indian or Asiatic (Elephas maximus), and the African (E. africanus), the distinctive characteristics of which are given under Elephant. See also Mammoth and Mastodon.

Extinct Proboscidea

Elephas.—The extinct representatives of the Proboscidea are of the greatest importance and interest, since they serve to connect the modern elephants with ungulates of more ordinary type. The Mammoth (Elephas primigenius) is treated in a separate article. Nearly allied is E. armeniacus of Asia Minor; but E. antiquus, of which the remains are abundant in many of the superficial formations of England and Europe generally, approximates in the structure of its molar teeth to the African elephant. It is represented in the Pleistocene of India by the closely allied or identical E. namadicus. Affinity with the African species is strongly marked in the case of the dwarf elephants of Malta (E. melilensis) and Cyprus (E. cypriotes); and the gigantic E. meridionalis, of the “forest-bed” of the east coast of England and the Upper Pliocene of the Val D'Arno, has likewise molars showing the broad lozenges of enamel-bordered dentine characteristic of the African type. These and other species indicate, however, that, so far as dental characters are concerned, generic separation of the African from, the Asiatic elephant is impossible. In North America the mammoth occurs in the far north, E. columbi, more akin to E. antiquus chiefly in the Central United States, and E. imperator (allied to E. meridionalis) in the south. The oldest representatives of this group are E. hysudricus and E. planifrons of the Lower Pliocene of Northern India; the latter of which developed premolars vertically replacing the anterior teeth of the molar series.

From E. planifrons there is an almost complete transition to the ridge-toothed elephants, such as E. ganesa, E. insignis,