Chapter IX: The Order Ungulata (7)

The neck is very short. The limbs are long and stout, and remarkable for the great length of the upper segment (especially the femur) as compared with the distal segment, the manus, and pes. 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 typical Ungulates, the popular fallacy that the joints of the Elephant’s leg bend in a contrary direction to that of other mammals has arisen. 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. The fibula also is quite distinct 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 typical Ungulates in its flatness, in the absence of a 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 Artiodactyles. Of the five toes present on each extremity (see Fig. 98), the middle one is somewhat the largest, and the lateral ones smallest, and generally wanting (especially in the hind foot) the complete number of phalanges. The ungual 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 Rodents and other lower orders of mammals, the cerebellum being entirely behind and uncovered by the cerebrum, but the hemispheres of the latter are richly convoluted.

The Proboscidea are exclusively vegetable feeders, living chiefly on leaves and young branches of forest trees and various kinds of herbage, which they gather and convey to their mouth by the 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 by it many of the functions of the lips of other animals are performed. By its means the Elephant is enabled to drink without bending the head or limbs; the end of the trunk being dipped into the 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 wild animals have been frequently observed fanning themselves with leafy boughs held in the trunk. The species are at present limited in their geographical distribution to the Ethiopian and Oriental regions, but they formerly had a far more extensive range.

_Family_ ELEPHANTIDÆ.

Cheek-teeth succeeding one another in an arc of a circle, and portions of only two, or at most three, of the hinder teeth in use at any one time. Premolars frequently lost, and in any case of no functional importance.

_Elephas._[279]—Dentition: _i_ ¹⁄₀, _c_ ⁰⁄₀, _dm_ ³⁄₃, _m_ ³⁄₃ = 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. The molars composed of numerous flattened enamel-covered plates or ridges of dentine, projecting from a common many-rooted base, surrounded and united together by cement, and extending straight across the crown, without (in most forms) any median division into inner and outer columns. 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. Premolars nearly always wanting. Skull of adult very high and globular. Mandible ending in front in a short, deflected, and spout-like symphysis. Vertebræ: C 7, D 19-21, L 3-4, S 4, C 26-33.

The existing species of the genus differ so much that they have been referred by some writers to distinct genera; fossil forms show, however, such a transition from the one to the other that it is scarcely possible to regard them even as the representatives of distinct groups.

In the well-known Indian or Asiatic Elephant (_E. indicus_) the average number of plates of the six successive molar teeth is expressed by the “ridge-formula,” 4, 8, 12, 12, 16, 24. The plates are compressed from before backwards, the anterior and posterior surfaces (as seen in the worn grinding face of the tooth, Fig. 181) being nearly parallel. Ears of moderate size. Upper margin of the end of the proboscis developed into a distinct finger-like process, much longer than the lower margin. Five nails on the fore feet, and four (occasionally five) on the hind feet.

This species inhabits in a wild state the forest lands of India, Burma, the Malay Peninsula, Cochin China, Ceylon, and Sumatra. The elephants from the last-named islands, presenting some variations from those of the mainland, have been separated under the name of _E. sumatranus_, but the distinction has not been satisfactorily established. The appearance of the Asiatic Elephant is familiar to all. Though rarely breeding in captivity, it has been domesticated from the most remote antiquity, and is still extensively used in the East as a beast of burden. In the wild state it is gregarious, associating in herds of ten, twenty, or more individuals, and though it may, under certain circumstances, become dangerous, it is generally inoffensive and even timid, fond of shade and solitude and the neighbourhood of water. The height of the male at the shoulder when full grown is usually from 8 to 10 feet, but occasionally as much as 11. The female is somewhat smaller.

In the African Elephant (_E. africanus_) the molars (Fig. 182) are of coarse construction, with fewer and larger plates and thicker enamel. Ridge-formula: 3, 6, 7, 7, 8, 10. The plates not flattened, but thicker in the middle than at the edges, so that their worn grinding surfaces are lozenge-shaped. Ears very large. The upper and lower margins of the end of the trunk forming two nearly equal prehensile lips. But three hoofs on the hind foot. This species now inhabits the wooded districts of the whole of Africa south of the Sahara, except where it has been driven away by human settlements. Fossil remains of Pleistocene age, undistinguishable specifically, have been found in Algeria, Spain, and Sicily. It was trained for war and show by the ancient Carthaginians and Romans, and recent experience of the species in captivity in England shows that it is as intelligent as its Asiatic relative, if not more so, while surpassing it in courage, activity, and obstinacy. Nevertheless, in modern times, no people in Africa have been sufficiently civilised or enterprising to care to train it for domestic purposes. It is hunted chiefly for the sake of the ivory of its immense tusks, of which it yields the principal source of supply to the European market, and the desire to obtain which is rapidly leading to the extermination of the species. In size the male African elephant often surpasses that of Asia, but the female is usually smaller. The circumference of the fore foot is half the height at the shoulder, a circumstance which enables the hunters to judge from the footprints the exact size of the animals of which they are in pursuit. The African Elephant also differs from its Indian congener in having tusks in both sexes, whereas in the latter the male only is so armed. Moreover, the eye is relatively larger, the forehead more convex, and the colour somewhat darker. Whereas the Indian Elephant frequents the depths of forests and seldom leaves their shade during the daytime, the following account by Sir Samuel Baker indicates different habits in the African species. This traveller observes: “In Africa, the country being generally more open than in Ceylon, the Elephant remains throughout the day either beneath a solitary tree or exposed to the sun in the vast prairies, where the thick grass attains a height of from nine to twelve feet. The general food of the African Elephant consists of the foliage of trees, especially mimosas. Many of the mimosas are flat-headed, about thirty feet high, and the richer portion of the foliage confined to the crown. Thus the Elephant, not being able to reach to so great a height, must overturn the tree to obtain the coveted food. The destruction caused by a herd of Elephants in a mimosa forest is extraordinary, and I have seen trees uprooted of so large a size that I am convinced no single elephant could have overturned them. I have measured trees four feet six inches in circumference and about thirty feet high uprooted by elephants. The natives have assured me that the elephants mutually assist each other, and that several engage together in the work of overturning a large tree.”

_Extinct Species of Elephant._—Abundant remains of Elephants are found embedded in alluvial gravels, or secreted in the recesses of caves, into which they have been washed by streams and floods, or dragged as food by Hyænas and other carnivorous inhabitants of these subterranean dens. Such remains belonging to the Pleistocene and Pliocene periods have been found in many parts of Europe, including the British Isles, in North Africa, throughout the North American continent from Alaska to Mexico, and extensively distributed in Asia, where the deposits of the sub-Himalayan Siwalik Hills, and equivalent deposits in the Punjab, Perim Island,[280] and Burma, belonging to the earliest Pliocene, are rich in the remains of Elephants of varied form. These species are chiefly known and characterised at present by the skulls and teeth; some of the latter resemble the existing Indian and some the African type, but the majority are between the two, and make the distinction between the two existing species as of generic importance quite impracticable. Others again approach so closely in the breadth and coarseness of the ridges and paucity of cement to _Mastodon_ as to have been placed by some zoologists in that genus. These form the subgenus called _Stegodon_ by Falconer, and may be regarded as a distinct group of the genus.

Among the best known extinct Elephants is _E. primigenius_, the Mammoth,[281] very closely resembling the existing Indian species, and one of the most recently extinct and extensively distributed of all the fossil forms. Probably no animal which has not survived to the historic period has left such abundant and well-preserved evidence of its former existence. The discovery of immense numbers, not only, as in the case of most extinct creatures, in the form of fragmentary bones and teeth, but often as more or less nearly entire carcases, or “mummies,” as they may be called, with the flesh, skin, and hair _in situ_, in the frozen soil of the tundras of Northern Siberia, has for a long time given great interest to the species, and been the cause of many legendary stories among the natives of the lands in which they occur. Among these one of the most prevailing is that the Mammoth was, or still is, an animal which passes its life habitually in burrows below the surface of the ground, and immediately dies if by any chance it comes into the upper air.

Of the whole group the Mammoth is in many respects, as in the size and form of the tusks, and especially the characters of the molar teeth, the farthest removed from the primitive Mastodon-like type, while its nearest surviving relative, _E. indicus_, has retained the slightly more generalised characters of the Mammoth’s contemporaries of more southern climes, _E. columbi_ of America, and _E. armeniacus_ of the Old World, if, indeed, it can be specifically distinguished from them.

The tusks or upper incisor teeth were doubtless present in both sexes, but probably of smaller size in the female. In the adult males they often attained the length of from 9 to 10 feet measured along the outer curve. Upon leaving the head they were directed at first downwards and outwards, then upwards and finally inwards at the tips, and generally with a tendency to a spiral form not seen in other species of Elephant. Different specimens, however, present great variations in curve, from nearly straight to an almost complete circle.

It is chiefly by the characters of the molar teeth that the various extinct modifications of the Elephant type are distinguished. Those of the Mammoth (Fig. 185) differ from the corresponding organs of allied species in the great breadth of the crown as compared with the length, the narrowness and close approximation of the ridges, the thinness of the enamel and its straightness, parallelism, and absence of “crimping,” as seen on the worn surface, or in a horizontal section of the tooth. Dr. Falconer gave the prevailing “ridge-formula” as 4, 8, 12, 12, 16, 24. Dr. Leith Adams, working from more abundant materials, has shown, however, that the number of ridges of each tooth, especially those at the posterior end of the series, is subject to very great individual variation, ranging in each tooth of the series within the following limits: 3 to 4, 6 to 9, 9 to 12, 9 to 15, 14 to 16, 18 to 27, excluding the small plates called talons at each end of the tooth. Besides these variations in the number of ridges or plates of which each tooth is composed, the thickness of the enamel varies so much as to have given rise to a distinction between a “thick-plated” and a “thin-plated” variety—the latter being most prevalent among the specimens from the Arctic regions, and most distinctively characteristic of the species. From the specimens with thick enamel plates the transition to the other species or varieties mentioned above, including _E. indicus_, is almost imperceptible.

The bones of the skeleton generally more resemble those of the Indian Elephant than of any other known species, but the skull differs in the narrower summit, narrower temporal fossæ, and more prolonged incisive sheaths required to support the roots of the enormous tusks. Among the external characters by which the Mammoth was distinguished from either of the existing species of Elephant was the dense clothing, not only of long coarse outer hair, but also of close woolly under hair, of a reddish-brown colour, evidently in adaptation to the colder climate which it inhabited. This character, for a knowledge of which we are indebted to the well-preserved remains found in Northern Siberia, is also represented in the rude but graphic drawings of prehistoric age found in caverns in the south of France.[282] In size different individuals varied considerably, but the average height does not appear to have exceeded that of either of the existing species of Elephant.

The geographical range of the Mammoth was very extensive. There is scarcely a county in England in which some of its remains have not been found either in alluvial deposits of gravel or in caverns, and numbers of its teeth are from time to time dredged up from the bottom of the sea by the fishermen who ply their trade in the German Ocean, having been washed out of the water-worn cliffs of the eastern counties of England. In Scotland and Ireland its remains are less abundant, but they have been found in vast numbers at various localities throughout the greater part of Central Europe (as far south as Santander in Spain and Rome), Northern Asia, and the northern part of the American continent, though the exact distribution of the Mammoth in the New World is still a question of debate. It has not hitherto been met with in any part of Scandinavia or Finland.

In point of time, the Mammoth belongs exclusively to the Pleistocene epoch, and it was undoubtedly contemporaneous with man in France, and probably elsewhere. There is evidence to show that it existed in Britain before, during, and after the glacial period.

As before indicated, it is in the northern part of Siberia that its remains have been found in the greatest abundance, and in quite exceptional conditions of preservation. For a very long period there has been from that region a regular export of Mammoth ivory in a state fit for commercial purposes, both eastward to China and westward to Europe. In the middle of the tenth century an active trade was carried on at Khiva in fossil ivory, which was fashioned into combs, vases, and other objects, as related by Abu’l Kásim, an Arab writer of that period. Middendorff reckoned that the number of tusks which have yearly come into the market during the last two centuries has been at least a hundred pairs, and Nordenskiöld, from personal observation, considers this calculation as probably rather too low than too high. They are found at all suitable places along the whole line of the shore between the mouth of the Obi and Behring Straits, and the farther north the more numerous do they become, the islands of New Siberia being now one of the most favourite collecting localities. The soil of Bear Island and of Liachoff Islands is said to consist only of sand and ice with such quantities of Mammoth bones as almost to compose its chief substance. The remains are not only found around the mouths of the great rivers, as would be the case if the carcases had been washed down from more southern localities in the interior of the continent, but are imbedded in the frozen soil in such circumstances as to indicate that the animals had lived not far from the localities in which they are now found, and they are exposed either by the melting of the ice in unusually warm summers or by the washing away of the sea cliffs or river banks by storms or floods. In this way the bodies of more or less nearly perfect animals, often standing in the erect position, with the soft parts and hairy covering entire, have been brought to light.

References to the principal recorded discoveries of this kind, and to the numerous speculations to which they have given rise, both among ignorant peasants and learned academicians, will be found in Nordenskiöld’s _Voyage of the Vega_ (English translation, vol. i. 1881, p. 398 _sq._) and a series of papers in the _Geological Magazine_ for 1880 and 1881, by H. H. Howorth, as well as in a separate work on the Mammoth by the same writer. For the geographical distribution and anatomical characters, see Falconer’s _Palæontological Memoirs_, vol. ii. 1868; Boyd Dawkins, “_Elephas primigenius_, its Range in Space and Time,” _Quart. Journ. Geol. Soc._ xxxv. p. 138 (1879); and Leith Adams, “Monograph of British Fossil Elephants,” part ii., _Palæontographical Society_ (1879).

_E. antiquus_, of the European Pleistocene, has a lower ridge-formula than in the Mammoth, the molars being narrower, and approximating to those of the African Elephant in structure. Small allied forms occur in the rock-fissures and caverns of Malta, and have been described as _E. mnaidriensis_ and _E. melitensis_; some of the individuals of the latter not exceeding 3 feet in height. The European _E. meridionalis_ is a southern form of somewhat earlier age, very common in the Upper Pliocene of Italy and France, and also in the so-called Forest-bed of the Norfolk coast. It attained very large dimensions, its height being estimated at upwards of 15 feet. The ridge-formula is lower than in _E. antiquus_, the molars are broad, with the worn enamel-discs generally expanded in the middle, and the enamel itself is crenulated.

Elephant remains are very abundant in the Pleistocene and Pliocene deposits of India, those from the latter beds being the oldest representatives of the genus. Of these the Pleistocene _E. namadicus_ appears closely allied to _E. antiquus_, from which it is distinguished by a bold ridge across the forehead. Among the Pliocene forms _E. hysudricus_ may be an ancestral type allied to the Indian Elephant; while _E. planifrons_ is closely related to _E. meridionalis_, although retaining the ancestral feature of developing premolars.

The Stegodont group is peculiar to the eastern parts of the Old World, and, as already observed, connects the true Elephants intimately with the Mastodons. The molars (Fig. 179, II) are characterised by the lowness of the ridges, while the intervening valleys may have but little cement, and there may be a more or less distinct longitudinal groove in the crown dividing each ridge into an inner and an outer moiety. In species like _E. insignis_ the ridge-formula is nearly the same as in _E. meridionalis_, but in _E. clifti_ some of the molars carry only six ridges, and premolars were present, so that we thus have such a complete transition to the next genus that it is very difficult to know where to draw the line between the two.

_Mastodon._[283]—Dentition: _i_ ¹⁄₁₋₀, _c_ ⁰⁄₀, _dm_ ³⁄₃, _m_ ³⁄₃. Upper incisors large, as in _Elephas_, sometimes with longitudinal bands of enamel, more or less spirally disposed. Lower incisors variable; when present comparatively small and straight, sometimes persistent, sometimes early deciduous, and in some species never present. Grinding surface of molars with transverse ridges, the summits of which are divided more or less into conical or mammillary cusps, and often with secondary or additional cusps between and clustering against the principal ridges; enamel thick; cement very scanty, never filling up the interspaces between the ridges. The third, fourth, and fifth cheek-teeth (_i.e._ the last milk-molar, and the first and second molars) having the same number of ridges,[284] which never exceeds five.

In the upper jaw the incisors, though of large size, were apparently never so much curved as in some species of Elephant, and they often have longitudinal bands of enamel, more or less spirally disposed upon their surface, which are not met with in Elephants. Lower incisors were present throughout life in some species, which have the symphysis of the lower jaw greatly elongated to support them (as in _M. angustidens_, _M. pentilici_, and _M. longirostris_). In the common North American species (_M. americanus_) the mandibular symphysis is short, but it may have a small incisor on one side. In other species no inferior tusks have been found, at all events in adult life (see figure of _M. arvernensis_).

The molar teeth increase in size from before backwards, but as many as three of these teeth may be in place in each jaw at one time. There is in many species a true vertical succession, affecting either the third, or the third and second, or (in _M. productus_) the first, second, and third of the six molariform teeth. These three are therefore reckoned as milk-molars, and their successors as premolars, while the last three, which are never changed, correspond to the true molars of those animals in which the typical dentition is fully developed. The study of the mode of succession of the teeth in the different species of Mastodons is particularly interesting, as it exhibits so many stages of the process by which the very anomalous dentition of the modern Elephants may have been derived by gradual modification from the typical heterodont and diphyodont dentition of the ordinary mammal. It also shows that the anterior molars of Elephants do not correspond to the premolars of other Ungulates, but to the milk-molars, the early loss of which in consequence of the peculiar process of horizontal forward-moving succession does not require, or allow time for, their replacement by premolars. It must be noted, however, that, in the Mastodon in some respects the least specialised in tooth-structure, the _M. americanus_ of North America, no vertical succession of the molars has yet been observed, although vast numbers of specimens have been examined.

The Mastodons have fewer ridges on their molar teeth than the Elephants; the ridges are also less elevated, wider apart, have a thicker enamel-covering, and scarcely any cement filling up the space between them. Sometimes (as in _M. americanus_) the ridges are simple transverse wedge-shaped elevations, with straight or concave edges. In other species the summits of the ridges are more or less subdivided into conical cusps, and may have accessory cusps clustering around them (as in _M. americanus_, see Fig. 187). When the apices of these are worn by mastication, their surfaces present circles of dentine, surrounded by a border of enamel, and as the attrition proceeds different patterns are produced by the union of the bases of the cusps, a trilobed or trefoil form being characteristic of some species (Fig. 188).

As already mentioned, certain of the molariform teeth of the middle of the series in Mastodons have the same number of principal ridges, those in front of them having fewer and those behind a greater number. These teeth were distinguished as “intermediate” molars by Dr. Falconer, and are three in number, namely the last milk-molar and the first and second true molars (or the third, fourth, and fifth of the whole series). The number of ridges on these intermediate molars is nearly always three or four, and the tooth in front has usually one fewer and that behind one more, so that the ridge-formula of most Mastodons can be reduced either to 1, 2, 3, 3, 3, 4, or 2, 3, 4, 4, 4, 5. The former characterises the section called _Trilophodon_ (of which an intermediate molar is shown in Fig. 188), and the latter that called _Tetralophodon_ by Dr. Falconer. These divisions are very useful, as under one or the other all the present known species of Mastodon can be ranged, but observations upon a larger number of individuals have shown that the number of ridges upon the teeth is not quite so constant as implied by the formulæ given above. Their exact enumeration is even difficult in many cases, as “talons” or small accessory ridges at the hinder end of the teeth occur in various stages of development, until they take on the character of true ridges. Transitional conditions have also been shown, at least in some of the teeth, between the trilophodont and the tetralophodont forms, and again between the latter and what has been called a “pentalophodont” type, which leads on towards the condition of dental structure characteristic of the true Elephants.

The range of the genus _Mastodon_ in time was from the middle of the Miocene period to the end of the Pliocene in the Old World, when it became extinct; but in America several species—especially the one best known, owing to the abundance of its remains, which has been variously called _M. americanus_, _M. ohioticus_, and _M. giganteus_—survived to a late Pleistocene period.

The range in space will be best indicated by the following list of some of the better known species. (1) Trilophodont series—_M. angustidens_,[285] _borsoni_, _pentelici_, _turiensis_, from Europe; _M. falconeri_ and _pandionis_, from India; _M. americanus_, _obscurus_, and _productus_, North America; and _M. cordillerum_ and _humboldti_, South America. (2) Tetralophodont series—_M. arvernensis_, _M. longirostris_, from Europe; _M. latidens_, _sivalensis_, and _perimensis_, from India; _M. mirificus_, from North America. _Mastodon arvernensis_ and _M. longirostris_, together with a trilophodont species, occur in the crag-deposits of Norfolk and Suffolk.

_Family_ DINOTHERIIDÆ.

An extinct family distinguished from the _Elephantidæ_ by the whole series of permanent cheek-teeth being in use at the same time.

_Dinotherium._[286]—Dentition of adult: _i_ ⁰⁄₁, _c_ ⁰⁄₀, _p_ ²⁄₂, _m_ ³⁄₃ = 22; all present at the same time, there being no horizontal succession, but the premolars replacing milk-teeth in the ordinary manner. The presence or absence of upper incisors has not yet been clearly ascertained. Lower incisors large, conical, descending, and slightly curved backwards, implanted in a greatly thickened and deflected beak or prolongation of the symphysis. In section they do not show the decussating striæ characteristic of Mastodons and Elephants. Crowns of molars carrying strong transverse, crenulated ridges, with deep valleys between, much resembling the lower ones of the Tapirs. Ridge-formula of the permanent molar series: 2, 2, 3, 2, 2. The three ridges of the first true molar are constant in both upper and lower jaws, although it is quite an anomalous character among Proboscideans for this molar to have more ridges than those which come behind it. The last milk-molar has also three ridges, the penultimate but two. The cranium is much depressed, with comparatively little development of air-cells. The remainder of the skeleton is imperfectly known, but apparently agrees in its general character with that of the other Proboscideans.

Remains of _Dinotherium giganteum_, an animal of elephantine proportions, strikingly characterised by the pair of huge tusks descending nearly vertically from the front of the lower jaw, were first discovered at Eppelsheim, near Darmstadt, and described by Kaup. They have since been met with in various Lower Pliocene and higher Miocene formations in the south of Germany, France, Greece, and Asia Minor. Closely allied forms also occur in the Lower Pliocene and Upper Miocene of India, but none are known from America.

_Suborder_ AMBLYPODA.

_Uintatherium._[287]—Among the most remarkable of the comparatively recent discoveries in the higher Eocene formations of the western states of North America has been one of a group of animals of huge size, approaching that of the largest existing Elephants, presenting a combination of characters quite unlike those known among other recent or extinct creatures, and of which there were evidently many species living contemporaneously, but all of which became extinct before the close of the Eocene period. To form some idea of their appearance, we must imagine animals very elephantine in general proportions and in the structure of their limbs. The feet had five short toes. The tail, as in the Elephants, was long and slender, but the neck, though still short, was not so much abbreviated as in the Proboscideans, and there is no evidence that these animals possessed a trunk. The head differed greatly from that of the Elephants, being long and narrow, more like that of a Rhinoceros, and, as in that animal, was elevated behind into a great occipital crest, and it had developed upon its upper surface three pairs of conspicuous, laterally diverging protuberances—one pair in the parietal region, one on the maxillaries in front of the orbits, and one (much smaller) near the fore part of the elongated nasal bones. Whether these were merely covered by bosses of callous skin, as the rounded form and ruggedness of their extremities would indicate, or whether they formed the bases of attachment for horns of still greater extent, like those of the Rhinoceros or of the Cavicorn Ruminants, can only be a matter of conjecture. There were no upper incisors, but usually three on each side below, of comparatively small size, as was also the lower canine. A huge, compressed, curved, sharp-pointed canine tusk, very similar in form and position to that of the Musk-Deer, descended from each side of the upper jaw. These were present in both sexes, but very much smaller in the female, as was also the flange-like process of the lower jaw by which they were guarded. Behind these, and at some distance from them, were on each side above and below six cheek-teeth, of comparatively small size, placed in continuous series, each with a pair of oblique ridges conjoined internally and diverging externally in a V-like manner, and provided with a stout basal cingulum. The normal dental formula was therefore _i_ ⁰⁄₃, _c_ ¹⁄₁, _p_ ³⁄₃, _m_ ³⁄₃ = 34; and the dentition had thus already attained a remarkable degree of specialisation, although the brain was smaller and more rudimentary in characters than in almost any other known mammal. In its comparative length and the absence of a third trochanter the femur of these animals resembles that of the Proboscidea. The first discovered evidences of the existence of animals of this group were described by Leidy in 1872, under the name of _Uintatherium_ (from the Uinta mountains, near which they were found). Subsequently the names _Dinoceras_, _Tinoceras_, _Loxolophodon_, etc., have been applied to various members of the group, but the characters by which they are distinguished do not seem of sufficient importance to allow of their separation from the type genus _Uintatherium_.[288]

_Coryphodon._[289]—Another interesting form referred to this suborder is _Coryphodon_, which appears to connect the _Uintatheriidæ_ with the most primitive Perissodactyla. It was first described by Owen in 1846 from a fragment of a jaw from the London Clay. Other remains were afterwards discovered in France, and lately in great abundance, indicating many species from the size of a Tapir to that of a Rhinoceros, in the Lower and Middle Eocenes of New Mexico and Wyoming in the United States. _Coryphodon_ had forty-four teeth; the canines of both jaws were large and sharp pointed, and the molars had strongly pronounced oblique ridges. The general proportions were those of a Bear, but the tail was of moderate length, and the feet short and wide. The femur had a third trochanter; and the cranium was devoid of protuberances. The genus should be regarded as the type of a distinct family _Coryphodontidæ_.

_Suborder_ CONDYLARTHRA.

The term Condylarthra has been proposed by Professor Cope for a number of generalised and mostly comparatively small Ungulates, which were probably allied both to the Perissodactyla and Artiodactyla, but present characters separating them from those divisions as commonly defined. In the structure of the carpus and tarsus these forms (which are chiefly known to us from the Eocene of the United States) come nearer to the Hyracoidea than to any other existing type. As a rule they have the full dental formula; the molars are brachydont, generally bunodont, and in many instances also tritubercular; while the premolars are always simpler than the molars.

The humerus is quite peculiar among Ungulates in having an entepicondylar foramen; the femur has a third trochanter; and the form and relations of the astragalus are similar to those obtaining in the Carnivora. The feet are usually furnished with five functional digits, of which the ungual phalanges are pointed. In many respects the skeleton of these remarkably generalised Ungulates approximates so decidedly to a Carnivorous type as to have led palæontologists to conclude that the Ungulata and Carnivora are branches of an original common stock.

In this work space only permits of allusion to a few of the more important types of this group. _Periptychus_, which occurs in the lowest Eocene of New Mexico, is a bunodont type readily distinguished by the vertical flutings of the premolars, and the small size of the incisors and canines. It has been suggested that this genus is closely related to the stock of the bunodont Artiodactyla. Of greater interest is the genus _Phenacodus_, which is regarded as the lowest factor in the series from which the modern Horse has been evolved, where it holds the position immediately below _Hyracotherium_ or, _Systemodon_ (see p. 374). One of the species was about the size of a Bull-dog, while another might be compared to a small Leopard. The structure of the cheek-teeth is such as might readily be modified into that obtaining in _Hyracotherium_; all the feet had five fully developed digits, and the tail was long. _Meniscotherium_ and _Hyracodontotherium_ are more specialised forms of somewhat later age, with a lophodont dentition: the latter genus being European.

_Suborder_ TOXODONTIA.

In addition to the _Macraucheniidæ_ and certain other forms noticed under the head of the Perissodactyla, the Tertiaries of South America have yielded some very remarkable forms of mammalian life, the nature and affinities of which have greatly puzzled all zoologists who have attempted to unravel them.

_Nesodon_ and _Toxodon_.—Among these _Nesodon_, from Patagonia, has the full typical Eutherian number of teeth; the crowns of the incisors being short, and the molars having a complex rhinocerotic type of structure somewhat intermediate between _Homalodontotherium_ (p. 412) and the following genus _Toxodon_. The typical species of _Nesodon_ was about as large as a Sheep, but nothing more is known of it than the teeth and portions of the skull.

_Toxodon_ is an animal about the size of a Hippopotamus; it was first discovered by Darwin, and many specimens have since been found in Pleistocene deposits near Buenos Ayres, and described by Owen, Gervais, and Burmeister. The teeth consist of large incisors, very small lower canines, and strongly curved molars, all with persistent roots, the formula being apparently _i_ ²⁄₃, _c_ ⁰⁄₁, _p_ ⁴⁄₃, _m_ ³⁄₃ = 38. The cranial characters exhibit a combination of those found in both Perissodactyles and Artiodactyles, but the form of the hinder part of the palate and the absence of an alisphenoid canal belong to the latter; and the tympanic, firmly fixed in between the squamosal and the exoccipital, ankylosed to both, and forming the floor of a long upward-directed meatus auditorius, is so exactly like that of the Suina that it is difficult to believe it does not indicate some real affinity to that group. These characters seem to outweigh in importance those by which some zoologists have linked _Toxodon_ to the Perissodactyla, and the absence of the third trochanter and the articulation of the fibula with the calcaneum tell in the same direction. According to the recent observations of Ameghino the hind feet were certainly tridactylous, and the front feet probably so. The earlier allied genera _Protoxodon_ and _Adinotherium_ are definitely known to have tridactylous front and hind feet, which conform to the Perissodactylate type, the bones of the proximal and distal rows of the carpus interlocking. _Acrotherium_, which has similar feet, differs from all other Ungulates, and indeed from all Eutherians except some individuals of the existing carnivorous genus _Otocyon_, in having eight cheek-teeth, five of which have been reckoned as premolars.

_Typotherium._—_Typotherium_ (Fig. 192), also called _Mesotherium_, from the same locality as _Toxodon_, was an animal rather larger than a Capybara, and of much the same general appearance. Its skeleton is completely known, and shows a singular combination of characters, resembling _Toxodon_ or a generalised Ungulate on the one hand, and the Rodents, especially the _Leporidæ_, on the other. In the presence of clavicles it differs from all known Ungulates, and in having two pairs of lower incisors from all Rodents. The teeth are _i_ ¹⁄₂, _c_ ⁰⁄₀, _p_ ²⁄₁, _m_ ³⁄₃ = 24.

From the Tertiaries of various parts of South America a number of forms more or less closely allied to _Toxodon_ and _Typotherium_ have been recently described, but as many of them are very imperfectly known, and there is much doubt as to their generic position, it will be unnecessary to refer to them further.

It will thus be seen that, although our knowledge of many of these forms is still very limited, we may trace among them a curious chain of affinities, which would seem to unite the Ungulates on the one hand with the Rodents on the other; but further materials are required before we can establish with certainty so important a relationship, one which, if true, would alter materially some of the prevailing views upon the classification of mammals.

_Group_ TILLODONTIA.

Here may be noticed a remarkable group of animals, called by Marsh, Tillodontia, the remains of which are found abundantly in the Lower and Middle Eocene beds of North America. They seem to combine the characters of the Ungulata, Rodentia, and Carnivora. In the genus _Tillotherium_ of Marsh (probably identical with the previously described _Anchippodus_ of Leidy) the skull (Fig. 193) resembled that of the Bears, but the molar teeth were of the Ungulate type, while the large incisors were very similar to those of the Rodents. The dental formula is _i_ ²⁄₂, _c_ ¹⁄₁, _p_ ³⁄₄, _m_ ³⁄₃. The first pair of incisors was very small; the upper molars were tritubercular, while the lower ones had crescentoid ridges as in _Palæotherium_. The skeleton resembled that of the Carnivores, but the scaphoid and lunar bones were distinct, and there was a third trochanter on the femur. The feet were plantigrade, and each had five digits, all with long pointed claws. In the allied genus _Stylinodon_ all the teeth were rootless. Some forms were as large as a Tapir.

These, with other more or less closely allied animals, such as _Calamodon_ and _Psittacotherium_, constituting a group called Tæniodonta, are included by Cope in his large order Bunotheria, to which also the existing Insectivora are referred. The dentition of some of these forms makes a remarkable approximation towards a Rodent type, while it has been suggested that there are also signs of remote Edentate affinities. The constantly increasing knowledge of these annectant forms adds to the difficulty so often referred to in this work of establishing anything like a definite classification of the heterodont mammals. An incisor tooth from the Swiss Eocene has recently been referred to _Calamodon_.

_Bibliography of Ungulata._—In addition to the works and
memoirs mentioned under the different sections of the order,
the following may be referred to:—W. Kowalevsky, “Monographie
des genus Anthracotherium,” _Palæontographica_ 1873; Id. “Sur
l’Anchitherium aurelianense et sur l’histoire paléontologique
des Chevaux,” _Mém. de l’Acad. Imp. des Sciences de St.
Pétersbourg_, 1873; Id. “On the Osteology of the Hyopotamidæ,”
_Philosophical Transactions_, 1873; L. Rütimeyer, “Versuch
einer natürlichen Geschichte des Rindes.” etc., _Neue Denks.
der allgem. Schweiz. Gesellsch. für Naturwissenschaften_, 1867;
Id. “Die Rinder der Tertiär-Epoche,” _Abhand. der Schweiz.
Paläont. Gesellsch._ 1877 and 1878; Id. “Beiträge zu einer
Natürliche Geschichte der Hirsche,” _ibid._ 1880-1881; C. J.
Forsyth-Major, “Beiträge zur Geschichte der Fossilen Pferde,”
_ibid._ 1880; M. Schlosser, “Beiträge zur Kenntniss der
Stammesgeschichte der Hufthiere und Versuch einer Systematik
der Paar- und Unpaarhufer,” _Morph. Jahrb._ 1886; E. D. Cope,
“The Perissodactyla,” _Amer. Natural._ 1887; M. Pavlow, “Études
sur l’histoire paléontologique des Ongulés,” _Bull. Soc. Imp.
Naturalistes Moscow_, 1887-1890. W. B. Scott and H. F. Osborn,
“The Mammalia of the Uinta Formation,” _Trans. Amer. Phil.
Soc._ vol. xvi. (1889).