Chapter IX
UNGULATA--CONDYLARTHRA--AMBLYPODA--ANCYLOPODA--TYPOTHERIA--TOXODONTIA-- PROBOSCIDEA--HYRACOIDEA
ORDER IV. UNGULATA
The existing members of this order can be readily grouped into the Hyracoidea, Proboscidea, Perissodactyla, and Artiodactyla, each of which divisions has quite the value of an order, and all of which are sharply marked off from each other. But as the discovery of so many fossil forms has to a great extent rendered these demarcations less sharp, it is better to regard all these groups as not more than sub-orders of a larger "Order" Ungulata. Even when this conclusion has been necessarily arrived at from a consideration of the more ancient groups of Ungulate animals, the definition of such an order remains a difficult matter for the systematist. For the earliest of these forms, more particularly the Ancylopoda, the Amblypoda, and the Condylarthra, whose peculiarities will be dealt with at length subsequently, are not by any means easily differentiated from the primitive Carnivorous mammals of that date, the Creodonta; these latter, moreover, fade into the Marsupials through the so-called Sparassodonta of Professor Ameghino. To confine ourselves to the Ungulates, we may perhaps define them as terrestrial animals with hoofs rather than claws or nails, and chiefly, if not entirely, vegetarian in habit. The teeth are bunodont or lophodont, the tendency to the production of the latter type being always marked. The walk, although plantigrade in the older types, becomes more and more digitigrade, except in such survivals from antiquity as Hyrax. There is, too, as we pass from the ancient types to the modern, a gradual perfection of the limbs as running {196} and not climbing or grasping organs; the number of toes becomes reduced, and culminates twice (in the horse and in the Litopterna) in one toe on each foot; at the same time the ulna becomes rudimentary and fuses with the radius, and the fibula in the hind-limb undergoes a like reduction. The clavicle is absent even in some of the oldest types; its presence in Typotherium[120] is highly remarkable. The tail too, an organ which is long in some of the early forms, gets short in their modern derivatives.
FIG. 110.--An early Ungulate. Phenacodus primaevus. x 1/12. (After Osborn.)
FIG. 111.--Series of metacarpals and metatarsals of Camelidae, to show secular and progressive increase in size. From left to right the species are Protylopus petersoni, Poebrotherium labiatum, Gomphotherium sternbergi, Procamelus occidentalis. F, Fore-foot; H, hind-foot; III, IV, third and fourth metapodials. (After Wortman.)
Coupled with the increasing perfection of the foot as an organ used merely for the support of the body, certain interesting changes have taken place in the arrangement with regard to each other of the several bonelets of the wrist and ankle. It has been held by Cope and others that the truly primitive disposition of these bones was that presented to us by certain early types, such as Meniscotherium or the existing elephant or Hyrax. In these animals there is (see Fig. 112) a serial {197} arrangement of these bones, the distal bones only, or very nearly only, articulating with the corresponding bones in the upper series. In the modern types (cf. Fig. 113) there is, on the other hand, an interlocking, so that the bones of the distal series articulate with two of those of the proximal series. By this is produced, as it would appear, a much firmer foot, less liable to "give" under pressure, and thus more fitted for an animal that runs. It is the same principle as that adopted in the laying of bricks. The actual stress and strain of impact has been held responsible for those changes. An equally ingenious and possibly truer explanation of the undoubted facts has lately been advanced by Mr. W. D. {198} Matthew.[121] He has pointed out that in some ancient Ungulates the carpus is not serial but interlocking, even in forms which belong to the earliest Eocene groups, such as the genus Protolambda among the Amblypoda. Now in the fore-foot of Meniscotherium and the living Hyrax there is a separate centrale which is wanting in the greater number of Ungulates. The absorption, that is the practical dropping out of this bone, would restore to an interlocking carpus the serial arrangement; while on the other hand, by the fusion of this bone with the scaphoid, the interlocking disposition would be maintained.
FIG. 112.--Bones of the manus A, of the Indian Elephant, Elephas indicus. x 1/8. B, of the Cape Hyrax, Hyrax capensis. x 1. c, Cuneiform; cc, centrale; l, lunar; m, magnum; p, pisiform; R, radius; td, trapezoid; tm, trapezium; s, scaphoid; u, unciform; U, ulna. (From Flower's Osteology.)
FIG. 113.--Bones of the manus A, of Rhinoceros, Rhinoceros sumatrensis. x 1/5. B, of Pig, Sus scrofa. x 1/3. Letters as in Fig. 112. (From Flower's Osteology.)
The gradual perfecting of the fore- and hind-limbs as running organs has been put down to the advent of the grasses, and the formation of large plains covered with this herbage. The same reason would also be in harmony with the equally gradual change in the shape of the molar teeth, from a tubercular form calculated for a mixed or even a carnivorous diet, to the flatter crushing surfaces exhibited by the lophodont teeth of later Ungulates. Strong {199} canines would in the same way cease to be useful, and even become encumbrances to such grazing creatures; and their disappearance is one of the salient features in the history of the Ungulata, that is of the modern representatives of the order. The extraordinary hypertrophy of these teeth in such a line as that of the Amblypoda, which has left no descendants, was one of the reasons perhaps for the decay of those great pachyderms of mid-Tertiary times; their excessive armature became an encumbrance, since it was not accompanied by improvements in other necessary directions. Some of the features of the Tertiary Ungulates have, however, been dealt with in our general sketch of the mammalian life during that epoch, and need not be again referred to here. Of existing Ungulates there are no clear indications of the descent of the Elephants or of the Hyracoidea. Their structure proclaims these two divisions to be of ancient descent, and not to be modern twigs of the Ungulate stem. As to the Perissodactyla and the Artiodactyla we cannot bring them together nearer than in quite early Tertiary times. The order Condylarthra seems to be the starting-point of both these sub-divisions. Euprotogonia has been considered to be an ancestor of the Perissodactyle branch, and Protogonodon or Protoselene of the Artiodactyla. If this be true, {200} the likenesses which Titanotherium shows to the Artiodactyla must be either purely superficial and secondary, or a cropping out of ancient characters which had been dormant for many generations.
HORNS.--The Ungulata are the only order of mammals which possess horns; as they are on the whole a more defenceless group than the Carnivora, it may be that the horns are a counterpoise to the teeth and claws of the latter; need for defence and for armature in the combats with their own kind for the favours of the does has led to a different kind of protective and aggressive mechanism. Horns as weapons are, however, particularly effective in this group wherever they exist. A Ruminant is most frequently a large and heavy animal without the agility and litheness of the Carnivore. It is precisely to this sort of animal, where weight is an important consideration, that horns are the most suitable weapons. This is further shown by the fact that although the general term horn is used to describe the weapons of the Ungulate mammals, there is more than one kind of structure included under this general term; it is indeed probable that the extreme terms in the series of horns have been independently acquired by their possessors. There is but little in common between the horns of a Giraffe and of a Rhinoceros. In the Rhinoceros we have one or two horns, in the latter case one placed behind the other, which are purely epidermic growths; they may indeed be regarded as matted masses of hair, borne, it is true, upon a boss of bone, which however is not a separate structure. The Giraffe supplies us with the simplest term in that series of horns which are partly epidermal and partly bony. The paired horns of this animal have often been contrasted with those of the Deer, for example; but there is no fundamental difference between them. In the Giraffe a pair of bony outgrowths, originally separate from the skull which bears them, but ultimately ankylosed to it, are covered by a layer of entirely unmodified skin. A distinction of undoubtedly practical importance is usually drawn between the Hollow-horned Ruminants, i.e. Oxen, Goats and Antelopes, and the Deer tribe. There is nevertheless no fundamental distinction. In the Antelopes there is a core of bone, the "os cornu" as it has been termed, which is covered by a horny layer, the horn proper, variously modified in shape and size according to the genus or species. In the Deer there is the {201} same os cornu, which may however be branched, but which is in the same way covered by a layer of modified integument; this is known as the "velvet"; it only lasts for a certain period, and is then torn off by the exertions of the animal itself, leaving behind the bony core, which is popularly termed the horn. It will be clear that here is only a difference of comparative unimportance; the same essential features are present in both groups of animals, but the modification of the epidermis has progressed along different lines. Both can be referred back to the primitive conditions seen in the paired horns of the Giraffe. Even the difference, such as it is, is bridged over by the Antelope Antilocapra, where the os cornu is bifid and the horn is periodically shed, as is the velvet of the stag; but in the stag the bony part of the horn is also shed, a state of affairs which has no parallel in the Hollow-horned Ruminants. The great Sivatherium may conceivably be an annectant form between the two types of compound horns, i.e. those of the Antelope and those of the Deer. This creature had two pairs of horns, of which, naturally, only the bony cores remain; the hinder pair of these were branched. But although so far they resemble the Deer's horns rather than the Antelope's, Dr. Murie has thought that they were covered by a horny sheath and not by soft skin as in the Deer. In any case these horns were apparently never shed, which is a point of likeness with the Antelope and of difference from the Deer. Apart therefore from the nature of the covering of the bony cores, there are good grounds for looking upon them as intermediate between those of the Deer and those of the Antelopes.
The horns of the Ruminants are frequently a secondary sexual character; this is especially the case with the Deer. The Reindeer is, however, an exception, both the stags and the does having horns. That they are associated with the reproductive function is shown by their being shed after the period of rut, the destruction of the velvet at that period, and also by the effect upon the horns which any injury to the reproductive glands produces. Some useful facts upon this latter head have been amassed by Dr. G. H. Fowler,[122] who noticed in a series of stags, horns showing various degrees of degeneration in the antlers produced by varying degrees and periods of gelding. From the facts {202} here collected it is clear that a direct effect is produced. If we are to regard horns as secondary sexual appendages which have been subsequently handed on to the female by heredity, we should expect to meet with examples of animals now horned in both sexes, of which the earlier representatives had the horns confined to one sex. This is most interestingly shown by the extinct and Miocene Giraffe, Samotherium, of which the male alone had a pair of short horns, while the skull of the female was entirely hornless; the modern Giraffa, as is well known, has horns in both sexes.
It is interesting to note that the existing Perissodactyles and Artiodactyles are to be distinguished by their unpaired or paired horns. But while there are no Artiodactyles with unpaired horns (save occasional sports) the Perissodactyles have more than once tried, so to speak, paired horns, which ultimately proved fatal to them. The Rhinoceros Diceratherium apparently inherited and improved upon the small paired horns of Aceratherium, but it has left no descendant. The paired horned Titanotheria offer another instance of the same apparent incompatibility between the Perissodactyle structure and the persistence of paired horns.
SUB-ORDER 1. CONDYLARTHRA.
This group is characterised by the following assemblage of characters. Extinct, often plantigrade Ungulates, with five-toed limbs. Bones of carpus and tarsus not always interlocking, but sometimes lying above each other in corresponding positions. The humerus has an entepicondylar foramen. Dental formula quite complete; the molars brachyodont and bunodont. The premolars are simpler than the molars. The canines are small. As with other early types, the zygapophyses are flat and do not interlock. The astragalus is like that of the Creodonta. This group was American and European in range, the remains of its rather numerous genera being of Eocene time. The best-known genus is Phenacodus, of which some account will be given before discussing the, in many cases, more fragmentary remains of other allied forms.
The genus Phenacodus was first described so long ago as 1872, from a few scattered teeth. Since then several nearly complete skeletons have been obtained, and we are in full possession of {203} the details of its osteology. It was not a large creature (see Fig. 110, p. 196), about 6 feet in length, with a small head. The feet were more or less plantigrade, and five-toed. The last phalanges of the toes show that they carried hoofs and not claws; yet the fore-feet look a little as if they could be used as grasping organs. The third digit of both hind- and fore-feet exceeds the others, and thus a Perissodactyle-like foot characterised this Eocene creature. The tail is exceedingly long, and must have reached the ground as the animal walked. This is of course by no means an Ungulate character. Still, in the totality of its organisation the animal was decidedly Ungulate, though Professor Cope spoke of Phenacodus as not merely an ancestral Ungulate but as the parent form of Insectivores, Carnivores, Lemurs, Monkeys, and Man himself! The scapula indeed is from its breadth and oval contour rather like that of a Carnivore. The clavicles as in other Ungulates are absent. The femur is Perissodactyle rather than Artiodactyle in the presence of a third trochanter. The creature had fifteen pairs of ribs and five or six lumbar vertebrae. The two bones of the leg which lie below the femur are perfectly distinct and separate. A cast of the brain-case shows that the cerebral hemispheres were smooth and small, the cerebellum of course completely uncovered and nearly as large as the cerebrum. The olfactory lobes were also large. The complete skeleton of Phenacodus has lately been excavated more fully from the enveloping matrix by Professor Osborn,[123] and mounted in what is regarded as the natural position of the beast. It appears that though five-toed it went upon the three middle toes only, and furthermore that of these the middle one was the more prevailing, so that Phenacodus was distinctly "Perissodactyle," at least in habit. Moreover its "long hind-quarters, the long powerful tail ... are reminiscent of Creodont ancestry." The genus was European and American in range.
Meniscotherium ( = Hyracops[124]) comprises several forms of about the size of a fox; they are both European and American in range. The teeth are more distinctly Ungulate in form than those of Phenacodus, with a [125]-shaped outer wall. The skull is described as possessing "indifferent, primitive characters," permitting a comparison with those of Opossums, Insectivores, and {204} Creodonta. It has, as in Phenacodus, no orbital ring. The humerus resembles that of a Carnivore rather than that of an Ungulate. The carpus and tarsus are serial. The fibula articulates with both the calcaneum and the astragalus, which is not the case with Phenacodus. It is suggested that these animals are ancestral forms of the Chalicotheres. In the brain the hemispheres do not cover the cerebellum.
More primitive apparently than Phenacodus was the less-known genus Euprotogonia, or Protogonia[126] as it has been called. The best-known species is E. puercensis, so called from its occurrence in the Puerco beds of the American Eocene. It was a slender, long-limbed creature, smaller than Phenacodus, with a long and heavy tail as in that animal. Like Phenacodus it was semiplantigrade, and shows more likenesses to the Creodonta. The skull is only known by a part of the lower jaw with teeth, and by the teeth of the upper jaw. The vertebrae are not entirely preserved, but enough remain to show that the animal had a tail of 16 or 17 inches, which is a considerable length when compared to its height, about a foot at the rump. In the fore-limb the most noteworthy point is that the ulna has a convex posterior border as in the Creodonts, the same border in Phenacodus being concave. The humerus is slender, with less-marked tuberosities. The fifth digit seems to have been less reduced. The phalanges seem to have borne horny sheaths somewhat intermediate between hoofs and claws. The pelvis is described as being, as is also that of Phenacodus, rather like that of the Creodonta. The right hind-limb is known in all its details. It appears that the bones are not serial but interlocking; this, however, on the views with regard to the relations of these two forms of tarsus mentioned on p. 198, does not militate against regarding Euprotogonia as the ancestor of the genus Phenacodus. The third toe is the pre-eminent one, the animal thus being Perissodactyle. The lateral digits are larger than in Phenacodus, and the metatarsals and the phalanges are slightly curved, which is again a Creodont character as compared to the perfectly straight corresponding bones of Phenacodus. It seems evident that this animal is to be looked upon as a more ancient type than Phenacodus, even if not as its actual ancestor.
Another group of the Condylarthra contains the genus Pertipychus and some others. Periptychus has the full dentition {205} of forty-four teeth, the molars being of course bunodont, with the three chief tubercles most developed. The bones of the tarsus interlock and are not serial, as they are in many other members of the Condylarthra. The astragalus has a shorter neck than in Meniscotherium, for example. It has in this a likeness to the same bone in the Amblypoda, to the primitive members of which, such as Pantolambda, this animal bears much resemblance. "Astragali and many skeletal bones of Periptychus rhabdodon and Pantolambda bathmodon are almost indistinguishable," observes Mr. Matthew. The fore-feet of this genus are unknown, but it would seem that it was plantigrade from the evidence of the hind-feet. There are several species of the genus.
Possibly, but not at all certainly, the Mioclaenidae, with the genera Mioclaenus and Protoselene, are to be referred to this same order of primitive Ungulates. It is only necessary to mention them here, because they show very clearly the primitive form of dentition of these early Eocene mammals. The teeth are quite complete and unbroken by a diastema. The canines are but little pronounced. The molars are not strictly tritubercular, but have a prevailing trituberculy. The nature of the feet is not known. Since the genus Protoselene, as its name denotes, shows an indication of a commencing selenodonty, it has been suggested that this group is the stock whence the Artiodactyles have been derived.
In any case, whether the particular comparisons that have been made as to the relationship of various forms of Condylarthra are valid or not, it seems to be plain that this group represents the earliest Ungulate stock, but little differentiated from the contemporaneous Creodonts.
SUB-ORDER 2. AMBLYPODA.
This group of extinct mammals has the following principal characteristics:--
They are large, semiplantigrade Ungulates, of heavy build and apparently elephantine gait. The dentition is for the most part complete as in other ancient groups, and the canines are in the later forms big tusks. The back teeth are brachyodont and ridged (lophodont). Both radius and ulna in the fore-limb, and tibia and fibula in the hind-limb, are well developed. The bones {206} in the carpus are alternating in position. The toes are five in both feet, and are very short. There is a hint of commencing "perissodactylism" in the fore-feet at any rate. The brain is small and the hemispheres smooth.
The Amblypoda, or Amblydactyla, are so called on account of their short and stumpy feet and toes. They were held by Professor Cope to be on the direct line of ancestry of both Perissodactyles and Artiodactyles, a view which is on the whole not accepted at present.
FIG. 114.--Skull of Protolambda bathmodon. x 3/4. e.a.m, External auditory meatus; m, mastoid; m.f, mastoid foramen. (After Osborn.)
As is the case with other groups, the Amblypoda commenced existence as a sub-order with relatively small forms such as Pantolambda, the most ancient type known, which is in many respects a transition between the later forms and other groups of mammals such as the Creodonta.[127] The race culminated and ended in the giant Dinoceras and Coryphodon, and spread into the Old World. In spite of their smooth and diminutive brain, these mammals were able to hold their own and to multiply into many species and genera; in this they were perhaps aided by their formidable tusks and by the horns which many of them possessed. The teeth seem to imply an omnivorous diet, which was quite possibly an additional advantage in the struggle for existence. It does not seem to be necessary to divide off the Dinoceratidae into a sub-order equivalent to the Coryphodontidae as was done {207} by Professor Marsh; the numerous points in common possessed by the members of both families forbid their separation more widely than as families.
The earliest types of Amblypoda belong to the genus Pantolambda, of which the species P. bathmodon was about four feet in length. As restored it seems to have had proportionately short fore- and hind-limbs, and it had a long tail. It was apparently plantigrade, and would have had not a little likeness to a carnivorous type. The skull has no air cavities, such as are developed in the later types from the Lower Eocene, e.g. Coryphodon; Pantolambda is from the basal Eocene. The frontal bones show no trace of the horns that are developed in subsequent forms; the nasals are comparatively long; the zygomatic arch is slender. The molar teeth are in the primitive form of trituberculy, and the premolars, as is so often the case with primitive animals, are unlike the molars in form, being less markedly selenodont. As to the vertebral column, the dorsal vertebrae appear to have had short spines, which argues, as it does also in the case of the larger and heavier Coryphodon, a feebleness in the development of ligaments and muscles supporting and moving the head. The scapula seems to have the same peculiar leaf-like form that it has in the later Coryphodon.[128] This primitive type shows an entepicondylar foramen in the humerus. It is interesting to observe that the posterior border of the ulna is convex, as in the Creodonts, and in the early Condylarthrous form Euprotogonia. In the subsequently-developed Amblypoda, as in the later Condylarthra, that bone acquires a concave outer border. In the carpus the os centrale is distinct. In the femur the third trochanter is well formed; it gradually dies out in later Amblypoda. The fibula articulates with the calcaneum. This species, according to Osborn, "typifies the hypothetical Protungulate, being more primitive than either Euprotogonia or Phenacodus."[129]
FIG. 115.--Skeleton of Coryphodon radians. x 1/10. (After Osborn.)
The genus Coryphodon is known by a large number of species, of which the first was discovered in this country, and was represented merely by a jaw with some teeth. This was named by Sir R. Owen C. eocaenus, and was dredged up from the bottom of the sea off the Essex coast. A second specimen consisted of a single {209} canine tooth only, and was brought up from a depth of 160 feet during the making of a well at Camberwell. More abundant remains have since been found in North America.
This genus had a large head, and in some specimens traces of the "horn cores," so marked in the related Dinoceras, are to be noticed. The skull is broad behind and narrowed in front; the roofing bones show the cellular spaces so characteristic of the Elephant. The jugal bone, however, is not, as it is in the Elephant, placed in the middle of the somewhat massive zygomatic arch. As in some other primitive Ungulates (e.g. Phenacodus) there are twenty dorso-lumbar vertebrae, of which fifteen bore ribs.
The scapula seems to have possessed a peculiar leaf-like form, swelling in the middle and ending almost in a point above. It has a well-marked spine, and the acromion projects much. The fore- as well as the hind-feet are in a state of transition between plantigradism and digitigradism. It was at one time held that the animal was digitigrade as to the fore-feet and plantigrade as to the hind-feet. Though, as has been pointed out, it is a fact that the hind-feet are often on a different plane of evolution from the fore-feet, it seems that this amount of difference does not characterise any Ungulate, not excepting the genus now under consideration.
The toes are very spreading. The pelvic girdle is of great strength and broadness. The femur, as in the Perissodactyles, has a well-developed third trochanter; but whereas in this particular the hind-limb is Perissodactyle, it is Artiodactyle in the fact that the tibia and the fibula articulate with the astragalus and calcaneum. The ridged teeth have given the name to the genus.
A curious feature in the structure of the genus are the slender spines of the dorsal vertebrae, which contrast with the enormous ones of some other Ungulates--more curious in this genus, which is of heavy build, than in the lighter Pantolambda. The back of the animal is short, and the limbs are very spreading, so that the gait was doubtless shuffling. The large head, and short and heavy limbs and limb girdles added probably to its cumbrous walk or trot. The canines are great tusks, and spread out on both sides of the mouth.[130]
The late Professor Cope, in 1874, described the probable appearance of the Coryphodon in the following words:--"The general appearance of the Coryphodons, as determined by the skeleton {210} probably resembled the bears more than any living animals, with the important exception that in their feet they were much like the elephant. To the general proportions of the bears must be added the tail of medium length. Whether they were covered with hair or not is of course uncertain. Of their nearest living allies, the elephants, some were hairy and others naked.... The movements of the Coryphodons doubtless resembled those of the elephant in its shuffling and ambling gait, and may have been even more awkward from the inflexibility of the ankle."
The most recent members of this sub-order come from the Middle Eocene beds, and are chiefly referable to the genus Dinoceras, with which Tinoceras and Uintatherium are at least very nearly related, if not identical. These creatures were of great size, larger than the earlier types which have been considered. They show a certain superficial resemblance to the Titanotheriidae, on account of the massive horn cores upon the skull. These horn cores are large upon the maxillae and the parietals, and are paired; on the nasals are smaller horns. The bones of the skull have air cavities. The incisors of the upper jaw are absent; the canines are enormous tusks, and the lower jaws are flanged downwards near the symphysis where these tusks border them. Contrary to what is found in the older types, where the position of the condyle of the lower jaw is normal, this prominence faces backwards in the Dinocerata. The same shortness of the spines of the dorsal vertebrae prevails in this group as in the other Amblypoda, though it is perhaps hardly so marked. The scapula has not the peculiar acuminate form that exists in Coryphodon, but is triangular and broad above. The limbs are elephantine, in that the angle between the humerus and the femur respectively, and the bones which follow, is not marked. The hind-limbs are especially straight. The tail is short as compared with that of the primitive Amblypoda. The Dinocerata are purely digitigrade. The entepicondylar foramen has, as in the Coryphodonts, disappeared. The os centrale of the carpus has become fused, and no longer exists as a separate bone. The fibula no longer articulates with the calcaneum, but both that bone and the ulna are well developed. The genus Astrapotherium is placed among the Amblypoda by some authorities.[131]
{211}
SUB-ORDER 3. ANCYLOPODA.
The history of the discovery of the members of this order is very instructive as illustrating the dangers of laying too much classificatory importance upon detached fragments of animals. So long ago as 1825 terminal phalanges of a new creature were found in the Miocene of Eppelsheim, and sent to Cuvier. Cuvier named them "Pangolin gigantesque," deeming them, on account of their general form and cleft terminations, to pertain to the Edentata. In the same bed some seven years later were found certain teeth clearly of an Ungulate character, to which the generic name of Chalicotherium was applied. It was subsequently discovered that the teeth and the claws belonged to the same animal, and, later, further remains turned up which disclosed a creature having the anomalous composition of an Ungulate with decisively Ungulate teeth, but with the feet to a large extent like those of an unguiculate animal. The same confusion of characters occurs also, it will be remembered, in the distinctly Artiodactyle Agriochoerus (see p. 331). Indeed the feet of the latter when first discovered were erroneously, as it now appears, referred to the present order of Ungulates under the name of Artionyx. It is probable that the genus Moropus of North America is a member of this group, and that it is probably congeneric with a somewhat different type of Ancylopod known as Macrotherium. It is also clear that Anisodon, Schizotherium, and Ancylotherium, if not congeneric with either of the two recognised genera, are at least very close to them.
Chalicotherium has a skull which recalls that of some of the earlier Ungulates; it has, however, no incisors at all, and no canines in the upper jaw; this feature has led to the belief that the animal is related to the Edentata, and that it is in fact a link between them and the Ungulata. The molars, like those of the Perissodactyla, are of the buno-selenodont type. It also agrees with that group (to which it has been approximated by several writers) in the. tridactyl manus and pes, and in the characters of the tarsus. But although tridactyl, the axis of the limb passes through the fourth digit. Chalicotherium is not mesaxonic, as are the Perissodactyles. Moreover, it has no third {212} trochanter, and the unguiculate claws have already been referred to. As to the latter, which are short, it is not the end phalanx but the first which is retracted; thus Chalicotherium differs markedly from both Carnivorous and Edentate types; for in the former it is the last phalanx which is retracted, while in the Edentates the same phalanx is flexed downwards. The limbs of Chalicotherium are nearly of the same size, and the animal seems to have been stout and quadrupedal.[132]
Macrotherium, like the last genus, seems to have been common to both New and Old Worlds. It is to be distinguished by a number of characters. It is supposed to have been "semi-arboreal and fossorial"; the fore-limbs are much longer than the hind, the relative proportions of the radius and tibia being 70 to 29. The ulna was distinct from the radius, whereas in Chalicotherium the two are coalesced, or nearly so. Young specimens appear to possess a full set of incisors; whether this is the case or not with Chalicotherium is not known.[133]
Homalodontotherium is sometimes placed in the group.
SUB-ORDER 4. TYPOTHERIA.
It is a little difficult to be confident that the Typotheria are rightly referred to the Ungulata, since they contradict two important Ungulate rules. They have clavicles, which are elsewhere missing, and the thumb looks as if it were opposable.[134] An Ungulate is essentially a running animal, and has no need of a grasping finger. Still Typotheria are placed by most within the Ungulate series, though their undoubted likenesses to other groups, especially to the Rodentia, are admitted, and indeed emphasised. Cope places them definitely with the Toxodonts.
The Typotheria are an extinct group of smallish beasts, confined, like the Toxodontia, to South America, a region which during the Tertiary period, and into the Pleistocene, abounded with strange and varied types of Ungulate animals.
The earlier forms of Typotheria may be exemplified by some {213} account of the genus Protypotherium. This animal was of about the size of a Hyrax, which indeed it resembles in several points of structure. The teeth have the primitive number of forty-four, and they are close set, leaving no diastema; the molars are rootless and grow persistently; they are simple and Rodent-like in surface pattern. The shape of the lower jaw is like that of Hyrax, being rounded in outline posteriorly; there is no projecting angle as in the Rodents, and this remark applies to the Typotheria in general. The aspect of the Rodent lower jaw is characteristically different from that of Hyrax and the forms under consideration.
Some other characters of these early forms of Typotheria can be gathered from an inspection of other genera. In Icochilus both hand and foot were five-toed, and, as in ancient Ungulates generally, the bones of the wrist and of the ankle are serially and not alternately arranged. Moreover, an os centrale is present in the carpus. Both thumb and big toe were opposable. The skull has a remarkably Rodent-like appearance, but the palate is not so narrowed as in these animals.
In the more recent forms of Typotheria the dentition has become reduced. The canines are lost, and as the incisors are reduced also, to one on each side of the upper, and two on each side of the lower jaw, the likeness to a Rodent skull is increased. There is also evidence of a modification from the more primitive forms in the loss of one premolar or even more, in the alternating bones of the carpus, in the disappearance of the centrale, and in the loss of a toe upon the hind-foot. In these more recent forms the fibula articulates with the astragalus instead of with the calcaneum. Typotheria of these more recent forms may be illustrated by the typical genus Typotherium. It has the reduced dental formula I 1/2 C 0/0 Pm 2/1 M 3/3; the molars are simple in pattern, and much like those of Toxodon. The upper incisors are powerful and curved, but are surrounded by a layer of enamel, which is not limited to the anterior face, as it practically is in Rodents. The sacrum is composed of a large number of vertebrae--some seven--a state of affairs which recalls the Edentata. The shoulder blade is not Ungulate in form. It has a strong spine, with an acromion and a well-developed metacromion. The terminal phalanges are enlarged and hoof-like.
In the genus Pachyrucos there are three premolars, otherwise {214} the formula is the same as in Typotherium. The animal seems to have had nails rather than hoofs. The thumb was opposable. The fibula is fused below with the tibia, whereas in the last genus these two bones are quite separate from each other.
SUB-ORDER 5. TOXODONTIA
The group Toxodontia,[135] like so many others, is exceedingly hard to define. Nor are its limits any easier to mark out than many others of the groups of Ungulates. It will be best perhaps to give an account of Toxodon, and of a few types which seem to lie near it in the system, and then to indicate how far they resemble or depart in structure from other Ungulates. Toxodon itself is known from complete skeletons. It lived in Argentina during the "Pampean" period, which seems to be of the Pleistocene age. A large number of species, however, have been described, some of which seem to go farther back in time, and to have existed during the Miocene period further south in Patagonia.
The size of this creature was about that of a large Rhinoceros; it has a bulky body and a large head, which was borne low down, on account of the bending downwards of the anterior vertebrae; in this aspect the figures of the skeletons recall Glyptodon and similar Edentates. The beast was discovered by Darwin, and originally described by Owen. "During his (Mr. Darwin's) sojourn in Banda Oriental," writes the Rev. H. Hutchinson, "having heard of some 'giants' bones' at a farmhouse on the Sarandis, a small stream entering the Rio Negro, he rode there, and purchased for the sum of eighteenpence the skull which has been described by Sir R. Owen. The people at the farm-house told Mr. Darwin that the remains were exposed by a flood having washed down part of a bank of earth. When found, the head was quite perfect, but the boys knocked the teeth out with stones, and then set up the head as a mark to throw at." The whole of the Pampean area is a valley of dry bones, and the remains of Toxodon are abundant there. The skull of Toxodon is not unlike that of a horse in general aspect; but the orbit is not separated from the temporal fossa. The premaxillae are furnished above with a slight protuberance directed towards {215} the free end of the nasals, which may be related to the presence of a short proboscis. The zygomatic arch is strong and broad: the mandibles are provided with a long symphysis. The dental formula is I 2/3 C (0-1)/1 Pm 4/(3-4) M 3/3. The teeth are prismatic and hypselodont, growing from persistent pulps. The molar teeth are slightly arched in form, whence the name of Toxodon, "bow teeth." The strong chisel-shaped incisors suggest the Rodents and Hyrax. The cheek teeth, moreover, are by no means unlike those of Rodents in their pattern. They are at any rate not at all like those of existing Ungulates. The small size of the canine and of the first premolar produces a diastema in the tooth series. The sacrum consists of five vertebrae, and the ischium does not articulate with it.
The shoulder blade has a strong spine, but only a rudimentary acromion; nor is the coracoid well developed. The radius crosses the ulna, as in the Elephant; the whole fore-limb is shorter than the hind-limb, which must have exaggerated the hang-dog expression of the creature when alive. The elements of the carpus interlock in the modern fashion. Those of the tarsus, however, are primitive in lying below each other without alternation. The carpus has a centrale. The fibula articulates with the calcaneum. The femur has no third trochanter. There are three toes to all the limbs. It is clear that this assemblage of characters will not allow the placing of Toxodon in any living Ungulate order. If the middle toes appear by their slight pre-eminence to approach the Perissodactyle form, the peculiar surface contour of the molar teeth, letting alone the absence of a third trochanter on the femur, will not permit this classification.
Allied to Toxodon is the genus Nesodon. It was so named from an "island lobe" on the inner side of the upper molars. This creature, smaller than Toxodon, also differs from it in the fact that the dentition is complete, and in the pattern of the molars, which is rather more complex. There is still the slight projection upon the premaxillary bones, but the nostril is directed more forwards than in Toxodon. The zygoma, too, is massive. The second pair of incisors in the upper jaw and the outer (third) pair in the lower jaw form biggish tusks in the adult. These and the molar teeth are, however, finally rooted, and do not grow, as in Toxodon, from persistent pulps. The genus is from the older Tertiary of Patagonia. Five or six species have been described. Some are as large as a Rhinoceros, others as small as a Sheep. {216}
There is no doubt about the close alliance of the two genera just referred to. It is more doubtful whether Homalodontotherium and its allies should be placed, as they often are, in the neighbourhood of the Toxodonts. Homalodontotherium owes its name to its even row of teeth without a diastema. It was a creature of equally large size with Toxodon, and also came from the Tertiary strata of Patagonia. The teeth are the typical forty-four, and the molars like those of a Rhinoceros; they are, however, brachyodont and not hypselodont as in Toxodon. This genus, however, shows an important difference from the Rhinocerotidae and from the other Toxodontia in the fact that it was five-toed, and that the bones of the carpus and tarsus are set in relation to each other in the linear serial fashion.
Undoubtedly a near relative of Homalodontotherium is Astrapotherium. This creature was of equal bulk, and was also Patagonian in range. The teeth are reduced in number, but the animal was provided, like a Wild Boar, with great tusks, which were, however, formed by the incisors. This animal is very imperfectly known; it is the form of the molars and the large size of the incisors which have led to its association with the Toxodontia. As to the resemblance of the teeth of this genus and of Homalodontotherium to those of Rhinoceros, it is difficult to regard it as evidence of near affinity. The likeness is probably to be looked upon as a case of parallelism in development. Exactly the same explanation is possibly to be given to the likeness which the teeth of Toxodon and Nesodon show to Rodents, or even to Edentates. As to their affinities Zittel observes:--
"The entirety of their osteological characters argues for the Toxodon a separate position in the neighbourhood of the Perissodactyla, Proboscidea, Typotheria, and Hyracoidea. The relations to the Rodentia rest mainly upon the converging development of the teeth, not upon true relationship."
SUB-ORDER 6. PROBOSCIDEA.
Large vegetable-feeding animals, usually scantily covered with hair, and with the nostrils and upper lip drawn out into a long proboscis. Digits five on both limbs. Femur and humerus not bent upon lower leg and fore-arm in a position of rest. Skull {217} with abundant air cavities in the roofing and other bones. The incisors are developed into long tusks, which exist in the upper jaw alone, in the lower jaw alone, or in both jaws. There are no canines. The molars are lophodont. The clavicle is absent. The femur has no third trochanter. The bones of the carpus are serially arranged and do not interlock. The stomach is simple. The brain has much convoluted cerebral hemispheres, but the cerebellum is completely uncovered by them. The intestine is provided with a wide caecum. The testes are abdominal. The teats are pectoral in position. The placenta is non-deciduate and zonary. There are two venae cavae superiores.
The position of the limbs in the Elephant tribe is unique among living animals: their straightness that is to say, and the absence or very slight development of angulation at the joints of the limb bones. This same feature has been observed in the extinct Dinocerata and in the Titanotheria. It must not, however, be assumed from the resemblance to these ancient forms that there is much affinity between them and the Proboscidea, or that the latter have retained an ancient feature of organisation. The oldest Ungulates for the most part, and the Creodonts to which they are undoubtedly related, have much bent limbs. It must be considered, therefore, that the arrangement obtaining in the Elephants is purely secondary. Professor Osborn has put forward the reasonable view[136] that the vertical limbs of all these colossal creatures are due to "an adaptation designed to transmit the increasing weight" of these animals. The huge bulk of the body is better borne by vertical pillars than by an angulated limb. Other points, however, such as the exposure of the cerebellum, the two venae cavae, the five digits, and the absence of a third trochanter, argue a low position for the Proboscidea in the Eutherian group.
The group can be readily divided into two families, the Elephantidae and the Dinotheriidae. We will commence with the former.
The Elephants proper, ELEPHANTIDAE, differ from the Dinotheriidae in, and are characterised by, a number of anatomical features. They possess long tusks (incisors) either in both jaws, or, if only in one jaw, in the upper. The molar teeth are very large--so large that only a few of them are simultaneously in use. There are but three definable genera of Elephantidae, of which {218} Elephas alone survives. This genus also includes many extinct forms, both American and European, as well as Asiatic and African. The entirely extinct genera are Stegodon and Mastodon. The group is clearly one dwindling towards extermination. From the Middle Miocene downwards these great "pachyderms" have existed; and from the Miocene up to Pleistocene times they were almost world-wide in range and numerous in species.
The genus Elephas comprises usually large, but occasionally (the pygmy Elephant of Malta) quite small forms. The external features of the genus differ slightly in different species, and will therefore be described in relation to those species which we shall notice here. The vertebral formula is C 7, D 19-20, L 3-5, Sa 4-5, Ca 24-30, or even more.
The bodies of the vertebrae are remarkable for their shortness and for the very flattened articular surfaces.
FIG. 116.--A section of the cranium of a full-grown African Elephant, taken to the left of the middle line, and including the vomer (Vo) and the mesethmoid (ME); an, anterior, and pn, posterior narial aperture. x 1/12. (From Flower's Osteology.)
The skull is large and massive. Its large and heavy character is, as has been stated in the definition of the sub-order, due to the immense development of air cavities in the diploe; the diameter of the wall of the skull is actually greater than that of the cranial cavity. These cavities are not obvious in the young animal. They are most conspicuous in the roofing bones of the skull, but are seen elsewhere, and thicken the basis cranii, {219} the maxillae, and so forth. This state of affairs, together with the presence of the huge tusks, has, as it were, pushed back the nasal orifices to near the top of the skull in a very Whale-like fashion. As in the Cetacea, the nasal bones are limited in size, and the premaxillae send up processes to join the frontals and the nasals. There is a straight and somewhat slender zygomatic arch, but the orbit is not separated from the temporal fossa. The malar bone is small, and, as in Rodents, forms the middle part of the zygoma. This is not the case with most Ungulata. The symphysis of the mandibles forms a spout-like rim. The scapula has a narrow prescapular, but a very wide postscapular region. The spine has a strong process projecting backwards from near its middle; this is a point of likeness to certain Rodents. No Elephant has a clavicle. The most remarkable feature about the fore-limb is the separation and crossing of the radius and ulna. The arms of these animals are permanently fixed in the position of pronation. The foot is short, and the bones of the carpus are serially arranged. There are, however, traces of a commencing interlocking of these bones in many forms. The hind-feet are somewhat smaller than the fore-feet, and the tibia and fibula are both developed.
As to the teeth, this genus is to be distinguished from allied forms by the presence of tusks in the upper jaw only. These tusks have no bands of enamel such as characterise those of Mastodon. They are incisors. There is, however, a trace of the former enamelling in the shape of a patch at the tip, which soon wears away. The molar teeth of Elephas are so large that the jaws cannot accommodate more than at the most two and a part of a third at a time. These are gradually replaced by others to the number of three, the replacement of teeth suggesting that of the Manatee. Each molar is deeply ridged, the interstices between the ridges being filled up with cement. As the tooth wears away, therefore, the surface continues to be flat. Each ridge consists of a core of dentine surrounded by a coat of enamel. The number of these ridges varies greatly from species to species. The Indian Elephant is one of those which have the greatest number of plates in a single tooth, as many as twenty-seven.[137] Of the six molars which {220} eventually appear, the first three are considered to correspond to premolars. But successional teeth are rare in the genus; that is to say as far as concerns the molars, for the tusks have their milk forerunners. As to the molars it is apparently only E. planifrons which certainly shows a milk dentition. In Mastodon and older types a milk dentition is commoner.
The viscera of the Elephant have been examined by many zoologists. The latest paper, dealing chiefly with the African species, but containing facts about its Indian congener also, is quoted below.[138] The Elephant is remarkable in possessing, in addition to the three usual pairs of salivary glands present in mammals, a fourth, situated in the molar region, and opening on to the cheek by many pores. This gland is especially well developed in Rodents. There is a gland which may be mentioned in this connexion, though it opens externally between the eye and ear, known as the temporal gland; its use does not seem clear. The thoracic cavity of the Elephant, as may be inferred from the large number of ribs, is very large as compared with the abdominal.
The stomach is simple in form, and the epithelium of the oesophagus does not extend into it as is the case with the Horse and Rhinoceros. A gland or a collection of smaller glands occurs in the stomach, and recalls the "cardiac gland" of the Wombat and the Beaver, also that of the Giraffe. The large intestine is long, rather more than half the length of the small intestine. The caecum is well developed in these animals. The liver has a very simple form, being but slightly lobulated. It is actually only bilobed, but it is important to notice that this division does not correspond to the two halves of the liver. As shown by the attachment of the suspensory ligament, one half consists of the left lateral lobe alone, the other half embracing the remaining primary lobes. The simplicity of the liver looks like an archaic character. No Elephant has a gall-bladder. The lungs again are simple in form through their slight lobulation. Each half in fact is without subdivisions, and is of a triangular form. In this the Elephants resemble the Whales, as in the simple liver. In both cases probably the likeness is due to the permanence of primitive features of organisation. The brain[139] of the Elephant {221} has hemispheres which are extremely well convoluted; but they leave the cerebellum entirely uncovered. This suggests a brain which is a great specialisation of a low type. The brain has been particularly compared with that of the Carnivora, with which group the Elephants agree in the characters of the placenta. It is, however, always a matter of the very greatest difficulty to compare the brains of mammals belonging to different orders.
There are but two living species of Elephant, of both of which we shall now proceed to give some account. Only a few of the rather numerous fossil forms can be touched upon here.
The African Elephant, E. africanus, has been sometimes referred to a distinct genus or sub-genus, Loxodon, by reason of the lozenge-shaped areas on the worn grinding-teeth. It lives, as its name denotes, in Africa. This species has a number of external features which enable it to be distinguished from the Oriental Elephant. The head slopes back more, and has not the two rounded bosses which give so wise a countenance to the Indian species. The ears are very much larger. The tip of the trunk has a slight triangular projection on both the lower and the upper part of the circumference of the aperture. There are four nails on the fore-feet and three on the hind. As in the Indian form, the toes are all bound together, and do not appear for any part as free digits. A thick pad of fat, etc., makes the animal when alive look as if plantigrade, whereas it is, as a matter of fact, digitigrade. In internal features the most prominent difference from E. indicus is in the molar teeth, which are ridged by much fewer ridges. The outside number for a single tooth in the present species is 10 or 11. In Elephas indicus on the other hand there are as many as 27.
The African Elephant, thinks Sir Samuel Baker, reaches a height of about 12 feet, and it will be remembered that the notorious "Jumbo" was found to be 11 feet high at the shoulder. The tusks are found in both sexes, as in the Indian beast, but are relatively larger in the female in the species now under consideration. It is also a rather more active creature, and is more savage;[140] however it can be tamed, as is shown by several {222} specimens which have been and are in the possession of the Zoological Society, and other proprietors. It was apparently used in the past. Certain Carthaginian coins are stamped with a figure of the African Elephant; but in Africa no attempts are now made to utilise this creature except for food and ivory.
FIG. 117.--African Elephant. Elephas africanus. x 1/56. (After Sir Samuel Baker.)
The meaning of an Elephant as an emblem. upon a coin appears to be eternity, and there is no question but that the {223} Elephant is a long-lived animal. It is said that it hardly reaches proper maturity before forty, and that 150 years is not beyond probability in the way of longevity. Even longer periods have been assigned to it.
The tusks of the Elephant are by no means necessarily sexual adornments, used for fighting purposes only. The African Elephant is a most "industrious digger," and grubs up innumerable roots as food. It appears to be a fact that during these operations the right tusk is mainly used, and in consequence that tusk is shorter as well as thinner than the other. Two average tusks would weigh respectively 75 and 65 lbs., the latter of course being the weight of the more worn right tusk. These weights, it should be observed, by no means indicate the limits to which finely-developed tusks can attain. The very heaviest tusk known to Sir Samuel Baker[141] weighed 188 lbs. This was sold at an ivory sale in London in the year 1874. The pace of the African Elephant, says the same authority, is at most at the rate of fifteen miles an hour at first, and of course in a furious rush. This pace cannot be kept up for more than two or three hundred yards, after which ten miles an hour is a better approximation to the rate which can be kept up for long distances.
The Indian Elephant, Elephas indicus (or Euelephas indicus, if the genus Loxodon is to be accepted), is better known and has been longer known than the African. It occurs in India and Ceylon, and in some of the Malayan islands, the Elephants of which latter parts of the world have been regarded as a distinct form, an apparently unnecessary procedure.
FIG. 118.--Indian Elephant. Elephas indicus. x 1/54. (After Sir Samuel Baker.)
This species does not stand so high at the shoulder as the African; its back is more rounded in the middle. The trunk has but one pointed tip; there are five nails on the fore- and four on the hind-feet. As this species comes from India and the East, it has been longer as well as better known than the African form. Thus many of the stories and legends that have congregated round Elephants apply really to this form. As is well known, the Indian Elephant is much used as a beast of burden, and for other purposes where its huge strength renders it invaluable. But its great drawback as a servant of man is its great independability. On the one hand we have furious, vicious, and generally unreliable {224} Elephants, and on the other perfectly docile creatures, who obey the slightest hint from their driver. Huge though the Elephant is, it is frequently a timid beast. Sir Samuel Baker relates how one which he was riding fairly bolted at the sight of a Hare. To {225} be bolted with by an Elephant is far from pleasing, though a rather exciting event. It makes for the nearest jungle at once, being, much more than the African species, an inhabitant of forest. And in rushing through the dense undergrowth, the occupiers of the Elephant's back are apt to be swept off or cut to pieces by innumerable thorns.
Elephants, no doubt of the Indian species, were used by the Persians in battle, and from fifteen which were captured at the battle of Arbela some notes were drawn up by Aristotle. In stating that the animal reaches an age of 200 years, the naturalist and philosopher was probably not very far out. The mode of Elephant-catching as related by Aristotle is that pursued at the present day. Then, as now, tame Elephants were made use of as decoys. Pliny,[142] who was apt to confound fact and fiction in a somewhat inseparable tangle, had something to say about Elephants, both Indian and African. Serpents, he thought, were their chief enemies, which slew them by coiling round them and thrusting their heads into the trunk, and so stopping respiration. In Europe Elephants were first seen in the year B.C. 280. Pyrrhus used them in his invasion, and copying his example the Romans themselves learnt to use Elephants. The first Elephant seen in England arrived in the year 1257, presented by the King of France to Henry III. It was kept in the Tower (for long afterwards a menagerie), and died at twelve years of age. Much use of the Elephant has been made in symbols. We have spoken of the African Elephant on Carthaginian coins as an emblem of eternity. The Oriental Elephant resting on the back of a tortoise and supporting the world is the same idea; and it is instructive to note that remains have been found in the Siwalik Hills of a tortoise which would have been actually big enough to support the creature, even "Jumbo," who weighed 6-1/2 tons. Another symbol is that of an Elephant upon whose back is a child with arrows; this occurs on a medal of the Emperor Philip. It can perhaps hardly signify the eternity of a strong human feeling!
The intelligence of the Elephant has been both exaggerated and minimised. Perhaps the most elaborate attempt to endow the beast with unusual mental perceptions is that of Aelian, who related that an Elephant carefully watching his keeper, wrote after him with his trunk letters upon a board. That the animal does {226} possess a good deal of brains, seems to be shown by the way in which a well-trained animal will obey the slightest sign of the mahout in India. According to Sir Samuel Baker, localities which produce in abundance particular kinds of fruit are remembered, as well as the time at which the fruit will be at its best. Stories of revenge, which are numerous enough, attest, so far as their data are to be accepted as accurate, the power of memory possessed by the Elephant.
In spite of their longevity, however, Elephants, unlike Rome, have not been built for eternity. We can only find two living species; but in past times Elephants were very numerous. They commenced, so far as we know, in the Miocene.
The existing forms are known in a fossil, or at least sub-fossil state, from diluvial deposits; and it is interesting to note that the African Elephant had formerly a wider range than now. Its bones (described as E. priscus) have been met with in Spain and Sicily.
One of the best known of completely-extinct Elephants is the Mammoth, E. primigenius. This great Elephant in most respects more nearly approached the existing Indian Elephant. The teeth have quite as numerous plates. The tusks were enormous, reaching a maximum length of 15 feet; they were much curved upwards as well as outwards. A large tusk weighs as much as 250 lbs. The Mammoth was of exceedingly wide range. Not only was it found in various parts of Europe, but it was especially abundant in Siberia, as is exemplified by the fact that for the last two hundred years as many or more than 100 pairs of tusks annually have been sold from that region. It also occurred in America together with forms at least not far removed from it, such as E. columbianus. Mammoths have been more than once found as entire carcases in the frozen soil of Siberia. The first was discovered in the year 1799, and rescued some years later for the St. Petersburg Museum. This example showed that the Mammoth, unlike existing Elephants, was covered with thick wool mingled with long and more bristly hairs of some 10 inches in length. The softer wool formed a kind of mane beneath the neck, which hung down as far as the knees. Another carcase was discovered later by Lieut. Benkendorf, who did not save it, but was nearly swept along with it into the sea by a flood. These creatures died in the position in which they were found by being bogged when in search of vegetation or water. {227}
How primeval man, with his inferior weapons, slew the Mammoth is not easy to understand; but that they were contemporaneous is clearly shown by associated remains, and by the notorious sketch of the Mammoth on a piece of its own ivory, in which curved tusks and a forehead like that of an Indian Elephant are plainly to be seen. Although it was only so recently as the year 1799 that an example of this great creature was actually studied on the spot, and removed to St. Petersburg, the existence of Mammoths and of ivory is a matter of much more ancient knowledge. M. Trouessart relates[143] that fossil ivory was known to the Greeks. Theophrastus spoke of ivory imbedded in the soil, and the tusks were recovered by the Chinese. It is a curious fact that the Chinese described and figured the Mammoth as a kind of gigantic Rat. The likeness between the elephantine molar and that of Rodents has been commented upon; but the existence of its tusks below the level of the ground led the Chinese Natural Historians to consider that the ways of life of the Mammoth were those of the Mole. As to the carcases themselves, the Chinese said that the flesh was cold, but very healthy to eat. This expression can hardly be explained, except upon the view that fresh carcases were known to that people long before they were known to us of the Western world. The value of the Mammoth ivory was known to antiquity; the famous Haroun-al-Raschid gave to King Charlemagne not only a pair of living Elephants, but a "horn of Licorne," which seems undoubtedly to have been a name for the tusks of the Mammoth. For in an account of the sacred treasures of Saint Denis, published in the year 1646, the author states this to be the fact.
The causes of the disappearance of the Mammoth are not easy to understand. Some held that it was a naked animal like the existing Elephants, and that the lowering of the temperature in Siberia proved fatal; it is, of course, now certain that it was clothed with dense woolly hair. Along with the bogged corpses of the great pachyderm, numerous trunks of pine-trees have been found, together with associated remains of other animals now extinct in that neighbourhood. Thus it is plain that Siberia was once covered by mighty forests, through which the Mammoth roamed. The decay of these forests, upon whose branches the Elephant fed, as is attested by the remains of pine leaves found {228} in the interstices of its teeth, was the signal for the disappearance of their most colossal inhabitant.
The large number of remains of this and of other extinct species of Elephas in this country gave rise to the supposition that they were Elephants brought over by Caesar to aid in the subjugation of these islands. The Rev. J. Coleridge (father of the poet) pointed out that though Caesar in his Commentaries made no mention of any such importation of Elephants, a passage in the Stratagems of Polyaenus expressly mentions that Cassivelaunus was confronted by the Romans with an Elephant clad in a coat of mail, by whose aid the crossing of the Thames was effected. At the time that attention was called to this (1757) it was not popular to hint at the possibility of fossils. So that fact, conveniently historical, served to explain away a difficulty. It is remarkable that the Elephant, common enough of course in Asiatic monuments, actually occurs in English architecture. Mr. Watkins, from whose interesting work (Natural History of the Ancients) a good many of the facts detailed here are drawn, tells us that the church of Ottery St. Mary has an Elephant's head sculptured on one of its pillars. The same ornament appears in Gosberton Church, Lincolnshire. Whether this has anything to do with a reminiscence of formerly existing Elephants is a hard question to answer. In this figure of an Elephant the trunk has a spiral representation, and the trunk of an Elephant is believed by some to be intended by the common "so-called Pictish ornamentation" in Scotland; this spiral alone is to be seen constantly. If it is a reduction of an Elephant to its simplest terms, it is highly interesting as an almost undoubted survival of remembrance of Elephants. For at such a period we cannot use the memories of Crusaders or others who may have visited the East to explain the facts. The sculptured Elephants' heads might conceivably be so explained.
The name Mammoth, thinks Mr. Watkins, may be derivable from the Arabic word Behemoth. He quotes a writer, who first described the beast in 1694, as using the two words indifferently. The Arabs, moreover, were then as they are now great ivory traders; and in the ninth and the two succeeding centuries explored the confines of Siberia, as they now do the forests of Africa, for ivory. The "Behemoth" of Job "eateth grass as an ox.... He moveth his tail like a cedar" (the Hippopotamus has a much more {229} stumpy appendage). "Behold, he drinketh up a river, and hasteth not" is surely much more suggestive of the copious draughts of an Elephant than the possibly equally copious but not so visible libations of a Hippopotamus.
The most ancient of the true Elephants (genus Elephas) is E. meridionalis. It is of the African type, i.e. the plates of the molar teeth are not abundant, and are not so many as in the existing E. africanus. It seems to have been one of the largest of Elephants, standing 4 metres high. Its remains are abundant in Europe, and are known also from England. Like this species E. antiquus is also of the African type. It was contemporary with man. Certain dwarf or "pony" races found in caves in Malta, and called Elephas melitensis or E. falconeri, are believed to belong to this species. Mr. Leith Adams, who described these[144] remains, placed them in two dwarf species called by the names used above, and found associated with them a larger form, which he referred to E. antiquus. The existence of these animals in Malta seems to argue at least its former larger dimensions, and the presence of more abundant fresh water. The remarkable swimming capabilities of the Elephant do not necessarily imply either a former absence of land connexion or, on the other hand, its existence. Nor as a third possibility can it be suggested that the dwarf size argues an island of limited dimensions, when we bear in mind the huge tortoises of the Galapagos and some other islands. It is important to notice that Elephants of the African type (Loxodon) were not formerly absent from India. E. planifrons was one of these.
The genus Stegodon is so called from the fact that the molar teeth, seen in longitudinal section, present a series of roof-shaped folds, the interstices between which are not, or are, imperfectly filled up with the cement which in Elephas reduces the surface of the teeth to a level plane. This genus is exclusively Asiatic, and is Miocene to Pleistocene in time range. The number of ridges on the molars is small, not more than two. The incisors (tusks) have no enamel; the skeleton generally is like that of Elephas, between which and Mastodon the present genus is intermediate. Among the four or five species is S. ganesa (called after the Indian Elephant-headed divinity), with tusks 10 feet long, to be seen at the British Museum of Natural History. {230}
The last genus of the family Elephantidae is Mastodon, so called from the structure of the molar teeth. These are provided with but few transverse ridges, not more than five, so that their structure is intermediate between those of Dinotherium and those of Stegodon. Between the ridges are sometimes isolated, boss-like protuberances (whence the name of Mastodon), produced by a subdivision of the ridges. There is either but little or no cement between the ridges. This genus differs from nearly all other Elephantidae by the possession of milk molars, which occasionally persist throughout life, the permanent dentition in those cases being a mixture of milk and permanent teeth, as has been (erroneously) stated of the Hedgehog.[145]
The tusks (incisors) are sometimes present in both jaws, and as they have, during youth at any rate, a coating of enamel, the likeness to the chisel-shaped incisors of Rodents is patent. In connexion with the implantation of incisors in the lower jaw, many species have a prolongation of the bones of that part of the skeleton. In the bones, generally, there is not very much difference from Elephas, but the forehead is a little less pronounced. The genus existed from the Miocene and became extinct in the Pleistocene. It was nearly world-wide in range, being known from all four continents. Naturally with this very wide range was associated a large number of species. Zittel enumerates no less than thirty-two.
This genus is the only one of the Elephantidae which extended its range into South America, where the remains of two species occur. The bones of these great Elephants have attracted attention for some centuries. They were often held to be the bones of giants (as they actually were!), and in one case were ascribed to a deceased monarch, Teutobochus. The American Indians considered that equally gigantic men lived who were able to combat these great Proboscideans. There are legends of the Mastodons as living animals, which is quite probable, considering their geological age. There is a curious parallelism between the legends of two such widely-separated localities as North America and Greece. Buffon relates how among the Indians of Canada there was a belief that the Great Being destroyed both Mastodons and men of equal proportions, with thunderbolts. With this we may perhaps compare the story of the destruction of Typhoeus by Zeus, who {231} also used thunderbolts. One of the giants was not slain, but was compelled to stand and bear up the heavens. Atlas holds thus the position of the Elephant supporting the globe of Indian mythology.
FIG. 119.--Dinotherium giganteum. Side view of skull, 1/15th natural size. Miocene, Germany. (After Kaup.)
The genus Dinotherium, sole representative of the family DINOTHERIIDAE, differs in a number of important particulars from the true Elephants. In the Elephants, if there is but a single pair of incisors, these are found in the upper jaw; in Dinotherium there is apparently but a single pair, but these are implanted in the lower jaw, the symphysis of which is much prolonged and greatly bent downwards, so that the tusks emerge at right angles to the long axis of the head, and are even bent backwards. The molar teeth are five in number on each side of each jaw and are bi- or tri-lophodont, not unlike those of the Tapir. There is no cement in the valleys between the ridges of these teeth, and there is a regular succession, the premolars being two and the molars three.[146] All the teeth are in use at the same time, {232} their small size enabling them to be accommodated in the jaw together. The skull of Dinotherium is lower than that of Elephas or Mastodon. The bones of the skeleton generally are like those of Elephas.
Though a suggestion of marsupial bones attached to the pelvis has been discredited, there is no doubt that Dinotherium occupies the most primitive position among the Proboscidea; but at the same time it cannot be regarded as the ancestor of Elephants, as it is so much specialised in various ways. The incisors for one thing forbid this way of looking at the creature. It is an ancient genus found in beds of Miocene age in Europe and Asia. It is not known from America. The creature was larger than any Elephant. Eighteen feet in length has been assigned to it. The enormous weight of the lower jaw and tusks seems to argue that it was at least partially aquatic in habit, and that it may have used these tusks for grubbing up aquatic roots or for mooring itself to the bank. At first there were naturalists who considered it as an ally of the Manatee, and the skull is not unsuggestive of that of the Sirenia.
Pyrotherium has been referred to the Proboscidea; but our knowledge of that form is limited to a few teeth from Patagonian rocks of an uncertain age.[147] They are simple bilophodont molars, very like those of Dinotherium. A tusk has been found in the neighbourhood of these teeth which may possibly belong to the same animal; but it is uncertain.
SUB-ORDER 7. HYRACOIDEA.
This group of small mammals contains only one well-marked genus which is usually named Hyrax, although Procavia seems to be the accurate term. Popularly these creatures are known as Coneys. They have a singular resemblance to Rodents, the short ears and much reduced tail, besides the squatting attitude adopted, contributing to this merely skin-deep likeness. They agree with other Ungulates in the structure of the molar teeth, which are much like those of Rhinoceros; in the absence of a clavicle; in the absence of an acromion; in the reduction of the digits of the limbs to four digits in the manus and three in the pes. On the {233} other hand they differ from most Ungulates in the incisors growing from persistent pulps, a point in which they resemble the Rodentia. The muffle also is split as in those animals. The Hyracoidea are peculiar in the fact that in addition to the caecum at the junction of the small and large intestines, there are a pair of caeca (bird-like in being paired) some way down the large intestine. The dorsal vertebrae are unusually numerous, 22. The adult dentition according to Woodward,[148] who has recently examined the matter, is I 1/2 C (1/0) Pm 4/4 M 3/3, while the milk dentition is I 3/2 C 1/1 Pm 4/4.
FIG. 120.--Cape Hyrax. Hyrax capensis. x 1/8.
The inclusion of the canine of the permanent set of teeth in brackets signifies that it is the milk canine which occasionally persists. It should further be remarked about the teeth that they are both hypselodont and brachyodont, the extremes being connected by intermediate forms. Another peculiarity of the genus is the dorsal gland, which is covered with hair of a different colour to that covering the body generally. This is present in all species.
The genus Hyrax (the most recent authority on the subject, Mr. Oldfield Thomas,[149] only allows one genus) is limited in its range to Ethiopian Africa and to Arabia, including Palestine, It does not reach Madagascar. Mr. Thomas allows fourteen species with two or three sub-species. {234}
Some of the Coneys live in rocky ground, while others, formerly placed in the genus Dendrohyrax, frequent trees, in holes in which they sleep. The Coney of the Scriptures is familiar, who is "exceeding wise," though a "feeble folk." But the further observation that he "cheweth the cud but divided not the hoof," is obviously entirely wrong. As to the wisdom, it is said that this beast is too wary to be taken in traps; while the suggestion of chewing the cud is, according to Canon Tristram, to be interpreted in the light of a habit of working and moving its jaws which the animal has. The traveller Bruce kept one in captivity to see if it did really chew the cud, and found that it did!
* * * * *
{235}
Chapter X
UNGULATA (continued)--PERISSODACTYLA (ODD-TOED UNGULATES)--LITOPTERNA
SUB-ORDER 8. PERISSODACTYLA
FIG. 121.--Bones of the manus A, of Tapir (Tapirus indicus). x 1/5. B, of Rhinoceros (Rhinoceros sumatrensis). x 1/5. C, of Horse (Equus caballus). x 1/8. c, Cuneiform; l, lunar; m, magnum; p, pisiform; R, radius; s, scaphoid; td, trapezoid; tm, trapezium; u, unciform; U, ulna; II-V, second to fifth digits; V in B, and II and IV in C, represented by rudimentary metacarpals. (From Flower's Osteology.)
These Ungulates derive their name, which is that given by the late Sir Richard Owen, from the fact that the middle digit of the hand and foot is pre-eminent. As will be seen from Fig. 121, the axis of {236} the limb passes through the third finger, which is larger than any of the others, and is symmetrical in itself. In this the present group contrasts with the Artiodactyla, where the axis is not "mesaxonic," but where there are two digits, on either side of the axis, which are symmetrical with each other. This arrangement of the limbs is highly characteristic, but appears to be not quite universal. In the Titanotheres, which form a group of the Perissodactyles, the fore-limbs are not quite accurately mesaxonic. Nor on the other hand can all Ungulates which show the Perissodactyle condition be safely included in the present group. The ancient Condylarthra and the Litopterna show precisely the same state of affairs. But other features in their organisation lead to their separation from the Perissodactyles, of which, however, the Condylarthra are probably ancestors. The Litopterna on the other hand, which possess even one-toed members like Equus, are believed to represent a case of parallelism in development. The number of functional toes varies from four to one. In the ankle joint the astragalus either does not, or does only to a comparatively slight extent, articulate with the cuboid as well as with the navicular bone. Moreover the fibula when present does not as a rule articulate with the calcaneum. In the opposed group of Artiodactyles the precise reverse of these conditions obtains. It is usually stated as part of the definition of this group that they do not possess horns of the type of those met with in the Cervicornia and Cavicornia. But the strong bony bosses on the skull of many Titanotheres, so curiously reminiscent of those of the not nearly related Dinoceras and Protoceras, may well have supported horns of the Ox and Antelope pattern.
FIG. 122.--Bones of the manus of Camel (Camelus bactrianus). x 1/8. c, Cuneiform; l, lunar; m, magnum; R, radius; s, scaphoid; td, trapezoid; u, unciform. (From Flower's Osteology.)
The teeth of the Perissodactyles are lophodont, more rarely bunodont. The selenodont Artiodactyle form of molar is not met with. The dental formula, moreover, is at least near the {237} complete one, the more modern forms as usual being the more deficient in numbers of teeth.
The dorso-lumbar vertebrae are as a rule twenty-three; but the extinct Titanotheres are again an exception; for, at least in Titanotherium, there are but twenty of these vertebrae--an Artiodactyle character. The femur has a third trochanter. There are so few recent Perissodactyles that an enumeration of the distinguishing characters of the viscera may very probably be useless for purposes of classification. But the living genera at any rate are to be separated from the living Artiodactyles by the invariable simplicity of the stomach coupled with a very large and sacculated caecum. The liver is simple and not much broken up into lobes, and the gall-bladder is always absent. The brain is well convoluted. The teats are in the inguinal region. The placenta in this group is of the diffused kind.
FIG. 123.--Anterior aspect of right femur of Rhinoceros (Rhinoceros indicus). x 1/2. h, Head; t, great trochanter; t', third trochanter. (From Flower's Osteology.)
The living Perissodactyles belong to three types only, indeed to three genera only (in the estimation of most), which are the Horses, Tapirs, and Rhinoceroses. But taking into account the extinct forms, they may be divided primarily (according to Professor Osborn) into the four following groups:--(1) Titanotherioidea, including but one family, Titanotheriidae; (2) Hippoidea, including the families Equidae and Palaeotheriidae; (3) Tapiroidea, with two families, Tapiridae and Lophiodontidae; and (4) Rhinocerotoidea with families Hyracodontidae, Amynodontidae, and Rhinocerotidae. It is conceivable, according to the same writer, that the Chalicotheres (here treated of as a separate sub-order, Ancylopoda) should be added to the Perissodactyle series.
FIG. 124.--Side view of skull of Horse with the bone removed so as to expose the whole of the teeth. c, Canine; Fr, frontal; i^1, i^2, i^3, incisors; L, lachrymal; m^1, m^2, m^3, molars; Ma, malar or jugal; Mx, maxilla; Na, nasal; oc, occipital condyle; Pa, parietal; pm^1, situation of the vestigial first premolar, which has been lost in the lower, but is present in the upper jaw; pm^2, pm^3, pm^4, remaining premolars; PMx, premaxilla; pp, paroccipital process; Sq, squamosal. (After Flower and Lydekker.)
FAM. 1. EQUIDAE.--This family, which includes the living Horse, Zebras, and Asses, as well as a number of extinct genera agreeing with those types in structure, may be defined by the possession of but one functional toe, the two lateral ones being mere splints, or but little more. The molar teeth are hypselodont, and {238} the premolars, with the exception of the first, resemble the molars in their pattern. The orbit is completely surrounded by bone. The incisors are chisel-shaped, with a pit on the free surface. The canines are rudimentary if present. The radius and ulna are fused, as are the tibia and fibula. Although for the sake of uniformity a family, Equidae, is here separated from its allies, it is quite impossible owing to the full state of our knowledge of this group to draw a really hard-and-fast line between this family and the Palaeotheriidae. We shall deal presently with the conjectured pedigree of the Horse, which naturally involves that family, and which presents an unbroken series from four-toed Perissodactyles to the present one-toed Horse, the various bones and teeth becoming modified in the course of the descent "with the regularity of clockwork." We are compelled to draw the line at functional second and third toes; directly these are no longer used the animal is a Horse in the strict sense! This is irrational and regrettable, but necessary for practical purposes, if {239} we are to continue the plan of defining the various families of Mammalia.
The genus Equus[150] contains not only the Horse, but the Asses and Zebras. The genus is to be distinguished as regards external characters by the following features:--The body is thickly clothed with hair; there is a more or less bushy tail and mane; the colours are apt to be disposed in stripes of black or blackish upon a yellowish brown ground; this is of course best seen in the Zebras, but the wild Asses also have some traces of it, if only in the single cross-bar of the African Wild Ass, and it is even "reversionary" in the domestic Horse at times. There are no horns upon the forehead or elsewhere; the fore-limbs or both pairs have a callous pad upon the inside, which is possibly to be looked upon as an aborted gland, possibly originally of use as secreting some odorous substance calculated to enable strayed members of the herd to regain their companions. The terminal phalanx of each of the (functionally) single digits is enclosed in a large horny hoof.
The main internal features of structure which divide this genus of Perissodactyles from the Rhinoceros or the Tapir, or from both, are: the existence of strong incisors, three on each side of each jaw; there are canines, but these are small and do not always persist in the full-grown mare. They are popularly known as "tusks" or "tushes." The first of the four premolars (the "wolf tooth") is small and quite rudimentary; it is often absent. As there are three molars, the present genus has the "typical" number of the Eutherian dentition, i.e. forty-four. In the skull the orbit is--as it is not in Tapirs and Rhinoceroses--completely encircled by bone. There is but one functional finger and toe on each hand (Fig. 121 C) and foot; the second and third digits are represented by mere splints, one of which may as an abnormality be enlarged, and reach nearly as far as the well-developed digit. There are even occasionally traces of digit number two.
The Horse, E. caballus, is to be distinguished from its congeners by the small callosities on the hind-limbs which it possesses in addition to the larger ones on the fore-limbs. The hairy covering of the tail is more abundant, as is also the mane. The head too is proportionately smaller, and the general contour {240} more graceful. Though Zebra markings are not usual upon E. caballus, there are plenty of examples of--what we may perhaps in this case term--a "reversion" to a striped state. The celebrated "Lord Morton's mare,"[151] whose portrait hangs in the Royal College of Surgeons, is an interesting case of this. It was as a matter of fact thought to be an example of that rather doubtfully-occurring phenomenon, "telegony." Its history is briefly this. The animal was the offspring of a mare that had previously produced to a male Quagga a hybrid foal. Afterwards a second foal was produced by the same mare to an Arab sire. This foal, the one in question, was striped, and hence was thought to be an example of male prepotency. But instances are known of unquestioned Horses which show the same stripes, such as a Norway pony which had not even seen a Zebra!
A last remnant of the naked palm of the hand and sole of the foot is left in the shape of a small bare area, smaller in the Horse than in the Asses, known technically as the "ergot," the term being that of the French veterinarians. As already mentioned, the Horse differs from the Asses and Zebras in the fact that the hind-limbs have callosities on the inner side. They are known as "chestnuts," and their nature has been much disputed. It has been suggested that they are the last rudiment of a vanished toe; but in all probability they are, as already suggested, traces of glandular structures, which are common, upon the limbs in many animals (see above, p. 12).
It is a singular fact that there are apparently no wild Horses of this species. The case is curiously analogous to that of the Camel, which also is only known as feral or domesticated. Why the Horse should have become extinct as a wild animal, considering that when it does run wild it can thrive abundantly, is impossible to understand. Sir W. Flower thinks[152] that "the nearest approach to truly wild horses existing at present are the so-called Tarpans, which occur in the Steppe country north of the sea of Azov between the river Dnieper and the Caspian. They are described as being of small size, dun colour, with short mane and rounded obtuse nose." But he adds that there is no evidence to prove whether they are really wild. In favour, however, of their possibly being wild and indigenous European Horses, may be {241} mentioned the fact that their general build and appearance is highly suggestive of the wild Horses sketched by primitive man upon ivory.
A really wild Horse, and possibly the ancestor of the European domestic Horse, is E. przewalskii of the sandy deserts of Central Asia. This animal has been believed to be a mule between the Wild Ass and a feral Horse; but if a distinct form, and probability seems to urge that view, it is interesting as breaking down the distinctions between Horses and Asses. The species possesses the four callosities of the Horse, but has a poorer mane and an asinine tail.
There is no question that the Horse has been a domestic animal for very many centuries. Hieroglyphics appear to show that the Egyptians had not originally domesticated the Horse; it seems to have been first introduced among them by the Hyksos or Shepherd Kings.[153] Whatever the date may be, it is certain that considerably anterior to the Egyptians the Assyrians and Phoenicians possessed Horses. In Western Europe the date of the introduction of the Horse seems to have been during the bronze epoch. Lord Avebury[154] has pointed out that out of eighteen cases of graves in which the remains of Horse were found, twelve contained metal implements, i.e. 66 per cent. This does not of course prove that the Horse was domesticated at that period, but it throws doubt upon the earlier occurrence of the Horse in abundance. The Horse, however, does occur on the Continent associated with the remains of man during the Quaternary period.[155]
Messrs. Cuyer and Alix enumerate between fifty and sixty domesticated races of Horse, not counting the supposed wild varieties which have been already referred to. These may be further subdivided; for instance, under the race "pony" we may distinguish the Irish, Scotch, and Shetland varieties, all of which, however, according to Sanson, have originated in Ireland. They are used, remark the authors above quoted, "par les jeunes filles des lords pour leurs promenades." The Arab, the Barb, the Suffolk Punch, etc., are among the numerous races of domestic Horses, into which to enter properly would require another volume, and that of large size. {242}
The Asses and Zebras differ from the Horse in the characters mentioned under the description of Equus caballus. In addition to these may be pointed out a feature to which attention has been directed by Mr. Tegetmeier.[156] According to him the period of gestation in the Horse is only eleven months; in the others more than twelve.
FIG. 125.--Asiatic Wild Ass. Equus onager. x 1/20.
Opinions as to the number of species of Asses differ. On the most liberal estimate there are three Asiatic and two African species. The best known of the Asiatic Wild Asses is the Onager, E. onager. It is of a uniform yellowish, "desert" colour, with a dark stripe along the middle of the back, and is found in Persia, the Punjab, and the country of Cutch. The creature is of great swiftness; it has been stated to be untameable, but Mr. Tegetmeier makes the absolutely opposite statement that the Ass occasionally "becomes so tame as to be troublesome"! The Syrian Wild Ass, E. hemippus, hardly, if at all, differs from this.
FIG. 126.--Nubian Wild Ass. Equus africanus. x 1/20.
The Kiang, E. hemionus, seems to have more claims to distinctness. In the first place it has a more limited and a {243} different distribution; it is confined to the high tablelands of Thibet at an elevation of 15,000 feet and upwards. In correlation with this habitat it has a thicker and more "furry" coat, which is, moreover, of a darker shade than that of the Onager. This coat is shed in the summer, and replaced by one which is not so dark in hue. It is an interesting fact that the African Wild Asses approach to the zebra type in having at least traces of stripings. There are apparently two species. The best known, the Nubian Ass, E. africanus, is probably the parent of the domestic donkey. It has a dorsal longitudinal stripe, and another across the shoulder--in legend the marks of the Saviour. The matter of the name of this Ass seems difficult to decide. It has been called also E. asinus and E. taeniopus. It has been observed that this animal has a great aversion to water, and a delight in rolling in the dust--both of which characteristics argue a desert existence. But on the other hand the Kiang will plunge boldly into streams, yet it would seem to be the descendant of a purely desert form. The Ass is a longer-lived {244} animal than the Horse. Mr. Tegetmeier calls attention to a donkey living in 1893 which had been ridden fifty-five years previously. The Horse, on the other hand, lives not much more than twenty-five years.
A second species of African Wild Ass, E. somalicus,[157] is distinguished by its greyer colour, by the absence of the shoulder stripe, by the very faint development of the dorsal stripe, and by the presence of numerous cross stripes upon the legs. It has, too, smaller ears, and a longer and more flowing mane. Mr. Lort Phillips, an experienced naturalist and traveller, saw a herd of these Wild Asses in Somaliland, which he regarded as being of quite a new species. A living example in the Zoological Society's Gardens led Mr. Sclater to an identical conclusion, which was supported, as he pointed out, by the fact that this Ass has a different range to the African or Nubian Wild Ass.
Of the Zebras three species are usually allowed; these are E. zebra, the "Mountain" or "Common" Zebra, E. burchelli, E. grevyi, as well as E. quagga. Professor Ewart thinks that the Common Zebra, Burchell's, and the Quagga are not very distinctly marked off from each other. No one, however, has any doubt of the distinctness of E. grevyi. This latter differs from the rest in its larger size, in the large head and ears, and in the marked hairiness of the ears. It would seem to be a primitive type of Zebra, if the fact that the occasional reversion of hybrids to a parent form be allowed; for Professor Ewart found a cross-bred Zebra to present several characteristics in the face-marking of this, the finest of the Zebra tribe. Only four specimens of E. grevyi have been exhibited alive in Europe--two in Paris, and two in the Zoological Society's Gardens in London. The latter were presented to Queen Victoria by King Menelek of Abyssinia. The species was named by Professor A. Milne-Edwards in honour of a late President of the French Republic, from an example also sent by King Menelek.
The Common Zebra has closer and darker stripes than Burchell's, but not quite so close as in E. grevyi. It has also a very characteristic arrangement of stripes on the withers in the form of a gridiron. This latter is wanting in both the other species. In E. grevyi, in fact, this part of the back is white. E. zebra has also a dewlap in front. E. burchelli has fewer and broader {245} stripes, and between them lie in many cases shadow-stripes of a faint brown.
FIG. 127.--Burchell's Zebra. Equus burchelli. x 1/20.
All these animals, and the Quagga too, are absolutely confined to Africa. Mr. R Crawshay,[158] in describing what he considered to be a new variety, remarked upon the curiosity of E. burchelli. "They remain out in the sun on the plains all day long, not retiring into covert at all. They are then an intolerable nuisance to any one in pursuit of other game; indeed this may be said of them at all times. If once they notice you, they draw in and mob you in their curiosity--only, however, when one takes no interest in them, for when they fancy they are the object of the intruder's attention, no animals are more watchful and cunning in safeguarding themselves. If only their curiosity were manifested in silence it would not so much matter, but it vents itself in snorts and thundering stampedes, which puts every beast within earshot on the qui vive."
Whether Burchell's Zebra[159] can be further subdivided into species or sub-species appears to be doubtful. Dr. Matschie considers that Equus boehmi may be regarded as a valid form, and in addition to this two sub-species, E. burchelli granti and {246} E. burchelli selousi, have been proposed for what are at most local races. But it is at present far from certain whether their distribution favours this subdivision.
The Quagga was more striped than is sometimes represented in illustrations. According to Dr. Noack, from whose paper[160] upon the animal I quote here, the transverse stripes reached back as far as the buttocks; they were, however, completely absent from the legs. The animal is, as every one knows, probably completely extinct. In the year 1836 it was still abundant; in 1864 the last specimen ever exhibited was received by the Zoological Society. Mr. W. L. Sclater thinks that it may have survived in the Orange River Colony as late as 1878, but admits that any certainty is difficult, as it was frequently confounded by the Boers with Burchell's Zebra. Its rarity is emphasised by the fact that it is not mentioned in the recent work of that most skilful of hunters, Mr. F. Selous. Gaudry places the Quagga nearest of all living Equidae to the Hipparion gracile of Pikermi.
FOSSIL EQUIDAE.--The existing Equidae all belong to the genus Equus, though there are some who would (quite unnecessarily) divide off the Zebras as a genus Hippotigris. The genus Equus itself goes back in time to the Pliocene, during which epoch there lived in India E. sivalensis, the same species according to some with the E. stenonis of Europe. None of these species, Old World or New, are easily to be separated from E. caballus. But many names have been given to them. It is of course perfectly conceivable that they may have differed among themselves as much as do the existing Zebras and Asses, the separation of which would be hardly possible did we know their bones only. There are, however, extinct genera, undoubtedly related so closely to Equus as to be placed in the same family, though clearly separable as genera. Hipparion is one of these genera; its remains are known from Europe, Asia, and North Africa, from beds of Miocene and Pliocene times. A large number of different species have been described. It was a beast of about the size of a Zebra. The principal characters are that each foot has three toes, of which, however, the two side ones are smaller than the central toe. There is a marked round fossa on the maxillary bone, a feature shared by the South American Onohippidium.[161] The pattern of {247} the molar teeth is, too, a little different from that of Equus. Protohippus of the North American Pliocene is also three-toed, but the two additionally-developed toes are smaller than in Hipparion. Other forms are dealt with below in connexion with the ancestry of Perissodactyles. It is a curious fact about Hipparion, which is not now regarded as on the direct line of equine descent, that the edges of the enamel plates of the molars may show a complicated folding very like that presented by that clearly terminal form of Perissodactyle life, the gigantic Elasmotherium. This is indicative of high specialisation, which ended in extinction.
ANCESTRY OF THE HORSES.--The LOPHIODONTIDAE and the PALAEOTHERIIDAE are two of the most interesting extinct families of Perissodactyles; for among them we find what would appear to be the ancestral forms of both the existing Tapirs and Horses. The Rhinoceroses also would seem to be derivable from the Palaeotheriidae. The very vagueness of the characters of these creatures, considered from a classificatory point of view, has led to much diversity in their placing. This though gratifying to the evolutionist is tiresome to the writer who wishes to give a methodical account of their various characters. It will be best perhaps not to attempt an accurate placing or to reconcile conflicting opinion, but to give some salient features of osteology which lead to the belief in their relationship to existing groups of Perissodactyles. A book upon the history of mammals would be incomplete without some account of that well-ascertained series of forms which seem to connect these primitive Perissodactyles with the modern Horse. Equus, in fact, is not only the "show horse" of the doctrine of evolution, but also the "stalking horse."
In the Eocene of both Europe and America are met with a number of forms from which we may start. Hyracotherium, regarded on the one hand as the type of a sub-family of the Equidae themselves, and on the other as a member of the family Lophiodontidae, was a small-sized animal, three feet or so in length; it possesses the complete Eutherian dentition with a slight diastema. The orbits are not separated from the temporal fossa; the fore-limbs were four-toed, the hind three-toed, with moderately long metapodia, especially on the hind-feet. The shoulder blade {248} has a well-marked coracoid process. The radius and ulna are separate; so too are the tibia and fibula. Eohippus, belonging to the same sub-family, is slightly more primitive; for the hind-feet have a rudiment of digit I. Orohippus is a little nearer to the Horses in that the molar teeth have acquired a little further advance towards the equine type. Instead of the tubercles of the teeth remaining for the most part separate, they have fused into a set of ridges, of which, however, the pattern is less complex than in the modern Horses. In other respects Orohippus is much like Hyracotherium. Pachynolophus seems to be but a synonym.
The next stage is shown by Mesohippus, a Lower Miocene form, usually referred to the neighbourhood of Palaeotherium. It has nearly lost one of the toes of the fore-foot, a rudiment only remaining; the metapodials, at any rate of the fore-feet, seem to be slightly increased in length. The orbit is not encircled by bone, but there is a strong process from the frontal, which nearly meets the zygomatic arch.
Anchitherium, from the Upper Miocene, is not far removed in structure from the last-mentioned form; it is a trifle nearer the existing Horse in several points. The ulna is further reduced and fused with the radius below: the rudiment of digit V is still more rudimentary; the two lateral digits are smaller in proportion to the central one than they are in Mesohippus; the fibula is fused below with the tibia. From this form to Equus is a small series of steps, characterised by the still further reduction of all the digits except III, by the still further reduction of the already rudimentary ulna and fibula, and by the increasing depth of the molar teeth, which are of course, in Equus, hypselodont.
Another interesting conclusion may seem to follow when we consider the geographical range of the ancestral Horses. Hyracotherium and Pachynolophus occurred both in the Old and New World. From them may have arisen the Horses of both hemispheres. After that point there is a division. Mesohippus is American, and we get at Equus in that continent through Desmatippus and Protohippus. On the other hand there are no remains known of Mesohippus in Europe; and unless subsequent researches prove the existence of Mesohippus, we have to rely upon forms which are placed with Anchitherium and Hipparion.
It seems that in America the next genus in the direct line of equine descent to Mesohippus is Miohippus. It is smaller in {249} size than Anchitherium, to be considered immediately. The odontoid process of the axis is just beginning to assume the characteristic spout-like shape of that of the existing Horse and many modern Ungulates. The median digit of both fore- and hind-limbs has become greatly enlarged as compared with the corresponding digit of earlier forms.
It is held, however, that Anchitherium is not on the direct line of descent either in America or in Europe, in both of which it occurs. Its teeth are in some respects less Horse-like than in some of the more ancient genera, to which the converse would be expected on the descent theory. Its hoofs are much elongated and flattened, a mark of specialisation and not appropriate to a creature holding an intermediate position in the equine series. Both the American (A. equinum) and the European species (A. aureliense) are of very large size, larger than its successors, and such "alternations in bulk are unlikely."
The genus Desmatippus of Professor Scott[162] fills in the gap between Miohippus and Protohippus. The molars and premolars are brachyodont, but there is a thin deposit of cement in the tooth valleys, leading towards the more complete filling of these valleys with cement, which is found in Protohippus. This genus of Horses, of which there is at present but one species, D. crenidens, was three-toed, and "the lateral digits, so far as can be judged by fragmentary remains, were still fairly developed, and though much more reduced than in Miohippus, appear to be somewhat less so than in Protohippus."
To recapitulate, the following is the probable series of equines in America--Mesohippus, Miohippus, Desmatippus, Protohippus.
The development of the limbs of the Horse shows a most interesting series, of stages, which correspond in part to the ancestral forms which palaeontology seems to prove to be the line of the descent of our existing Equidae. This matter has recently been elucidated by Professor Ewart, who details the following facts and comparisons:--
In the youngest embryo (about 20 mm. in length) the humerus is somewhat curved, and considerably longer than the radius and carpus taken together. The first-named bone is shorter in the adult, and the proportions of that bone in the young as well as its curvature are suggestive of that ancient {250} Ungulate Phenacodus (see p. 202). In the next stage (an embryo of 25 mm.) the humerus has slightly decreased in proportionate length, and has come to be more like that of Hipparion. In both of these embryos it should be noted that the ulna is complete and separate from the radius. In the second of the two it has more distinctly acquired the form which it will possess in the adult. The second metacarpal--one of the splint bones of the adult--is tipped with a small nodule of cartilage, which is clearly the representative of one or more of the phalanges belonging to that digit.
FAM. 2. TAPIRIDAE.--The Tapirs may be distinguished from the Horse and from the Rhinoceros tribe by a few characters, which are as follows:--
The dentition is generally the full one of forty-four teeth. The premolars in the more ancient forms are unlike the molars, but like them in more recent forms. The molars of the upper jaw have two crests parallel and united by an outer crest. The fore-feet have four, the hind-feet three toes.
The family is fully as ancient as that of the Equidae, but the specialisation of the toes never advances so far. The modern representatives of the order are, so far as the feet are concerned, in the condition of very early representatives of the equine stock. Nor do the teeth of the Tapirs ever reach the complicated pattern of that presented by at least the modern Horses, or indeed of the Palaeotheres. Apart from this it is not an easy matter to distinguish accurately between these several families, including the Lophiodontidae, which, as already mentioned, is placed nearer to the Tapiridae than to the Palaeotheriidae. Indeed the differentiation of these two families, the Tapiridae and the Lophiodontidae, seems to be a matter of the greatest difficulty. The difficulty is well emphasised by the fact that naturalists disagree most profoundly as to the relations of various genera of extinct Tapir-like animals. For Mr. Lydekker the genus Lophiodon includes also the American genera Isectolophus and Systemodon, which are placed by Zittel in the sub-family Tapirinae as opposed to Lophiodontinae, which contains Lophiodon and Helaletes. The existing Tapirs can be differentiated from the existing Horses with great ease, as the following account of the existing genera will show.
{251} [Illustration]
FIG. 128.--American Tapir. Tapirus terrestris. x 1/10.
The genus Tapirus is now met with only in South and Central America, and in the Malay Peninsula and the islands of Java and Sumatra. This animal is in many respects the most ancient of existing forms referable to the Perissodactyle order. It has four toes on the front-feet, though only three on the hind-feet. The number of teeth is 42--nearly the typical Eutherian number. The Tapirs are always moderately-sized animals, entirely covered with hair, and usually of a brownish-black colour. The Malayan Tapir is, however, banded broadly with white--a single band; the young of the Tapir is spotted, and striped with white. The nose and upper lip conjoined are produced into a short trunk, precisely comparable with that of the Elephant. As in the Rhinoceros--and in this both contrast with the other existing Perissodactyle genus Equus--the temporal fossa is not separated from the orbit by bone. Of existing Tapirs there are at any rate T. terrestris,[163] T. roulini (the "Tapir Pinchaque" of Cuvier), T. dowi and T. bairdi in America (the last two being sometimes separated into a distinct genus, Elasmognathus, on account of the prolongation of the ossified mesethmoid), and T. indicus in the East. The tapir, probably T. terrestris, is described by Buffon as "a dull and gloomy animal." It is certainly mainly nocturnal in habit. The name terrestris was given by Linnaeus, who placed it in the same genus as {252} Hippopotamus amphibius; hence the epithet applied to the Tapir. But as a matter of fact it loves marshy neighbourhoods, and is in a way amphibious. This does not of course apply to the Andesian T. roulini, which inhabits the cordillera of Ecuador and Colombia. The distribution of existing Tapirs is, as is so often the case, restricted when compared with that of their extinct congeners and allies. In Europe the remains of the genus Tapirus are abundant from Pliocene strata, and its remains are there known from as far back as the Miocene. The genus is thus one of the very oldest forms of Mammalia at present inhabiting the earth.
FIG. 129.--Malayan Tapir. Tapirus indicus, young. x 1/10. (From Nature.)
The Malayan Tapir is to be distinguished from the American (T. terrestris--the other species have not been dissected) by the greater development of the valvulae conniventes in the intestine, the absence of a moderator band in the heart, and the less elongated caecum, which is sacculated by only three bands, there being four in T. terrestris.[164] The animal frequents the most retired spots among the hill woods, by which habit it seems {253} largely to escape the Tiger, its most formidable foe in those regions of the world. Its quickness of senses enables it also to slip away with rapidity. It can proceed at a great pace when disturbed, and can readily push its way through obstacles. The young animal, like that of the American species, is dark brown with yellowish spots. It is stated by Mr. H. N. Ridley that the young animal lies during the hot part of the day under bushes, in which situation "its coat is so exactly like a patch of ground flecked with sunlight that it is quite invisible." It is interesting to note that here, as with some other animals, it is the young that are especially protected by such mechanisms. Moreover, some of the spots are round and some are more elongated, so that the resemblance to spots of sunlight which come in a direct and in a slanting direction is greatly increased. Even the colours of the adult are not so conspicuous when it is in its native haunts as might be supposed. The breaking up of the ground colour into tracts of two different colours prevent it from striking the eye so plainly as if it were of one colour throughout. "When lying down during the day it exactly resembles a grey boulder, and as it often lives near the rocky streams of the hill jungles, it is really nearly as invisible then as it was when it was speckled."[165]
FAM. 3. RHINOCEROTIDAE.--This family is to be distinguished from the preceding by a number of characters, which though not universal are general. In the first place, there are commonly horns, or a horn, consisting of what appears to be an agglomeration of hair-like structures fixed upon a roughened patch of bone on the surface of the nasals. The incisors are diminished or defective, and the upper canines are often wanting. The molars and premolars are alike. The fore-feet are four- or three-toed, but are functionally tridactyle; the hind-feet are three-toed. The skeleton in this family is massive, and the limbs relatively short. The skull, as in the Tapirs, has a confluent orbit and temporal fossa. The upper lip is generally more or less prehensile; the body is as a rule--to which the Pleistocene Hairy Rhinoceros is of course an exception--rather sparsely covered with hair. In this feature the Rhinocerotidae contrast both with the Tapiridae and the Equidae. The family in reality contains but one existing genus, though three have been instituted, viz. {254} Rhinoceros, Ceratorhinus, and Atelodus. As there are so few existing species the subdivision of animals which agree in so many and such highly-characteristic features seems to be an unnecessary procedure. The existing Rhinoceroses are but a fragment of the total number of known forms from past epochs. The family is very markedly on the wane.
The genus Rhinoceros is characterised by its heavy build and thick, almost smooth, skin--smooth, that is to say, so far as concerns the slight development of hair--which is often thrown into folds. There is one or there are two horns on the fore-part of the head, which are, as has already been pointed out, structures sui generis, and not exactly comparable with the horns of other living Ungulates. There are three nearly equal toes on both fore- and hind-limbs. The canine teeth of existing species have disappeared; the incisors are, or are not, present; the molars and premolars are three and four in each half of each jaw.
The visceral anatomy of the Rhinoceros has been much investigated so far as concerns the Asiatic forms. A curious feature, which serves to discriminate some of the Asiatic species from others, is to be seen in the small intestine. In Rh. indicus[166] this gut is furnished with numerous long cylindrical narrow outgrowths "like tags of worsted"; in the allied Rh. sondaicus these tags are present, but are flatter and broader; while in the two-horned Rh. sumatrensis there are no tags at all, but only smooth valve-like folds. Another mark by which these species can be distinguished depends upon the variation in the presence or absence of certain glands imbedded in the integument of the foot--the so-called "hoof glands." These occur in Rh. indicus and Rh. sondaicus, but are absent in Rh. sumatrensis.
Sir W. Flower[167] studied some years since the skull features which serve to differentiate the existing forms.
In Rh. sumatrensis the two long downward processes of the squamosal bone, termed respectively post-glenoid and post-tympanic, do not unite below the auditory meatus. In this the species in question agrees with the African forms but not with the one-horned Asiatic species, where the two processes completely fuse. Again, another character, though perhaps less important, {255} is the sloping backwards instead of forward of the occipital crest in all two-horned species, whether African or Asiatic.
The Asiatic Rhinoceroses have, what the African animals have not, functional incisor teeth throughout life. It has been proposed on these and other grounds to separate generically the African and Asiatic forms.
FIG. 130.--Indian Rhinoceros. Rhinoceros indicus. x 1/40.
The Asiatic Rhinoceroses include three well-differentiated species, in all of which the skin is much thrown into folds. Rh. indicus is the largest form. It is one horned, and has enormous folds of skin at the neck and hanging over the limbs. So like artificial armour is this thick plating, that Albrecht Duerer may be excused for having given the beast the appearance of being actually mail-plated in a sketch which he made of a specimen sent over to the King of Portugal in 1513. This particular beast, one of if not the first sent over to Europe, proved so intractable in disposition that the king sent it as a present to the Pope. But "in an access of fury it sunk the vessel on its passage"! The horn of this and of other species was held until almost our times to have medicinal and other more curious values. So recently as 1763 it was gravely asserted that a cup made of its horn would fall to pieces if poison were poured into it. "When wine is poured therein," wrote Dr. Brookes in the year referred to, "it will rise, ferment, and seem to boil; but when {256} mixed with poison it cleaves in two, which experiment has been seen by thousands of people." John Evelyn also wrote of a well in Italy which was kept sweet by a Rhinoceros' horn. This species seems to be long-lived, even in captivity; a specimen now to be seen in the Zoological Society's Gardens has been there since the year 1864.
Rhinoceros sondaicus, the Rhinoceros of the Sunderbunds, has a much wider range than the last species or Indian Rhinoceros. This is unknown out of India itself, and is there limited to a small region; the Sondaic form is found in Bengal and in the Malayan Islands. It is a smaller species, and the armour has a tesselated appearance. The female generally, if not always, is hornless.
FIG. 131.--Sumatran Rhinoceros. Rhinoceros sumatrensis. x 1/15. (From Nature.)
The Sumatran species, Rhinoceros sumatrensis, is to be distinguished from the last two by its two horns. It is also covered {257} by a much thicker coat of hairs, which are sometimes blacker and sometimes redder. On account of its two horns it has been proposed to separate it from the other Oriental species into a distinct genus, Ceratorhinus. The animal has much the same range as the last species, but extends to Borneo. A variety of this species with hairy ears, from Assam, has been separated as a distinct form, under the name of Rh. lasiotis, by Mr. Sclater. The animal upon which that species was founded was until quite recently living in the Zoological Society's Gardens.
FIG. 132.--Hairy-eared Rhinoceros. Rhinoceros lasiotis. x 1/30.
There are only two certainly-known species of Rhinoceros in Africa. These are the White Rhinoceros (Rh. simus) and the Black Rhinoceros (Rh. bicornis). The origin of the names is not easy to understand, since the "white" animal is, if anything, darker in colour than the Black Rhinoceros. It is stated, however, that in past years the specimens of Rh. simus found in the south-west of Cape Colony were "paler and whiter in colour than those in the north-east." At present there are no grounds for distinguishing the species by their colour characters. But they are plainly distinguishable on other grounds. Rhinoceros simus has a square upper lip, and in relation to this crops the herbage upon the ground. Rh. bicornis has a prehensile upper lip projecting beyond the lower, and in a corresponding fashion feeds principally upon the branches of shrubs, It has been pointed out by Mr. {258} Coryndon[168] that the calf of Rh. simus "always runs in front of the cow, while the calf of Rh. bicornis invariably follows its mother." Both animals of course have two horns, and upon the varying proportions of the horns a large number of "species" have been made in the past. It is stated that the longest horn of the "White Rhinoceros" known measures 56-1/2 inches; while that of R. bicornis is shorter, 40 inches being apparently the maximum. But the animal is smaller.
FIG. 133.--Head of Rhinoceros bicornis.
The possible third African species of Rhinoceros[169] has been provisionally named after Mr. Holmwood, and is based upon two horns 41 and 42 inches long, which may be abnormal horns of Rh. bicornis; but they are thinner and have a smaller pedicel.
EXTINCT RHINOCEROTIDAE.--The existing Rhinoceroses are thus confined to Africa, to certain parts of the continent of Asia, and to some of the large islands lying to the south of that continent. But formerly the genus, and allied genera, had a wider range. As far back as the Miocene we meet with remains of Rhinoceroses closely allied to existing forms. The more ancient forms have, as is natural, more ancient characters. Thus in Rh. schleiermacheri of the Miocene, canines appear to have been present. The Miocene Aceratherium, primitive in the absence of horns as its {259} name denotes,[170] had also canines and, in one species, six incisors in the lower jaw. This Aceratherium had, moreover, four toes in the fore-feet. In the Miocene and later the Rhinoceros existed in Europe and America. There was even a purely northern form, the Rh. tichorhinus, which possessed a woolly covering and had the same range as the Mammoth. This Rhinoceros was two-horned.
The post-Pliocene and European Elasmotherium was a colossal rhinocerotine creature. This great beast had two horns and a body 15 feet long. Its limbs are not known, and as the teeth are different from those of Rhinoceroses in general, it may not have belonged to this group at all, though Osborn is inclined to derive it from Aceratherium, admitting at the same time that the evidence is "decidedly slender." The teeth in fact are like those of a Horse in being hypselodont and prismatic in form. As to the two horns, they were apparently not exactly like those of typical Rhinoceroses; there was an enormous horn posteriorly, supported on a huge boss of bone, and in front of this a roughened spot suggests a smaller or at least a much more slender horn.
It is important to notice that fossil Rhinoceroses belonging to the restricted genus Rhinoceros were in Europe invariably two-horned; it is only in India, where they still exist, that one-horned forms are met with in a fossil state.
FIG. 134.--Skeleton of Hyracodon nebrascensis. x 1/12 (After Scott.)
The Rhinoceroses of America were mostly hornless. Diceratherium is an exception; but in many cases it had two parallel not successive horns, and these were, to judge from the slight prominences, but feeble in development, and perhaps hardly exactly comparable with the formidable weapons of the Old-World forms. Aceratherium tridactylum, with indications of paired horns, may be ancestral to Diceratherium. The American forms have weak and slender nasals in correspondence with the absence of horns; the sagittal crest is retained in contradistinction to the great flattened surface of the skull in the horned Rhinoceroses. Aceratherium of both divisions of the globe probably represents the ancestral group of the horned and the hornless forms. This being the case it is highly interesting to note a distinct convergence in the quite {260} separate American genera towards the European horned genera. A genus sometimes united with Aceratherium, but still differing from it in some points, is Aphelops (Teleoceras).[171] This animal is more nearly approximated to "the modern standard" of Rhinoceroses than is its possible ancestor Aceratherium. The skeleton in general is more robust, even surpassing that of modern forms, and approaching the Hippopotamus. There is a reduction in the upper incisors, which are limited to two pairs, and the lower molars {262} are reduced to five. The lower incisors are only two. The sagittal crest is less marked; the fifth digit is reduced to a tiny nodule representing the metacarpus. It had a small nasal horn. There are numerous other details of likeness to modern Rhinoceroses in this creature, which has only community of descent with them from the older hornless forms, such as Aceratherium and Caenopus. In the genus Peraceras the upper incisors are as completely gone as in the living African Rhinoceroses.
FIG. 135.--Skeleton of Aphelops (Teleoceras) fossiger. x 1/15. (After Osborn.)
The most ancient rhinocerotine types[172] are the Hyracodonts and the Amynodonts. They both date from the Eocene, and became extinct in the succeeding Oligocene. Hyracodon[173] (Fig. 134) was "an agile, light-chested, and rather long-necked" type, resembling a Horse in build. There were no horns present, but the hoofs were more like those of the Horses than of the existing Rhinoceroses. These animals were apparently plain dwellers and defenceless, which is held to account for their compact hoofs and outward similarity to a Horse. The genus is Oligocene. The dental formula is I 3/3 C 1/1 Pm 4/3 M 3/3.
It is surmised by Professor Scott that the number of dorso-lumbar vertebrae was twenty-three or twenty-four. The radius and ulna are complete and separate bones, but the latter is somewhat reduced. There are four metacarpal bones, of which, however, the fifth is much reduced. The animal is only three-fingered. The tibia and the fibula are distinct, and show no tendencies towards fusion; but the fibula is much reduced. There are only three metatarsals and three toes. Had this line, which is to be regarded as a side branch of the Rhinoceros stem, not died out, it would probably have resulted, thinks Professor Scott, in monodactyle--very Horse-like types. It is later than the next genus to be described, Hyrachyus, of which it is possibly a descendant. An intermediate type, Triplopus, appears to bind together Hyracodon and Hyrachyus.
In Hyrachyus agrarius the skull is long and narrow, the facial region being markedly longer than in existing Rhinoceroses. The mastoid portion of the periotic bone is widely exposed upon the outer face of the skull, which is, as has been said, not the case with the existing genus Rhinoceros. The dentition is the complete Eutherian dentition of forty-four teeth. The upper {263} molar teeth are strikingly like those of the genus Rhinoceros. The fore-feet are pentadactyle, but functionally tetradactyle; the hind-feet tridactyle. The ulna is less reduced than in Hyracodon, and the dorso-lumbar vertebrae are twenty-five.
FIG. 136.--Skeleton of Metamynodon planifrons. x 1/22. (After Osborn and Wortman.)
The Amynodonts were short, heavy types, probably marsh-haunting in habit, and possibly with a proboscis like that of the Tapir. The orbit is higher than it is in the purely terrestrial {264} Hyracodonts, and it is suggested that when swimming it was raised above the surface as with the Hippopotamus. "This feature," observes Professor Osborn, "with the long curved tusks, undoubtedly used in uprooting, suggests the resemblance between the habits of these animals and those of the hippopotami." There were no horns in the Amynodonts. The face is shorter than in the Hyracodonts, and the mastoid is covered as in recent Rhinoceroses. The canines are very strongly developed into tusks, but the incisors show signs of disappearance. We know of the genera Amynodon, Metamynodon, and Cadurcotherium. All except the last, which is European, are American in range.
FAM. 4. TITANOTHERIIDAE.--These Oligocene Ungulates, often attaining to large dimensions, are nearly peculiar, so far as is at present known, to the North American Continent, and are at least most abundant in it.[174] Many generic names, such as Titanotherium, Brontotherium, Brontops, Titanops, and Menodus, have been given to them; but a recent study of the entire material accessible for description or already described has led Professor Osborn to the opinion that there was but a single genus, to which the name Titanotherium must be applied. Of this genus there are some thirty well-characterised species, of which the gradual evolution can be traced from the lowest strata of the White River beds where their remains occur. An entire skeleton of T. robustum enables us to understand the osteology of these forms and to compare them with other Perissodactyles. This animal was more than 13 feet long, standing some 7 feet 7 inches in height. It seems to have presented during life the aspect of a Rhinoceros with perhaps a touch of Elephant. The skull is not unlike that of a Rhinoceros in general dimensions and shape; but there are a pair of apparent horn cores anteriorly, which are smaller in the more ancient forms and acquire a large size, a forward direction with a divergence of the two in the later forms. A glance at the accompanying figures of skulls (Fig. 137) of early and later Titanotheres will exhibit the changes in this particular which the skulls underwent in the lapse of time occupied by the deposition of these Oligocene beds. The nasals are short in the later, longer in the more early species, such as T. heloceras and T. coloradense. The zygomatic arch projects much, and is "shelf-like" in the later forms, the skull thus getting an immense breadth, which, {266} together with the long and divergent horn cores, must have given to the living animal a most bizarre appearance. It is an interesting fact that this animal, though a Perissodactyle, agrees with the Artiodactyla in the nineteen dorso-lumbar vertebrae, of which seventeen bear ribs.
FIG. 137.--Three figures showing the cranial evolution of Titanotherium. Upper figure, T. trigonoceras; middle figure, T. elatum; lower figure, T. platyceras. (After Osborn.)
The genus further agrees with the Artiodactyles in the structure of the carpus. The toes of the fore-limb are four, those of the hind-limb three; but while the hind-limb is undoubtedly Perissodactyle in the arrangement of its component parts, the fore-limb shows a hint of an Artiodactyle mode of structure. This limb is paraxonic, the axis of the limb passing between the two middle digits. It may be that this genus represents more nearly than any other Perissodactyle or Artiodactyle the primitive stem from which both have diverged, though, of course, it is not old enough to be very near to the actual ancestor. The molar dentition is the typical one; the incisors seem to vary as to their presence or absence, and, if present, in their numbers. In comparing the older with the more recent forms it is noteworthy that there has been an increase of size exactly as there has been during the evolution of the Camels and some other groups of Ungulates. As already mentioned, the size of the horn cores also increases until it culminates in the extraordinary species, T. platyceras and T. ramosum, in which these are half as long as the skull, flattened in form, and connected at their bases by a "web" of bone. Arrived at this amount of specialisation the genus Titanotherium apparently exhausted its capacities for modification and ceased to be. The many generic names may be explained by sexual differences on the one hand and an incomplete knowledge of connecting links on the other.[175]
Palaeosyops is somewhat like a Tapir in build, the skull especially resembling that of the Tapir. As in Titanotherium the molar teeth, instead of having an outer wall formed by fused cusps, have a [176]-shaped outer wall on one side and two or one cusps on the opposite side. It is, moreover, an Eocene form, and in correspondence with its greater age is more primitive in some points of structure, for example, in the absence of horns and in the full dental formula. The fore-limbs are four-toed, the hind {267} three-toed. It was intermediate between a Tapir and a Rhinoceros in size. It has been shown, too, from casts of the interior of the skull, that the cerebral hemispheres are much less convoluted than were those of Titanotherium.
Related to Palaeosyops is another primitive Titanothere, the genus Telmatotherium. This is also Eocene, from the Uinta Basin, the uppermost of Eocene strata. The skull of these creatures was rather elongated, and not unlike that of a Titanothere in general aspect. The dentition was complete and the canines not very large. The horns, which acquire so prodigious a development in the later Titanotheres, are just recognisable in at any rate many species of this genus Telmatotherium, the name being thus by no means an apt one. Better was that proposed by Dr. Wortman, of Manteoceras or "prophet horned." The horns are small elevations upon the frontals just at the junction of these with the nasals, and, indeed, lying partly upon the latter bones. In T. cornutum the horns are chiefly borne upon the very long nasals, whose size contrasts with the same bones in the more highly-developed Titanotherium. It appears to be quite possible that Titanotherium was evolved from the genus Telmatotherium.[177]
SUB-ORDER 9. LITOPTERNA.
Whether the MACRAUCHENIIDAE should be considered as a separate group of Ungulata is a matter of dispute. Cope placed them in a special order of Ungulates which he called Litopterna. Zittel, on the other hand, regards them as definitely Perissodactyles. One curious point of resemblance to existing Horses is shown--that is the presence of a pit in the incisor teeth. This matter seems to be so important as to need a placing of these forms in the neighbourhood of the Perissodactyles, even of the Equidae; it is so peculiar a character, and apparently so little related to any obvious similarity in way of life, that it seems to mark a special affinity. Not so the fact that in Macrauchenia at any rate the orbit was entirely surrounded by bone as in the Horse. We find that condition so frequently acquired in many groups,--a development from an earlier condition where the cavity for the lodgment of the eye is in continuity with the temporal {268} fossa, that it cannot be regarded as anything more than a mark of specialisation. It is, in fact, the case that the Macraucheniidae are in many points specialised, while retaining many primitive features of structure.
The chief primitive features are: the non-alternating positions of the wrist- and ankle-bones; these, of course, interlock in the Perissodactyles of to-day and in many extinct families. Then the absence of a diastema in the tooth series, coupled with the presence in Macrauchenia of a complete dentition. The small brain may be referred to the same category. Macrauchenia must have been a strange-looking animal. It walked upon three toes on each limb; the skull was Horse-like in general form, but the nostrils are removed to a point about as far back as in the Whales or nearly so, the nasal bones being correspondingly reduced. This it is thought argues a proboscis. The humerus is particularly compared by Burmeister[178] to that of a Horse. The radius and ulna though both well developed are fused. The neck is long, and, as in the Camel, the vertebral arteries run inside the neural arches. Since the fore-legs seem to have been rather longer than the hind-legs, though only very slightly, and the neck was long, the animal may have presented some likeness to the Giraffe. It is interesting to note that in the proportions of humerus to ulna this animal is more Lama-like than Horse-like. On the other hand, the proportions of femur to tibia are more Horse-like. The remains of the creature are limited to South America, and to quite superficial deposits. It is evidently a specialised type, and has pursued a course parallel to that of the Horse. Much nearer to the Horse however, but apparently by convergence only, is the genus Thoatherium, usually placed in a separate family, the PROTOROTHERIIDAE. In this creature, which has many archaic characters, the toes are reduced to one in each foot. In an allied form, Protorotherium, we have the two lateral toes diminishing just as in Anchitherium.
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