Equidae

Equidae, the family of perissodactyle ungulate mammals typified by the horse (Equus caballus); see Horse. According to the older classification this family was taken to include only the forms with tall-crowned teeth, more or less closely allied to the typical genus Equus. There is, however, such an almost complete graduation from the former to earlier and more primitive mammals with short-crowned cheek-teeth, at one time included in the family Lophiodontidae (see Perissodactyla), that it has now become a very general practice to include the whole “phylum” in the family Equidae. The Equidae, in this extended sense, together with the extinct Palaeotheriidae, are indeed now regarded as forming one of four main groups into which the Perissodactyla are divided, the other groups being the Tapiroidea, Rhinocerotoidea and Titanotheriide. For the horse-group the name Hippoidea is employed. All four groups were closely connected in the Lower Eocene, so that exact definition is almost impossible.

In the Hippoidea there is generally the full series of 44 teeth, but the first premolar is often deciduous or wanting in the lower or in both jaws. The incisors are chisel-shaped, and the canines tend to become isolated so as in the now specialized forms to occupy nearly the middle of a longer or shorter gap between the incisors and premolars. In the upper molars the two outer columns of the primitive tubercular molar coalesce to form an outer wall, from which proceed two crescentic transverse crests; the connexion between the crests and the wall being imperfect or slight, and the crests themselves sometimes tubercular. Each of the lower molars carries two crescentic ridges. The number of toes ranges from four to one in the fore-foot, and from three to one in the hind-foot. The paroccipital, postglenoid and post-tympanic processes of the skull are large, and the latter always distinct. Normally there are no traces of horn-cores. The calcaneum lacks the facet for the fibula found in the Titanotheroidea.

In the earlier Equidae the teeth were short-crowned, with the premolars simpler than the molars; but there is a gradual tendency to an increase in the height of the crowns of the teeth, accompanied by increasing complexity of structure and the filling up of the hollows with cement. Similarly the gap on each side of the canine tooth in each jaw continues to increase in length; while in all the later forms the orbit is surrounded by a ring of bone. A third modification is the increasing length of limb (as well as in general bodily size), accompanied by a gradual reduction in the number of toes from three or four to one.  
  
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Fig. 1.—a, Side view of second upper molar tooth of Anchitherium (brachyodont form); b, corresponding tooth of horse (hypsidont form).  
  
All the existing members of the family, such as the domesticated horse (Equus caballus) and its wild or half-wild relatives, the asses and the zebras, are included in the typical genus. In all these the crowns of the cheek-teeth are very tall (fig. 1, b) and only develop roots late in life; while their grinding-surfaces (fig. 2, b and c) are very complicated and have all the hollows filled with cement. The summits of the incisors are infolded, producing, when partially worn, the “mark.” In the skull the orbit is surrounded by bone, and there is no distinct depression in front of the same. Each limb terminates in one large toe; the lateral digits being represented by the splint-bones, corresponding to the lateral metacarpals and metatarsals of Hipparion. Not unfrequently, however, the lower ends of the splint-bones carry a small expansion, representing the phalanges.

Remains of horses indistinguishable from E. caballus occur in the Pleistocene deposits of Europe and Asia; and it is from them that the dun-coloured small horses of northern Europe and Asia are probably derived. The ancestor of these Pleistocene horses is probably E. stenonis, of the Upper Pliocene of Europe, which has a small depression in front of the orbit, while the skull is relatively larger, the feet are rather shorter, and the splint-bones somewhat more developed. In India a nearly allied species (E. sivalensis), occurs in the Lower Pliocene, and may have been the ancestor of the Arab stock, which shows traces of the depression in front of the orbit characteristic of the earlier forms. In North America species of Equus occur in the Pleistocene and from that continent others reached South America during the same epoch. In the latter country occurs Hippidium, in which the cheek-teeth are shorter and simpler, and the nasal bones very long and slender, with elongated slits at the side. The limbs, especially the cannon-bones, are relatively short, and the splint-bones large. The allied Argentine Onohippidium, which is also Pleistocene, has still longer nasal bones and slits, and a deep double cavity in front of the orbit, part of which probably contained a gland. Onohippidium is certainly off the direct line of descent of the modern horses, and, on account of the length of the nasals and their slits, the same probably holds good for Hippidium.  
  
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Fig. 2.—a, Grinding surface of unworn right upper molar tooth of Anchitherium; b, corresponding surface of unworn molar of young horse; c, the same tooth after it has been some time in use. The uncoloured portions are the dentine or ivory, the shaded parts the cement filling the cavities and surrounding the exterior. The black line separating these two structures is the enamel or hardest constituent of the tooth.  
  
Species from the Pliocene of Texas and the Upper Miocene (Loup Fork) of Oregon were at one time assigned to Hippidium, but this is incorrect, that genus being exclusively South American. The name Pliohippus has been applied to species from the same two formations on the supposition that the foot-structure was similar to that of Hippidium, but Mr J.W. Gidley is of opinion that the lateral digits may have been fully developed.

Apparently there is here some gap in the line of descent of the horse, and it may be suggested that the evolution took place, not as commonly supposed, in North America, but in eastern central Asia, of which the palaeontology is practically unknown; some support is given to this theory by the fact that the earliest species with which we are acquainted occur in northern India.  
  
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Fig. 3.—Successive stages of modification of the left fore-feet of extinct forms of horse-like animals, showing gradual reduction of the outer and enlargement of the middle toe (III).  
  
a, Hyracotherium (Eocene).

b, Mesohippus (Oligocene).

c, Anchitherium (Miocene).

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d, Hipparion (Pliocene).

e, Equus (Pleistocene).  
  
Be this as it may, the next North American representatives of the family constitute the genera Protohippus and Merychippus of the Miocene, in both of which the lateral digits are fully developed and terminate in small though perfect hoofs. In both the cheek-teeth have moderately tall crowns, and in the first named of the two those of the milk-series are nearly similar to their permanent successors. In Merychippus, on the other hand, the milk-molars have short crowns, without any cement in the hollows, thus resembling the permanent molars of the under-mentioned genus Anchitherium. From the well-known Hipparion, or Hippotherium, typically from the Lower Pliocene of Europe, but also occurring in the corresponding formation in North Africa, Persia, India and China, and represented in the Upper Miocene Loup Fork beds of the United States by species which it has been proposed to separate generically as Neohipparion, we reach small horses which are now generally regarded as a lateral offshoot from the Merychippus type. The cheek-teeth, which have crowns of moderate height, differ from those of all the foregoing in that the postero-internal pillar (the projection on the right-hand top corner of c in fig. 2) is isolated in place of being attached by a narrow neck to the adjacent crescent. The skull, which is relatively short, has a large depression in front of the orbit, commonly supposed to have contained a gland, but this may be doubtful. In the typical, and also in the North American forms these were complete, although small, lateral toes in both feet (fig. 3, d), but it is possible that in H. antilopinum of India the lateral toes had disappeared. If this be so, we have the development of a monodactyle foot in this genus independently of Equus.

The foregoing genera constitute the subfamily Equinae, or the Equidae as restricted by the older writers. In all the dentition is of the hypsodont type, with the hollows of the cheek-teeth filled by cement, the premolars molariform, and the first small and generally deciduous. The orbit is surrounded by a bony ring; the ulna and radius in the fore, and the tibia and fibula in the hind-limb are united, and the feet are of the types described above. Between this subfamily and the second subfamily, Hyracotheriinae, a partial connexion is formed by the North American Upper Miocene genera Desmatippus and Anchippus or Parahippus. The characteristics of the group will be gathered from the remarks on the leading genera; but it may be mentioned that the orbit is open behind, the cheek-teeth are short-crowned and without cement (fig. 1, a), the gap between the canine and the outermost incisor is short, the bones of the middle part of the leg are separate, and there are at least three toes to each foot.

The longest-known genus and the one containing the largest species is Anchitherium, typically from the Middle Miocene of Europe, but also represented by one species from the Upper Miocene of North America. The European A. aurelianense was of the size of an ordinary donkey. The cheek-teeth are of the type shown in a of figs. 1 and 2; the premolars, with the exception of the small first one, being molar-like; and the lateral toes (fig. 3, c) were to some extent functional. The summits of the incisors were infolded to a small extent. Nearly allied is the American Mesohippus, ranging from the Lower Miocene to the Lower Oligocene of the United States, of which the earliest species stood only about 18 in. at the shoulder. The incisors were scarcely, if at all, infolded, and there is a rudiment of the fifth metacarpal (fig. 3, b). By some writers all the species of Mesohippus are included in the genus Miohippus, but others consider that the two genera are distinct.

Mesohippus and Miohippus are connected with the earliest and most primitive mammal which it is possible to include in the family Equidae by means of Epihippus of the Uinta or Upper Eocene of North America, and Pachynolophus, or Orohippus, of the Middle and Lower Eocene of both halves of the northern hemisphere. The final stage, or rather the initial stage, in the series is presented by Hyracotherium (Protorohippus), a mammal no larger than a fox, common to the Lower Eocene of Europe and North America. The general characteristics of this progenitor of the horses are those given above as distinctive of the group. The cheek-teeth are, however, much simpler than those of Anchitherium; the transverse crests of the upper molars not being fully connected with the outer wall, while the premolars in the upper jaw are triangular, and thus unlike the molars. The incisors are small and the canines scarcely enlarged; the latter having a gap on each side in the lower, but only one on their hinder aspect in the upper jaw. The fore-feet have four complete toes (fig. 3, a), but there are only three hind-toes, with a rudiment of the fifth metatarsal. The vertebrae are simpler in structure than in Equus. From Hyracotherium, which is closely related to the Eocene representatives of the ancestral stocks of the other three branches of the Perissodactyla, the transition is easy to Phenacodus, the representative of the common ancestor of all the Ungulata.

See also H.F. Osborn, “New Oligocene Horses,” Bull. Amer. Mus. vol. xx. p. 167 (1904); J.W. Gidley, Proper Generic Names of Miocene Horses, p. 191; and the article Palaeontology.

(R. L.*)

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