Fig. 64.—The Great Anteater (Myrmecophaga jubata). (From Sclater, List of Animals in Zoological Society’s Gardens, 1883, p. 190.)
Tamandua.[101]—This genus closely resembles the last in anatomical structure, but the head is much less elongated, the fur is short and bristly, the tail tapering, prehensile, with the under side throughout and the whole of the terminal portion naked and scaly. The stomach is similar to that of Myrmecophaga, but with the muscular pyloric gizzard not quite so strongly developed. There is a distinct ileo-colic valve and a short globular cæcum. The fore foot has a very large claw on the third toe, moderate-sized claws on the second and fourth, a very minute one on the first, and none on the fifth, which is entirely concealed within the skin. The hind foot has five subequal claws. Vertebræ: C 7, D 17, L 2, S 5, C 37. There are very rudimentary clavicles.
Fig. 65.—Tamandua Anteater (Tamandua tetradactyla). From Proc. Zool. Soc. 1871, pl. xliii.
The Tamandua (Fig. 65) is much smaller than the Great Anteater, and differs essentially from it in its habits, being mainly arboreal. It is an inhabitant of the dense primeval forests of South and Central America. As different individuals vary much in their coloration, it is possible that there may be more than one species. The usual colour is yellowish-white, with a broad black lateral band, covering nearly the whole of the side of the body.
Fig. 66.—Cæca of the Two-toed Anteater (Cycloturus didactylus). i, Ileum; c, colon.
Cycloturus.[102]—The skull is much shorter even than in Tamandua, and is arched considerably in the longitudinal direction. It differs from that of the other members of the family mainly in the long canal for the posterior nares not being closed by bone below, as the greater part of the palatines and the pterygoids do not meet in the middle line. The mandible has a prominent, narrow, recurved coronoid, and a well-developed angular process; it is strongly decurved in front. Vertebræ: C 7, D 16, L 2, S 4, C 40. Ribs remarkably broad and flat. Clavicles well developed. Manus remarkably modified, the third digit being greatly developed at the expense of all the others, and having a stout short metacarpal and but two phalanges, of which the most distal is large, compressed, pointed, and much curved, and bears a very strong hook-like claw. The second digit has the same number of phalanges, and bears a claw, but is very much more slender than the third. The fourth is represented only by the metacarpal and one nailless phalanx, the first and fifth only by very rudimentary metacarpals. The pes is also completely modified into a climbing organ. The hallux is rudimentary, consisting of a metatarsal and one phalanx, concealed beneath the skin; but the other four toes are subequal and much curved, with long pointed compressed claws. The tuber calcanei is directed towards the plantar surface, and parallel with it and extending to about double its length is a greatly elongated sesamoid ossicle. These together support a prominent calcarine cushion, to which the nails are opposed in climbing. Stomach pyriform, with muscular walls, but no distinct gizzard-like portion, as in the foregoing genera. Commencement of the colon provided with two small cæca (Fig. 66), resembling those of many birds, narrow at the base, and rather dilated at their terminal blind ends, and communicating with the general cavity by very minute apertures. Tail longer than the body, tapering, bare on the under surface, and very prehensile. Fur soft and silky.
This genus has also but one species certainly known, the Little or Two-toed Anteater (C. didactylus), an animal not larger than a Rat, of a general yellowish-colour, and exclusively arboreal in its habits. It is a native of the hottest parts of South and Central America.
The greater part of the skin strongly ossified. On the back and sides the union of numerous quadrate or polygonal scutes forms a hard shield, usually consisting of an anterior (scapular) and posterior (pelvic) solid portion (which overhang on each side the parts of the body they respectively cover, forming chambers into which the limbs are withdrawn), and a variable number of rings between, connected by soft flexible skin so as to allow of curvature of the body. The top of the head has also a similar shield (cephalic), and the tail is usually encased in bony rings or plates. The outer or exposed surfaces of the limbs are protected by irregular bony scutes, not united at their margins; but the skin of the inner surface of the limbs and under side of the body is soft, and more or less clothed with hair. Hairs also in many species project through apertures between the bony scutes of the back. The ossified dermal scutes are everywhere covered by a layer of horny epidermis. Teeth numerous, simple, of persistent growth, and usually monophyodont, but in one genus (Tatusia) a succession of teeth has been observed. Zygomatic arch of skull complete. Cervical vertebræ with extremely short, broad, and depressed bodies. The atlas free, but the second and third, and often several of the others, ankylosed together both by their bodies and arches. Lumbar vertebræ with accessory zygomatic processes, and very large metapophyses, supporting the bony carapace. Clavicles well developed. A third trochanter on the femur. Tibia and fibula ankylosed at their distal extremities. Fore feet with strongly developed, curved claws, adapted for digging and scratching—three, four, or five in number. Hind feet plantigrade, with five toes, all provided with nails. Tongue long, pointed, and extensile, though to a less degree than in the Anteaters. Submaxillary glands largely developed. Stomach simple. Uterus simple. Placenta discoidal, deciduate. The brain is generally characterised by the large size of the olfactory lobes (Fig. 57), and the slight development of sulci on the hemispheres; the sylvian fissure being represented only by a very open and shallow angle. From the earliest stage of development the stapes is stirrup-shaped, thus showing a nearer affinity to the higher mammals than is presented by the Sloths.
The animals of this family are commonly called Armadillos, a word of Spanish origin, having reference to their armour-like covering. The existing species are all of small or moderate size. They are mostly, though not universally, nocturnal in their habits, and are all omnivorous, feeding on roots, insects, worms, reptiles, and carrion. Armadillos are harmless and inoffensive creatures, offering no resistance when caught, their principal means of escape from their enemies being the extraordinary rapidity with which they can burrow in the ground, and the tenacity with which they retain their hold in their subterranean retreats. Notwithstanding the shortness of their limbs they can run with great rapidity. Most of the species are esteemed good eating by the natives of the countries in which they live. They are all inhabitants of the open plains or the forests of the tropical and temperate parts of South America, with the exception of one species (Tatusia novemcincta), which ranges as far north as Texas. Of the existing genera, Chlamydophorus stands apart from the rest in the formation of its external covering; but in all other respects Tatusia is the most aberrant form, exhibiting a peculiar type of structure of the fore feet, which in all the others show modifications, though in very varying degrees, of a single and different type.
The reproductive organs of the Dasypodidæ differ from those of the Sloths and Armadillos in the presence of a largely developed copulating organ in the male, and of a simple vagina of corresponding length in the female. The testes are still abdominal, although not in the same position; and the penis still wants both the glans and bulb. The uterus is nearly or quite as simple as in the Sloths and Anteaters; and there is no reason to believe that the placentation is essentially different from that obtaining in the other groups.
Subfamily Chlamydophorinæ.—In most anatomical characters, especially the structure of the fore foot, this little group resembles the Dasypodinæ; but it differs remarkably from all other known Armadillos, living or extinct, in the peculiar modification of the dermal armour.
Chlamydophorus.[103]—Teeth ⁸⁄₈₋₉, subcylindrical, somewhat compressed, moderate in size, smaller at each end (especially in front) than at the middle of the series. Skull broad and rounded behind, pointed in front. Muzzle subcylindrical and depressed. A conspicuous rounded, rough prominence on the frontal bone, just before each orbit. Tympanic prolonged into a tubular auditory meatus, curving upwards round the base of the zygoma. Vertebræ: C 7, D 11, L 3, S 10, C 15. Upper part of head and trunk covered with four-sided horny plates (with very small thin ossifications beneath), forming a shield, free, and overhanging the sides of the trunk, and attached only along the middle line of the back. The plates are arranged in a series of distinct transverse bands, about twenty in number between the occiput and the posterior truncated end, and not divided into solid thoracic and pelvic shields with movable bands between. The hinder end of the body is abruptly truncated and covered by a vertically-placed, strong, solid, bony shield, of an oval (transversely extended) form, covered by thin epidermic plates. This shield is firmly ankylosed by five bony processes to the hinder part of the pelvis. Through a notch in the middle of its lower border the tail passes out. The latter is rather short, cylindrical in its proximal half, and expanded and depressed or spatulate in its terminal portion, and covered with horny plates. The dorsal surfaces of the fore and hind feet are also covered with horny plates. The remainder of the limbs and under surface and sides of the body beneath the overlapping lateral parts of the dorsal shield are clothed with rather long, very soft, silky hair. Eyes and ears very small, and concealed by the hair. Extremities short. Feet large, each with five well-developed claws, those on the fore feet very long, stout, and subcompressed, the structure of the digits being essentially the same as those of Xenurus and Priodon. Nipples two, pectoral. Visceral anatomy closely resembling that of Dasypus, the cæcum being broad, short, and bifid.
The Pichiciago (C. truncatus), a small burrowing animal, about 5 inches long, inhabits the sandy plains of the western part of the Argentine Republic, especially the vicinity of Mendoza. Its horny covering is of a pinkish colour, and its silky hair snow white. It is rare, and its habits are but little known. A second species, C. retusa, from Bolivia, has been described by Burmeister. It is of rather larger size, and has the dorsal shield attached to the skin of the back as far as its edge, instead of only along the median line.
Subfamily Dasypodinæ.—Fore feet usually with all five digits developed and with nails, though the first and fifth may be suppressed. The first and second long and slender, with the normal number and relative length of phalanges. The others stout, with short broad metacarpals, and the phalanges greatly reduced in length and generally in number by coalescence. The ungual phalanx of the third very large, that of the others gradually diminishing to the fifth. Dasypus, as now restricted, has the most normal form of manus, but the modifications so markedly developed in all the others (and culminating in Tolypeutes) are foreshadowed, as it were, in it. Ears wide apart. Mammæ one pair, pectoral.
Dasypus.[104]—Teeth ⁹⁄₁₀ or ⁸⁄₉, of which the anterior in the upper jaw is usually implanted in the premaxillary bone. The series of teeth extends posteriorly some distance behind the anterior root of the zygoma, almost level with the hinder edge of the palate. They are large, subcylindrical, slightly compressed, diminishing in size towards each end of the series; the anterior two in the mandible much smaller, and more compressed than the others. Cranial portion of the skull broad and depressed. Facial portion triangular, broad in front and much depressed. Auditory bulla completely ossified, perforated on the inner side by the carotid canal, and continued externally into an elongated bony meatus auditorius, with its aperture directed upwards and backwards. (In all the remaining genera of Dasypodinæ the tympanic bone is a mere half ring, loosely attached to the cranium.) Mandible with a high ascending ramus, broad transversely-placed condyle, and high slender coronoid process. Vertebræ: C 7, D 11-12, L 3, S 8, C 17-19. Head broad and flat above. Muzzle obtusely pointed. Ears of moderate size or rather small, placed laterally, far apart. Body broad and depressed. Carapace with six or seven movable bands between the scapular and pelvic shields, each plate, or scute, being marked by a regular ellipse formed of widely separated punctures. Tail shorter than the body, tapering, covered with plates forming distinct rings near the base. Fore feet with five toes; the first much more slender than the others, and with a smaller ungual phalanx and nail; the second, though the longest, also slender. The third, fourth, and fifth gradually diminishing in length, all armed with very strong, slightly curved, compressed claws, sloping away from an elevated rounded inner border to a sharp, outer, and inferior edge. The hind foot rather short, with all five toes armed with stout, compressed, slightly curved, obtusely pointed claws—the third the longest, the second nearly equal to it, the fourth the next, the first and fifth shorter, and nearly equal.
To this genus belongs one of the best known-species of the group, the Six-banded Armadillo or Encoubert (D. sexcinctus) of Brazil and Paraguay. A very similar species, D. villosus, the Hairy Armadillo, replaces it south of the Rio Plata. There are also two very small species—D. vellerosus, from the Argentine Republic and North Patagonia, and D. minutus from La Plata. The latter differs from the other three in having no tooth implanted in the premaxillary bone. Remains apparently referable to D. villosus occur in the Pleistocene cavern-deposits of Brazil.
Xenurus.[105]—Teeth ⁹⁄₉ or ⁸⁄₈, of moderate size and subcylindrical. The most posterior placed a little way behind the anterior root of the zygoma, but far from the hinder margin of the palate. Cranium somewhat elongated, much constricted behind the orbits, and immediately in front of the constriction considerably dilated. Mandible slender; coronoid process very small and sharp-pointed, sometimes obsolete. Vertebræ: C 7, D 12-13, L 3, S 10, C 18. Head broad behind. Ears rather large and rounded, wide apart. Movable bands of carapace 12-13; the scutes being marked by an obscurely granular sculpture. Tail considerably shorter than the body, slender, and covered with nearly naked skin, with but a few small, scattered, dermal bony plates, chiefly on the under surface and near the apex. On the fore feet the first and second toes are long and slender, with small claws and the normal number of phalanges; the other toes have but two phalanges; the third has an immense falcate claw; the fourth and fifth similar but smaller claws. The hind feet are comparatively small, with five toes, bearing small, triangular, blunt nails; the third longest, the first shortest. The best known species of this genus, the Tatouay or Cabasson, X. unicinctus, is, after Priodon gigas, the largest of the group. It is found, though not abundantly, in Surinam, Brazil, and Paraguay, its remains occurring in the Pleistocene cavern-deposits of Brazil. Others, X. hispidus and lugubris, have been described, but little is as yet known of them.
Priodon.[106]—Teeth variable in number, and generally differing on the two sides of each jaw, usually from 20 to 25 on each side above and below, so that as many as 100 may be present altogether; but as life advances the anterior teeth fall out, and all traces of their alveoli disappear. The series extends as far back as the hinder edge of the anterior root of the zygoma. The teeth are all very small; those in the anterior half of each series being strongly compressed, with flat sides and a straight free edge; the posterior ones are more nearly cylindrical, with flat truncated, free surfaces. Vertebræ: C 7, D 12, L 3, S 10, C 23. Head small, elongated, conical. Ears moderate, ovate. Carapace with 12-13 movable bands. Tail nearly equal to the body in length, gradually tapering, closely covered with quadrangular scales, arranged in a quincunx pattern. Fore feet with five toes, formed on the same plan as those of Xenurus, but with the claw of the third of still greater size, and that of each of the others, especially the fifth, proportionately reduced. Hind foot short and rounded, with five very short toes, with short, broad, flat, obtuse nails. The only known species, the Great Armadillo (P. gigas), is by far the largest of existing members of the family, measuring rather more than 3 feet from the tip of the nose to the root of the tail, the tail being about 20 inches long. It inhabits the forests of Surinam and Brazil. The powerful falcate claws of its fore feet enable it to dig with great facility. Its food consists chiefly of termites and other insects, but it is said to attack and uproot newly-made graves for the purpose of devouring the flesh of the bodies contained in them.
Tolypeutes.[107]—Teeth ⁹⁄₉ or ⁸⁄₉, rather large in proportion to the size of the skull, the hinder end of the series reaching nearly to the posterior margin of the palate. Vertebræ: C 7, D 11, L 3, S 12, C 13. Ears placed low on the sides of the head, rather large, broadly ovate. Carapace with its scapular and pelvic shields very free at the sides of the body, forming large chambers into which the limbs can be readily withdrawn. Only three movable bands; sculpture of scutes in the form of subconcentrically arranged granules. Tail short, conical, covered with large bony tubercles. The fore feet formed on the same type as in the last genus, but the peculiarities carried out to a still greater extent. The claw of the third toe is very long and falcate, the first and fifth greatly reduced and sometimes wanting. On the hind foot the three middle toes have broad, flat, subequal nails, forming together a kind of tripartite hoof; the first and fifth much shorter, with more compressed nails.
The Armadillos of this genus have the power of rolling themselves up into a perfect ball, the shield on the top of the head and the tuberculated dorsal surface of the tail exactly fitting into and filling up the apertures left by the notches at either end of the carapace. This appears to be their usual means of defence when frightened or surprised, as they do not burrow like the other species. They run very quickly, with a very peculiar gait, only the tips of the claws of the fore feet touching the ground. Three species are described:—T. tricinctus, the Apar; T. conurus, the Matico; and T. muriei. Remains apparently referable to T. conurus are of not uncommon occurrence in the Brazilian cavern-deposits.
Subfamily Tatusiinæ.—This group contains but one genus, Tatusia.[108] Teeth ⁸⁄₈ or ⁷⁄₇, very small subcylindrical. The first and second subcompressed, the last considerably smaller than the others. They present the remarkable peculiarity (elsewhere found among Edentates, so far as is yet known, only in Orycteropus) of all being, with the exception of the last, preceded by two-rooted milk teeth, which are not changed until the animal has nearly attained its full size. Vertebræ: C 7, D 9-11, L 5, S 8, C 20-27. Head narrow, with a long, narrow, subcylindrical, obliquely-truncated snout; pterygoids meeting in the middle line below the nasal passage. Ears rather large, ovate, and erect, placed close together on the occiput. Carapace with seven to nine distinct movable bands; sculpture on scutes consisting of pits arranged in a V-shape. Body generally elongated and narrow. Tail moderate or long, gradually tapering; its dermal scutes forming very distinct rings for the greater part of its length. Fore feet with four visible toes, and a concealed clawless rudiment of the fifth. Claws all long, slightly curved, and very slender, the third and fourth subequal and alike, the first and fourth much shorter. Hind feet with five toes, all armed with strong, slightly curved, conical, obtusely-pointed nails. The third longest, then the second and fourth; the first and fifth much shorter than the others.
Fig. 67.—The Peba Armadillo (Tatusia novemcincta).
This genus differs from all the other Armadillos in having a pair of inguinal mammæ, in addition to the usual pectoral pair, and in producing a large number (four to ten) of young at a birth, all the others having usually but one or two.
The Peba Armadillo, T. novemcincta (Fig. 67), is a well-known species, having an extensive range from Texas to Paraguay. It is replaced in the more southern regions of South America by a smaller species, with shorter tail, the Mulita (T. hybrida), so called from the resemblance of its head and ears to those of a mule. T. kappleri is a large species from Surinam.
A rare Armadillo from Peru described under the names of Cryptophractus pilosus and Praopus hirsutus, but which evidently belongs to Tatusia, is of some interest owing to the thick coat of hair with which it is covered. This animal appears to be closely allied to T. novemcincta, from which it mainly differs by having the whole of the carapace covered with a thick coating of light brown, fine, but rather stiff hair, about an inch and a half in length. Similar hair is found on the cheeks, the proximal portions of the limbs, and (although less abundantly and shorter) on the under surface of the body. The cephalic shield, snout, feet, and the tail, with the exception of the root, are bare. The coating of hair on the back and sides completely conceals the carapace, except near the margin of the scapular region; but by separating the hairs the bands and scutes are rendered visible.[109]
In the Pleistocene cavern-deposits of Brazil have been found remains of T. novemcincta, and also of T. punctata, which appears to be an extinct form nearly allied to T. kappleri, but of somewhat larger size.
Extinct genera.—In addition to remains referable to existing genera, the above-mentioned deposits have also yielded evidence of the former existence of extinct generic types of Armadillos, some of which attained very large dimensions. Of these Eutatus was a large form distinguished from all existing genera by the circumstance that the whole of the carapace was composed of movable bands, which were thirty-three in number. Dasypotherium was a still larger form, furnished with eight teeth, of which the second seems to have been larger than the others; this genus is regarded as connecting the modern Armadillos with the next one. The gigantic Chlamydotherium, the scutes of which are common in the Brazilian caves, is considered to have been as large as a Rhinoceros; the carapace has several movable bands, but the teeth approximate in structure to those of the next family, so that the genus tends to connect the Armadillos with the Glyptodonts.
Fig. 68.—Tooth of Glyptodon from the side, and from the grinding surface. (After Owen.)
In the Pleistocene cavern-deposits of Brazil, but still more abundantly in the fluviatile deposits which cover the country in the neighbourhood of Buenos Ayres, are found the remains of some of the most remarkable forms of mammals yet discovered, the Glyptodonts, which may be regarded as forming a separate extinct family. They differ from the existing Dasypodidæ in their large size, and in having the carapace composed of a solid piece (formed by the union of a multitude of bony dermal scutes) without any movable rings, and in usually having also a ventral piece or plastron. The facial portion of the skull is very short. A long process of the maxillary bone descends from the anterior part of the zygomatic arch. The ascending ramus of the mandible is remarkably high. The teeth are ⁸⁄₈ in the known species, all much alike, having two deep grooves or flutings on each side, so as to divide them into three nearly distinct lobes (Fig. 68). The vertebral column is almost entirely ankylosed into a solid tube, and there is a complex joint at the base of the neck, to allow of the head being retracted within the carapace. The limbs are very strong, and the feet short and broad, resembling externally those of an elephant or tortoise. This family is mainly characteristic of the southern half of the American continent, but some species of the type genus ranged into Texas and Mexico. Many species of the family have been described and figured, especially by Burmeister (in the Annales del Museo publico de Buenos Aires), among which the following may be noticed. Hoplophorus is characterised by the sculptured and frequently thin scutes of the carapace, those of the periphery being flat, and not raised into prominences. The caudal sheath has several overlapping movable rings at the base, and ends in a long subcylindrical terminal tube similar to the one represented with the carapace of Glyptodon in Fig. 69, which in all probability really belongs to the genus under consideration. Each foot has four complete digits, and the humerus has an entepicondylar foramen. Most of the species are of medium size. Part of a caudal tube from Uruguay described as Eleutherocercus indicates, however, a much larger allied form, in which the tail appears to have had a number of stout bristles protruding from the joints between the scutes. Panochthus comprises very large Glyptodonts, distinguished by the great thickness of the scutes of the carapace, which are ornamented with tubercles. The termination of the caudal sheath forms a tube bearing large radiated tubercles. Euryurus is distinguished by the radiate sculpture of the scutes of the carapace. Dœdicurus, of which one species was about twelve feet in length, also has a rugose sculpture on the carapace; but the termination of the caudal tube is expanded into a club-like shape, flattened from above downwards, and covered with tubercles mingled with a few large radiate discs, which, as in Panochthus, probably carried horny spines in the living condition. The typical genus Glyptodon has each scute of the carapace ornamented with a rosette-like sculpture, the peripheral scutes being raised into conical prominences (Fig. 69). The caudal sheath, instead of being like the one represented in the figure, was entirely composed of a series of movable rings, ornamented with large tubercles. The manus had five digits, and the pes four; and there was an entepicondylar foramen to the humerus. A species of this genus, which attained very large dimensions, was made the type of Schistopleurum, on the supposition that the tail of Glyptodon was of the type represented in Fig. 69. The genus Thoracophorus, of the Pleistocene of South America, as well as Carioderma, of the Pliocene of Texas, differ from all the preceding in having the scutes of the carapace in the form of disconnected nodules. Glyptodonts also occur in South American beds of earlier age than the Pleistocene, some of these forms having enamel bands on the teeth. “Why such a form as the Glyptodon should have failed to keep his ground is,” as the late Professor W. K. Parker remarks, “a great mystery; nature seems to have built him, as Rome was built, for eternity.”
Fig. 69.—Glyptodon clavipes (Pleistocene, South America). From Owen. The tail is incorrectly restored, and it is probable that the figured portion belongs to Hoplophorus. The left lower corner shows an upper and a lower view of the skull, and the right a section of the caudal sheath.
Covered externally (except the under surface of the body and inside of the limbs) with large imbricated horny scales, and scattered hairs growing in the intervals. No teeth. Tongue long, vermiform, and protractile. No accessory articular processes to the lumbar vertebræ, but the anterior zygapophyses largely developed and deeply concave, completely embracing the semicylindrical surfaces of the posterior zygapophyses. Limbs short, with five complete digits on each foot. Scaphoid and lunar bones of carpus united. Uterus bicornuate. Placenta diffused and non-deciduate. All the existing forms belong to the Ethiopian and Oriental regions of the Old World. The absence of additional articular processes to the lumbar vertebræ is a character in which this and the following family differ from all the preceding forms.
Manis.[110]—Skull somewhat of the form of an elongated cone, with the small end turned forwards; very smooth and free from crests and ridges. No distinction between the orbits and temporal fossæ. The zygomatic arch usually incomplete, owing to the absence of the jugal bone. No distinct lachrymal bone. Palate long and narrow. The pterygoids extend backwards as far as the tympanics, but do not meet in the middle line below. Tympanic ankylosed to the surrounding bones, and more or less bullate, but not produced into a tubular auditory meatus. Rami of mandible very slender and straight, without any angle or coronoid process. From near the anterior extremity of the upper edge a sharp, conical, tooth-like process projects upwards and outwards. No clavicles. No third trochanter to the femur. Ungual phalanges bifid at their terminations. Caudal vertebræ with very long, strong transverse processes and numerous chevron bones. Tongue long, vermiform, flattened towards the tip; its retractor or sterno-glossal muscles arising from the hinder extremity of the immensely prolonged ensiform cartilage of the sternum. Stomach with thick lining membrane and muscular walls, and a special gland near the middle of the great curvature, consisting of a mass of complex secreting follicles, the ducts of which terminate in a common orifice. No cæcum. A gall-bladder. Head small, depressed, narrow, pointed in front, with a very small mouth-opening. Eyes and pinna of ear very small. Body elongated, narrow. Tail more or less elongated, convex above, flat underneath. The whole of the upper surface of the head, the upper surface and sides of the body, the whole of the tail, and the outer sides of the extremities covered with large, overlapping, horny scales, but usually with a few stiff hairs growing between and projecting beyond them. The sides and under surface of the head, the under surface of the body, and the inner sides of the limbs without scales, but with a rather scanty covering of hair. Limbs short. In walking the dorsal surface and outer sides of the phalanges of the two outer digits of the front feet alone rest on the ground, the points of the nails turning upwards and inwards. The third toe the longest, with a powerful compressed curved claw; the second and fourth with similar but smaller claws, that of the pollex often almost rudimentary. Hind feet plantigrade, with the hallux very short, and the four other toes subequal, with moderate, curved, subcompressed nails.
The reproductive organs of Manis are of a totally different type from those of the families already noticed. The testes lie in the inguinal canal; and the penis is external and well developed. The uterus is truly bicornuate, the vagina not divided, and the placenta diffused and non-deciduate. All the organs and fœtal membranes are, indeed, formed very much on the plan of those of the Ungulates, without any trace of the special peculiarities obtaining in the typical American Edentates.
The animals of this genus, which includes all the existing forms, are called Pangolins or Scaly Anteaters, and are all of small or moderate size, terrestrial and burrowing, and feed mainly on termites. Several of them can climb trees. Their length varies from 1 to 5 feet. They can roll themselves up in a ball when in danger. Their peculiar elongated form, short limbs, long, gradually-tapering tail, and scaly covering give them on a superficial inspection more the appearance of reptiles than of mammals. The species are not numerous, and may be divided into two groups distinguished by a few not very important external characters; these groups also coinciding with the present geographical distribution of the genus. These two groups, according to Mr. O. Thomas, may be distinguished as follows.
The Asiatic pangolins are characterised by having the central series of body-scales continued quite to the extreme end of the tail, by having many isolated hairs growing up between the scales of the back, and by their small external ears. They all have a small naked spot beneath the tip of the tail, which is said to be of service as an organ of touch. There are three species, viz. Manis javanica, ranging from Burma, through Malacca and Java, to Borneo; M. aurita, found in China, Formosa, and Nipal; and the common Indian Pangolin, M. pentadactyla, distributed over the whole of India and Ceylon. The African species have the central series of scales suddenly interrupted and breaking into two at a point about 2 or 3 inches from the tip of the tail; they have no hair between the scales, and no external ear-conch. The following are the four species belonging to this group:—the Long-tailed Pangolin (M. macrura), which has a tail nearly twice as long as its body, and containing as many as forty-nine caudal vertebræ, being the largest number known among mammals; the White-bellied Pangolin (M. tricuspis), Fig. 70, closely allied to the last, but with longer and tricuspid scales, and white belly hairs. These two, like the Indian species, have a naked spot beneath the tail tip, a character probably correlated with the power of climbing, and they are, moreover, peculiar in having the outer sides of their fore legs clothed with hair, all the other species being scaly there as elsewhere. Lastly, the Short-tailed and the Giant Pangolins (M. temmincki and gigantea), both of which have their tails covered entirely with scales, and evidently never take to arboreal habits. All the four species of the second group are found in the West African region, one only, M. temmincki, extending also into south and eastern equatorial Africa.
Fig. 70.—The White-bellied Pangolin (Manis tricuspis).
According to Professor W. K. Parker,[111] who remarks upon the peculiarly aberrant nature of the group, the horny scales of the Pangolins really consist of cemented hairs. This writer states that “in the early embryo lozenge-shaped tracts of skin are seen all over its body, with lines of thinner cuticle between. Under the microscope, sections of these thicker tracts show that they are composed of fine hairs, cemented together by a copious growth of epidermic cells; here and there larger hairs are seen, but these fail to reach the surface, turning again towards the inside, like nails driven into wood that is too hard for their points.”
The same author also observes[112] that there are occasional instances of the presence of eight cervical vertebræ in the Pangolins—a feature which has been considered to indicate some former genetic connection between this family and the Sloths.
The following account of the habits of Manis tricuspis is given by Mr. L. Fraser in his Zoologia Typica:—
“During my short residence at Fernando Po I succeeded in procuring two living specimens of this animal. The individuals, judging from the bones, were evidently not adult; the largest measured 30 inches in length, of which the head and body were 12 inches and the tail 18 inches. I kept them alive for about a week at Fernando Po, and allowed them the range of a room, where they fed upon a small black ant, which is very abundant and troublesome in the houses and elsewhere. Even when first procured they displayed little or no fear, but continued to climb about the room without noticing my occasional entrance. They would climb up the somewhat roughly hewn square posts which supported the building with great facility, and upon reaching the ceiling would return head foremost; sometimes they would roll themselves up into a ball and throw themselves down, and apparently without experiencing any inconvenience from the fall, which was in a measure broken upon reaching the ground by the semi-yielding scales, which were thrown into an erect position by the curve of the body of the animal. In climbing, the tail, with its strongly pointed scales beneath, was used to assist the feet; and the grasp of the hind feet, assisted by the tail, was so powerful that the animal would throw the body back (when on the post) into a horizontal position, and sway itself to and fro, apparently taking pleasure in this kind of exercise. It always slept with the body rolled up; and when in this position in a corner of the building, owing to the position and strength of the scales, and the power of the limbs combined, I found it impossible to remove the animal against its will, the points of the scales being inserted into every little notch and hollow of the surrounding objects. The eyes are very dark hazel, and very prominent. The colonial name for this species of Manis is ‘Attadillo,’ and it is called by the Boobies, the natives of the island, ‘Gahlah.’ The flesh is said to be exceedingly good eating, and is in great request among the natives.”
The Indian species is said to live in pairs, and to give birth to one or two young at a time in the spring. Their burrow reaches a depth of some twelve feet, and terminates in a large chamber, which may be as much as six feet in diameter. A faint hiss appears to be the only sound emitted by these animals.
Remains of a large species of Manis, which are indistinguishable from the corresponding bones of the existing West African M. gigantea, are found fossil in cave-deposits in the Karnul district of Madras. This is one among several instances of the close connection between the Pleistocene and Pliocene mammalian fauna of India with the existing African fauna.
Palæomanis.[113]—The lower Pliocene deposits of the Isle of Samos, in the Turkish Archipelago, have yielded remains of a Pangolin fully three times the dimensions of M. gigantea, upon the evidence of which the genus Palæomanis has been established.
External surface scantily covered with bristle-like hairs. Teeth numerous, apparently heterodont, diphyodont, and of peculiar and complex structure, being traversed by a number of parallel vertical pulp-canals. Lumbar vertebræ with no accessory zygapophyses. Femur with a third trochanter. Fore feet without pollex, but all the other digits well developed, with strong moderate-sized nails, suited to digging, the plantar surfaces of which rest on the ground in walking. Hind feet with five subequal toes. Mouth elongated and tubular. Tongue subvermiform. Uterus bicornuate. Placenta broadly zonular. Feeding on animal substances. Terrestrial and fossorial in habits. Now mainly limited to the Ethiopian region.
Orycteropus.[114]—The total number of permanent teeth appears to be from eight to ten in each side of the upper, and eight in the lower jaw; but they are never all in place at one time, as the small interior teeth are shed before the series is completed behind. In the adult they number usually five on each side above and below, of which the first two are simple and compressed, the next two larger and longitudinally grooved at the sides, the most posterior simple and cylindrical. The last three in either jaw having no milk-predecessors, may be regarded as true molars. The structure of all these teeth is quite peculiar among mammals, though resembling that of some fishes. Their summits are rounded before they are worn; their bases do not taper to a root, but are evenly truncated and continually growing. Each tooth is made up of an aggregation of parallel dental systems, having a slender pulp-cavity in the centre, from which the dentinal tubes radiate outwards, and being closely packed together each system assumes a polygonal outline as seen in transverse section. The small anterior teeth have milk-predecessors which are fully noticed below. Skull moderately elongated. The facial portion subcylindrical and slightly tapering. The zygoma complete and slender. The palate ends posteriorly in the thickened transverse border of the palatines, and is not continued back by the pterygoids. The tympanic is annular, and not ankylosed to the surrounding bones. The mandible is slender anteriorly, but rises high posteriorly, with a slender recurved coronoid, and an ascending pointed process on the hinder edge below the condyle, which is small, oval, and looks as much forwards as upwards. Vertebræ: C 7, D 13, L 8, S 6, C 27. The large number of lumbar vertebræ is peculiar among Edentates. Tongue less vermiform than in Myrmecophaga, being thick and fleshy at the base, and gradually tapering to the apex. The salivary apparatus is developed much in the same manner as in that genus, but the duct of the submaxillary gland has no reservoir. The stomach consists of a large subglobular cardiac portion, with a very thick, soft, and corrugated lining membrane, and a smaller muscular, pyloric part, with a comparatively thin and smooth lining. There is a very distinct ileo-cæcal valve, and a considerable-sized cæcum; also a gall-bladder. Head elongated, with a tubular snout, terminal nostrils, and small mouth-opening. Ears large, pointed, erect. Tail nearly as long as the body, cylindrical, very thick at the base, tapering to the extremity.
The reproductive organs and placentation of Orycteropus are formed upon a principle unknown in the more typical Edentates, or, in combination, in any other mammals. Thus the testes, in the one described example, were inguinal, but appeared to descend, at all events temporarily, into a scrotum; but the penis is scarcely larger than that of the Great Anteater. The uterus is still more fully bicornuate than in Manis, with its two lateral chambers opening separately into the vagina, as in certain Rodents. The placenta is broadly zonary, but it is not known whether it is deciduate or not. It might readily be derived from the diffused placenta of Manis by the abortion of the fœtal villi at the two poles of the ovum.
The Orycteropodidæ have long been regarded as widely different from other Edentates, their presumed affinity with the Manidæ being more or less problematical; but the discovery recently made by Mr. O. Thomas[115] that they have a milk-dentition still further emphasises their aberrant nature. According to this observer, it appears that there are normally no less than seven milk-teeth in the upper jaw, the hindmost of which is far larger than the others, having a rudimentary crown, and a distinct anterior and posterior root. The other milk-teeth are styliform, the four anterior ones being very minute, and separated from one another by equal intervals; the foremost of all is situated immediately behind the premaxillo-maxillary suture. In the mandible only four milk-teeth have hitherto been detected, of which the hindmost has the comparatively complex form found in the corresponding upper tooth. None of these milk-teeth appear, however, to cut the gum, so that the whole set is entirely functionless. Under the microscope these milk-teeth show signs of possessing a commencement of the remarkable histological structure found in the permanent teeth.
Mr. Thomas remarks that since “the three large posterior teeth of Orycteropus, already distinguished by their more molariform shape, do not have milk-predecessors, while all the small teeth anterior to them do, and in addition the last milk-tooth is markedly different from those in front of it, we ought apparently no longer to look upon this animal as an homodont, but instead to consider it as an originally heterodont form in which the incisors and canines have been suppressed to allow free play to the mobile vermiform tongue.
“But important as a knowledge of the presence of a milk-dentition in Orycteropus is, it does not at present render any easier the difficult questions as to the phylogeny and systematic position of that animal. Although called an Edentate, it has always been recognised as possessing many characters exceedingly different from those of the typical American members of the order. It has in fact been placed with them rather on account of the inconvenience of forming a special order for its reception than because of its real relationship to them. Now, as they are either altogether toothless, or else homodont and monophyodont (apart from the remarkable exception of Tatusia), it seems more than ever incorrect to unite with them the solitary member of the Tubulidentata, toothed, heterodont, and diphyodont, and differing from them in addition by its placentation, the anatomy of its reproductive organs, the minute structure of its teeth, and the general characters of its skeleton.
“But if Orycteropus is not genetically a near relation of the Edentates, we are wholly in the dark as to what other mammals it is allied to, and I think it would be premature to hazard a guess on the subject. Whether even it has any special connection with Manis is a point about which there is the greatest doubt, and unfortunately we are as yet absolutely without any palæontological knowledge of the extinct allies of either. Macrotherium even, usually supposed from the structure of its phalangeal bones, to be related to Manis, has lately proved to have the teeth and vertebræ of a perissodactyle Ungulate, and one could not dare to suggest that ancestors of Manis, or Orycteropus were to be sought in that direction. Lastly, as the numerous fossil American Edentates do not show the slightest tendency to an approximation towards the Old World forms, we are furnished with an additional reason for insisting on the radical distinctness of the latter, whose phylogeny must therefore for the present remain one of the many unsolved zoological problems.”
The Aard-Varks (Earth-Pigs) as these creatures are commonly termed, from the name bestowed on them by the Dutch Boers of the Cape, are of nocturnal habits, sleeping during the day in their burrows, which are usually found in the neighbourhood of the tall hills or mounds made by termites. Indeed, wherever these hills are abundant it is stated there is a good chance of finding an Aard-Vark, the food of these animals consisting almost exclusively of termites and ants.
Two existing species are recognised, namely the Cape Aard-Vark (O. afra) from South Africa, and another (O. æthiopicus) from the north-eastern parts of Africa, ranging into Egypt. An extinct species has been described from the Lower Pliocene of the Isle of Samos, in the Turkish Archipelago, differing from the existing forms by the larger proportionate size of the lateral metatarsals.