Handbook to the Primates/Lemurinae
THE TRUE LEMURS. SUB-FAMILY III. LEMURINÆ.
The third sub-family of the Lemuridæ contains the True Lemurs, which are characterised by the possession of a soft, thick, and woolly fur, the head rounded behind, with a specially elongated muzzle. They have small and oval ears, with the exterior aspect covered with long hair, but the inside naked, except round the margin. Their hind-limbs do not show so great a disproportionate length compared to that of the fore-limbs, as in the next sub-family, the Indrisinæ. The ankle-bones (tarsus) are only slightly elongated, and their toes are not united by a membrane. Their long and bushy tail is sometimes longer and sometimes shorter than the body. The females produce one or two, nearly naked, young at a birth, the mammæ being either two or four in number. The skull presents a central ridge on the frontal bone, and its facial portion is much elongated, the inter-orbital space being depressed and wider, and the orbits also directed somewhat outward and less straightforwardly than in several of the genera already noticed. The maxillary bones are generally much reduced, and the incisor teeth carried by them not unfrequently entirely aborted. The teeth in this Sub-family vary in number from 32 to 36, the dental formula being I (0-2)2, C 11, P 33, M 33. The foot is slightly elongated by the lengthening of the naviculare bone of the ankle, the others being short. In the wrist (carpus) the central bone (centrale) may be present or absent; its absence, however, is a character which is met with otherwise only in Man, the Chimpanzees, and the Endrina and some other Lemurs, to be described later on. The cæcum is not markedly developed.
The external coloration of the species of this Sub-family is remarkably variable, the variation being chiefly in the upper portion of the hairs, as their base is generally slate-grey.
The sub-family Lemurinæ embraces four genera: the True Lemurs (Lemur), of which there are now eight recognised species; the Hattock (Mixocebus), with a solitary species; the Gentle-Lemurs (Hapalemur), containing two species, and the Sportive-Lemurs (Lepidolemur), with seven species. Some of the most elegantly coloured species in the Animal Kingdom belong to this group. They are gregarious, and most of them arboreal, though some are not so. They form rather an exception to the general rule among Lemurs, in not being nocturnal. They feed during the morning and evening, emitting loud cries as they move about, and during the heat of the day, they often lie stretched out in the sun; at night they rest with their long tails coiled about them. In their mode of progression they are more quadrupedal than most of the other Lemuroids; they jump, walk, or run on all fours. Their food consists of fruits, birds' eggs, birds and insects. Their infants are carried about close to, and concealed amid, the hair of their mother's breast; when older they cling to her back.
The True Lemurs are all inhabitants of Madagascar and of the adjacent Comoro Islands. They are unknown on the African continent.
THE TRUE LEMURS. GENUS LEMUR.
Prosimia, Brisson, Regn. Anim., p. 220 (1756).
Lemur, Linn., Syst. Nat., i., p. 44 (1766).
Varecia, Gray, P. Z. S., 1863, p. 135.
This genus contains the typical Lemurs, in their most restricted sense. They are characterised by having a very Fox-like head, and an elongate and tapering face, shelving on each side of the nose. A long fringe of hair surrounds their chin and cheeks. They have all large and tufted ears, and large eyes, with superciliary ridges rising higher than the forehead. Their tail is always half as long as the body at least. The fore-limbs are somewhat shorter than the hind-limbs, and both the wrist and ankles are haired. The ankle is not elongated, nor is the great toe as large as in the next family—the Indrisinæ. On the outside of the palm of the hand and under the base of the fingers are situated fleshy pads, giving them greater grasping power. The True Lemurs have only one pair of mammæ, which are situated on the breast.
In the skull the facial region is much elongated, its measurement from the anterior margin of the orbit forward being greater than the longitudinal diameter of the orbit, and the space between the eye-sockets is narrow and depressed. The bony palate is short, extending back only to the posterior end of the median molar. The posterior portion of the ear-capsules (the mastoidal and squamosal regions) is not inflated—a character which separates this genus from Galago. The pre-maxillary bones are large and protrude in front, if the skull be viewed from the side. The angle of the lower jaw is not produced downwards and backwards. In some species a large maxillary sinus projects into the anterior part of the orbit; in some also the foramen rotundum does not coalesce with the sphenoidal fissure (see page 11), but has a distinct opening. The teeth are of the normal number, namely thirty-six. In the upper jaw the incisors are small, sub-equal, and situated anteriorly to the canines and are not in contact with each other, or with the latter. The canines are very large, tusk-like, and set in an excavated notch on the jaw. All the pre-molars have one main cusp to the outside; the anterior pre-molar, however, has a supplementary minute front cusp, while the median has in addition one large interior cusp; both it and the posterior pre-molars are vertically taller than their anterior fellow. The molars have two inner cusps, and two main outer cusps with a supplementary minute fore cusp, as well as two cusps on the ridge joining the fore and hind outer cusps; the posterior molar—the smallest of the three—is, however, larger than the posterior pre-molar, and has only the front inner cusp and no supplementary external cusp. The lower jaw shows a gap between the canine and the anterior pre-molar. The anterior pre-molar, which is vertically taller than the rest, is edged and cutting, taking the place of a tusk; the anterior and median pre-molars are also separated by a small space; the latter, which is equal in vertical height to the posterior, has an inner cusp and a low cusped heel. The molars have two outer main cusps, of which the front one is more developed than the hind one, and two inner cusps, often with an intermediate cusp between them; the pair of fore and the pair of hind cusps are joined by transverse ridges, and the two outside cusps by a backwardly directed semicircular ridge; the posterior molar is four-cusped.
The dorsal and lumbar vertebræ together do not exceed twenty in number.
The hind portion of the cerebellum is large, which points to intellectual inferiority in the True Lemurs as compared with the Apes.
The species of this genus are all confined to the island of Madagascar and some of the smaller adjacent islands. They are gregarious, living in large companies in the forests, feeding on fruits, insects, and such small animals, birds, and lizards as they may capture. Like the Howlers of S. America and the Gibbons of the East Indies, they are very noisy. Their agility is wonderfully great, and is displayed chiefly in the evening. During the brighter hours of the day they sit somnolent, either alone with their heads buried between their arms, their tail coiled round the neck, or in twos or threes embracing each other with their arms. In walking they use their fore-limbs less as hands, and more as feet than do the members of the next family—the Indrisinæ—both when on the ground, as well as when climbing among the trees.
I. THE RUFFED LEMUR. LEMUR VARIUS.
Lemur macaco, var. Schreber, Säugeth., p. 142, pl. 40 B (1775).
Lemur macaco et L. ruber, Geoffr., Ann. Mus., xix., p. 159 (1812).
Lemur varius, Is. Geoffr., Cat. Méth. Primates, p. 71, no. 2 (1851); Schl., Mus. Pays. Bas., vii., p. 301 (1876); Milne-Edwards et Grandid., H. N. Madag., Mamm., Atlas, pls. 123-129 (1690).
(Plate VII.)
Characters.—Face and top of head black; a stripe over the eyes, ridge of nose and tip of nostrils, creamy-white; a patch on the shoulder, the inside of the fore-legs, the inner surface of body, a patch on the front of the thighs, the inner side of the limbs, and the feet, black; tail black, washed with white on the upper surface; rest of body creamy-white.
PLATE VII.
THE RED-RUFFED LEMUR.
The Ruffed or Variable Lemur derives its name from the remarkable variability of its external markings: so much is this the case, indeed, that not a few of them have been described as distinct species. This variability appears to be entirely individual, and is by no means constant. The Black-mantled variety has the back of the neck, the shoulders and interscapular region entirely black. Another form has the ears, the ruff, and a bar across the muzzle extending over and in front of the eyes, joining the ruff, pure white; the fore-arms, legs, a bar across the buttocks joining the thighs greyish-white; face, legs, and tail black; a ring encircling the body like a belt between the fore- and hind-limbs, yellowish-white; rest of body dark reddish-brown. A third variety has the ears, ruff, and outer side of the arms and legs pure white; the flanks rusty-red, the rest of the body black.
The Red-ruffed Lemur (L. ruber) is a very well-marked variety of the same species, and may easily be recognised by the ears, ruff and whole upper surface of body being dark rusty-red, with the outer surface of thighs and legs white; or, the ears, ruff and whole upper surface (except a white patch on the back of the neck) may be dark brown, with a white garter on each ankle; otherwise it may be entirely black. It is this variety which we have figured on Plate VII.
Distribution.—Throughout the north-east of Madagascar.
Habits.—The Ruffed Lemur, called by the natives "Varikossi," has a loud, harsh and powerful voice, which can be heard for a long distance.
II. THE BLACK LEMUR. LEMUR MACACO.
Lemur macaco, Linn., S. N., i., p. 44 (1766); Schl. Mus. Pays. Bas., vii., p. 302 (1876); Milne-Edwards et Grandid., H. N. Madag., Mamm., pls. 131, 132 (1890).
Lemur niger, Schreb., Säugeth., pl. 40 A (1775).
Lemur leucomystax, Bartlett, P. Z. S., 1862, p. 347, pl. xli. (female).
Varecia nigra, Gray, P. Z. S., 1863, p. 136.
Characters.—Ears tufted, with long hairs continuing down the side of the neck to the angle of the mouth.
Male.—Entirely black.
Female.—Formerly described as a distinct species, and known as the White-whiskered Lemur (L. leucomystax). Face and lips black, darkest on the nose, round the eyes and hinder part of the head; forehead blackish-grey; whiskers and ear-tufts white, almost concealing the ears. General colour of body rich ferruginous brown, darker on the middle of the back; arms, legs and neck reddish-yellow; tail whiter; throat, under side of body and inner side of limbs creamy-white.
There is a considerable amount of variation in this species. Some individuals have the lower back and base of tail white; the belly greyish-white, the feet brown, and the toes black. In others the black frontal spot is wanting, the back of the head being reddish-white; the basal half of the tail is dark orange-red, remainder of the body rich rusty-brown. On the fore-arm is a cluster of stiff hairs, which occurs in association with a large underlying sweat-gland, whose function is not yet understood.
Distribution.—The north-west coast of Madagascar.
Habits.—The special habits of this species of Lemur are unknown, but in all probability they agree with those of the group in general, as given under the heading of the genus. It is said to utter a coarse grunting call-note.
The young males are born black like the father, and the young females have the colour of the mother. Dr. Sclater has observed that in specimens in confinement in the Zoological Gardens, in London, the female carried her young one transversely across her belly, its long tail passing round her back and then round its own neck.
III. THE MONGOOSE LEMUR. LEMUR MONGOZ.
Lemur mongoz, Linn., S. N., p. 44, no. 2 (1766); Scl., P. Z. S., 1871, p. 231, figs, 1, 2; Schl., Mus. Pays. Bas., vii. p. 312 (1876); Milne-Edw. et Grandid., H. N. Madag., Mamm., pls. 133-153 (1890).
Lemur anjuanensis, Geoffr., Ann. Mus., xix., p. 161 (1812).
Prosimia melanocephala, Gray, P. Z. S., 1863, p. 137, pl. xviii.
Prosimia xanthomystax, Gray, P. Z. S., 1863, p. 138, pl. xvii.
Characters.—Fur woolly and thick; eyelashes long; some long bristles behind the angle of the mouth; face long; no ear-tufts and whiskers, but a sub-auricular patch of long hair; some long hairs on the digits; tail bushy.
Male.—Head, face, streak across the crown of head and down the forehead brownish-black; ears of the same colour, white-fringed; cheeks and a spot on the sides of the forehead iron-grey; sub-auricular cheek-patch white, slightly washed with rufous; rest of upper surface reddish-grey; tail darker; chest and under side rufous-grey.
Female.—Rufous-brown above; neck and shoulders white; throat white; frontal spot black; face whitish.
The colour of the fur in this species varies to an extraordinary degree, and before this fact was recognised, a number of supposed species, founded on the colour of the animals alone, were described. In course of time, however, as specimens were obtained in greater number, it became evident that the variation was only in the colour of the fur, and that there was none in their anatomical and osteological structure to warrant their being considered distinct species. They have, therefore, all been now classified by Professor Milne-Edwards and M. Grandidier in their great work on the Natural History of Madagascar, as so many varieties of one species, Lemur mongoz. Of these varieties, the most important are:—
THE RED-FOOTED LEMUR. LEMUR RUFIPES.
Male.—Face in front of a line above the eyes, dark reddish-brown; hands and feet bright rufous-brown; under side of body and inner side of limbs reddish-grey.
Female.—Wrist and ankles with adjacent part of limbs above brownish-red.
THE RED-FRONTED LEMUR. LEMUR RUFIFRONS.
Male.—Grizzly, washed with rufous; fore-arms, hands, feet, haunches, outer side of legs, and top of the head between the ears, rufous.
Female.—Grizzly brown; top of head grizzly black; patch over and round the eyes greyish-white.
THE GREY-HEADED LEMUR. LEMUR CINEREICEPS.
Face and frontal spot black; cheeks, sides, top of head, side of neck, and outside of ears grey; rest of body orange-red.
THE COLLARED LEMUR. LEMUR COLLARIS.
Male.—Head blackish-brown; cheeks, sides of throat, mark over eyes, and base of ears, yellowish-grey, washed with orange-red or rufous; a spot at the side of the nose, grey; chin, throat, and under side of the body, blackish-grey.
Female.—Centre of nose black; sides of nose, chin, cheeks, including the eyes, ears, sides of throat, iron-grey, slightly flushed at the lower side of the neck under the ears with reddish-orange. Specimens from the island of Mayotte (L. mayottensis, Schl.) differ from L. collaris in having a blackish spot over the root of the tail.
THE RUFOUS LEMUR. LEMUR RUFUS.
Has a yellowish-white frontal band and whiskers.
THE BLACK-FACED LEMUR. LEMUR NIGRIFRONS.
Has a brownish-black band over the forehead, including the eyes; muzzle, patch on top of head including the ears, the side of the head below the ears, sub-auricular tufts, throat and under surface, grey.
THE WHITE-FACED LEMUR. LEMUR ALBIFRONS.
Forehead, top of head, ears, throat, and chest white.
Pure albino varieties are also quite common.
Distribution.—The Mongoose Lemur with its numerous varieties is found throughout the island of Madagascar, in Mayotte, and in Anjuan or Johanna Island, one of the Comoro group.
Habits.—Gregarious and diurnal, feeding on fruits, insects, and small animals.
IV. SCLATER'S LEMUR. LEMUR NIGERRIMUS.
Lemur nigerrimus, Scl., P. Z. S., 1880, p. 451, figs. 1 and 2; Milne-Edw. et Grandid., H. N. Madag., Mamm., pls. 154, 155 (1890).
Lemur macaco (nec L.), Scl., P. Z. S., 1878, p. 1016.
Prosimia rufipes, Gray, Ann. N. H., 1871, p. 339 (female).
Characters.—Face covered with short hair; ears nude and without tufts; nose-pad and lower lips nude. Similar to L. macaco, but larger and more intensely black, with a raised crest of upstanding hair on the head, formed by the longer fur of the body terminating arcuately on the forehead. External ears pinkish flesh-colour. Eyes blue, turning to green. Length, 16 inches; tail, 20 inches.
Female.—(Prosimia rufipes of Gray) Brown; eyes brownish-yellow.
Distribution.—Cap d'Ambra, N. Madagascar.
Nothing is known of the habits of this species.
V. THE WHITE-HANDED LEMUR. LEMUR ALBIMANUS.
Le Maki aux pieds blancs, Audebert, H. N. Singes, p. 10, pl. 1 (1797: male).
Lemur albimanus, Is. Geoffr., Ann. Mus., xix., pp. 161-169 (1812); Milne-Edw. et Grandid., Hist. Nat. Madag., Mamm., Atlas, pls. 156, 157, 162-164, 165, figs, 1 and 2 (1890).
Lemur mongoz (nec L.), Schl., Mus. Pays. Bas., vii., p. 312 (1876, pt.).
Characters.—Nose sharp and Dog-like; eyes oblique; ears, except the central portion, haired.
Male.—Face, anterior to a line over the forehead, cheeks, snout (except a greyish wash on its sides and the upper lip) umber-brown; rest of head, neck, down to the middle of the back, and fore-limbs, grey; margins of ears, chin, and under surface of body white; rest of back and hind-limbs umber-brown; tail darker, except for a short distance at the base; upper surface of hands and feet grey. The nose varies in different species in the amount of grey colouring, and the forehead and face in depth of brown. Some specimens also have an arcuate black band over the forehead from one outer corner of the eye to the other.
Female.—Greyish-black; nose grey; rest of face washed with brick-red, deeper on the forehead, cheeks, ears, and sides of neck, fainter in tint on the upper back; lower back and tail darker, except at the base, where it is washed with reddish-yellow. Hands and feet greyish-white. The colour of the face varies much in different specimens, being deeper or lighter rufous. The arcuate band from the corners of the eyes over the forehead varies in breadth and depth of colour.
Distribution.—Madagascar; the precise locality unknown.
VI. THE CROWNED LEMUR. LEMUR CORONATUS.
Lemur coronatus, Gray, Ann. and Mag. N. Hist., x., p. 257 (1842); Schl., Mus. Pays. Bas., vii., p. 313 (1876); Milne-Edwards et Grandid., Hist. Nat. Madag., Mamm., Atlas, pls. 158-161, 165, 166.
Lemur chrysampyx, Scheurm. Mém. Cour. Acad. Brux., xxii., p. 6 (1848 = female).
Prosimia coronata, Gray, P. Z. S., 1863, p. 138.
Characters.—Tips of ears naked; tail a little more than the length of the body.
Male.—Face, nose, and region round the eyes greyish-white; cheeks and forehead rufous or yellowish-red; a conical spot in the centre of the head between the eyes, dark brown or black, intruding sometimes on the rufous of the forehead; ears white; inner side of limbs and under side of body greyish-white; tail rufous at base, the upper side blackish, and the under side lighter; rest of body sienna-grey.
Female.—Upper side entirely grey, washed with yellowish cream-colour on the middle and lower part of the back, and on the upper side of the tail; long black hairs present in the tail; the under side entirely silvery-grey; fur at base black, the tips grey or silvery; instead of the black spot on the forehead there is a golden yellow-hooped, or widely V-shaped, bar above the eyes, narrower in the centre over the nose.
Albino specimens are sometimes found, which are entirely white, except for the golden bar over the eyes.
VII. THE RED-BELLIED LEMUR. LEMUR RUBRIVENTER.
Lemur rubriventer, Is. Geoffr., C. R., xxxi., p. 876 (1850); Schl., Mus. Pays. Bas., vii., p. 311 (1876); Milne-Edw. & Grandid., Hist. Nat. Madag., Mamm., Atlas, ii., pls. 167-170 (1890).
Lemur flaviventer, Is. Geoffr., tom. cit., p. 876 (1850).
Characters.—Inner margins and outside of ears haired, the interior nude.
Male.—Face, a line down the forehead, and snout dark maroon-brown; a ring round the eyes cobalt-blue; rest of head and cheeks reddish-brown; upper side of body speckled reddish-brown, darker on the lower back; tail almost black, with long white hairs distributed throughout its length; feet rufous; under side of body pale.
Female.—Like the male, but having the cheeks whitish; a narrow ring round the eyes pale blue; upper surface umber-brown, washed with reddish-yellow; under side and inner sides of limbs yellowish; ruff reddish-chestnut.
Young.—Head entirely rufous; nose black.
Distribution.—Madagascar.
VIII. THE RING-TAILED LEMUR. LEMUR CATTA.
Lemur catta, Linn., S. N., i., p. 45, no. 4 (1766); Schl., Mus. Pays. Bas., vii., p. 314 (1876); Milne-Edw. et Grandid., Hist. Nat. Madag., Mamm., Atlas, pls. 171-172 (1890).
Characters.—Inside of ears naked; no ruff round the face; top of head greyish-black; face, rest of head, lower surface of body, and inner side of the limbs pearl-grey; upper surface sienna-grey. Tail pearl-grey, banded with from ten to twelve black rings, distinguishing it from all other Lemurs, which have the tail of one colour. Length of body and tail together, 40 inches.
On the fore-arm above the wrist-joint there is, in both sexes, a comb-like bony outgrowth (becoming in old males a prominent spur) continuous with the palm of the hand by means of a narrow strip of black, hairless skin; near it there is a cluster of long stiff hairs over an underlying sweat-gland, the function of which is still unknown.
Distribution.—This species inhabits chiefly the rocky and treeless regions of the south and south-western borders of the Betsileo province of Madagascar. It is, however, not entirely confined to these treeless districts, for it has been recorded as occurring in bands of some numbers in the neighbouring forest regions.
Habits.—The Ring-tailed Lemur—one of the handsomest species of the genus and the only one in which the tail is not uniformly coloured—is of gentle manners, active, and graceful. According to the notes of the Rev. G. A. Shaw, as recorded in a paper in the Zoological Society's "Proceedings," it lives among the rocks where a few stunted trees occur, and over this rocky ground it can easily travel, in places where it is impossible for the natives, although bare-footed, to follow it. The palms of its hands and feet are smooth and leather-like, enabling the animal to apply them firmly to the wet rocks. This Lemur feeds on bananas and wild figs. In the winter its chief sustenance consists of the prickly-pear, peeling off the spiny skin with its long upper canines. According to the same observer, this Lemur rarely drinks water; indeed, it is said that the species living in the west of Madagascar, including two kinds of White Lemur, subsist without water, while those on the east coast invariably drink water with their meals. When fighting, the Ring-tailed Lemur scratches vigorously and strikes out with its hands.
THE HATTOCKS. GENUS MIXOCEBUS.
Mixocebus, Peters, M. B. Akad. Berlin, 1874. p. 690.
This genus contains but one species, whose characters are therefore those of the genus also.
THE HATTOCK. MIXOCEBUS CANICEPS.
Mixocebus caniceps, Peters, M. B. Akad. Berlin, 1874, p. 690, pl. i., pl. ii. (Skull.)
Characters.—Snout sharp, with a naked nose-pad; eyes very large; ears very short, rounded, higher than broad, scarcely appearing beyond the fur, and sparsely covered with short hair; limbs long, the digits with unkeeled nails; tail as long as the body, or slightly longer; inter-maxillary bones more prominent than in the species of the next genus, and containing a small incisor tooth on each side; no inter-parietal bone; upper canine not vertically longer than the grinders; the upper pre-molar and molar series of teeth arranged to converge but slightly anteriorly, forming, as seen from the front, a somewhat convex line, differing in this from some species of Lepidolemur, in which these teeth are arranged in a nearly straight line.
Top of head grey, the base of the hairs Mouse-grey, with black or white tips; a triangular patch on the middle of the head, darker; band on the sides and middle of the nose dark brown, widening out on the forehead and over the eyes; a dark ring round the eyes, merging into the dark brown colour of the nose; front border of the ears, a patch behind the latter, the lips, chin, sides of cheek, and chest a creamy- or yellowish-white; throat grey; upper side of the body, outside of the limbs, and dorsal end of the tail, rufous-grey; back portion of the upper part of the thigh, the hinder part of the belly, and the greater part of the upper side of the tail yellowish-rufous; the upper side of hands dark brown, of the feet yellowish-grey; extremity of tail blackish-brown. Length of body, 12½ inches; tail, 13½ inches.
Distribution.—Confined to Madagascar.
Habits.—The habits of the Hattock, as the natives name this animal, are quite unknown.
THE GENTLE-LEMURS. GENUS HAPALEMUR.
Hapalemur, Is. Geoffr., Cat. Méth. Primates, p. 74 (1851).
This genus has been constituted for two species of a specialised type of Lemur, characterised by a globose head, a short muzzle, with a tapering nose and short hairy ears. The hind-limbs are longer than the fore-limbs, the feet short and broad, and the tail hairy and equal in length to the body. The female has four teats, two on the breast, or on the shoulder, and two on the abdomen.
In regard to their skeletal characters, the facial portion of the skull is short and narrow in front—the nasal bones being arched—and the brain-case rounded. The cranium presents no elevated frontal crests, as among the members of the next genus (Lepidolemur). The pre-maxillary bones are very small. The hind margin of the bony palate, which dilates posteriorly, does not extend behind the mid-line of the last molar. The squamosal region of the skull and the outer and posterior—the mastoidal—portion of the ear-capsules (periotic bones), is not inflated in the members of this genus. Their lower jaw is very characteristic, being massive in front and possessing a very long symphysis (or line of junction of its two halves), its angle being also very large, and produced downward, inward, and backward, even more than in the genus Indris. The naviculare bone of the ankle (tarsus) is relatively short, thus differing from the same region in Microcebus and in Galago; the carpus (or wrist) has no central (os centrale) bone.
In Hapalemur the teeth are of the normal Lemurine number, viz., 36; but the dentition as a whole is peculiar and characteristic. Each series of teeth is very uniform and equal, and those anterior to the molars are serrated. In the upper jaw the incisors are very small, sub-equal, and situated close together; the posterior tooth on each side being (when the skull is viewed from the side) internal to and touching the canines. The canines are small, and the gap between them and the anterior pre-molar is very small. The anterior pre-molar is slightly taller vertically than its median fellow, and stands close up to it without an interval; it has one main (and sometimes one rudimentary) outer cusp; the posterior pre-molar, which closely resembles a molar, and is often the largest tooth in the jaw, having one inner cusp united by ridges to its two outer cusps. The molar teeth are sub-equal to the hindmost pre-molar, and have one front inner and two outer cusps, without an oblique ridge between them, and also a well-developed cingulum, cusped externally. Of the lower teeth, the anterior and median pre-molars are set obliquely, the median having three outer and two inner cusps (the two inner being united to the two hind outer by ridges). The posterior pre-molar is quite molariform, and, with the molars, presents three outer and two (or three) inner cusps, of which the two inner are united by ridges to the outer hind cusps, while transverse ridges unite the main outer and inner cusps together. The molars are cingulate towards the outside.
PLATE VIII.
THE GREY GENTLE-LEMUR.
The brain is narrower and shallower than that of the genus Lemur, and presents no specially close resemblance to the same organ in the Indrisinæ or the Lorisinæ.
I. THE GREY GENTLE-LEMUR. HAPALEMUR GRISEUS.
Lemur griseus, Geoffr., Mém. sur les Makis. Mag. Enc., i., p. 48 (1796).
Hapalemur griseus, Is. Geoffr., Cat. Méth. Primates, p. 74 (1851); Mivart, P. Z. S., 1864, p. 613 (Skull); Schleg., Mus. P. B., vii., p. 361 (1876).
Hapalemur olivaceus, Is. Geoffr., Cat. Méth. Primates, p. 75 (1851); Schl., Mus. P. B., vii., p. 316 (1876).
Cheirogaleus griseus, Giebel., Säugeth., p. 1018 (1856); V. der Hoeven, Tijds. Natuurl. Gesch., p. 38, pl. i., fig. 1 (1844).
Hapalolemur griseus, Scl., P. Z. S., 1863, p. 161; Gray, P. Z. S., 1863, p. 828, pl. lii.
(Plate VIII.)
Characters.—Fur long and soft, not woolly; ears short, hairy, with long black vibrissæ between them; tail bushy, and as long as the body; general shade above greyish Mouse-colour, washed with rufous and speckled with black on the crown, back and external surface of limbs; shoulders and fore-limbs bluish-grey; cheeks, throat, breast, and inner side of limbs ochraceous white; under side of body whitish-yellow; tail and hands grey, washed with black. Body and tail equal, 15 inches in length.
Facial portion of skull short; brain-case rounded; lower jaw shorter and higher than in Lemurs generally; great toe large and broad; on the inner side of both arms close to the wrist occurs a rough patch (extending down to the bare skin of the palm) corresponding to a gland beneath, in the male, spine-like, while in the female hairy processes are present, together with a tuft of long hairs; external to this patch is a callous pad; mammæ opening on the shoulder; intestine large; cæcum small.
Young.—Reddish-yellow below.
Distribution.—The Grey Gentle-Lemur inhabits the eastern side of the Betsileo province of Madagascar.
Habits.—The "Bokombouli," as the natives name this animal, is the smallest of any of the True Lemurs. It is nocturnal, and lives, according to the Rev. G. A. Shaw, among the bamboos in the higher-level forests of the island. Its lower incisors are used as scrapers, and nearly all its teeth are serrated and very effective in cutting off the bamboo shoots, on which it feeds. To enable it to grasp smooth surfaces, such as the stems of the bamboo and other trees it frequents, it possesses a broad pad under each great toe.
II. THE BROAD-NOSED GENTLE-LEMUR. HAPALEMUR SIMUS.
Hapalemur (Prolemur) simus, J. E. Gray, Cat. Monkeys, Brit. Mus. App., p. 133 (1870); id. P. Z. S., 1870, p. 828, pl. lii., pp. 829, 830, figs. 1-4 (Skull).
Prolemur simus, J. E. Gray, P. Z. S., 1872, p. 851.
Hapalemur simus, Beddard, P. Z. S., 1884, p. 392; Jentink, Notes Leyd. Mus., vii., p. 33 (1885).
Characters.—Nose broad and truncated; ears short, covered with long hair on the outside and along the margin inside.
Very similar to H. griseus; head and upper back dark reddish-grey, faintly washed with rufous; sides of head, neck, and region round the eyes lighter; sides of nose and region between the eyes black; ears dirty grey; lower back, sides of body, and outer surface of limbs sooty-grey, with here and there a wash of rufous; the patch on the end of the rump and upper part of the base of the tail uniform pale yellowish rust-colour; remainder of tail sooty-grey; from the chin to the chest yellowish-grey; under side of body and inner side of arms pale sooty-grey.
No spines on the fore-arm above the wrist as in H. griseus. In the skull, the nose is broad, square, and truncated; the pre-maxillæ very small; the lower jaw weak and narrow in front.
Distribution.—Only known from Madagascar.
Habits.—The habits of the Broad-nosed Lemur are said to differ in no respect from those of the foregoing species.
THE SPORTIVE-LEMURS. GENUS LEPIDOLEMUR.
Lepilemur, Is. Geoffr., Cat. Méth. Primates, p. 75 (1851).
Lepidolemur, Peters, M. B. Akad. Berlin, 1874, p. 690 (1874).
This genus contains, according to Dr. Forsyth Major, as many as seven species. This excellent comparative anatomist has made a very careful revision of the group, and the present writer has gratefully to acknowledge from him many valuable notes incorporated under this section, as well as his kindness in supplying for publication the diagnoses of his new species.
Dr. Major divides these seven species into two series:—(A) a group of four larger species, and (B) a group of three smaller species.
The members of this genus are smaller than the True Lemurs of the genus Lemur. Their head is conical and short, their ears large, round, and membranaceous, and the tail is shorter than the body. In this latter character and in their shorter limbs they differ from Mixocebus. The fourth finger and toe are the longest digits of their respective extremities, the nails of all are keeled, and that of the great toe is very large and flat.
In the skull, the muzzle is longer than the longitudinal diameter of its orbit in the series of larger species (Section A); in the smaller species (Section B) the muzzle is shorter.
Their dentition presents several important characters. The series of upper molars and pre-molars form almost a straight line, both sides being almost parallel, or only slightly convergent towards the front. In the upper jaw the incisors are wanting; the canines are very large and grooved internally, and have a posterior heel. There is no gap between them and the anterior pre-molar, which last is vertically taller than the rest, and has one cusp to the outside, whereas the median and posterior have an inner cusp as well. The anterior and median molars have the inner hind cusp rudimentary, but the cingulum rises into a minute cusp, both at the fore and hind edge; the posterior molar is three-cusped. The whole of the cheek-teeth gradually broaden and decrease in vertical height from before backward as far as the median molar. In the lower jaw the anterior pre-molars are large, canine-like, and decumbent, and have a strong process on their anterior margin (resembling that in the corresponding tooth in Indris); the median and posterior pre-molars have one external cusp, and the latter tooth one interior cusp in addition. The anterior and median molars have a rudimentary fifth cusp, which is large in the posterior molar.
The pre-maxillæ are very much reduced, so that the teeth they usually carry are generally wanting. The bony palate is short, its hind margin extending back only to the middle of the median molar; its anterior foramina are small; and it differs from that of Microcebus and Chirogale in having its posterior perforations small. The angle of the lower jaw is produced downwards and backwards. The mastoid portion of the ear-capsules (periotic bones) as well as the squamosal are markedly enlarged and swollen, in this respect differing from the skulls of Lemur and Hapalemur. The ridges in the temporal bone unite into a frontal (sagittal) ridge, and the space between the orbits is depressed; a depression is also present on the cheek in front of the lachrymal foramen. The foot is slightly elongated by the lengthening of the naviculare bone of the ankle (tarsus), the thin bones of which are short. In the wrist (carpus) there is no os centrale or central bone, which is otherwise invariably present in the Primates, except in Man, the Chimpanzees, the Gentle-Lemurs, and the Endrina.
The Sportive-Lemurs are confined to Madagascar and are nocturnal and arboreal creatures, feeding on leaves and fruits.
In Group A (the larger species) are included: 1, The Weasel-like Lemur (L. mustelinus); 2, the Red-tailed Sportive-Lemur (L. ruficaudatus); 3, Edwards' Sportive-Lemur (L. edwardsi); and 4, the Small-toothed Sportive-Lemur (L. microdon). Group B (consisting of the smaller species) comprises: 1, The Round-headed Sportive-Lemur (L. globiceps); 2, Grandidier's Sportive-Lemur (L. grandidieri); and 3, the White-footed Sportive-Lemur (L. leucopus). With the exception of the two first-named species, all the others are here made known for the first time by Dr. Forsyth Major. Very little is recorded of the habits of these animals. They are so rare that at present the various species are known from a few skins or alcoholic specimens in European museums. They are said to be inhabitants only of the forest-country, nocturnal in their habits, sleeping coiled up in some retreat all day, but issuing forth at night, at which time they are very agile in their movements.
SECTION A. (SPECIES MAJORES.)
I. THE WEASEL-LIKE SPORTIVE-LEMUR. LEPIDOLEMUR MUSTELINUS.
Lepilemur mustelinus, Is. Geoffr., Cat. Méth. Primates, p. 76 (1851); Schl. et Pollen, Faun. Madag., Mammif., p. 10, pls. 4, 6, fig. 3; Schl., Mus. P. B., vii., p. 317 (1876).
Lepilemur dorsalis, Gray, Cat. Monkeys, Brit. Mus. App., p. 135 (1870).
Characters.—Fur soft and woolly; ears rounded, naked excepting at the base behind; muzzle elongated. Above, reddish-grey. Face and cheeks grey; throat white; under side of body and inner side of limbs, pale grey; tail short-haired, the posterior third dark brown. Length of body, 14 inches; and tail 10 inches.
Skull large and massive; the brain-case small and inflated; facial region long, differing in this character from L. ruficaudatus; orbits very large, thus differing from the three remaining species of the larger group (A); the process of the maxilla intervening between the nasal and lachrymal bones; molar teeth large.
Distribution.—This species occurs in the north-east of Madagascar, and, according to Grandidier, in the north-western corner of the island.
Habits.—The "Fitili-ki," as the natives have named this animal, is found in the forests in small companies. It is nocturnal in its habits, feeding on leaves and fruits.
II. THE RED-TAILED SPORTIVE-LEMUR. LEPIDOLEMUR RUFICAUDATUS.
Lepilemur ruficaudatus, Grandidier, Rev. et. Mag. de Zool., 1867, p. 256.
Lepilemur pallidicauda, Gray, P. Z. S., 1872, p. 850.
Lepilemur mustelinus (nec. Is. Geoffr.), Schl., Mus. P. B., vii., p. 317 (in part).
Characters.—Smaller than the last species; head much broader than it is long; snout short and conical; ears ovate, exposed, short-haired; tail long, thicker at the end, and covered with softer and longer hairs. Fur pale or reddish-grey; head dark brown; the shoulders and outer side of the arms grey, washed with brown; chin, breast, and inner side of limbs and under side of body whitish; upper side of the base of the tail rather dark brown, this colour extending further down in the tail of the female; rest of the tail uniform pale brownish or greyish-red.
Skull very broad compared with its length, more massive, and showing a shorter muzzle than in L. mustelinus; orbits smaller than in any of the other species in Group A.
Distribution.—South-western Madagascar.
III. MILNE-EDWARDS' SPORTIVE-LEMUR. LEPIDOLEMUR EDWARDSI.
Lepidolemur edwardsi, Forsyth Major.[1]
Characters.—"Similar to L. ruficaudatus; upper part of head grey; ears membranaceous, but encircled on the inner and posterior side by an incomplete belt of dark brown colour, which distinguishes the species from L. ruficaudatus; shoulders and outer side of the fore-limbs reddish-brown. Back greyish-brown, lighter on the outer side of the hind-limbs; an uninterrupted dark dorsal streak from the middle of the back to the centre of the forehead is very conspicuous between the shoulders. Breast, inner sides of the fore- and hind-limbs, and lower surface of the body greyish-white.
"The skull long and narrow; molars and pre-molars large, especially transversely; orbits small, yet larger than in L. ruficaudatus; the mastoidal portion of the ear-capsules and squamosal region of the skull conspicuously inflated. Bony palate more elongate than in L. mustelinus; par-occipital process present."
Distribution.—Betsako, north-west of Madagascar.
IV. THE SMALL-TOOTHED SPORTIVE-LEMUR. LEPIDOLEMUR MICRODON.
Lepidolemur microdon, Forsyth Major.
Characters.—"Somewhat similar to the Weasel-like Lemur (L. mustelinus) in coloration, but having the back and the outer portion of the shoulder and fore-limbs bright chestnut, passing into russet on the back (darker between the shoulders), on the outer parts of the hind-limbs and tail, as well as on the top of the head, where it is washed with greyish. A dark, dorsal stripe from the centre of the forehead to the middle of the back, where it is darkest. Breast and under surface of body yellowish-grey.
"Skull markedly distinguished from that of the other species by the small size of the molars; pre-molars not diminished in size; a depression at the base of the nasals; the bony palate more elongated than in L. mustelinus."
Distribution.—The eastern districts of the Betsileo province, Central Madagascar.
PLATE IX.
THE WHITE-FOOTED SPORTIVE-LEMUR.
SECTION B. (SPECIES MINORES.)
V. THE ROUND-HEADED SPORTIVE-LEMUR. LEPIDOLEMUR GLOBICEPS.
Lepidolemur globiceps, Forsyth Major.
Characters.—"The smallest of the Sportive-Lemurs. Similar to Lepidolemur ruficaudatus, but less rufous down the fore-limbs; the tail drab colour.
"Skull very characteristic; the brain-case broad, high, and globose, the facial region short; the premaxillæ more reduced than in any other species; the external auditory channel very large; the occipital region less vertical than in the species of Section A."
Distribution.—Ambulisatra, south-west Madagascar.
VI. GRANDIDIER'S SPORTIVE-LEMUR. LEPIDOLEMUR GRANDIDIERI.
Lepilemur mustelinus, Gray (nec Geoffr.), P. Z. S., 1863, p. 144.
Lepidolemur grandidieri, Forsyth Major.
Characters.—"General colour cinnamon; head greyish; an indistinct median dorsal streak from the forehead along the back; inner side of the limbs and under side of the body yellowish-grey.
"Skull remarkable for the large size of its orbits, and for the anterior convergence of its upper dental cheek-series being greater than in the other members of the group."
Distribution.—North-west Madagascar.
VII. WHITE-FOOTED SPORTIVE-LEMUR. LEPIDOLEMUR LEUCOPUS.
Lepidolemur leucopus, Forsyth Major, Ann. and Mag. Nat. Hist., xiii., p. 211 (1894).
(Plate IX.)
Characters.—Ears large, long, membranaceous; tail shorter than the body. Upper side Chinchilla-grey, with an indistinct median brownish stripe from the neck to the root of the tail. Top of head brownish-grey, with a darker median stripe; cheeks and chin whitish. Ears encircled by a broad ring of whitish hair. Neck, shoulders, and upper parts of the fore-arm pale rufous. Breast and belly greyish-white; inner surfaces of the hind-limbs and the heels pure white.[2] Tail greyish, with a rusty tinge. Length, 12 inches; tail, 10½ inches.
The skull is longer and broader than that of L. grandidieri; the mastoidal portion of the ear-capsules and the adjacent squamosal region very largely inflated; bony palate elongated; dental cheek-series short; molar teeth small and slender, distinguishing this species from L. grandidieri, their small size also separating it from L. globiceps.
Distribution.—This species is at present known only from Fort Dauphin in the south-east of Madagascar. [Type in British Museum.]