1. EPOMOPHORUS. 13 elevations on each side and by ill-defined semicircular toothed lines, as in Cynonycterts straminea. The first undivided ridge is straight, and is placed immediately behind the canines; the second is similar, and corresponds to the first premolars; the third is represented by a large oval papilla on each side, opposite the middle back tooth ; the fourth, fifth, and sixth by similar but much smaller and less elevated papillee, succeeded by transverse semicircular toothed lines behind (Plate II. fig. 4).
Upper incisors small, equally distant, the outer incisor on each side faliing out im old indwiduals, so that the dentition then resembles that of E. comptus; lower incisors also small and separated, unequally trifid; premolars and molars as in EB. gambianus. The adult female is considerably smaller than the male; and the following Table exhibits the relative proportions of an adult male and female respectively :— Epomophorus franqueti. Adult g.jAdult 2.
5:8 2:15 0°85 0:95 3°35 15 third finger, metacarpal 25 + 55 a Ist ph. ...... 16 - 5 = Qnd ph......cccseercerseeeee eae 2:2 3 fifth finger, metacarpal.............:::ceeeeeeeseeneeees é 2°45 as es few Usb aplse gustaiaecee secon ee seseae pees 1:25 1-2 i 5 gl) eM PU ac snes chante Goevoseummaneeawneess 13 1-2 gg PADI weevettecacencosceesseevelsvecteeans set aesenteceneces 1-5 1-4 _ FOOL ace eS a ae EO BS dE RR 0-9 0-9 Hab. West Africa (Sierra Leone, Gold Coast, Lagos, Gaboon). This species is easily recognised by the very large size of the shoulder-pouches and remarkable development of the hairs lining them (in the adult male), as well as by the comparatively wide interfemoral membrane, by the total absence of the tail, by the deciduous upper outer incisors, and by the peculiar form of the palateridges.
a. 9 ad., al. Sierra Leone. Purchased.
b. fo ad, al. Gaboon. ; ce. 2 imm,, al. M. Lidth de Jeude.
5. Epomophorus comptus.
Epomophorus comptus, Allen, Pr. Ac. Nat. Sct, Philad. 1861, p. 158. Muzzle shorter than in Z. gambianus; in adult females the ear is considerably longer than the muzzle ; outer and inner margins of the ear-conch equally convex in lower two thirds, the tip narrowly rounded off, the upper third of the outer margin slightly concave
14 PTEROPODID.Y.
beneath the tip; nostrils opening laterally, separated by a wide and shallow emargination ; upper hp with a slight vertical groove. Wings from the distal extremity of the first phalanx of the second toe. Interfemoral membrane as in &. yambianus; no trace of a tail externally.
Fur (so far as can be determined from an inspection of specimens preserved in alcohol) somewhat similar in colour and distribution to that of £. gambicnis.
The palate has three straight undivided ridges in front, and four or five ill-defined semicircular toothed ridges (as in Oynonycteris straminea) behind; the first undivided ridge is placed slightly behind the canines, the s2cond between the single pair of upper premolars ; the third between the first upper molars; while the first semicircular ridge is divided, and corresponds to the last upper molars (Plate IT. fig. 5).
Upper incisors, consisting (in the adult animal) of a single unicuspidate pair only, placed in the centre of the premaxilla (Plate II. fig. 5), and separated from each other by a slight interval (immature individuals have small outer incisors placed near the larger inner pair); lower incisors nearly equal in size, forming a straight row across, the outer incisors equally bifid, the middle pair unequally bifid, their outer cusps shortest.
Length (of an adult 2) about 5!-0, head 1-9, eye from tip of nostril 0!-7, ear 0'-95, forearm 3-4, thumb 1-45; third finger— » metacarp. 2!-35, Ist ph. 1-55, ond ph. o 4; fitth finger—m-ta- . carp. 2!-3, Ist ph. 1!/-2, 2nd ph. 1/2: tibia 1/3, foot 0-8. Hab. West Africa (Lagos, Gaboon).
This species is undoubtedly closely allied to E. franqueti, with © which it agrees generally in structure, but is distinguished by the presence of a single pair of upper incisors in the adult animal (in E. franqueti the outer incisors are wanting in very old individuals . only), by the form of the palate-ridges, and by the mode of ae ment of the wings to the toes. :
Males of this species have probably shoulder-glands similar "to those of #. franquetz, to which it is most closely related; but I am unable to describe them, as I have not yet had an opportunity of examining a male specimen.
a. 2 ad.sk., al. Elloby District, Gaboon. H. Ansell, Esq. [P.]. 6. Epomophorus pusillus.
Epomophorus schoensis, Tomes, P. Z.S. 1860, p. 56, 1861, p. 12, figs. 4, 4a.
Epomophorus pusillus, Peters, MB. Akad. Berl. 1867, p. 870. Much smaller than any other known species of the genus; the length of the forearm scarcely more than half the length of that of EL. macrocephalus.
Muzzle rather short, the eye (in the adult female) placed almost midway between the ear and the end of the nose; ears nearly
2. PTEROPUS. 15 double the length of the muzzle, shaped as in Z. gambianus ; nostrils rather widely separated, the shallow emargination between them continuous with a groove dividing the lower lip; lips thickened near the extremity of the muzzle and folded near the angle of the mouth.
Wings from the base of the second toe; tail extremely short and thin *, Fur yellowish brown above ; beneath, paler brown, the extremities of the hairs greyish. The wing-membrane is covered almost as far outwards as a line drawn from the elbow to the knee, also the humerus and three fourths the forearm and the legs as far as the ankles ; the feet are naked.
The palate is very peculiarly marked, and quite on a different plan from that of any other species of the genus. A broad but shallow Y-shaped groove is formed by the union of two grooves, each commencing opposite the single premolar on each side, and uniting in the centre and continued backwards as a single central groove which terminates in the vaulted roof of the palate behind. (Plate IT. fig. 6.)
Length (of an adult 9), head and body 2'-3, head 1!-2, eye from tip of nostril 0-45, ear 0-6, forearm 2'-0, thumb 0!-85; third finger—metacarp. 1!-35, Ist ph. 1-0, 2nd ph. 1/25; fifth finger— metacarp. 1/25, Ist ph. 0'"65, 2nd ph. O'"7; tibia .0'-85, foot 0-55.
Hab. West Africa (Yoruba, Gambia, Lagos, Gaboon). This and the preceding species approach the species of Cynonycteris and Cynopterus more closely than others in the shape of the head ; indeed, dried skins appear at first sight to belong to species of these genera.
a,b. 9 imm. & ad. sks. Abeokuta, Yoruba. F. Nicholson, Esq. [P.]. ce. & ad, al. Lagos. Purchased.
d. 2 imm., al. Gaboon. H. Ansell, Esq. [P.].
2. PTEROPUS.
Pteropus, Brisson, Quadrup. p. 153 (1756); Geoffroy, Ann. du Muséum, xv. p. 90 (1810); Llkiger, Prodr. Syst. Mammal. p. 118 (1811) ; Peters, MB. Akad. Berl. 1867, p. 821; Dobson, Journ. Asiat. Soc. Beng. 1873, p. 196; Monogr, Asiat. Chiropt. p. 15 1876).
een et Spectrum, Gray, Catal. Monkeys & Fruit-eating Bats, pp. 100, 102 (1870).
Muzzle long, narrow, and cylindrical ; nostrils projecting by their inner margins, the extremity of the muzzle deeply emarginate between them; upper lip with a vertical groove in front, bounded laterally by rounded naked ridges; second finger with a distinct claw ; metacarpal bone of the middle finger shorter than the second ® Hitherto I have found no male of this species among the different specimens examined by me, and consequently I am unable to state whether shoulderpouches exist or not.
16 PLEROPODID.E.
finger; wing-membrane attached to the back of the first phalanx of the second toe; tail none ; interfemoral membrane deeply emarginate behind, in some species scarcely developed in the centre: penis with a distinct bone.
sie 4 1—1 3—3 2—2 Dentition. Ine. 5, ¢. j=, PM. gy M. E> 1— 5-8 Upper incisors in a semicircular row, in a plane anterior to and separated on each side from the canines; lower outer incisors close to the canines, the inner pair generally separated by a slight interval and smaller; first upper premolars generally very small and deciduous (larger than the middle lower incisorsin P#. leucopterus and Pt. molossinus only).
Skull elongated, the cranio-facial axis almost in the same right line, the brain-case but slightly vaulted and raised above the faceline, with (in most species) a prominent sagittal crest ; facial portion compressed ; cerebral cavity contracted in front; frontals small, with long postorhital processes, the bases of which are pierced by a foramen ; premaxillaries small, close together in front; bony palate produced backwards behind the last molar almost as far as the middle of the zygomatic arches; basioccipital wide between the auditory bulle ossee ; paroccipital processes long. Range. From the Comoro Islands in the Mozambique Channel to the Navigators’ Islands in the Pacific Ocean, extending throughout the Malagasy Subregion, the Oriental Region f, and the Australian Region (except the Sandwich Islands, Ellice’s group, Gilbert’s group, Tokelau, the Low Archipelago, and New Zealand).
Pt. edwardsii is found abundant in Madagascar, in the Seychelle Islands, and in the Comoro Islands (the latter are scarcely 200 miles distant from the African coast), and yet not a single species of this genus has been discovered in Africa (see remarks on the distribution of the genus Preropus in the Introduction). This genus includes the largest known species of Bats. In one of the species, at least, the body attains a length of 12 inches, the forearm of nearly 9 inches, and the outstretched wings measure about 5 feet across. They are the only Bats in which the fur of the back of the neck and shoulders differs conspicuously in colour and quality from that of the back. In most of the species, also, a more or less conspicuous tuft of thickened unctuous reddish-yellow hairs (generally about half an inch in diameter) exists.on either side of the neck near the point of origin of the antebrachial membrane, * In a specimen of Pteropus medius from Nipal (in the collection) an additional last upper molar exists on both sides, so that the number of back teeth is 6—6 6—6" Chiroptera, it appears evident that the three anterior teeth in each side of the upper jaw in Pleropus should be considered premolars, in which case the normal number of true mclars would be 4 + Pteropus pselaphon is found in Bonin Island, and Pt. dasymallus inhabits Southern Japan (Kiousiou Island), which has been included doubtfully by Mr. Wallace in the Palzarctic Region, and most probably really belongs to the Oriental Region.
As the number of true molars never exceeds = in any other genus of
2. PTEROPUS. 17 more developed in the male than in the female, and corresponding to the position of glands beneath. These glands secrete an oily substance on which the peculiar musky odour of the animals depends, and which is remarkably abundant during the breeding-season. The fur of the adult male, especially that covering the back of the head and neck, appears to be always more brightly coloured, and the differently coloured portions of the fur contrast much more strongly together than in the female of the same age*. But although the colour of the tippet in the female is darker or less brilliantly coloured than that of the male, it is rarely as dark as that of the fur of the back; and this difference in colour between the fur of the neck and that of the back is, perhaps, due to “ protective mimicry ”’t.
The teeth of all the species agree in number and in general characters, but differ remarkably in certain species in relative development. In most instances the premolars and molars are narrow, with an external and internal cusp separated by a longitudinal groove, which soon disappears with age (in some cases even before the animal has attained maturity) by the internal cusp becoming worn down. In Pt. leucopterus, Pt. aneiteanus, Pt. molossinus, and Pt. jubatus the premolars and molars reach their greatest development, in the firstnamed in general size, in the other three species in the presence of additional basal cusps and ridges; while in Pt. scapulatus and Pt. macrotis these teeth are reduced to the smallest size known in the genus, so that they resemble closely those of the genus Macroglossus. Such differences in the development of the teeth are evidently due to corresponding differences in the nature of the food ; for we find the molar teeth reduced to the most rudimentary condition in those species of the family which are known to feed on pulpy fruits, such as the long-tongued Macroglossi; these, it may be remarked, are inhabitants of the tropical parts especially of the regions through which the Pteropodide are distributed, where soft * See the remarks on this in the writer’s paper, ‘On Secondary Sexual Characters in the Chiroptera,” P. Z. 8S. 1873, p. 241. + At first sight it might appear that this remarkable contrast of colours would render the animal more conspicuous to passing enemies, and, consequently, more subject to their attacks when hanging in an exposed position, as, for example, from the branches of a lofty tree. But any one who has seen a colony of these Bats suspended from the branches of a banyan tree or from a silk-cotton tree, must have been struck with their resemblance to large ripe fruits; and this is especially noticeable when they hang in clusters from the leaf-stalks of the cocoanut-palm, where they may be easily mistaken for a bunch of ripe cocoanuts. Hanging close together, each with his head bent forwards on the chest, his body wrapped up in the ample folds of the large wings, and the back turned outwards, the brightly-coloured neck and head are presented to the view and resemble the extremity of a ripe cocoanut, with which the larger species of this genus closely correspond in size. i It is worthy of note that this remarkable contrast in colour between the fur of the neck and back is much less developed in the smaller species of the genus, and not at all in the comparatively aber 4 smaller species of the other genera of this family, which, by their small size, are less easily seen, and are enabled to hide themselves in places inaccessible to the larger species. Cc
18 PTEROPODID.E.
fruits are always obtainable ; and there are also found the largest species of the genus, while the smallest, and those provided with long woolly hairs extending thickly upon the extremities appear to be confined to the small wind-swept oceanic islands situated towards the extreme northern or southern limit of the distribution of the family.
So — are the number of species of this genus, and so closely are they related to one another by different characters, that it is extremely difficult to arrange them according to their natural affinities or to divide the genus into sections. At first sight it might appear possible to divide them according to the quality and distribution of the fur, or by the form of the skull and teeth; but the occurrence of such forms as Pteropus molossinus, which has the quality of fur and ears characteristic of one section, the distribution of fur of another, and the teeth of a third, or of Pt. poliocephalus, which resembles one group of species in the quality and distribution of the fur, while it agrees with a very different section in the development of the ears and teeth, renders a natural arrangement of the species in a linear series quite impossible.
The following Synopsis must therefore be considered chiefly as an index to the species, the position of each species with regard to the others not necessarily indicating its special affinities, which are pointed out in the general description. Synopsis of the Species.
a. Fur soft and woolly; ears generally shorter than oa muzzle, more or less concealed by the long ur, a’. Legs densely clothed as far as the ankles or backs of the feet with erect woolly hair (naked in Pt. molossinus only).
a", Fur of the back nearly as long as that of the neck; ears shorter than the muzzle.
a’. First upper premolar very small or deciduous; premolars and molars without prominent cusps.
a‘, Ears very short, clothed with long hairs, and concealed by the long fur of the head ; legs clothed above and beneath. 1, Larger: forearm 62; ears narrow, acutely pointed: a wiwecdoweawneintn Pt. vulgaris, p. 23. 2. Smaller: forearm 3-7; ears shorter, Tp. 24, obtuse; fur exceedingly fine and soft. Pt. rubricolls L'. Ears longer, half clothed, projecting , slightly beyond the fur of the head; legs half naked beneath.
b°, Legs clothed as far as the ankles. 3. Collar not well defined, passing gradually into the light-coloured fur of [p. 25. the packs tiscuet written nee Pt. dasymallus, yellowish white ..............., Pt. formosus, p.2€.
2. PTEROPUB.
ce’. Legs clothed to the base of the claws. 5. Collar scarcely distinguishable by colour from the fur of the back ; fore- BPM Oss eee see ets ghdsiiacaveaene's 6, Collar pale yellowish white above, very distinct; forearm 64 ..... 6". First upper premolar persistent, as large as one of the inner lower incisors; molars acutely tubercular; third lower premolar with an antero-internal broad projection. ce’, Ears slightly shorter than the muzzle, nearly naked; legs clothed to the ankles. 7. Collar bright reddish brown, shoulders pale yellow; forearm 4'"5..... arate: ce". First upper premolar persistent, as large as (or larger than) one of the lower outer incisors; second lower premolar with an antero-internal basal projection. d‘, Kars scarcely longer than the fur of the head, nearly naked ; legs naked. 8. Collar scarcely distinguishable by colour from the fur of the back; shoulderglands with tufts of bright yellow hairs; forearm 3'"5.............06. b". Fur of the back slightly appressed, unlike that of the neck; ears as long or longer than the muzzle.
ad". First upper premolar small or deciduous ; molars rather narrow.
9. Ears much longer than the muzzle, attenuated above and pointed; legs clothed above and beneath ; forearm 6’"2...... e'". First upper premolar persistent, nearly equal to one of the upper incisors in cross-section; molars very broad and strong. 10, Ears as long as the muzzle, obtusely pointed; legs clothed, above, as far as the ankles; fur and membranes very pale-coloured; forearm 6”............ b. Fur straight, rarely woolly on the legs; ears generally as long or longer than the muzzle. b'. No premolar or molar with an anterior basal projection. .
b". First upper premolar persistent, as large as one of the lower incisors, o'". Fur of the back appressed, on the interfemoral and legs erect, woolly. 11. Ears shorter than the muzzle; the last upper molar as large as one of the lower outer incisors; fur ferruginous brown; forearm 5B... cece eect eens ees 12. Ears slightly longer than the muzzle; the last upper molar very small, scarcely as large as one of the lower inner incisors ; fur yellowish brown; foream 45 ... c'. First upper premolar minute, generally deciduous. ce", Fur of the back straight, directed backwards, Dp.
. Pt. lombocensis, 19 p.26.
Pt. pselaphon, . Pt, vetulus, p. 27.
[p.29.
Pt. aneiteanus, [p.30.
Pt. molossinus, [p. 31.
Pt. poliocephalus, [p.32.
Pt. leucopterus, Pt. raynert, p. 83.
34.
o2
PTEROPODID.
scarcely appressed, on the interfemoral and legs similar.
c‘, Wings from the sides of the back; general colour of the fur and membranes brown. 18. Ears shorter than the muzzle; the concave surface of the conch thinly clothed with short hairs ; tibia covered almost to the ankles; premolars and molars with posterior basal projections; forearm 5". 14. Kars slightly longer than the muzzle, the concave surface of the conch nearly covered with soft fur, tibize half-clothed ; premolars and molars with posterior basal rojections; forearm 4’"9 ............ 15. Ears scarcely as long as the muzzle, naked; tibie clothed almost to the ankles; premolars and molars without basal projections ; forearm 4"5........ d‘, Wings from the back near the spine; fur pale-coloured, silky, generally with more or less distinct facial streaks; legs clothed to the ankles.
d°, With distinct facial streaks. a. Facial streaks black or dark brown. 16. Muzzle half surrounded by a black streak behind the eyes; forearm Orsseralcracosanatseiepeteraeteaneesites 17. Muzzle completely surrounded by a black streak behind the eyes; fore- APM AAS sire necator ss co5 raster aries B. Facial streaks white.
Me scerevecassysucieeoi uses nce:
Oeseetel anata alera ca SG y. No distinct facial streaks. 19. Ears as long as the muzzle; fore- OA a sco gees plover ade nieve eee . a". Fur of the back short, closely appressed (in adults) ; tibie naked or very thinly clothed with a few scattered hairs. e*, Molars very narrow, but slightly raised above the gum; first upper premolar minute, persistent.
20. Ears slightly longer than the muzzle; molars exceedingly narrow; forearm 5'-4 21. Ears much longer than the muzzle; molars larger ; forearm 5'"4 .......... J‘. Molars well developed ; first upper premolar minute, generally deciduous. J°. Wings from the back close to the spine ; back generally clothed by a narrow longitudinal line of short fur, or quite naked. f°. Interfemoral membrane very narrow in the centre, concealed by the fur. a, Earsnotlongerthanthe muzzle, obtuse. 22. Smaller: forearm 4”-5 ; membranes alo brown> sii ceencehs Geatels 23, Larger: forearm 7"; membranes black [p. 35, Ft. samoensis, [p.36.
Pt. rodricensis, Pt. brunneus, p.37.
[p.38.
Pt. personatus, [p.89.
Pt. capistratus, Pt. wallacet, p. 40.
[p.40.
. Pt. temminckii, [p.41.
Pt. scapulatus, Pt, macrotis, p.42.
Pt. griseus, p. 44.
{p. 44.
Pt. melanopogon,