1, EPOMOPHORUS. 5 The Bats of this genus are easily distinguished by their remarkably large and long head, very expansible lips, and by the invariable white tuft of hair which adorns the margins of the ears in front and behind in both males and females of every known species. The males are larger than the females, and have longer and narrower heads, and in most species are provided with peculiar glandular pouches, which are situated in the integument of the side of the neck near the point of the shoulder. ‘These pouches are rudimentary or quite absent in females. Jn the males they are lined with a glandular membrane, from which long coarse yellowish-white hairs arise, which project by their extremities from the mouth of the sac, and, when the pouches are everted, form conspicuous epaulet-like tufts on the shoulders. In the single species, #. monstrosus, in which these shoulder-pouches are not developed, the males are distinguished by most peculiar foliaceous processes of the integument of the front of the muzzle, of which rudiments only occur in the females (seo Plate I. figs. la & 2).
The fur is very similar in all the species, being rather short and of a peculiar light yellowish or cinnamon-brown colour with a greyish tinge ; the abdomen in many species with a large oval patch of whitish hairs, and in eyery species, even in the aberrant and very peculiar EZ. monstrosus, the bases of the outer and inner margins of the ear are adorned with tufts of white hairs.
The Epomophori live on figs, and probably on other soft fruits, the juicy contents of which their voluminous lips and capacious mouths enable them to retain and swallow without loss while the process of mastication is going on.
The different species agree so closely together in external form and in the colour of the fur that it is extremely difficult to distinguish them by characters derived from an examination of their external structure. Fortunately we have, in the form and arrangement of the palate-ridges, very reliable characters by which specimens preserved in alcohol may be at once determined, and these characters are set forth in the synoptical Table at p. 7, and illustrated by the figures on Plate IJ. In the case of dried skins the discrimination of the species is attended with considerable difficulty ; but the form of the ears, the presence or absence of the tail, and especially the measurements of the bones of the extremities afford the most reliable characters. E. lubiatus has been described from dried skins, and the only specimens known are skins also. Jt is therefore uncertain whether it is a distinct species or a small variety only of some of the larger species of the genus.
Synopsis of Subgenera.
I. Shoulder-pouches not developed in either males or females; front of the muzzle with rominent cutaneous foliaceous expansicns ; molars with distinct external cusps ...... Hyprignaruvs, p. 7.
6 1. PTEROPODIDA, II. Shoulder-pouches well developed in males, rudimentary or quite absent in females ; lips pendulous and folded on the sides (especially in adult males), but without broad cutaneous expansions in front ; molars with smooth slightly grooved crowns without CUSPS eee eigie ole yaa ote ere SHO gna chew wien sn EpoMoPHoRws, p. 7. Subgenus Hypsienataus.
1. Epomophorus monstrosus.
Hypsignathus monstrosus, Allen, Proc. Acad. Nat. Set. Philad. 1861, . 156; Peters, MB. Akad. Berl. 1867, p. 870; Gray, Catal. Mon- Le and Fruit-eating Bats, p. 124 (1870).
Spyrocephalus labrosus, Murray, P. Z. 8. 1862, p. 8, pl. i. Considerably larger than any other known species of the genus. Head (in adult males) very large and cylindrical, the anterior extremity of the muzzle thick and obtuse: the distance from the eye to the end of the nose slightly more than half the length of the head ; on either side of the extremity of the muzzle (in adult males) a very prominent, deep, semicircular fold of integument, continuous above with the lower margin of the nostril, and below and behind with the horizontal margin of the upper lip; front margin of the upper lip papillate, fimbriated; corresponding margin of the lower lip similar, but thinner and divided by a V-shaped incision in the centre. In females the muzzle is conical and much shorter, and its upper surface scarcely elevated above the nasal apertures, the lower margins of which are connected with the sides of the lower lip by folds similar to those in the males, but very much less developed (Plate I. figs. 1, la, 2).
The palate has three undivided straight transverse ridges, whereof the first corresponds to the space between the canines and the first premolars, the second to the second premolars, the third to the molars; the fourth consists of two papille ; the remaining ridges are indistinct, arched forwards and finely toothed (Plate II. fig. 1). In the adult male, on either side of the neck, a strong integumentary band extends forwards from the point of origin of the antebrachial membrane to the side of the back of the head. These bands are not developed in females, and they appear to be supports to the head, which is so remarkably large in the adult male. Ears shorter than the muzzle, triangular, attenuated in their upper third, and subacutely pointed; the usual white patch of hairs along the anterior and posterior margin of the ear-conch. Interfemoral membrane 0'"4 deep in the middle; no trace of a tail; wings from the base of the second phalanx of the second toe. Upper surface of the muzzle nearly naked in the male, in the female covered with very short grey hairs; fur above and beneath rather short, but dense and very soft, greyish slate-brown, paler beneath. About three fourths of the forearm is covered with short fur,
1. EPOMOPHORUS. i and the fur of the back extends thickly upon the interfemoral membrane, concealing its posterior margin in the centre, and the legs are clothed as far as the ankles. Upper incisors small, in a semicircle, separated from each other and removed from the canines by a space on either side; lower incisors in pairs, separated by a slight interval in the centre; the third lower premolar and the first molar have each three cusps externally, very distinct in the first molar especially ; the second and last molars have two external cusps. The adult female differs from the male not only in the smaller and differently shaped head and in the absence of the neck-bands described above, but also in being smaller in nearly all other dimensions, as the following Table of measurements * shows :— Ist ph. 2nd ph. Ist ph.
2nd finger, metacarp 4th finger, metacarp ’ ’ ” eye from nose.
tibia. foot.
ear.
forearm.
thumb.
” ’ ” ’ ” ” eS ’ Length, head.
16 | 1:25, 52/19 | 36 | 2-45) 3-4 | 3-4 | 1-65 55) 21 13 oo ne Adult ¢.
| Adult Q.] 27] 1:3] 1-1 | 47) 1-85] 34 | 235) 33 | 3:25) 1-60 bie 2:0 | 1:3 “i Hab. Western Tropical Africa (Gambia, Calabar, Gaboon). a. @ ad. sk. Gambia. Purchased. b. Q juv., al. W. Africa. Purchased. _ ec. f ad., al. Old Calabar. A. Murray, Esq. [P.]. (Type of Spyrocephalus labrosus, Murray.) d. 2 imm.,al. Neighbourhood of Cameroon Purchased. Mountains.
« § imm.,al. Elloby District, Gaboon. H. Ansell, Esq. [P.]. Subgenus EpomopHorvs. Synopsis of the Species. ‘A. With transverse palate-ridges in front. (Large Epomophori.) a. Palate-ridges very prominent, no semicircular toothed ridges behind; tail very small but distinct. a’. Muzzle very long and slender; the fifth palate-ridge divided, with a lozengeshaped cell in the centre ....-.-++- 1. E. macrocephalus, p. 8. * he measurements in all cases are in inches and decimal parts of an inch.
8 PTEROPODID.E.
b'. Muzzle shorter and broader, the fifth palate-ridge simple, divided in the :
centre by a narrow incisor ........ 2. E. gambianus, p. 10. b. Palate-ridges less prominent, with semicircular toothed ridges behind; no tail externally.
c'. Muzzle shorter than in EF. gambianus ; the third palate-ridge developed on ; the sides only ....... 0. eee eee eee 3. E. franqueti, p. 12. d'. Muzzle short; the third palate-ridge complete, undivided ; upper incisors (in adults) two only .............. 4, E. comptus, p. 13. B. No transverse palate-ridges. (Small Epomophori.)
e'. Muzzle much shorter and comparatively broader than in any of the preceding species; palate with a longitudinal -shaped groove ........ cece eee 5. E. pusillus, p. 14. Two other species have been recognised, Epomophorus wahlbergii, Sundevall, and £. labsatus, Temminck (see p. 11). The former appears to me to differ only from &. gambianus in the greater hairiness of the extremities; the latter seems to be distinguished chiefly by its smaller size. Both are represented as yet by dried skins only, so that an examination of their palate-ridges is impossible.
2. Epomophorus macrocephalus.
Pteropus macrocephalus, Ogilby, P. Z. S. 1835, p. 101; Wagner, Suppl. Schreb. Sduyeth. i. p. 867 (1844).
Pteropus megacephalus, Swainson, Nat. Hist. and Class. Quadrup. p- 92 (1835).
Pteropus epomophorus, Bennett, P. Z. S. 1835, p. 149. Epomophorus whitei, Bennett, Trans. Zool. Soc. 1835, vol. ii. p. 38, pl. 6 Epomophorus macrocephalus, Tomes, P. Z. S. 1860, p. 50, 1861, p. 11, figs. 1, la; Peters, MB. Akad. Berl. 1867, p. 869. Head very long and narrow ; muzzle nearly half its length, much longer than the ear ; the eye is less than half the distance from the ear that it is from the end of the muzzle; nostrils rather prominent and diverging, opening laterally with a wide emargination between ; the upper lip rounded in front, divided infericrly by a narrow shallow groove, which is not continuous with the emargination between the nostrils above; sides of the lip much expanded and folded in old males, with a prominent fleshy glandular (?) cushion on each side, half internal, near the angle of the mouth. Ears simple, oval, much longer than broad, rounded off above, the inner margin convex to the tip, the outer margin similarly convex in lower two thirds, flattened or slightly concave in upper third. Shoulder-pouches large, well developed in adult males, rudimentary in females and in young males.
Wing-membrane from the middle of the back of the first phalanx of the second toc; interfemoral membrane very narrow in the
1. EPOMOPHORUS. 9 centre ; tail very short and feeble, quite free from the interfemoral membrane, and concealed by the surrounding fur. Above yellowish brown, with a slightly greyish tinge; paler beneath ; the abdomen pale yellowish white ; a tuft of white hairs along the lower third of the outer and inner margins of the ear. Fur short, extending upon the humerus and fleshy part of the forearm and upon the adjoining wing-membrane, but the antebrachial membrane is nearly naked; the greater part of the interfemoral is covered with short fur, which also extends along the legs to the back of the feet, and short hairs are thinly spread over a large part of the wing-membrane between the humerus and the leg. Roof of the mouth much vaulted, especially posteriorly, with six very prominent palate-ridges, which are characteristic of the species: the first is straight, and is placed across the palate slightly behind the canines ; the second is slightly arched forwards, and corresponds to the first upper premolar; the third is close to the second, is more arched, and commences opposite the second premolar, or opposite the space between the first and second ; the fourth is separated by a greater interval, and corresponds to the molar teeth, though it does not extend as far as their bases laterally ; the fifth is placed at a still greater distance behind, and is very peculiarly shaped, being rhomboidal, hollowed out in the centre, and divided into two lateral triangular halves by a narrow groove; the sixth ridge is simple, divided in the centre, and placed at an equal distance behind (Plate IT. fig. 2).
The head is much shorter in females, which are also slightly smaller than the males in most measurements. In young males the head is also longer than in young females of the same age, though not in so marked a degree. In the following Table of comparative measurements these differences are shown :— eye from nose.
forearm.
second finger. fourth finger.
tibia. foot.
” ’ ” ’ ? ’ ’ ’ Length, head and body.
head. ear Adult g.| 63 | 26 | 09 | 12 | 35 | 59 | 45 | 13 | 0:85 Adult 9.| 50| 23 | 09 | 10 | 33 | 57 | 43 | 125] 0-85 Duma eo ac) BO | OR OB) SB 1 ss: |) eee | | Immat.@.| 38| 1:65] 08 | 065| 28 | 47 | 365] 1:05! O85 Hab. West Africa (Gambia, Guinea, Lagos). °
10 PTEROPODID.E.
E. macrocephalus may be at once distinguished by the very long and narrow muzzle, nearly half the length of the whole head, and by the peculiar form of the fifth palate-ridge. a-d. f& ad. sks. Gambia.
ef. dg & ad., al. West Africa, Mr. Argent’s collection. g. & ad., al. West Africa, 3. Epomophorus gambianus.
Pteropus gambianus, Ogilby, P. Z. 8. 1835, p. 100. ? Pteropus wahlbergii, Sunderall, Ofv. Vet. Akad. Stockh. 1846, p. 118. Epomophorus crypturus, Peters, Reise nach Mussambique, Sdugeth. p- 26, pl. v. & pl. xiii. figs. 1-6.
Head very long, nearly as long as in H. maerocephalus, but broader, and the length of the muzzle is comparatively less. Lips very pendulous, with remarkably large cheek-pouches in adult males (Plate LI. fig. 3). Ears oval, nearly as long as the muzzle (in the ¢); outer and inner margin of the ear-conch almost equally convex. Nostrils simple, their inner margins projecting slightly in front, separated by a deep emargination, which also divides the upper lip in front.
Shoulder-glands large, well developed in adult males, rudimentary in females and in young males. “ Wing-membrane, interfemoral membrane, and tail as in #. mucrocephalus.
Above yellowish brown, paler beneath; the abdomen in adult males with greyish oval patch.
Fur of the back moderately long and dense, covering the short interfemoral membrane behind and the wing-membrane along the sides of the body, extending also thickly upon the fleshy part of the forearms, and upon the legs to the backs of the feet. Roof of the mouth much vaulted, as in #. macrocephalus, but wider. Palate-ridges six, straight, very prominent: the first is placed slightly behind the canines, and has a slightly triangular projection in the centre of its posterior margin; the second and third ridges are simple, undivided, and close together, the furrow between them corresponds to a line drawn between the first premolars ; the fourth is opposite the first molar, and is half divided by a small incision in the centre; the fifth corresponds to the second or last molars, and is divided by a narrow incision in the centre into two half-ridges ; the sixth is behind the tooth-row at a considerable distance, and is similurly divided (Plate II. fig. 3 a). The muzzle is much shorter in females, which are also (as in EZ. macrocephalus) slightly smaller than the males in most measurements. In the following Table of comparative measurements, the sizes of an adult male and female are contrasted :-——
1. EPOMOPHORUS. 11 Epomophorus gambianus. Ad. g. | Ad. Q.
Length, head and body scssccecescsscavseescctsinssesesvacevsses 58 55 speORL:Aickdeteuaaeesste:sooncitesaneecasoaiessedanedexesosts2-42-0 » eye from tip of nostril 1... ce ceeeeceeeeeeeee ree 1:0 0-7 Bi OBIE Rca Heer Mate rdioncs aa cakerave ai aannanenaseteeees 0:95 0-85 » forearm 3-4 33 » thumb 1:55 1-4 , third finger, metacarpal 2°35 2°25 A is 1 15 15 ” ” ” 2 2:2 2°25 » fifth finger, metacarpal 2:25 2:15 - % aye. AA SUYPU. oe cavaeachusuavoestwcsnmcaseeuecane 11 1:2 i. ei, (eNO pbs. yetatcceaaceG ar waeestes Pecaatese Aci 1:15 fyRADAssissiaisisiseensrasacienvaasreuceneseagersinissaeedaaivawsanenn142 SieMOOGis.aneeaceedecaaoevouaesGocenenebediewvoceeeeantesy0-807 Hab. Equatorial and Southern Africa, extending even as far south as Port Elizabeth.
Specimens from the southern extremity of the African continent have their extremities more densely clothed with hair. Hpomophorus wahlbergit, Sundevall, was founded on an example from Natal, which appears to differ from E. gambianus only in the greater hairiness of the legs. . crypturus, Peters, was described from a female specimen, probably not full-grown, which, to judge from the description of the palate-ridges, appears to be identical with this species.
A specimen of an apparently immature male (to judge from the condition of the extremities of the finger-bones) from Benguela (marked “@” in the collection) is even slightly larger than the adult specimens from which the above measurements were taken, and the shoulder-pouches are not well developed.
a. imm. sk. Gambia.
b. ad., al. Gaboon. H. Ansell, Esq. [P.].
e. 2 imm., al. Benguela, Angola. I. F. Monteiro, Esq. [P.]. d. dé imm, al. Benguela, Angola. F. F. Monteiro, Esq. | P. ]. e. o ad., al. Angola.
f. Q juv., al Angola. ; a g,h. & ad., al. Shupanga, Zambesi. Dr. Kirk tet 2 9 imm,, al. Dar-es-Salanus. Dr. Kirk [C.].
jk. 6 & Q sks. Port Elizabeth.
lad. 8 3a. Epomophorus labiatus.
Pteropus labiatus, Temminck, Monogr. Mammal. ii. p. 83, pl. 39 €1835-41) ; Wagner, Suppl. Schreb. Stiugeth. i. p. 306 (1844), v. .608.
eehyenata labiatum, Temminck, Esquiss, Zool.. sur la eéte de Guiné, p. 68 Pteropus schoénsis, Riippell, Mus. Senckenb. iii. p. 131 (vide Peters, MB. Akad. Berl. 1867, p.869).
Epomophorus labiatus, Tomes, P. ZS. 1861, p. 11; Peters, t.c. Epomophorus anurus, Heuglin, Nova Acta Acad, Carol. Leop. Nat.- Cur, xxiii. p. 12 (1864).
12 PTEROPODID.E.
The type of #. labiatus in the Leyden Museum appears to me to be an example of an immature individual of a small variety of &. gambianus. No trace of palate-ridges exist in the dried stuffed skin, so that it is difficult to determine its relationship to &. gambianus with certainty. In the colour and distribution of the fur, in the form of the ears, and in the relative length of the bones of the extremities it agrees closely with that species.
The following are the measurements of the type :— Length, head 1-7, ear 0!'-72, eye from nostril 0"-75, forearm 2''-6, thumb 1!-2; third finger—metacarp. 1!"8, Ist ph. 1!/-1, 2nd ph. 1'"6 ; fifth finger—metacarp. 1!"7, Ist ph. 0/8, 2nd ph. 0-8 ; tibia 1'"1, foot 0!'-65.
Two skulls in the collection from Abyssinia and a skin evidently belong to this form; one of the skulls affords the following measurements :— Extreme length 1-8, breadth across zygomatic arches 1-0; from supraorbital foramen to end of nasals 0-9; length of bony palate 0"-95, width of same 0!-5, Hab, Abyssinia, Shoa.
a. ? ad. sk. Abyssinia. Purchased.
b. skull of a.
e. skull of specimen from same locality.
+. Epomophorus franqueti.
Epomophorus franqueti, Zomes, P. Z. S. 1860, p. 54, pl. 75. Epomops franqueti, Gray, Catal. Monkeys and Fruit-eating Bats, p- 126 (1870).
Muzzle shorter than in 2. gambianus, but, as in that species, longer in the male than in the female; ears as long as the muzzle (in the adult male); ear-conch rounded off above, the upper third of the outer margin faintly concave ; nostrils separated by a rather wide and shallow emargination, which is continued downwards as a small shallow groove dividing the upper lip in front ; shoulder-glands in the male large, with very long yellowish-white hairs projecting to a considerable distance, in the female quite absent; sides of the lips expansible, but not to such an extent as in 4. gambianus. Wing-membrane from the middle of the back of the first phalanx of the second toe; interfemoral membrane wider in the centre than in other species; no trace of a tail.
Above, cinnamon-brown ; beneath, similar, but paler, and the abdomen has a very distinct oval patch of whitish hairs in the centre. The fur extends outwards upon the membranes almost as far as a line drawn from the middle of the humerus to the knee, covers the interfemoral between the thigh to its posterior margin, and passes downwards thinly upon the legs.
The palate-ridges are peculiar, but less developed than in any other known species of the genus. The first two only are prominent and undivided ; the remaining ridges are represented by small oval