The Mammals of the Southern African Subregion

<>*«««««« W— J9cck fL-U clM c-A-l-i - 289 Elephant (Loxodonta africana) Olifant Wi THE MAMMALS OF THE SOUTHERN AFRICAN SUB REGION BY JOHN D. SKINNER M.Sc. Ph.D. F.I.Biol. F.Z.S. F.R.S.(SA) Director Mammal Research Institute University of Pretoria AND THE LATE REAY H.N. SMITHERS D.Sc. M.I.Biol. Senior Research Officer Mammal Research Institute University of Pretoria WITH COLOUR PLATES BY DICK FINDLAY University of Pretoria, Pretoria Republic of South Africa 1990 Order CHIROPTERA Bats The fossil record The earliest known fossil bat was discovered in the early Eocene deposits in America (Jepsen, 1966). In relative dating this is 75 million years after the appearance of the first mammals. From the structure of its teeth and skeleton it was certainly an insect eater although, unlike present day insect-eating bats, it had two claws on the forearm, one on the thumb, the other on the first finger, which is a characteristic of modern fruit eaters. In all respects, however, it closely resembles bats as we know them today but gives us no clues as to the origin of flight in the Order or their origin prior to this time. The oldest fossil bat known from Africa is Vampyravus orientalis which was recovered from the Oligocene deposits of Egypt, but whose relationships remain problematical. For a long time it was thought that bats were related to the Tupaiidae, the tree shrews, a Family of insectivores that occur in Asia and India, and to the Dermoptera, the flying lemurs of the Far East and that the Chiroptera and Insectivora evolved from a common, probably arboreal, ancestor (Anderson & Knox Jones, 1967). Pettigrew (1986) and Pettigrew, Robson, Hall & Mc Anally, (in press) have shown that the Megachiroptera have a Primate-like visual system and musculoskeletal adaptations of the flight apparatus, neither of which are found in the Microchiroptera. They believe that the Megachiroptera evolved from early Primates, which were moderate sized gliders, of which the other living descendants are the Dermoptera; the Microchiroptera from small, agile insectivores with forelimbs modified for gliding during leaps. This means that the power of flight must have evolved in parallel in two separate lines of mammals, one of them ancestral Primates. Four Families have been recognised from the early Miocene beds of East Africa. The fruit-eaters, the Pteropodidae, are represented by Propotto Ieakeyi, whose fossil remains were thought to be those of a lemur originally (Simpson, 1967). but were later shown to be those of a fruit bat (Walker, 1969). Its cheekteeth were set closer together than in modern forms, the molar teeth low with blunt cusps and lacking the longitudinal groove seen in our living pteropodids. Butler (1978) nevertheless, did not believe that the Pteropodidae were of African origin as it is in the Oriental Region that the greatest diversity is found. In the Microchiroptera the six Families that are represented in the Subregion occur in the fossil record as follows: The Emballonuridae, the sheath-tailed and tomb bats, are known from the early Miocene beds of Rusinga, on the shores of Lake Victoria, and in Europe from the early Oligocene. The Nycteridae, the slit-faced bats, are known from undescribed material from the Pliocene beds of East Africa (Butler. 1978). The Rhinolophidae, the horseshoe bats, are known from the early Eocene or early Oligocene of Europe and the late Pliocene deposits of Makapansgat, Transvaal (De Graaff, 1960a). The Hipposideridae occur in the middle Eocene beds of Europe, but no fossil remains have been recovered so far from the Subregion. The two largest Families of bats that occur in the Subregion, the Vespertilionidae and the Molossidae, which are known from fossil remains from the beginning of the Oligocene in Europe and from scanty remains from the MioPliocene of North Africa, are not known from East Africa until much later, in the early Pleistocene (Butler, 1978). Dentition Young bats have milk teeth like most young mammals. Those of the Megachiroptera are simpler than those of the Microchiroptera and are peg-like with hooked ends. In the Microchiroptera they are similarly hooked, but have one or two cusps lower down on the sides of the teeth below the primary cusp, which is located near the tip. It is thought that this hooking assists the young in clinging more firmly to the teat or fur of their mothers when being carried around. In some Families, e.g. the Rhinolophidae, milk teeth are present in the young prior to birth, but are reabsorbed before they are born (Spillman, 1927). Slaughter (1970) suggested that, as females of this Family have a pair of dummy teats situated low down on the belly not connected to the mammary glands, there may be less need for the clinging milk set (see Family Hipposideridae, Fig. III.l). While many species retain the maximum number of three incisor teeth on either side of the lower jaw, none of them have more than two on either side in the upper jaw. Which incisor has been lost in the course of evolution has been the subject of controversy among authorities. In many bats the cutting faces of the lower incisors are divided into two (bifid) or three (trifid) sections, which may be used like combs in grooming. The canine teeth are well developed and, in many insect-eating bats, have a broad collar near the base, better developed inside than on the front of the teeth. Slaughter (1970) suggested that it protects the gums from damage by the hard exoskeletons of beetles. Fruit-eating bats do not require this protection and have canines that are smooth to their bases. No bat has more than three premolar teeth on either side of both jaws, although the basic number for placental mammals is four. Again there is some controversy as to which has been lost, the most widely accepted theory being that it is the first (Miller, 1907). In the fruit-eating bats the molar teeth have flat faces, an adaptation to masticating soft food. In the insect-eaters, the cheekteeth are equipped with high cusps to deal with their harder insect diet. In the fossil record there are no types of dentition which are intermediate between the two widely different types. Location of prey Those who study and therefore have to handle bats grow to appreciate their cleanliness and the delicacy of the complicated structures of the nose and ears which are part, in many species, of the sophisticated mechanism known as echolocation which allows them to navigate and catch their prey in total darkness. Clicks or bleeps made by bats bounce off objects in their path and their positions are then located by perception of the echo from them. The time lag between the emission of the clicks or bleeps by the bat, their reception after bouncing back from the object ahead of them and the bat's reaction to the stimulus is something to the order of 1/100th of a second. The bat is both the transmitter of the impulses and their receiver. The outer ears are the amplifiers, the inner ear the receiver and booster, and the brain the computer that sorts, files and transmits the information to the body. Deprived of the use of one ear, a bat perceives direction but cannot pinpoint the object. The calls of bats are mostly far above the range of sound audible to the human ear, which is from about 20 vibrations a second, in some people with acute hearing, up to 18 000 vibrations per second. The sound a bat emits to locate obstacles and its prey may extend up to as much as 230 000 vibrations per second. Bats can vary the length of their calls, sending out pulses of less than 1/1000th of a second when nearing the prey, at the same time making adjustments to their course to compensate for the prey's violent evasions. Bats use different kinds of sound to obtain different kinds of information. A steady call allows them to estimate the 54 speed and direction of the prey. This system exploits the Doppler effect which causes the fall in pitch of an automobile horn as it passes the listener. Another call employs frequency modulation or change of pitch. Echoes of this call are heard slightly differently by the bat's two ears and its brain interprets the difference in terms of heading and range of the moving prey. Some bats improve on their accuracy by adding a harmonic call to the basic call or by raising the pitch of the call. In doing the latter they must sacrifice range since high frequency sound dissipates faster in air than low frequency. Using low frequency calls, bats can detect prey at up to about 10 m but with high frequency not over about 0,6 m (Fullard, 1981). Wing and flight membranes The flight membranes of a bat extend from the shoulder around the tips of the digits to the foot, continuing as the interfemoral membrane from the inside of the hind limbs and partially or wholly enclosing the tail vertebrae. The relationship of the membrane to the tail vertebrae is a most important character in distinguishing between the Families and the manner in which it attaches to the foot is constant within a genus or species. The membrane is elastic and contracts as the digits are folded together, and is supplied liberally with blood vessels. When at rest, the wings are folded against the body. Reference is made in the text to the bony structure supporting the membrane, the various parts of which are illustrated and named as follows: Key to the Suborders in the Subregion, after Hayman & Hill (1971) 1. Second digit on forearm terminating in a claw (Fig. V.l), margin of the ear forming a complete ring; ear tragus absent; interfemoral membrane greatly reduced, little more than a narrow band along the margin of the hind legs; tail absent or rudimentary (Fig. 45.1); cheekteeth simple, without well-developed cusp pattern (Fig. V.3) . . . Suborder Megachiroptera The fruit-eating bats 2. Second digit on forearm without a claw; margin of the ear not forming a complete ring; ear tragus generally present (absent in Family Rhinolophidae); interfemoral membrane and tail generally well developed; cheekteeth cuspidate, with generally well developed W pattern . . . Suborder Microchiroptera The insect-eating bats Fig. V.l. Wing of a fruit bat. Pteropodidae: Eidolon helvum; note claws on first and second fingers. Fig. V.2. Head of a fruit bat, Pteropodidae: Epomophorus sp. a upper arm or humerus b forearm c first digit or thumb d second digit or finger e third digit or finger f fourth digit or finger g fifth digit or finger h metacarpal i first phalanx j second phalanx k shin bone or tibia 1 thigh bone or femur Classification The Order Chiroptera is divided into two Suborders, the Megachiroptera, the fruit-eaters, and the Microchiroptera, the insect-eaters. Suborder MEGACHIROPTERA Fruit-eating bats All members of this Suborder belong to the Family Pterooccurring in the Subregion is simple, tapering and dog-like podidae, the general account given of the Suborder applying without noseleaves (Fig. V.2). to all members of this Family. The muzzle in all the members These are characterised by the fact that most of them have 55 two claws on the wing structure, one on the thumb, the other on the first or index finger (Fig. V.l). All Megachiroptera of the Subregion have these two claws, the Microchiroptera have only one. The ears of the Megachiroptera are simple tubular structures and lack the ear tragus possessed by all the Microchiroptera, with the exception of the members of the Family Rhinolophidae. There are other important features which differ in the two Suborders. Mossman (1937) for example, showed that the foetal membranes are fundamentally different, differences which involve the development of the yolk sac and the structure of the placenta. Most of the Megachiroptera rely on sight for orientation, the exception being the Egyptian fruit bat, Rousettus aegyptiacus, which has in addition powers of echolocation. They orient visually, when there is sufficient light to allow for this, but emit "clicks" with their tongues in total darkness which, picked up by the ears as the sound bounces back, give them powers of orientation. In the Microchiroptera the "clicks" are produced by the larynx and they rely on their powers of echolocation in total darkness for their orientation, their sight being less well developed than in the Megachiroptera. Although the Megachiroptera are classed generally as fruit-eaters, and for most of them fruit is their principal food, some of them eat flowers, flower buds and pollen and others lap nectar. There are no true nectar lappers in the Subregion (see Epomophorus gambianus), but they have interesting adaptations to their feeding habits. The long-tongued bat, Glossophaga soricina, of South America and other nectar lappers have extremely long thin tongues, the tips of which are equipped with long spinous papillae which act like absorbent brushes. Their muzzles are long and narrow and their molar teeth are to all intents and purposes nonfunctional. The Megachiroptera include among its members some of the world's largest bats, as well as some very small species. The flying fox, Pteropus neohibernicus, of New Guinea, has a wing span of some 1,8 m; the African Nanonycteris veldkampi a wing span of about 180 mm. The largest species that occurs in the Subregion is the straw-coloured fruit bat, Eidolon helvum, which has a wingspan of up to 0,66 m. The Megachiroptera tend to be gregarious, occurring in huge colonies. In the Subregion one of the species, Peter's epauletted fruit bat, Epomophorus crypturus, regularly occurs in colonies numbering hundreds and the cave dweller, the Egyptian fruit bat, Rousettus aegyptiacus, in thousands. In the Subregion eight species of Megachiroptera, or fruit-eating bats occur. As far as is known the strawcoloured fruit bat, Eidolon helvum, only visits the Subregion on migration, returning to the forested regions further north to have their young. They are not common in the Subregion. All eight members of the Family Pteropodidae that occur in the Subregion can be recognised by their dog-like faces (Fig. V.2), with elongated snouts. They are fairly large with total lengths, measured from the tip of their snouts to the end of their small tails, where they protrude sufficiently to be measured, of over 120 mm. The diagnostic feature used in differentiating between the four species of Epomophorus and the one species of Epomops is the arrangement and positioning of the ridges on the skin of the palate (Figs 40.1; 41.1). V. Family PTEROPODIDAE Key to the genera after Meester et al. (1986) 1. Ears with white basal tufts ... 2 Ears without white basal tufts ... 3 Fig. V.3. Pteropodidae, teeth of lower jaw, right hand side, (a) Eidolon helvum (b) Rousettus aegyptiacus. 2. Six prominent palatal ridges, one or two being postdental, and these are no different from inter-dental ridges; post-dental palate strongly concave posteriorly .. . Epomophorus Five palatal ridges, the last two post-dental, these being differentiated from interdental ridges, thick, and each bearing two triangular projections; post-dental palate flattened posteriorly .. . Epomops 3. Forearm generally 110-130 mm; colour generally tawny, with dorsal fur restricted to a narrow median band, sharply demarcated from naked wing membranes ... Eidolon Forearm 65-102 mm; ruff of stiff hair present or absent on lower neck of adult males .. . Rousettus Genus Epomophorus Bennett, 1836 Key to the species after Hayman & Hill (1971) (Measurements in mm) 1. One palatal ridge behind the last tooth (Fig. 40.1.a); length of forearm 77—89 in males, 72—86 in females . . . wahlbergi Two palatal ridges behind the last tooth (Fig. 40.1.b) ... 2 2. Fourth palatal ridge midway between the third and fifth (Fig. 40.1.b) ... 3 Fourth palatal ridge much nearer the third than the fifth; length of forearm 87-91 in males, 82 in females (Fig. 41.1.b) ... angoiensis 3. Length of forearm 87-93 in males, 81-86 in females .. . gambianus Length of forearm 81-85 in males, 79-80 in females ... crypturus No. 40 Epomophorus wahlbergi (Sundevall, 1846) Wahlberg's epauletted fruit bat Wahlberg se witkolvrugtevlermuis Plate 5 Colloquial Name Named after Johan August Wahlberg, explorer, hunter and outstanding collector of zoological material in southern 56 Africa. He was killed in 1856 by a wounded elephant near the Savuti in northeastern Botswana (Gyldenstolpe, 1934). Taxonomic Notes Hayman & Hill (1971) listed two subspecies: E. w. haldemani (Halowell, 1846) from the northern and western parts of their distributional range on the continent, and E. w. wahlbergi, mainly from the southern and eastern parts. There is considerable overlapping, both in characters and distribution and E. w. haldemani eventually may be shown to be untenable (Hayman & Hill, 1971). Description Buffy-brown to brown on the upper parts and pale buffybrown on the under parts. In the field it is impossible to distinguish Wahlberg's epauletted fruit bat from Peters' epauletted fruit bat, with which it so often associates. The males, as in Peters', have glandular pouches on the shoulders covered with long white hairs which, when open, resemble white rosettes. Both sexes have whitish patches at the base of the brownish ears. The only way the two species can be recognised readily is that in Wahlberg's there is only one ridge on the palate behind the last molars, whereas there are two ridges in E. crypturus (Fig. 40.1). Wahlberg's is slightly smaller than Peters', the comparative lengths of the forearms being 77-89 mm in males and 72-86 mm in females, whereas in Peters' it is 81-85 mm in males and 79-80 in females (Table 40.1). Table 40.1 Measurements (mm) and mass (g) of Wahlberg's epauletted fruit bats, E. wahlbergi Province, and in Angola. There are no records from northeastern Mozambique, north of the Zambezi River, but it is likely that they occur there, at least in the east, as they are recorded from adjacent parts of southern Tanzania and Malawi. Males Females X n Range X n Range TL 138 7 120-156 140 8 122-144 Hf c/u 23 7 20-25 21 8 18-23 E 23 7 20-27 24 8 22-28 F/a 84 3 81-85 78 8 68-85 Mass 93,9 5 67,7-113,9 112,3 4 94-140 .xdXj Fig. 40.1. Palatal ridges: (a) Epomophorus wahlbergi (b) E. crypturus. Distribution South of the Sahara, excluding the Southern African Subregion This species is confined to the continent south of the Sahara and has been recorded coastally in the Gabon; Central African Republic, and in parts of northern and southern Zaire. It occurs in Uganda; Kenya; Somalia; in the eastern parts of Tanzania; widely in Malawi and Zambia, excluding the southern parts of the country and parts of the Northern Southern African Subregion In this sector they tend to be confined to the eastern parts of the continent, penetrating westwards into the drier areas up river valleys such as those of the Zambezi and Limpopo rivers and their tributaries. They occur widely in Mozambique, south of the Zambezi River, penetrating into the eastern parts of Zimbabwe, where they also occur in the Limpopo River Valley as far west as its confluence with the Shashi River. In the Transvaal they occur in the Limpopo River Valley and in the eastern parts of the province penetrate westwards to Tzaneen in the Letaba District and southwards to the Groblersdal District. Their distribution in the Transvaal and in adjacent parts of Mozambique suggests that they eventually will be shown to occur in Swaziland, but there are no records to substantiate this at the moment. They occur at least in the eastern parts of Natal and narrowly along the coastal fringe as far as the Uitenhage District in the Cape Province. Habitat Tropical forest and evergreen riverine forests where there are fruit-bearing trees. While their occurrence lies largely within areas with a mean annual rainfall in excess of 700 mm, they penetrate up river valleys carrying evergreen forest into otherwise much drier country with a minimum annual rainfall as low as 250 mm (Limpopo Valley). Habits They hang up during the day in the dense canopy of evergreen trees, sometimes in colonies numbering dozens of individuals and often in association with Peters' epauletted fruit bat, E. crypturus, which is always in far greater numbers. In coastal cities in Mozambique they roost in evergreen trees in the cities' parks, and even in the trees along busy streets. Like E. crypturus they exhibit considerable local movements actuated by their search for food. At preferred feeding sites they rfiay settle in the cover of evergreen trees during 57 the day and then quite suddenly, when the supply is exhausted, move off elsewhere (Fenton, Brigham, Mills & Rautenbach. 1985). Food Soft and pulpy wild fruits such as wild figs, Ficus spp, are favoured in particular and they have also been observed to take mahobohobo, Uapaca kirkiana and U. sansibarica: mobola plum, Parinari curatellifolia and kudu berries, Pseudolachnostylis maprouneifolia. Among orchard fruits they will take guavas, plums and apricots. Their feeding habits are similar to those of Peters' epauletted fruit bat, E. crypt urus. Reproduction Gravid females with a single foetus have been taken in Zimbabwe in June and December which suggests that they may breed widely throughout the year but further information is required to substantiate this. No. 41 Epomophorus angolensis Gray, 1870 Angolan epauletted fruit bat Angola-witkolvrugtevlermuis Colloquial Name Named Angolan as it was described originally from a specimen taken at Benguela, Angola. Taxonomic Notes No subspecies have been described. Description Very similar externally to E. crypturus and E. wahlbergi, but slightly larger, the males with forearm lengths of 87-91 mm, the females about 82 mm. Males have a total length of about 190 mm; about 150 mm in females (Table 41.1). They have two palatal ridges behind the last molar teeth as in E. crvpturus and E. gambianus, but differ from them in that the third and fourth ridges from the front are nearer together than to the second and fifth (Fig. 41.1 .b). In other skull characters they are similar to E. gambianus and different from E. wahlbergi which has only one palatal ridge behind the last molar teeth. Table 41.1 Measurements (mm) of a series of female E. angolensis, from northern Namibia (Shortridge, 1934) Females X n Range HB 146 12 130-155 T 3 12 3-4 Hf c/u 22 12 21-23 E 24 12 23-26 A single male measured Hb 185; T 5: Hf c/u 23; E 28. Distribution Restricted in their occurrence to the southwestern parts of Angola and the northwestern parts of Namibia. Not occurring in the western desert areas of the country, except possibly where there is riverine forest, or rivers running through them. Not recorded east of 17°E. Habitat Little is on record regarding the habitat requirements of this species, but as they are a fruit-eating species, they are probably confined to areas of riverine and other types of evergreen forest where there are fruit-bearing trees. Habits Shortridge (1934) recorded that in Ovamboland they hung up singly from the bare branches of large Acacia sp trees near the Cunene River. Apart from this brief observation nothing seems to have appeared in literature concerning their habits. Food Like other species of the genus Epomophorus they probably live predominantly on wild fruits. Reproduction Shortridge (1934) recorded that newly-born young were found clinging to their mothers in September and October in Ovamboland, Namibia. No. 42 Epomophorus gambianus Gray, 1870 Gambian epauletted fruit bat Gambiaanse witkolvrugtevlermuis Colloquial Name So named as the species was described from a specimen from Gambia in West Africa. Taxonomic Notes Hayman & Hill (1971) listed two subspecies, E. g. gambianus from the northern parts of the species' range which is replaced in the southern parts by E. g. parvus described by Ansell (1960c) from Zambia; the limits of the two subspecies are not known. E. g. parvus has been described as being smaller than E. g. gambianus (Ansell, 1960c). Later Ansell (1978) thought that they might be conspecific with E. crypturus, the difference being in the proportion of the zygomatic breadth to the total length of the skulls in the males. In E. g. parvus it is less than half the total length of the skull, whereas in E. crypturus it is more than half. The matter remains unsolved and in the meantime Smithers (1983) followed Hayman & Hill (1971) in 58 riverine forest of the Zambezi River, the other in much drier woodland associated with a tributary of that river. Rosevear (1965) noted that they appeared to be indifferent to their vegetational surroundings, possibly because they are both fruit-eaters and nectar-suckers. Their occurrence in drier woodland, like other fruit bats, will be governed by the availability of food. Habits From the observations made in Zimbabwe by Thomas & Fenton (1978) they appear to be solitary in habit. In West Africa, Rosevear (1965) recorded that they occurred in colonies of 12 to 20, roosting during the day in clumps of bamboo and in the canopy of evergreen trees. In Zimbabwe, Thomas & Fenton (1978) noted that this species roosted in Natal mahogany, Trichilia emetica and sausage trees, Kigelia africana. One individual used five different trees in six days, another used a Natal mahogany on two consecutive days and then moved 150 m and 400 m between two sausage trees and a Natal mahogany. In Sierra Leone, Rosevear (1965) recorded that individuals in colonies hung singly, not close together like other members of the genus. They appear to be early movers in the evening, for they have been seen feeding as daylight fades. Food In Zimbabwe, Thomas & Fenton (1978) by telemetry recorded that the species began foraging less than 45 min. after dark. Individual bats visited the same Diospyros senensis shrub on two successive nights, spending between 90 min. and 105 min. in feeding on the fruits, with breaks of 10 min. and 15 min. while they rested in adjacent Natal mahogany trees, Trichilia emetica. They appeared to feed solitarily and to have access to this food resource for almost two hours before the arrival of the Egyptian fruit bat, Rousettus aegyptiacus. In West Africa, Rosevear (1965) recorded that in addition to eating wild fruits, they lapped nectar from flowers and may possibly eat the anthers and fleshy petals of some species. They will eat the fruit of wild figs, Ficus spp, and have been observed to eat bananas. In lapping nectar they visit each flower in turn for a quarter to three quarters of a minute, holding on by their feet and lapping the nectar with their long tongues. He recorded a specimen in the British Museum (Nat. Hist.) that was taken on the flower of a baobab, Adansonia digitata and they may also sip nectar from the flowers of the sausage tree, Kigelia recognising E. g. parvus as a valid subspecies, although Meester et al. (1986) do not agree and place it as a subspecies of E. wahlbergi. Description The Gambian epauletted fruit bat has a head and body of about 160 mm and a wingspan about 560 mm. The colour of the upper parts varies considerably from a sepia to a yellowish-cream, the under parts from off-white to about the same colour as the upper parts. The long, soft fur extends on to the upper and under parts of the upper arm and half of the forearm, and on to the upper parts of the legs, but only on to the proximal half of the legs underneath. It also extends sparsely on to the wing membranes, where on the upper surface it is similar in colour to the upper parts of the body, white on the under surface. They have the epaulettes on their shoulders like other members of the genus (see E. crypturus). The dark brown wings are short and broad with rounded tips. Most individuals have no noticeable tail, in others it may reach a length of about 5 mm. The muzzle, particularly in the male, is long and tapering, the nostrils enclosed in soft pads. They have two ridges on the palate behind the last molar teeth, the first subtriangular in shape, the second of normal form, divided into two sections one on either side of the midline (Fig. 41.1.a). The posterior portion of the palate is deeply hollowed out which is the main distinction between this genus and Epomops, in which it is shallow or flattish. a b Fig. 41.1. Palatal ridges: (a) Epomophorus gambianus (b) E. angolensis. Distribution South of the Sahara, excluding the Southern African Subreg ion They occur from Senegal and Gambia in West Africa eastwards in the High Forest and Guinea Savanna Zones to the Sudan and Ethiopia and southwards in northwestern Uganda and Zaire. They have been recorded in Zambia near the border with Angola, but not in the eastern sector, and no doubt in time will be shown to occur within Angolan limits in the southeast. In the extreme northeast of the country they are recorded in adjacent parts of Zaire. Southern African Subregion There are only two records of the occurrence of this species in the Subregion, both from Zimbabwe where a single specimen was taken in the riverine forest of the Zambezi River some 60 km up river from the Victoria Falls and another from the Sengwa Research Station in the northwest of the country (18 28 Aa). Habitat In the northern and western parts of their range Gambian epauletted fruit bats occur both in open savanna woodland and in forest (Rosevear, 1965). One of the two specimens taken in the Subregion was taken in the well developed 59 africana. Both these trees occur in the area in the Subregion from which they have been taken. Reproduction No information available. No. 43 Epomophorus crypturus Peters, 1852 Peters' epauletted fruit bat Peters se witkolvrugtevlermuis Colloquial Name Named after W.C.H. Peters who in 1852 published a monumental work on zoological material collected mainly in the Tete District of Mozambique. In the course of his career he was the first to describe eight new species of bats, six rodents, the porcupine, the bushy-tailed mongoose and Lichtenstein's hartebeest. Taxonomic Notes No subspecies are recognised. Description Peters' epauletted fruit bats are a common species in parts of the Subregion. The males are considerably larger than the females, adult males measuring about 150 mm in total length with mean masses of about 105 g and maximum masses of up to 140 g. The females measure about 120 mm in total length and have mean masses of about 76 g (Table 43.1). The colour of the body is very variable and although usually a brownish-buff on the upper parts, it may be pale buffy and, in the extreme, nearly white. Both sexes have white patches at the base of the funnel-shaped ears, but only the males have the epaulettes on the shoulders. These are sunken glandular pouches in the skin covered with long white hairs. When under stress, when vocalising or possibly under sexual stimulus, these pouches are everted, the white hair, which in this species is 9 mm long, forms the conspicuous white epaulettes. They share this feature with the other three species of Epomophorus, E. angolensis, E. gambianus and E. wahlbergi that occur in the Subregion. The under parts are lighter in colour than the upper parts and in adult males the lower part of the throat has a collar of russet brown which is less obvious in juveniles and in the females. They have two ridges on the palate behind the last molar teeth (Fig. 40.1.b). Table 43.1 Measurements (mm) and mass (g) of Peters' epauletted fruit bats, E. crypturus, from the northern parts of the Subregion (Smithers & Wilson, 1979) Males Females X n Range X n Range TL 149 12 130-170 122 12 110-125 Hf c/u 22 12 20-23 21 12 20-22 E 24 12 22-27 24 12 22-25 F/a 83 12 80-86 79 12 76-82 Mass 104,5 12 80-140 75,5 12 64-88 Distribution Confined to the southern part of the continent, being replaced northwards by the eastern epauletted fruit bat, E. anurus, which extends from the northern parts of the Rift Valley in eastern Zaire and Rwanda into Kenya, Tanzania, the southern Sudan and Ethiopia and westwards to Nigeria. South of the Sahara, excluding the Southern African Subregion They occur in southeastern Zaire and widely in Zambia and Malawi, while at present there are no records from northeastern Mozambique, east of Lake Malawi. They have been taken in the Tete District, north of the Zambezi River. While there are no records from southeastern Angola, they occur on the Zambian border and in the valley of the Okavango River in Botswana right to the Angolan border and no doubt occur in Angola, if only marginally. Southern African Subregion While there are no records from Namibia, as they occur in the valley of the Okavango River in Botswana, they may well be found to occur along the river into that country. In Botswana they are common throughout the valley of the Okavango River and in its delta south to Maun and on the Chobe River in the northeast. In Zimbabwe they occur throughout except in parts of the dry west and in the central parts of Mozambique, south of the Zambezi River. They are found along the borders of the Transvaal south to the Maputo District. In the Transvaal they occur in the north in the Limpopo River valley and in the east as far south as Gollela. This indicates a possible occurrence in Swaziland from which at the moment there are no records. Southwards there are isolated records from Natal and the eastern Cape Province. Habitat Peters' epauletted fruit bats are governed in their occurrence by the availability of fruit-bearing trees. They are associated predominantly with evergreen forests in the higher rainfall areas but occur in evergreen riverine forests deep into otherwise dry and unsuitable terrain. Their requirements are nowhere better exemplified than in northern Botswana where they are common throughout the Okavango Delta, with its rich riverine vegetation. They are not found in the drier associations of mopane, Colophospermum mopane, Acacia spp or Terminalia spp that surround it and which offer insufficient cover and food to satisfy their needs. Where there are isolated areas of slightly higher rainfall within otherwise drier country, such as in the Masvingo or Matopos Hills areas in Zimbabwe, where the run-off from the granite bosses supports an evergreen vegetation around their bases, they settle in these moister woodlands and move 60 into the dry surrounding country to preferred feeding sites. In parts man has provided additional suitable conditions for them in orchard development where fruits such as guavas, plums, mangoes and other soft fleshy fruits provide them with food and surrounding exotic trees such as Cyperus sp the deep foliage shelter which they require in which to roost. They are independent of surface water, obtaining all their moisture requirements from their food. Habits Peters' epauletted fruit bats are gregarious, occurring in colonies numbering up to hundreds of individuals. During the day they hang up by their feet, slightly spaced out one from another, on the thinner branches of evergreen trees where the thick foliage provides surrounding cover. In Zimbabwe they commonly use trees such as the wild fig, Ficus spp or the sausage tree, Kigelia africana, but they will use any evergreen tree provided it has thick foliage to provide cover and thin enough twigs to allow them to cling on to them. In Botswana they were found using the outer finer twigs of a clump of high growing bamboo in a garden fringing the Okavango River, strung out on these in such numbers that the poles were weighed down to within 3 m of the ground. The colonies are very noisy, the males vocalising with a repeated musical bark which is uttered predominantly when hanging, but also occasionally when on the wing. As they settle after a night's foraging, there is much bickering among members of the colony. The clawed first digit on the forelimb and wings are used to slash other members who attempt to hang up too close and it takes a considerable time for the colony to settle quietly for the day. Food The food of Peters' epauletted fruit bats consists almost entirely of wild or cultivated fruits which are soft and pulpy. The fruit of wild figs, Ficus sp; marula, Sclerocarya hirrea; mabola plum, Parinari curatellifolia; kudu berry, Pseudolachnostylis maprouneifolia; mahobohobo, Uapaca kirkiana; bird plum, Berchemia discolor; red milkwood, Mimusops zeyheri, and quinine tree, Rauvolfia caffra, are sought after. In feeding on small fruits they maintain their position on the outside of the canopy by wing flapping, pluck the fruit with their teeth and fly with it to a nearby feeding site. These sites are open branches of a size convenient for them to hang by the back feet. They fly to these carrying the fruit in their mouths, where it is manipulated by the claws on the first and second digits of the wings as they eat. In Zimbabwe the same bat would use the same feeding site over a period of a few nights and may well do so until the fopd source is depleted. They are wasteful feeders and when eating fruits such as mobola plums, the ground under the site gets covered in the discarded hard skins and pips and small bundles of chewed pulp as if, in eating the rather stringy pulp of this fruit, they are utilising the juices only. With very soft fruits such as figs they may cling to the fruit cluster or twigs and eat in situ or if the fruit becomes loosened in the process it is held in the mouth and carried to a feeding site. They are great raiders of orchard or garden fruits such as guavas, apricots, peaches and loquats and if pawpaws are left to ripen on the tree, they damage these by biting into them and scratching them as they cling. Apples, pears and other hard fruits remain untouched. Reproduction In a sample of 155 females taken throughout the year in Zimbabwe gravid females were observed from late July through to March. There was in this series a pronounced peak about August and from observations of juveniles, the main bulk of the young are born about September (Smithers & Wilson, 1979). In the Transvaal, Rautenbach (1982) observed lactating females with their young during November. A single young is produced at a birth. The young are born with their eyes closed, with a sparse coat of hair on the back and naked under parts. In the early stages of their lives they fix firmly on to one of the female's nipples, clinging to her with their clawed digits of the forearms and the claws on the feet and are carried by her while she is feeding. Later as they grow and become too large to be conveniently carried, she will leave them hanging in the roost. Twins are known. Genus Epomops Gray, 1870 Only one species of this genus occurs in the Subregion. No. 44 Epomops dobsonii (Bocage, 1889) Dobson's fruit bat Dobson se vrugtevlermuis Colloquial Name Named after G.G. Dobson who among his many other papers on bats published a Catalogue of the Chiroptera in the collection of the British Museum in 1878. Taxonomic Notes Hayman & Hill (1971) stated that the relationship of this species with others that occur in West Africa is not clear and the possibility of intergrading must be taken into account. The palatal ridge pattern is apparently not as fixed a character as was formerly supposed. In the meantime until further information becomes available it is considered as a valid species. Description Adult males have a head and body length of about 160 mm and forearms of 86 mm. The females are slightly smaller than the males (Table 44.1). The upper parts are greyish-brown, the under parts a drab cinnamon colour washed with grey. The males have dark greyish-brown throats, in the females the throats are greyish. The wings, ears and rhinarium are dark brown. The muzzle is elongate and broad, the sides nearly parallel. It is not, however, as elongated as in Epomophorus spp. They have three cheekteeth on either side of the upper jaw and five in the lower, the palate strongly arched from back to front and side to side. The post dental palate in this species has converging margins and is flat with five to seven palatal ridges posterior to the last tooth. In Epomophorus spp the post dental palate is almost parallel sided and is concave posteriorly and they have only up to two ridges posterior to the teeth. The hair that covers the glandular region on the shoulders of the males and forms the epaulettes is much longer than in E. crypturus, measuring up to 18 mm as against 9 mm in E. crypturus. It is in addition tinged yellow as against the pure white hair in E. crypturus, and when extended forms a much larger epaulette. Table 44.1 Measurements (mm) of male Dobson's fruit bats, E. dobsonii, from Zambia (Harrison, 1959) and females from Angola (Hill & Carter, 1941) Males Females X n Range X n Range HB 160 4 146-168 142 3 137-145 E 28 3 27-28 26 3 25-27 F/a 86 5 82-88,2 83 3 81,5-83,0 Distribution South of the Sahara, excluding the Southern African Subregion Occurs widely in central Angola and Zambia eastwards to the borders of Malawi from which country there are no 61 records at the moment, although they may occur there. Recorded from Katanga in southeastern Zaire, and from Rwanda and Tanzania. Southern African Subregion Represented from the Subregion by a solitary specimen from Kasane on the Chobe River in the extreme northeast of Botswana. Habitat Practically nothing is known about the habitat requirements of this species. In Angola they occur in savanna woodland but as they are fruit-eaters, this association must include fruit-bearing trees. The single record from the Subregion was taken in a colony of Peters' epauletted fruit bats, Epomophorus crypturus, in the riparian evergreen woodland of the Chobe River and certainly they will be associated in other parts of their distributional range with similar conditions. Habits Judging from the number of specimens in collections, E. dobsonii is nowhere so numerous as other fruit-eating bats such as E. crypturus or E. helvum. Food Apart from the fact that they eat fruit like other members of the Family, there is nothing specific on record. Reproduction Unknown. Genus Eidolon Rafinesque, 1815 No. 45 Eidolon helvum (Kerr, 1792) Straw-coloured fruit bat Geel vrugtevlermuis Colloquial Name They are not entirely straw-coloured, only on the shoulders and back, the rump and hind limbs normally being various shades of brown. Taxonomic Notes Hayman & Hill (1971) listed three subspecies, one from Madagascar, E. h. dupreaneanum Schlegel & Pollen, 1866; one from the African Continent, E. h. helvum, and one from Arabia, E. h. sabaeum Andersen, 1907. There have been differing opinions advanced, some authorities (Eisentraut, 1964) suggesting that no subspecies should be recognised; Andersen (1912) on the other hand believed that the Madagascar and Arabian forms should be recognised as distinct species. Description Straw-coloured fruit bats are by far the largest species of bats found in the Subregion. While it is difficult to judge the sexes in the field, there is a greater difference in size between them in this species than in the other fruit bats. Jones (1972) in West Africa recorded that the forearm in the males averages about 13% longer than in the females. The small number measured in the Subregion shows an average of 4% greater length in the males. They have a total length of about 190 mm with forearms in the males of about 116 mm, in females 113 mm and a mass of between 243 g and 280 g (Table 45.1). Although the colloquial name suggests that they are an overall pale golden-yellow colour, there is a considerable variation in colour between individuals and between one area of the body and another. On the upper parts the fur covers the head and shoulders and lies in a band down the mid-back extending on to the upper parts of the forearms, legs and narrow interfemoral membrane, but not on to the wing membranes. In some specimens there is a distinct yellow or orange collar on the throat which extends upwards on to the back of the neck but the remaining fur on the upper parts is either various shades of grey or brown, edged at the back of the shoulders and on the mid-back by a yellowish fringe. The fur on the rump and legs is often darker in shade than on the shoulders. The under parts are lighter in colour, in some the collar of slightly longer hair on the throat contrasts with the remainder, which is tinged paler yellow or orange. The skin underlying the collar on the throat is glandular and has been observed to secrete a sticky fluid with a musky smell (Allen, Lang & Chapin, 1917). Table 45.1 Measurements (mm) and mass (g) of straw-coloured fruit bats, E. helvum, from Zimbabwe (Smithers & Wilson, 1979) Males Females X n Range X n Range TL 190 5 165-205 189 6 181-207 T 15 5 14-18 15 6 11-20 Hf c/u 30 5 24-34 34 6 30-39 E 28 5 26-29 28 6 26-32 F/a 116 5 110-122 113 6 102-118 Mass 2 only 247,3 and 243,0 2 only 267,7 and 280,5 The naked wing membranes are dark blackish-brown, the wings long and pointed. Large specimens may have a wing span of up to 0,75 m. When resting the ends of the wings are folded back. The interfemoral membrane runs narrowly up the insides of the thighs and is only a few millimetres wide where it joins across the body. The tail which is about 15 mm long projects beyond this membrane for about half its length (Fig. 45.1). The first finger on the forearm is long and has a powerful curved claw which is used in clambering around in tree branches; the claw on the second digit is less well developed (Fig. V.l). The dental formula is: If C± Pf Mf = 34 Distribution As is shown under Habitat the distribution of this species has to be dealt with in a different manner from other species 62 Fig. 45.1. Tail and interfemoral membrane of Eidolon helvum. which are resident in the Subregion. The straw-coloured fruit bat is a migrant from its focus of distribution which lies to the north of the Subregion in the tropical forests of the continent and they are only seen on migration and not as residents. This means in effect that they are prone to turn up almost anywhere in the Subregion from time to time. At the moment most records are from the eastern and southern parts of the Subregion, for these areas with their higher rainfall than in the west, provide a better supply of wild fruits, their principal food. Nevertheless they have been recorded from the Subregion's most arid terrain, the Namib Desert in Namibia. In Zaire, Allen, Lang & Chapin (1917) observed that immense numbers journey about irregularly and then become abundant in regions from which they were previously absent. Even the vast colony at Kampala, Uganda, moves out during the months of June to September (Kingdon, 1974). They are recorded from sea level to altitudes of 2 000 in East Africa. South of the Sahara, excluding the Southern African Subregion Straw-coloured fruit bats occur in forests from Guinea in West Africa eastwards to Nigeria and throughout the tropical forests of Cameroun; Gabon; Zaire to the Rift Valley and parts of the Sudan; Uganda; Kenya; Tanzania and northern Angola. While it is impossible at the moment to clearly define the southern limits of the area in which they may be considered to be resident on a year round basis, this may well include parts of northeastern Zambia, Malawi and possibly northeastern Mozambique. South of this they are considered as occurring on migration, only remaining in areas for a shorter or longer period depending on the local availability of food. Southern African Subregion Of sporadic occurrence in Namibia; in Zimbabwe, as far west as the Matopo Hills, and in Mozambique, south of the Zambezi River, in the central and southern parts of the country. In the Transvaal they have been taken as solitary wanderers in the southwest of the province. They appear commonly in northeastern Natal and have been taken in the Orange Free State and the eastern Cape Province, with records from as far west, coastally, as the Bredasdorp district. Habitat Straw-coloured fruit bats occur in the Subregion in a wide variety of habitats from coastal forests in Natal and Mozambique to parts of the dry South West Arid Zone in Namibia. In the Subregion we only see them while they axe on migration, when they tend to scatter widely, not being truly resident and only remaining in an area if held there temporarily by the availability of a plentiful food supply. Their prime habitat is in the tropical forests of the continent which supply them with a variety of wild pulpy fruits during the greater part of the year, and it is the availability of food that determines their occurrence more than any other factors. Their prime habitat covers a very extensive area ranging from Guinea in West Africa eastwards to the Rift Valley in East Africa, a distance of some 5 000 km which is up to 650 km wide in parts. In their search for food resources they cover great distances, crossing country that is totally unsuitable for them and in which, were they to settle, they would perish. At times this movement is ill directed, for Rosevear (1965) recorded that they have been taken at sea some 250 km from the nearest land. He stated that in West Africa they have been found in the Sudan Savanna Zone which borders on the southern fringes of the Sahara, so that their sudden appearance in some of the more arid parts of the Subregion is not surprising. Habits Straw-coloured fruit bats are gregarious and subject to wide migration and movements in search of food supplies. The classic colony at Kampala, which roosts during the day in a grove of saligna gums, Eucalyptus saligna, was estimated by Ogilvie & Ogilvie (1964) to number some 200 000 individuals. They return to the grove about 07h00 in the morning and leave to forage about 18h30 in the evening. They hang in the grove in clumps of 1-50, chattering continuously, with some individuals always flying around in search of new places to hang. Fighting is common, contestants battering each other with their wings. In the Subregion they never occur in such huge numbers and colonies of up to 20 or 30 are exceptional. More often up to half a dozen is the rule and often single individuals are seen. Even in the Kampala colony, which was known prior to European settlement, they are subject to movement, for Kingdon (1974) recorded that they are absent from June to September, as they are too from other large colonies such as those at Avakupi in Zaire (Allen, Lang & Chapin 1917). In the forest habitat in West Africa, Rosevear (1965) noted that they tend to prefer clumps of very high trees to roost in, relic patches of ancient forest being preferred. In the Subregion they have a habit of hanging on power lines and pylons which accounts for the number of individuals that get electrocuted annually in Zimbabwe, where no less than seven of the 12 specimens in the collection of the National Museums were recovered after death by electrocution. Their huge wingspan of 0,75 m renders them prone to death in this manner when alighting to rest (Smithers & Wilson, 1979). Their wings are long and pointed, adapted to a rather slow but steady flight, interspersed with short periods of gliding which carry them over great distances. The powerful hooked claw of the thumb is used in clambering around branches 63 and in clinging on to trees while feeding. Rosevear (1965) described the evening flight of the colony as a remarkable sight. Most of them take off as if by a common signal, the flying colony when seen from a distance looks like a black cloud over the forest. Jones (1972) stated that, in roosting in trees, they tend to use the more sturdy branches, hanging with their wings half folded away from their bodies or along the sides of their bodies with the wing tips folded back. They remain alert and active during the day, hanging with their eyes open, ears erect and constantly moving. In addition to the daytime roosts, they rest from time to time during the night between bouts of feeding, hanging in trees. On taking to the wing they drop for distances up to 2 m from the roosts before flying. Food Digestion in the straw-coloured fruit bat is extremely rapid (Rodhain & Bequaert, 1916), which is probably the reason why stomachs examined from Zimbabwe and Mozambique were invariably empty. Allen, Lang & Chapin (1917) found the same state of affairs in the specimens they examined in Zaire. We have to rely on the observations of Rosevear (1965) in West Africa who reported on their feeding habits.. There they are a pest in banana plantations, a food which captive individuals eat with avidity. They eat the fibrous pulp of the fruits of the Borasses palm, Borassus sp, dates, and wild figs, Ficus spp. They will take large succulent flowers or flower buds such as those of the baobab, Adansonia digitota, and the silk cotton tree, Bombax sp. He recorded that they were reported to eat the fresh leaves of Erythrina sp, completely stripping the trees. Jones (1972) in West Africa reported that they eat mangoes, pawpaws, avocado pears, figs, passion fruit, custard apples and loquats. In feeding they hang by the claws on their thumbs and use both feet. They chew noisily, discarding fibrous material. In Zaire and West Africa the bats are eaten by the indigenous people, Allen, Lang & Chapin (1917) recording that at Leopoldville they saw bunches of them tied together being sold in the markets. Reproduction There are no indications at the moment that straw-coloured fruit bats breed in the Subregion. No specimens examined were gravid and it may be that they only breed within the limits of better developed and more extensive forested regions to the north. Allen, Lang & Chapin (1917) recorded that at Avakubi, in Zaire, from the presence of gravid females, they appeared to have a definite breeding season, the young being born about November. Rosevear (1965) stated that the young are carried about by their mothers until the stage when they are able to take care of themselves. The females usually have one young, occasionally two. Genus Rousettus Gray, 1821 This genus is represented on the continent by three species, two of which, R. aegyptiacus and R. angolensis occur in the Subregion. Members have a wide distribution: R. aegyptiacus is found from the Cape Province to Egypt and westwards to Senegal, extralimitally on the Mediterranean island of Cyprus, in parts of the Middle East and eastwards to Pakistan. The generic name Rousettus was applied to the group as the earliest known representatives were red in colour; the two species that occur in the Subregion, however, are brown or dark brown on the upper parts of the body. They are medium-sized bats with wingspans up to about 450 mm. The wing membranes are dark in colour, almost black. They have very short tails and narrow interfemoral membranes. In both species the eyes are large, as in other fruit-eating bats. The Egyptian fruit bat, R. aegyptiacus, has powers of echolocation which are brought into play in total darkness. This allows them to roost in caves that are totally dark, a shelter denied to other fruit-eating bats. Bocage's fruit bat, R. angolensis, on the other hand has no powers of echolocation and when using caves can only use parts that are adequately illuminated. The dental formula is: If Cj P| M§ = 34 and is the same as in the straw-coloured fruit bat, Eidolon helvum. The anterior upper premolar is very small and roughly equal in size to the upper incisor, whereas in E. helvum it is appreciably larger. The two upper and three lower molars and last premolar in R. aegyptiacus are distinctly longer than broad, whereas in R. angolensis they are only as long as broad. Key to the species after Hayman & Hill (1971) (Measurements in mm) 1. Wing membranes arise from the first toe; hair short and sleek; cheekteeth heavy; length of forearm generally above 89 and up to 96; adult males with no collar of bristly hairs on the throat .. . aegyptiacus Wing membranes arise from the second toe; hair longer and coarser; cheekteeth reduced in bulk; length of forearm less than 90; adult males with a distinct collar of bristly hairs on the throat . . . angolensis No. 46 Rousettus aegyptiacus (E. Geoffroy, i8io) Egyptian fruit bat Egiptiese vrugtevlermuis Plate 4 Colloquial Name The name derives from the fact that the species was described originally from a specimen from the Great Pyramid of Giza in Egypt. Taxonomic Notes Hayman & Hill (1971) listed four subspecies, only one of which, R. a. leachi (A. Smith, 1829), occurs in the Subregion. This was described from a specimen from "gardens about Cape Town". Description Egyptian fruit bats are the second largest species of fruit bat that occurs in the Subregion, only being exceeded in size by the straw-coloured bat, Eidolon helvum. Adults are about 150 mm in total length with masses of about 130 g. There are little or no differences between the sexes (Table 46.1). They have a wingspan of about 0,6 m. Their colour varies throughout their wide range, those from the Subregion are dark brown on the upper parts, sometimes tinged with slate-grey, the under parts smokegrey. The fur extends along the top of the forearm for about half of its length and along the top of the legs to the ankles. Underneath it extends half way along the forearm and only on to the upper parts of the legs. The rounded ears are naked except at their bases, the fur on the face is short and longer on the forehead and the upper parts of the body. The fur extends narrowly on to the dark brown wing membranes. The eyes are noticeably large. They have a distinct pale buffy or yellowish collar around the throat and neck which contrasts markedly on the upper parts with the general dark brown colour. The wings are rounded at the tips. 64 The dental formula is: IfQPfMf = 34 Table 46.1 Measurements (mm) and mass (g) of Egyptian fruit bats, R. aegyptiacus, from the northeastern parts of the Subregion (Smithers & Wilson, 1979) Males Females X n Range X n Range TL 150 17 140-162 150 19 140-162 T 18 14 15-22 19 17 16-24 Hf c/u 23 16 20-27 24 18 22-26 E 22 17 20-25 22 18 19-24 F/a 89 17 83-96 90 19 88-95 Mass 133,4 11 88,1-166,0 129,3 18 117,7-166, Distribution The Egyptian fruit bat has a very wide distribution from the Cape Province to Egypt and westwards to Senegal and beyond Continental limits to the countries at the eastern end of the Mediterranean, to Arabia, southeastern Iran, Baluchistan and west Pakistan. They do not occur in the Sahara or in the northwestern parts of the continent or in the southwestern sector except coastally. Northeastern Africa They occur in Egypt in association with the Nile Valley right to the coast of the Mediterranean. South of the Sahara, excluding the Southern African Subregion They occur from Senegal and are recorded from most of the countries eastwards to the Sudan and Ethiopia, in the High Forest Zone, Guinea savanna and in parts of the Sudan savanna. Southwards they are recorded widely in most countries to the borders of the Southern African Subregion. In Angola they have been taken in the central and northern parts of the country with the situation in the south not being known. In Zambia, although there are only a few records from the southeast, Ansell (1978) believed that eventually they would be shown to occur throughout the country. At the moment there are no records from Malawi or from northeastern Mozambique, but there is good reason to believe that they will be found there eventually. Southern African Subregion They appear to have an eastern distribution in the Subregion, not being recorded from Namibia, Botswana or the more arid parts of the Cape Province. They occur in the central and eastern parts of Zimbabwe, and in the central and southeastern parts of Mozambique, south of the Zambezi River. In the latter area they are absent from the more arid parts of the Banhine Flats in the west. In the Transvaal they are recorded from Pafuri in the northern parts of the Kruger National Park and from the Letaba and Barberton Districts. They occur in Natal and in the Cape Province narrowly westwards along the southern coast to Cape Town. Habitat Although from their distribution Egyptian fruit bats appear to be catholic in their habitat requirements, two factors at least are essential for them. As a fruit-eater the habitat must provide this and at the same time caves or similar structures must be available as cover in which to roost during the day. In Botswana and Namibia the semi-desert areas, while they have suitable caves, do not provide the necessary food, and in areas such as the Okavango Delta in Botswana where fruit is plentiful, the nature of the terrain is such that suitable caves are almost non-existent. So far they have not been recorded from the Drodtsky Caves in western Ngamiland which are certainly within reach of the food supply in the Okavango Delta and which are extensive enough to accommodate them. Where cover and a plentiful food supply occur together they are found in very large numbers. Habits Egyptian fruit bats are gregarious, occurring in colonies numbering up to thousands of individuals in caves or other similar structures. They hang up from the ceilings or the walls, usually by one foot, the wings closely folded around the body, and the other foot folded across the front of the body outside the enfolding wings. They pack tightly together in clusters, usually deep in the caves in total darkness. The colony is very restless and extremely noisy, chirping and chattering continuously, a noise which can be heard long before the cave entrance is approached. They avoid light, this sometimes resulting in continuous fighting for the darkest niches and crevices in a cave. Indeed, availability of dark places may act as a rough regulator of numbers in a cave (Herzig-Straschil & Robinson, 1978). As sundown approaches the restlessness and noise become more apparent as individuals start to groom themselves prior to the night's foraging. In the Tsitsikama National Park activity data indicated a different winter and summer pattern. In winter they became active on average 90 minutes after sunset, and activity ceased about 216 minutes before sunrise. In summer the period of activity was appreciably longer regardless of the shortened dark phase, starting 30 minutes after sunset and ending 12 minutes before sunrise. The delayed onset of emergence under the influence of moonlight in winter was quite pronounced (HerzigStraschil & Robinson, 1978). This latter was also suspected in the eastern Transvaal (Jacobsen & du Plessis, 1976). Rousette bats are the only species of fruit bat that have powers of echolocation, which allow them to move in total darkness. The system used is different from that found in Microchiroptera which emit high frequency sound from the mouth in the case of the Vespertilionidae, or from the complicated nasal structures in the Hipposideridae. In the case of the rousette bats Mohres & Kulzer (1956a) found that the Egyptian fruit bat was not inconvenienced when its eyes were covered and it was able to fly around in complete darkness, avoiding obstacles with ease. If, however, its ears were obstructed, it was helpless in complete darkness. The system of echolocation used is by repetitive tongue "clicks", those of lower frequency audible to the human ear, the echo of these being picked up by the ears which allows them to pinpoint obstacles ahead of them. The clicks are emitted through the corners of the mouth with the jaws closed. 65 The Egyptian fruit bats have a well developed eyesight and when returning to their caves, providing there is sufficient light, they will locate these by sight. As they enter they will switch to echolocation to find their way around and eventually to pinpoint the location of suitable hanging places. They therefore are fully equipped to find their way around either by sight or echolocation. Egyptian fruit bats are agile, fast fliers and can negotiate narrow openings between the branches of trees with ease. No. 47 Rousettus angolensis (Bocage, 1898) Bocage's fruit bat Bocage se vrugtevlermuis Food In feeding they will circle a fruiting tree, then hover before alighting and snuffling about on the branches to discover suitable fruit (Rosevear, 1965). Once secured, it may either be eaten in situ or conveyed in the mouth to a feeding site. The fruit is manipulated in one of the back feet, the other clinging to the branch while the food is eaten, or the food may be held against the chest by the back feet while it is eaten. The hard skin and seeds of fruit are discarded, the pulp chewed and any fibrous material dropped. Under favourite feeding sites quite large accumulations of discarded material may be found. Herzig-Straschil & Robinson (1978), in the eastern Cape, observed several individuals at a Cape ash tree (Ekebergia capensis) bearing ripe fruit in June and these bats also ate fruit from saffronwood (Cassine crocea), Bushman's poison (Acokanthera oppositifolia) and mistletoe (Viscum obscurum). In the Eastern Transvaal figs formed the basis of the bats' diet (Jacobsen & du Plessis, 1976), in particular the Cape fig (Ficus capensis), while F. petersii and F. sansibarica were also eaten at certain times of the year; other fruits utilised were from the Cape ash tree, water berry (Syzigium cordatum), red stinkwood (Prunus africana) and wild plum (Harpephyllum caffrum). In Zimbabwe, Thomas & Fenton (1978) recorded their eating the fruits of Diospyros senensis. Soft garden and orchard fruits are also eaten. Egyptian fruit bats have cheek pouches in which food can be stored temporarily. Lombard (1961) reported that, after eating, the hair on the chest which tends to become matted with the fruit juices, is thoroughly combed with the claws on the feet and the toes cleaned in the mouth. They also pick out fibrous material from between the teeth with the claws on the feet. Reproduction During the mating season, which in the Cape Province is from June to mid-September, Lombard (1961) noted that the males tended to form groups, ignoring the females and juveniles, which formed nursery groups on their own. Using Mutere's (1968) forearm age relationship it was estimated that birth in the Tsitsikama National Park occurs from October to June (Herzig-Straschil & Robinson, 1978), whereas in the eastern Transvaal the period was slightly shorter, with a peak in mid-summer (Jacobsen & du Plessis, 1976) and further north parturition is given as from September to December (Penzhorn & Rautenbach, 1988). A single young is produced at birth, occasionally twins, after a gestation period given by Kingdon (1974) as 105 to 107 days. In East Africa they breed twice a year and in Egypt throughout the year (Kingdon, 1974). The young are born with their eyes closed and their ears folded back, the eyes opening and ears erecting at about 10 days of age. At birth they cling tightly to the ventral surface of their mothers and are carried around by them for about six weeks during the suckling period. The mother then leaves the young bat in the cave while she forages, bringing back food for it. At first it licks the juices from her mouth and later eats mouthfuls for itself (Lombard, 1961). At this time of their lives the young bats usually lose weight and do not start to gain weight until they start to fly at about nine to 10 weeks of age. They recognise their young by smell as was shown in Kulzer's (1958) experiment when a female's own and a strange young were enclosed in bags. The mother correctly identified her own offspring as was shown by her frenzied attempts to release it. Colloquial Name Named after J.V. Barboza du Bocage who originally described the species Cynonycteris angolensis (=R. angolensis) from Angola. Taxonomic Notes Hayman & Hill (1971) listed three subspecies from the continent, only one of which, R. a. angolensis, occurs in the Subregion and then only marginally into Zimbabwe. Some doubt has been expressed in the past as to whether this species should be retained in the genus Rousettus or whether it deserves full recognition as a genus, Lissonycteris, on its own. It differs from R. aegyptiacus and other rousette bats in the possession by the males of a distinct neck ruff of coarse sticky hairs and the fact that it does not possess powers of echolocation. In addition it has a long, silky fur and the wing membranes arise from the second toe of the hind foot and not from the outside toe as in R. aegyptiacus; the cheekteeth are squarish, not rectangular, and the premaxilla is ossified across the front. As Meester et al. (1986) included it in the genus Rousettus, this view is followed. Description Slightly smaller than the Egyptian fruit bat, R. aegyptiacus, Bocage's, R. a. angolensis, has a head and body length of about 120 mm and a wingspan of about 400 mm. The relative size is perhaps best illustrated in the length of the forearm which in the Egyptian fruit bat measures 90-105 mm and in Bocage's fruit bat from the Subregion 81-85 mm (Table 47.1). The West African subspecies, R. a. smithi Thomas, 1908, is smaller still with a forearm length of 72-82 mm (Rosevear, 1965). The characteristic feature of male Bocage's fruit bats is the collar of stiff coarse hairs on the throat and sides of the neck, which forms whorls below the ears. It is dusky-orange in colour and is sticky, suggesting that it arises from a glandular area of skin. The colour of the upper parts in specimens from the Subregion is a rich brown, with the under parts lighter in shade than the upper, the collar in the males orange. The throat and sides of the neck in the females are sparsely haired. Table 47.1 Forearm and skull measurements (mm) of a series of four male and three female specimens of Bocage's fruit bat, R. angolensis, from Zimbabwe (Smithers & Wilson, 1979) F/a TL skull 83 43 Males Range 81-84 42-44 84 43 Females n Range 3 82-87 3 42-46 Distribution In the eastern part of the continent there are no records of this species between northern Tanzania and eastern Zimbabwe. It is likely however that they do occur, if only coastally, where there is suitable forest habitat. Their occurrence in the western Vila Pery District of Mozambique suggests that they may in time be taken in parts of the Dondo and Amatonga forests further east. South of the Sahara, excluding the Southern African Subregion They occur from Sierra Leone eastward in the High Forest Zone to Nigeria and from thence eastward to Ethiopia; southern Sudan; southern Uganda and southern Kenya. South of this they have been taken in Cameroun; Gabon; Congo Republic; Zaire and in northern Angola from whence at Pongo Andongo (9°40'S 15°40'E) the type specimen of R. angolensis was taken. They occur in the Mwinilunga 66 District on the Zaire border in Zambia and Ansell (1978) stated that they may also occur in the extreme northern parts of the country. Southern African Subregion They have been recorded from the Eastern Districts of Zimbabwe in the Inyanga and Mutare districts and south to Birchenough Bridge on the Sabi River and in adjacent parts of Mozambique, where they have been taken in the western Vila Pery District. These populations are far south of the general limits of their distribution. Habitat Like the Egyptian fruit bat the more important habitat requirements of Bocage's fruit bat are the availability of substantial shelter'in the form of caves or hollow trees in which to roost during the day and a plentiful food supply in the form of wild fruits. In West Africa they occur in the High Forest Zone (Rosevear, 1965) and they have also been taken in gallery forests, montane and lowland forest and in gardens in residential areas (Mutare, Zimbabwe). These types of habitat provide the necessary food supply and they can find roosting places in hollow trees if caves are not available in the vicinity. Unfortunately as all the specimens taken in the Subregion to date have been netted, there is no information on their roosting places. In West Africa they occur from sea level to over 900 m and in East Africa, on Mount Ruwenzori, to 1 800 m (Rosevear, 1965). In the Subregion they appear to be confined to lower altitudes of up to 1 200 m. Habits While no information is available from the Subregion, in West Africa, Eisentraut & Knorr (1957) recorded that these bats hang up near the entrance to caves, which one would expect in a species that has no powers of echolocation to navigate in the dark. They are gregarious, occurring in small colonies of up to about six together, and in the case of the Zimbabwe specimens from Mutare, were associated with far larger numbers of Peters' epauletted fruit bats, E. crypturus, feeding on guavas in an orchard. In other parts of eastern Zimbabwe eight were netted together in a forest clearing. In East Africa, Kingdon (1974) noted that they hang free in the roosting places and assume this posture immediately on alighting in trees, not crawling around among the branches like the straw-coloured fruit bat, Eidolon helvum, or the Egyptian fruit bat, R. aegyptiacus. Food In Mutare, Zimbabwe they were observed to eat wild figs, Ficus spp, and orchard fruits such as guavas and mangoes. Mangoes appeared to be chewed in situ. Kingdon (1974) stated that they stuff their cheek pouches full of food and then fly to a feeding site to eat at leisure. The food is manipulated by one foot as they hang from the other. Fibrous material is chewed and then discarded. Reproduction No information is available from the Subregion. Kingdon (1974) in East Africa observed that the stickiness of the collar increased and decreased from time to time, and thought it might be due to glandular secretion connected with sexual behaviour. Eisentraut (1956) in Cameroun found that females formed nurseries. Suborder MICROCHIROPTERA Insect-eating bats In the Subregion the Suborder Microchiroptera is represented by six Families, the Emballonuridae, the sheathtailed bats; the Nycteridae, the slit-faced bats; the Rhinolophidae, the horseshoe bats; the Hipposideridae, the leafnosed bats; the Vespertilionidae, a diverse Family which includes the serotines, the long-eared bats, hairy bats and others and the Molossidae, the free-tailed bats. Collectively 65 species occur in the Subregion. These are characterised by the possession of a single claw on the thumb of the forearm and, except in the case of members of the Family Rhinolophidae, a tragus in the ear. The differences in the foetal membranes between members of this Suborder and the Megachiroptera as pointed out by Mossman (1937) have been mentioned in dealing with that Suborder. The Microchiroptera rely, when it is totally dark, on echolocation for their orientation, many of them having complicated and most delicate structures around the nose which play a part in this process. As opposed to the "clicks" produced by the tongue of the only fruit bat that uses echolocation, the Egyptian fruit bat, Rousettus aegyptiacus, the "clicks" in the Microchiroptera originate in the larynx. Fenton (1975) showed, however, that some of the Microchiroptera that occur in the Subregion, including the common slit-faced bat, Nycteris thebaica, rely to some extent on vision for orientation when there is adequate light. Generally classed as insect-eaters, the Microchiroptera include species that eat fish, frogs, fruit, birds, drink blood and some are cannibals. Most of those that occur in the Subregion eat insects, although species such as the large slit-faced bat, Nycteris grandis, have been known to eat fish and frogs, albeit this is a departure from their normal insect diet. We have no blood-sucking bats in Africa, they are found only in the New World. 67 attitude. Ansell (1960a) recorded their association with trees in Zambia. Food Insectivorous. Reproduction In Zambia, Ansell (1960a) recorded a juvenile in January, but there is no information available from the Subregion. No. 57 Tadarida (Chaerephon) bivittata (Heuglin, i86i) Spotted free-tailed bat Gevlekte losstertvlermuis Colloquial Name So called from the tiny white and sometimes barely distinguishable flecks on the upper parts of the body. Taxonomic Notes No subspecies are recognised. Description They are about 100 mm in total length with a tail of 37 mm. The adult males have a mean mass of 15,4 g, females 14,6 g (Table 57.1). The upper parts are dark umber-brown and have a variable pattern of tiny white flecks or short stripes on the sides of the crown of the head, on the shoulders and sometimes on the flanks. The under parts may be the same colour as the upper parts or may be tinged with grey. The wing and interfemoral membranes are dark brown. Table 57.1 Measurements (mm) and mass (g) of spotted free-tailed bats, T. (C.J bivittata, from Zimbabwe (Smithers & Wilson, 1979) Males Females X n Range X n Range TL 105 10 102-108 105 21 100-111 T 37 10 34-40 37 21 34-40 Hf c/u 11 10 11 10 21 9-11 E 22 10 21-23 21 21 20-22 F/a 47 10 45-47 46 21 43-47 Mass 15,4 10 13,5-16,7 14,6 21 13,1-17,0 Distribution South of the Sahara, excluding the Southern African Subregion Recorded from Eritrea; Ethiopia; the southeastern Sudan southwards to Zambia. Not so far recorded from Malawi or Mozambique, north of the Zambezi River. Southern African Subregion Recorded from six widely scattered localities in Zimbabwe, Bulawayo marking its most southerly known limit of distribution at the moment. Habitat A savanna woodland species, occurring in Brachystegia woodland in Zambia and Zimbabwe. Habits A gregarious species, occurring in small numbers, not usually more than about six. Roosts during the day tucked away in the crevices of rocks, in caves and mine shafts. Food Insectivorous. Reproduction No information available from the Subregion. No. 58 Tadarida (Chaerephon) chapini (j.A. Allen, 1917) Pale free-tailed bat Bleek losstertvlermuis Colloquial Name So called because of their pale body colour and the whitish wing and interfemoral membranes. Taxonomic Notes Hayman & Hill (1971) listed three subspecies from the continent, T. c. chapini from Zaire and Uganda; T. c. lancasteri Hayman, 1938 from Zambia and Angola, and T. c. shortridgei (Thomas, 1926) from northern Namibia. Description A medium-sized species with a total length of about 150 mm, including a tail of just over 30 mm. The upper parts are pale cinnamon-brown, the hair off-white at the base, the Plate 4 46. Egyptian fruit bat, Rousettus aegyptiacus Egiptiese vrugtevlermuis 55. Angola free-tailed bat, Tadarida (Mops) condylura Angola-losstertvlermuis 67. Schreibers' long-fingered bat, Miniopterus schreibersii Schreibers se grotvlermuis 71. Temminck's hairy bat, Myotis tricolor Temminck se langhaarvlermuis 75. Banana bat, Pipistrellus nanus Piesangvlermuis 86. Cape serotine bat, Eptesicus capensis Kaapse dakvlermuis 90. Schlieffen's bat, Nycticeius schlieffenii Schlieffen se vlermuis 93. Lesser woolly bat, Kerivoula lanosa Klein wolhaarvlermuis 102. Geoffroy's horseshoe bat, Rhinolophus clivosus Geoffroy se saalneusvlermuis 112. Short-eared trident bat, Cloeotis percivali Drietand-bladneusvlermuis 80 under parts greyish-brown with a whitish band down the mid-belly and a narrow band of white along the fringe of the wing membranes next to the body, running from the forearm to the hind legs. The wing membranes are white, tinged yellow near the body, and between the forearms and the legs near the body are punctuated with tiny black specks. The interfemoral membrane is dark brown. The ears are joined by a band of skin across the top of the head and from this rises a crest of long hair, 12 mm in length, the basal half of the hair reddish-chestnut, the remainder white (Fig. 58.1). This long hair lies back on the head, reaching the base of the ears on either side. Just behind the base of this crest the hair on the head is white. The minute anterior upper premolars lie in the toothrow; the anterior lower premolar is smaller than the posterior. Fig. 58.1. Head: Tadarida chapini Distribution Too few specimens of this beautiful crested bat are known from too few localities to be able to assess the limits of their occurrence. South of the Sahara, excluding the Southern African Subregion Recorded from northeastern and eastern Zaire; Uganda; southern Ethiopia; northeastern Angola and northwestern and eastern Zambia. Kingdon (1974) stated that it was taken in Kenya, but did not give localities. Southern African Subregion Known only from northern Namibia, northwestern Zimbabwe and northern Botswana. Habitat Too little information is available to allow an assessment of the habitat in which they occur. In northeastern Zaire, the type locality of the species, Allen, Lang & Chapin (1917) observed that the two specimens were taken in an "open district" (open savanna woodland?) and not in the forest. One specimen was caught in a house, the other taken from the crop of a bat hawk, Macheirhamphus alcinus. In Botswana and Zimbabwe they were taken in mopane woodland. Food Insectivorous. Habits and Reproduction Unknown. No. 59 Tadarida (Chaerephon) pumila (Cretzschmar, 1830-1831) Little free-tailed bat Klein losstertvlermuis Colloquial Name The smallest of the molossids that occur in the Subregion. Taxonomic Notes Hayman & Hill (1971) treated the name pumila in its widest possible sense as the earliest name for a large group of the commonest African molossid bats whose relationships have been the subject of varying interpretations. They point out that attempts at subspecific separation based on colour are profitless. As here understood T. pumila includes both the dark and white winged forms, the latter previously known under the name T. limbata. Description The smallest of the free-tailed bats with a total length of about 90 mm including the tail which measures up to 39 mm (Table 59.1). As the species includes both the dark winged and light winged forms, the colour of these small bats can vary from those that have deep blackish-brown upper parts, the under parts slightly paler, to those that are browner above and whose slightly paler brown under parts have a broad irregularly edged band of white from the anus to the chest, this character coupled with having translucent white wing membranes. This band itself is variable and in some white-winged specimens is absent or barely discernible as a narrow white line. They all have a narrow band of hair on the wing membranes from the forearm to the thighs which varies in colour from white to tawny. The rounded ears are large for the size of their heads, are connected together across the top of the head by a band of skin and their outer edges are folded in upon the ears giving them a clumsy appearance. The antitragus is large and tends to conceal the tiny tragus which is asymmetrically bilobed at the tip (Fig. 56.1.a). 83 Table 59.1 Measurements (mm) and mass (g) of little free-tailed bats, T. (C.) pumila, from Botswana (Smithers, 1971) and Zimbabwe (Smithers & Wilson, 1979) Botswana Males Females X n Range X n Range TL 87 48 70-100 91 91 70-102 T - — F/a 37 10 36-39 37 12 36-38 Mass 11,5 20 10,3-13,9 12,1 65 10,1-16,1 Zimbabwe Males Females X n Range X n Range TL 93 10 90-98 95 20 90-100 T 36 10 35-39 35 20 31-39 F/a 38 10 37-39 38 20 37-39 Mass 11,1 11 10,1-12,7 11,3 12 10,1-14,8 Mass of gravid females Botswana Mass 13,6 Zimbabwe Mass 12,5 n Range 46 12,1-16,7 n Range 19 11,2-15,1 Distribution Extralimital to the continent Recorded from Yemen in southwestern Arabia. South of the Sahara, excluding the Southern African Subregion Recorded from Gambia; southern Niger; Nigeria; Chad; southeastern Sudan; Ethiopia; Eritrea; Uganda; Kenya; Tanzania; Congo Republic; Zaire; Angola; Zambia; Malawi and Mozambique, north of the Zambezi River. Southern African Subregion They occur throughout the Okavango Delta in northern Botswana, south to Lake Ngami; north and south of the plateau in Zimbabwe; widely in Mozambique, south of the Zambezi River; in the eastern Transvaal, Swaziland and northern Natal. There appears to be a break in their distribution westwards, the next records being in the southwest of the Cape Province and northwestwards to near Springbok in Namaqualand. Habitat Although apparently not occurring in the High Forest Zone of West Africa, they occur widely in the Guinea and Sudan savannas and in the forests of the Congo basin in Zaire (Rosevear, 1965). Southwards they are found in savanna woodland in Zambia and in dry mopane woodland at altitudes of less than 1 000 m in Zimbabwe. In the Cape Province they occur in the mountainous areas in the Cape Macchia Zone as well as in the more arid open country further north. They appear to be catholic in their choice of habitat. In Zimbabwe they showed a clear preference for lowveld conditions, never being taken on the plateau over the 1 000 m level. Habits Little free-tailed bats are gregarious, occurring in colonies in favourable areas in hundreds (Triangle, Zimbabwe) (Smithers & Wilson, 1979). Other colonies of 100 to 150 are known (Maun, Botswana) (Smithers, 1971). They roost during the day tucked away in crevices in brickwork under roofs, between the rafters and the corrugated iron roofing sheets, between the overlap of the sheets or in any other type of crevice that presents itself. Verschuren (1957) found them in Zaire in colonies of up to half a dozen in cracks in trees, the entrances to which were very narrow. The size of the colony may well depend on the amount of crevice space available. They appear to be indifferent to high temperatures, for the Maun colony lived in a badly ventilated attic where the temperature in the room underneath reached 40 °C and was certainly higher where they were roosting. About an hour before leaving the roost the colony becomes very restless, individuals squeaking and jockeying for position and generally being very noisy. Verschuren (1957) noted that the entrances to the crevices in trees were through small cracks. Both in the Triangle and Maun colonies the entrances to the attic were through very small apertures between the corrugated iron sheets and the brickwork which allowed only one or two to drop into flight at a time. They appear to forage singly, there being numerous records of solitary individuals entering houses. At the Savuti swamp in Botswana their solitary foraging habits were very noticeable. Their flight is fast and erratic, making them difficult to shoot as they flew past about 12 m above the ground with sudden swoops to within 3 m of it. A random sample of 165 adults, taken during the months of October to February showed a sex ratio of 45 males: 120 females, in August of 3 males: 12 females and in June of 1 male:4 females. Food Insectivorous. Reproduction In northern Botswana gravid females were taken in August and from October to February, by far the larger number in December. Many hairless young were seen in the colonies at Maun in February, some clinging to their mothers, others on their own in the roosts. This indicates that the young are born during the warm, wet summer months from about October to March. Van der Merwe, Rautenbach & Van der Coif (1986) showed that, in the Transvaal Lowveld, there were three peaks of parturition: in early November, late January and early April. The gestation period is about 60 days, females giving birth to a single young at a birth, with a birth mass of about 3,2 g. Females become sexually mature at the age of 5-12 months. The potential for mature females to produce up to three singletons per summer, is ascribed to their ability to undergo post-partum oestrus (Van der Merwe, Giddings & Rautenbach, 1987). 84 No. 64 Tadarida (Chaerephon) ansorgei (Thomas, 1913) Ansorge's free-tailed bat Ansorge se losstertvlermuis Southern African Subregion Recorded from four localities in Zimbabwe, one on the Mozambique border, which suggests that they will turn up there in due course; the Kruger National Park, and from the Mkuzi Game Reserve, Natal. Included in the first edition in the subgenus Tadarida, Meester et al. (1986) followed Freeman (1981) who showed the relationship of this species to other subgenus Chaerephon species. For convenience its original number (No. 64) is retained. Colloquial Name Named after Dr. W.J. Ansorge, a collector of zoological specimens, who collected specimens of this species in Malange, northern Angola. Taxonomic Notes No subspecies have been described. Hayman & Hill (1971), Largen, Kock & Yalden (1974) and Corbet & Hill (1980) placed this species in the subgenus Tadarida, a treatment followed here. Koopman (1975) and Freeman (1981) noted its relationship with T. bivitatta and the extralimital T. bemmeleni and placed it in Chaerephon, a treatment followed by Meester et al. (1986). Description Ansorge's free-tailed bats are a small species, their total length about 106 mm with a tail 36 mm long (Table 64.1). The upper parts of the body are dark umber-brown, the top of the head, neck and throat darker, almost black, and contrasting with the colour of the upper parts. The under parts are a shade lighter than the upper parts. The wing membranes are a translucent light brown, the ears large and rounded. They are very similar to the Egyptian free-tailed bat in colour, but they are smaller, a feature illustrated by the total length of the skull which in this species measures 18-20 mm in length, in the Egyptian free-tailed bat 19-21 mm. The braincase in this species is elevated, while it is flat in the Egyptian species. In this species the canines in the lower jaw are almost touching at their bases, whereas in the Egyptian species they are widely separated. Table 64.1 Measurements (mm) of two Ansorge's free-tailed bats, T. (C.J ansorgei TL T Hfc/u E F/a Male 102 36 — 20 43 Female 102 36 — 20 44 Habitat As this species usually roosts during the day in rock clefts, caves and mine adits the availability of this type of shelter may well be a primary habitat requirement, in addition to the availability of a plentiful food supply. Habits Ansorge's free-tailed bat, judging from the comments of Allen, Lang & Chapin (1917) who recorded a colony of hundreds in a pitch dark rock cleft in northeastern Zaire, is a gregarious species. They recorded how the people there suffocate the bats in their rocky roosting place, collecting basketfuls as food. In Zimbabwe they roosted in caves and mine adits. Like other closely related species, they probably pack tightly into crevices, which is suggested by Allen, Lang & Chapin's (1917) observations. Their colony had been in residence over a long period as there was a layer of dry guano on the floor over 0,3 m thick. This colony became very noisy when approached, individuals squeaking and jostling. Distribution This is not a common bat anywhere in its distributional range and many of the apparent blanks in its distribution may be due to its not having been collected so far. South of the Sahara, excluding the Southern African Subregion Recorded from Oku Lake, Cameroun; the Central African Republic; southern Sudan; Ethiopia; southeastern and northeastern Zaire; Kenya; western Uganda; northeastern Tanzania; Zambia and Malanje, Angola, the type locality. Food Insectivorous. Reproduction No information available. Subgenus Tadarida Rafinesque, 1814 In members of this subgenus the palatal emargination is deep and well developed, the ridging on the upper posterior molars forming a full W pattern (Fig. VII.4.a). 85 No. 60 Tadarida (Tadarida) lobata (Thomas, i89i) Big-eared free-tailed bat Grootoor-losstertvlermuis This species has very large ears which, like the wing membranes, are translucent. The ears arise well forward on the rostrum near the nostrils. The colour of the upper parts is cinnamon-brown with a conspicuous white spot on the shoulders, the under parts buffy, the buffy hair extending marginally onto the wing membranes. The upper lips are plain, not wrinkled. Peterson (1974) remarked that they appear to inhabit arid or semi-arid terrain. So far this species is only known from some 23 specimens from four localities in Kenya and one in Zimbabwe. Originally it was taken in Turkwell in northwestern Kenya and until 1970 remained known only from three specimens, the first described by Thomas in 1891, the second which was collected live in a house in Hatfield, Harare, Zimbabwe, some 2 000 km to the south, and the third from the Cherangami Hills, Kenya, collected by the Oxford University Expedition in 1969. Since then a specimen was collected in the Kapenguria district of Kenya and 19 from Maungu Hill in southeastern Kenya (Peterson, 1974). While the appearance of this species in Zimbabwe so far out of its usual range is surprising and may be a natural phenomenon in a species which is a fast, high flier, the possibility of its being transported in an aircraft cannot be ruled out. During the time it was taken there were regular air services from Nairobi to Harare, Zimbabwe, and as it is a species that uses substantial cover in which to roost, it could have made use of the shelter of an aircraft parked at night. Wiles & Hill (1986) recorded the transport of the North American small brown bat, Myotis lucifugus, to Guam, Mariana Islands, a distance of about 10 000 km by air. The transport of bats by ship is a well-known phenomenon, species reaching New Zealand from Japan, from Panama to the Bass Strait off southeastern Australia, and from Canada to Holland (Vaute, 1982). All the recent specimens were shot near water and therefore there is no information on their roosting places. Peterson (1974) suggested that, like their nearest relatives, T. fulminans, they probably used rock crevices. He recorded that the specimens taken at the Maungu Hill flew high over the water after dark, only three or four descending within 7 m of the ground. Their flight was fast and straight and while flying they uttered a distinctive single or double noted, loud squeak. No. 61 Tadarida (Tadarida) ventralis (Heuglin, i86i) Transvaal free-tailed bat Transvaalse losstertvlermuis Only 16 specimens of this species are known. The original specimen from which the species T. africana (Dobson, 1876) (T. ventralis) was described came from the Transvaal, no exact locality being given. No others have turned up in the province since that date. It is the rarest of the African molossids, single specimens being known from Ethiopia; eastern Zaire; Kenya; Malawi; the Transvaal and two localities in Mozambique; five from Katire in the southern Sudan and four from the Lake Manyara National Park in Tanzania (Table 61.1). It is also the largest of the three closely related species T. ventralis, T. lobata and T. fulminans. They average in total length 140 mm, with a forearm of 62 mm. The colour of the upper parts is variable. In the four specimens from Tanzania examined by Harrison (1971), three were umber-brown with a distinct gloss, the under parts lighter brown with a distinct median white streak of varying width. The fourth specimen had bright fulvous upper parts, almost orange-brown, the under parts fulvous, the streak white washed with fulvous, the flanks lighter in shade. This colour difference was not related to sex. In the middle of the throat they had a circular tuft of hair around a bare patch of skin, which may be glandular. In the fulvous specimen these hairs formed a beard and were ash-grey. In the other three specimens these hairs were white at the base with pale umber-brown tips. The colour of the Transvaal specimen is reddish-brown on the upper parts, 86 the under parts lighter in colour and brighter. The wing membranes are light brown. Its measurements (mm) are as follows: TL T Hf c/u E 147 61 12 25 Table 61.1 Measurements (mm) of a series of four specimens of the Transvaal free-tailed bat, T. (T.) ventralis, from Lake Manyara National Park, Tanzania (Harrison, 1971) Irrespective of sex TL X n Range TL 140 4 137-152 T 55 4 53-63 Hf c/u 12 4 11-13 E 20 4 18-24 F/a 62 4 61-65 Food Insectivorous. Habitat, Habits, Reproduction Unknown. No. 62 Tadarida (Tadarida) fulminans (Thomas, 1903) Madagascar large free-tailed bat Madagaskarse grootlosstertvlermuis Colloquial name Originally this species was described from Fianarantsoa, Madagascar. Taxonomic Notes No subspecies are recognised. Distribution Extralimital to the continent Recorded from eastern Madagascar. South of the Sahara, excluding the Southern African Subregion Recorded from northeastern Zaire; Rwanda; southern Kenya; northeastern Tanzania, Malawi and northeastern and southwestern Zambia. Southern African Subregion Recorded from four localities centred on Harare, Zimbabwe, and one locality in the Hwange National Park in the northwest; and in the Transvaal from Pafuri in the northern sector of the Kruger National Park (Pienaar, Joubert, Hall-Martin, de Graaff & Rautenbach, 1987). Madagascar large free-tailed bats are about 150 mm in total length, with a tail about 56 mm long and a mass of about 33 g (Table 62.1). The upper parts of the body vary in colour from reddish-brown to dark chocolate-brown. Individuals occur in which the upper parts are reddish; chocolatebrown, however, is the more usual colour. The under parts are similar in colour to the upper parts and have a distinct median white, irregularly edged band from the anus to the chest. The upper lips are plain, not wrinkled as in the Egyptian free-tailed bat. The skull has a well developed sagittal crest, the anterior upper premolars set in the toothrow, but slightly to the outside of it. The anterior lower premolar is only about half the height of the posterior premolar. Table 62.1 Measurements (mm) and mass (g) of a series of six male and eight female Madagascar free-tailed bats, T. (T.) fulminans, taken in Zimbabwe (Smithers & Wilson, 1979) Habitat A savanna woodland species, associated in Ethiopia with mountainous country and probably more dependent on rocky terrain than on the vegetational association, as they require rock crevices to roost in during the day. The specimens from Zimbabwe were all taken from areas of granite hills in the savanna woodland zone. Habits Madagascar large free-tailed bats are gregarious, but never occur in colonies of more than about 20. In the Chikupo caves, Zimbabwe, a colony packed into a crack in the granite, the opening of which was some 7 to 8 m above the floor and situated at the entrance. This allowed them a free fall in taking to flight. They left the roosting place at sundown, returning before first light in the morning. Their flight was fast and direct and they appeared to be high fliers. Males Females TL X n Range X n Range TL 147 6 145-150 143 8 138-150 T 56 6 54-59 55 8 53-57 Hf c/u 13 6 12-15 13 8 11-14 E 24 6 22-25 24 8 22-25 F/a 60 6 59-61 59 8 58-60 Mass 32,3 5 31,2-34,1 33,1 8 25,7-39,0 Food Insectivorous. Reproduction Six gravid females were taken at the Chikupo caves, Zimbabwe, in October, each carrying a single foetus. 87 No. 63 Tadarida (Tadarida) aegyptiaca (E. Geoffroy, 1818) Egyptian free-tailed bat Egiptiese losstertvlermuis Plate 5 Colloquial Name The original specimens were collected in Egypt during the Napoleonic wars by Geoffroy, who served under Napoleon. Taxonomic Notes Hayman & Hill (1971) listed two subspecies, T. a. aegyptiaca from Algeria and Egypt and mainly on the eastern parts of the continent to the Cape and T. a bocagei (Seabra, 1900) from the central and western regions. T. a. bocagei is slightly smaller and darker than T. a. aegyptiaca. Description Egyptian free-tailed bats have a total length of about 110 mm, tails 38 mm long and a mass of about 15 g (Table 63.1). The upper parts are dark sooty-brown, the top of the head and back of the neck darker still, nearly black. The under parts are generally the same colour as the upper parts, or they may be a shade lighter. In some individuals the throat is buffy, the mid-belly off-white, the flanks tinged grey or tawny. The wing and interfemoral membranes are a translucent light brown, the fur of the under parts spreading thickly but narrowly on to the wing membranes where it is generally the same colour as that on the under parts, but in some cases lighter and in exceptional cases white. The rounded ears are set closely together on the top of the head, the ear tragus larger than in the subgenera Chaerephon or Mops and not concealed by the antitragus. The braincase is broad and flat, tending to be depressed in the middle. It has no sagittal crest and there is no noticeable supraoccipital crest. The palatal emargination is large and reaches back to the level of the canines. The anterior upper premolars are minute and lie in the toothrow, the posterior premolar large. The ridge pattern on the molar is a full development of the W-pattern (Fig. VII.4.a). Table 63.1 Measurements (mm) and mass (g) of Egyptian free-tailed bats, T. (T.) aegyptiaca, from Zimbabwe (Smithers & Wilson, 1979) Males x n TL 112 11 T 38 11 Hf c/u 10 11 E 21 11 F/a 48 10 Mass 14.9 11 Distribution Although locally Egyptian free-tailed bats can be very numerous, records of their occurrence in some countries such as Angola and Zambia are restricted to a very limited range of localities. There remain large areas in the Subregion from which they have not been taken, such as parts of the Transvaal and Botswana, but there is no good reason to explain this, except lack of collecting. Extralimital to the continent Recorded from southwestern Arabia and India. North Africa Recorded from Algeria and Egypt, extending to the coast of the Mediterranean. South of the Sahara, excluding the Southern African Subregion There is a single record from central Nigeria, east of which they occur in the central and southeastern parts of the Sudan; Ethiopia; Eritrea; Somalia; northeastern Zaire; Uganda; Kenya and in northeastern Tanzania. There appears to be a break in distribution south, the next records being from eastern Zambia, on the Malawi border, and from the south of the country. They occur in central and southern Angola. There are no records at the moment from southern Tanzania, northeastern Mozambique or Malawi. Southern African Subregion They occur widely throughout the whole of the Subregion. Habitat In the southern parts of their distributional range they occur in all the vegetational associations available, excluding forest. They do occur, however, on forest fringes and can be as locally common in such areas as they are in the arid scrub savanna of the southern parts of Botswana or on the open grassland of the Makgadikgadi Pan and Lake Ngami. In the more arid areas in Botswana there was a noticeable association of this species with water holes, even with boreholes and reservoirs. At Tshane in southwestern Botswana the borehole was the only water available on a year round basis. Here they came to sip water from the surface of the round reservoir, 6 m in diameter. When the water dropped a metre they were able to reach the water surface, but unable to pull out at the other side and a large series was collected as a result. In dry country they were netted frequently over temporary pools. They may be dependent on water in arid areas, not only as a source of moisture, but for the richer insect life associated with it. Habits Egyptian free-tailed bats are gregarious, occurring in colonies numbered in dozens. They roost during the day in caves, rock crevices, under rock exfoliation, in hollow trees and behind the bark of dead trees. Shortridge (1934) stated that in Namibia the colonies can number in hundreds, and Herselman (1980) recorded several colonies numbering in hundreds in the Cape Province. In Zimbabwe, Wright (in litt.) reported that in the Pesu River gorge they roosted in nearly vertical cracks in the rock, 15 mm to 20 mm wide, packed in tightly. This colony was located by the noisy squeaking of its members. In western Zimbabwe they were Females Range X n Range 106-115 111 16 105-118 33-40 39 16 35-43 9-11 10 16 8-12 19-22 20 16 18-22 48-54 48 16 46-52 13,8-17.3 15.6 16 13,8-18.1 88 taken from hollow trees by smoking them out. They will also roost in the attics of houses, in crevices in the brick or in any other type of crevice available. Where they are found in caves they seek crevices and crannies in the rock in which to roost. Colonies have a very strong smell. Food Insectivorous. Reproduction In the Transvaal, Rautenbach (1982) took gravid females in September, October and December; Smithers & Wilson (1979), in Zimbabwe, a female with a single foetus in November and lactating females in December; and Wright (pers. comm.) recorded lactating females with young in northern Botswana in December. In the Cape Province, Herselman (1980) recorded heavily pregnant females in several localities in November. These data indicate that the young are born during the warm, wet summer months. Apparently maternity colonies are established as Herselman (1980) found all female colonies in November in two localities. No. 64 Tadarida (Tadarida) ansorgei (Thomas, 1913) Ansorge's free-tailed bat Ansorge se losstertvlermuis Freeman (1981) showed that this species is better placed in the subgenus Chaerephon and therefore it is transferred to follow T. (Chaerephon) pumila (No. 59), for convenience retaining its original number (No. 64) VIII. Family VESPERTILIONIDAE Vesper bats This is by far the largest Family of the insect-eating bats and the most diverse. In the Subregion three Subfamilies are recognised, comprising 10 genera and 29 species and including a range of woolly, long-fingered, serotine, longeared, hairy, house and other forms, making it difficult to apply a general colloquial name that is applicable to all members. Rosevear (1965) uses the name Vesper bats, borrowing it from the name for the Family which is derived from the Latin for a bat, vespertilio which seems appropriate. Members of the family occur throughout the world, except in the Arctic and Antarctic regions, where there is insufficient insect life to support them. Because of their diversity and the fact that many of the species can only be recognised by examination of their dental and cranial characters, it is impossible to present a key that can be used in the field to distinguish them. The head is devoid of noseleaves, the nostrils circular or crescent-shaped (Fig. VIII.1). The ears are separated widely with fairly conspicuous tragi varying in shape according to the species involved. Their tails are long, the vertebrae enclosed within the interfemoral membrane which is supported on either side by a calcaneum which arises from the ankle. Fig. VIII. 1. Head of Eptesicus capensis In the Subfamily Kerivoulinae the fur is erect and frizzy, the ends of the hair curled; in most of the other species the hair is adpressed to the body. The wings vary in shape from the long pointed form found in the Miniopterinae to others in which they are more rounded. In all of them the second finger has a metacarpal and one phalanx. There are no characters by which a Vespertilionid skull can be recognised, there being great diversity within the Family. Even the dentition is variable, the number of teeth varying according to the genus from 30 to 38. Some genera, for example Nycticeius, Scotophilus and Scotoecus have one incisor in the upper jaw while the remainder have two and. although the tooth characters are used in distinguishing the genera, they are very variable. In the genus Eptesicus there are no first upper premolars (Fig. VIII.2); in Pipistrellus on the other hand they are present (Fig. VIII.3), a character used to differentiate between the two genera. However, specimens of Eptesicus are known that possess this tooth on one side only and Pipistrellus that lack it or in which it is present again on one side only. Authorities are agreed, however, that it is convenient to retain the two genera (Ellerman & Morrison-Scott, 1951; Rosevear, 1965). Key to the Subfamilies (Meester et al., 1986) 1. Second phalanx of third digit about three times as long as first; braincase high and rounded . . . MINIOPTERINAE Second phalanx of third digit not especially elongated 2. Ears not funnel-shaped, without deep emargination below tip; tragus short and broad or long and narrow, but not sharply pointed; braincase not particularly high and rounded . . . VESPERTILIONINAE Ears funnel-shaped, with deep emargination below tip; tragus long, narrow, sharply pointed; braincase high and rounded . . . KERIVOULINAE Fig. VIII.2. Upper toothrow of Eptesicus capensis, with one upper premolar (a) and no anterior upper premolar. 89 Fig. VIII.3. Upper toothrow of PipisfreDus nanus to show small anterior upper premolar (a), between canine and posterior premolar. Subfamily MINIOPTERINAE Genus Miniopterus Bonaparte, 1837 Superficially most members of this genus appear very much alike but there is a wide size range which has in the past led to the description of over a dozen species. Hayman & Hill (1971) recognised only four species, three of which occur in the Subregion. These three species have overlapping forearm lengths and they cannot be separated on this basis alone, as is possible in other genera of bats. The mean length of the skull in fact remains the only criterion by which separation is possible at this juncture. Further study may well show that it is only possible to recognise two species, a large and a small, these being found occurring together in parts of their range (Hayman & Hill, 1971). Schreibers' long-fingered bat, M. schreibersii, which is very common in parts of the Subregion, has a very wide distribution from the Cape Province to Morocco and extralimitally in Europe, Asia, in the Australasian region and in Madagascar. In all members of the genus the braincase rises abruptly from the rostrum, as in the genus Kerivoula from which they can be distinguished, because in Miniopterus there are two normal-sized premolars and a third tiny vestigial premolar on either side of the upper jaw compared with Kerivoula which has three normal sized premolars. In addition, in Miniopterus the two anterior lower premolars are much smaller than the third compared with Kerivoula in which they are all approximately equal in size. (See Kerivoula Fig. 93.1). The dental formula is If Q P§ M| = 38 Van der Merwe (1985a) recorded the presence of a tiny, vestigial tooth lying between the upper canine and the following premolar on either side of the jaw in all three species of Miniopterus. He believed this to be P2 and that therefore the premolar following it should be P3 and not P2 as recorded by Miller (1907). The lower incisors are trifid, the anterior upper premolar much smaller than the posterior, which exceeds the molars in height. Key to the species after Meester et al. (1986) 1. Size larger, forearm 45-50 mm; skull length 16 mm and more ... inflatus Size smaller, forearm 42-47 mm; skull length c. 14-15 mm ... 2 2. Forearm 42-44 mm; skull length c. 14 mm; colour brownish black and/or russet . . . fraterculus Forearm 42-47 mm; skull length c. 15 mm; colour slatyblack ... schreibersii No. 65 Miniopterus inflatus Thomas, 1903 Greater long-fingered bat Groot grotvlermuis Colloquial Name The largest of the three species of long-fingered bats occurring in the Subregion. Taxonomic Notes Meester et al. (1986) listed two subspecies from the continent, only one of which occurs in the Subregion, M. i. ru/us Sanborn, 1936 which has a marginal distribution in the northeast. Description As the name implies this is the largest of the three species of long-fingered bats that are found in the Subregion. In average measurements of total length they are in fact the same size as Schreibers' long-fingered bat (Table 65.1), but when the lengths of the forearms, their masses and in particular the lengths of their skulls are considered, the size difference is noticeable. This species has a total skull length of 16 mm or more, Schreibers' about 15 mm and the lesser about 14 mm. The colour of the upper parts is deep chocolate or sootybrown, a little lighter on the under parts.-Like other members of the genus, the wings are long and pointed, the wing and interfemoral membranes very dark brown, almost black. Their ears are small with rounded tips, the ear tragus long and parallel-sided with a round tip and slightly expanded at the base of the outer edge (Fig. 65.1). Fig. 65.1. Ear tragus of Miniopterus inflatus In the skull the braincase rises abruptly from the rostrum as in Kerivoula, but can be distinguished from members of this genus as Miniopterus has only two premolars in the upper jaw as compared with Kerivoula which has three. They have three in the lower jaw, the two anterior ones smaller than the third, whereas in Kerivoula they are all about the same size. The lower incisors are trifid, the outer one indistinctly so. The anterior upper premolar is much smaller than the posterior one, which is taller than the molars. The dental formula is If Cy Pf M§ = 38 90 Table 65.1 Measurements (mm) and mass (g) of greater long-fingered bats, M. inflatus, from Zimbabwe (Smithers & Wilson, 1979) Irrespective of sex X n Range TL 112 4 111-116 T 56 4 56 Hf c/u 11 4 10-11 E 12 4 11-12 F/a 47 4 45-48 Mass 14,7 4 12,5-16,8 Skull TL 16 4 16 Distribution Very few records of the greater long-fingered bat are available and these are so scattered that it is impossible at the moment to assess their distributional limits. Food Insectivorous. Reproduction No information available. No. 66 Miniopterus fraterculus Thomas & Schwann, 1906 Lesser long-fingered bat Klein grotvlermuis South of the Sahara, excluding the Southern African Subregion Recorded from Cameroun; Gabon; Ethiopia; southern Somalia; Kenya; western and eastern Zaire; Uganda and from Kilwa Island, Lake Mweru, Zambia. Southern African Subregion Recorded from Mutare and the Zimbabwe National Monument, Zimbabwe, where in both localities they were taken in association with M. schreibersii. Habitat The availability of caves or mine adits and a plentiful food supply are probably the more important habitat requirements of this species; the type of vegetational association less important. Habits Practically nothing is known of the habits of this species. Where it occurs with Schreibers' long-fingered bat, M. schreibersii, in the Subregion it does so in very small numbers. Hanging in a cave with M. schreibersii, the size difference is not noticeable and it is only when the skulls are examined that this becomes apparent. Colloquial Name The smallest of the three species of long-fingered bats that occur in the Subregion. Taxonomic Notes No subspecies have been described. Description About 90 mm in total length with a tail of about 50 mm, the lesser long-fingered bat, as the name implies, is smaller than the commoner Schreibers' long-fingered bat, M. schreibersii (Table 66.1). Their upper parts are a rich russet-brown, the hair soft and woolly, the top of the head tending to be slightly darker than the back. The under parts are a rich light brown, darker on the throat. The wings are long and pointed with black membranes, the interfemoral membrane translucent dark brown. The light brown hair of the under parts extends sparsely onto the wing membranes between the elbow and the thighs. The ears are small for their size, strongly curved on their inner edges, the tips bluntly pointed. Table 66.1 Measurements (mm) of a male and female lesser longfingered bat, M. fraterculus, from the Transvaal (Rautenbach, 1982) Male TL T Hfc/u E F/a 98 48 8 8 42 Female 99 52 8 8 42 Distribution With the exception of records from southern Kenya and southern Malawi, all the other known records to date are from the Subregion. Southern African Subregion Recorded from Mutare, Zimbabwe; 25 km southwest of Covane, Inhambane District, Mozambique, south of the Zambezi River; the Barberton District, Transvaal; from the Natal midlands, and from localities in the southern and eastern parts of the Cape Province. Habitat Dependent on the shelter of caves or mine adits and a plentiful food supply, these factors may rank as their principal habitat requirements. Habits Like M. schreibersii and M. inflatus this is a cave dwelling species which occurs, like M. inflatus, alongside M. schreibersii, in small numbers. Nothing is known of its habits. Food Insectivorous. 91 Reproduction Bernard (1980a) showed that in Natal copulation and ovulation took place about mid-May to the beginning of June. This was followed by a period of delayed implantation of the embryo of some two and a half months, parturition occurring in December. The females carried a single foetus in the right uterine horn (Bernard, 1980b). Table 67.1 Measurements (mm) and mass (g) of Schreibers' longfingered bats, M. schreibersii, from the Transvaal (Rautenbach, 1982) Males Females X n Range X n Range TL 112 54 102-128 109 39 93-118 T 53 55 46-60 52 41 46-58 Hf c/u 10 55 8-11 10 35 8-11 E 11 56 8-13 11 38 9-12 F/a 45 25 42-48 45 12 44-47 Mass 10,1 33 6,0-13,0 10,2 19 7,8-12,6 Distribution Schreibers' long-fingered bat has extralimitally to the continent a very wide distribution extending eastwards from Europe through India and the Far East southwards through Malaya to northern and eastern Australia. Between the European and western North African populations there is a break in their distribution southwards, occurring again from Cameroun and Ethiopia south to the Cape Province. Extralimital to the continent Recorded from Spain and France eastwards to Turkey and south to Israel and from there through southern Asia; India and Sri Lanka east to Malaya and north to Japan and south to northern and eastern Australia. North Africa Recorded from Morocco; Algeria and Tunisia. South of the Sahara, excluding the Southern African Subregion Recorded from Cameroun; Central African Republic; southern Sudan; Ethiopia; northeastern, southeastern and southern Zaire; Rwanda; Uganda; southeastern Kenya; parts of Tanzania; Mozambique, north of the Zambezi River; Malawi; Zambia and Angola. No. 67 Miniopterus schreibersii (Kuhl, 1819) Schreibers' long-fingered bat Schreibers se grotvlermuis Plate 4 Southern African Subregion They occur in northern Namibia, about as far south as Gobabis; in northern, eastern and parts of southern Botswana: throughout Zimbabwe and Mozambique, south of the Zambezi River; in parts of the Transvaal; in Natal and the Orange Free State. In the Cape Province they occur Taxonomic Notes Meester et al. (1986) included all the material from the Subregion in M. s. natalensis (A. Smith. 1833). Description Schreibers' long-fingered bats have a total length of some 110 mm, with tails of about 50 mm and a mean mass of 10 g (Table 67.1). The upper parts are very dark brown, the under parts slightly lighter, especially towards the lower belly. The wings are long and pointed, the wing and interfemoral membranes very dark brown, nearly black. The ears are small for the size of the body and have rounded ends. The shape of the skull is typical for members of this genus (Fig. 67.1). Fig. 67.1. Skull of Miniopterus schreibersii 92 widely except in the central and in parts of the north of the province. Habitat Schreibers' long-fingered bats are cave dwellers and the availability of caves or other similar substantial shelter such as mine adits are an essential habitat requirement. These and a plentiful food supply may well govern their occurrence, for otherwise they are found in a wide range of vegetational associations and in areas with a mean annual rainfall as low as 300 mm to over 1 000 mm. The maternity caves in which the females give birth must provide the correct conditions of temperature (Van der Merwe, 1973c), and the availability of such caves must be an important habitat requirement. Dwyer (1966) showed that suitable maternity caves could attract females from an area with a radius of up to 300 km. Herselman (1980) showed that in the Cape Province banded individuals moved up to distances of 250 km to sites for winter hibernation. Habits Schreibers' long-fingered bats are gregarious. In a study in the Transvaal, Van der Merwe (1975) found that movements involve both long distance seasonal migrations as well as local inter-cavern movements. Annual migrations take place between caves situated on the southern Transvaal Highveld and caves situated in the northern Transvaal bushveld. These long distance migrations occur during the periods late winter and spring, for movement towards the bushveld caves, and late summer migrations for movement towards the Highveld caves (Van der Merwe, 1975). These long distance migrations range over 150 km and are undertaken by pregnant females en route to maternity caves situated in the northern Transvaal bushveld. Later the same females and their weaned young return to caves in the southern Transvaal Highveld where mating and hibernating colonies are formed. Females from various Highveld caves, and perhaps further afield, aggregate in the same maternity caves year after year to give birth and raise their young. In some of the maternity caves the number of adult females alone is more than 100 000 (Van der Merwe, 1973a; 1978). These north-bound migrations start at the end of July (Van der Merwe, 1973a; 1978) when the hibernation period comes to an end and the hibernating colonies start breaking up. Females do not necessarily migrate directly to the maternity caves but, in some cases, make use of pre-maternity caves in the northern Transvaal bushveld where they stay temporarily until the urge to give birth drives them to the maternity caves (Van der Merwe, 1973a). Many females, however, do not migrate immediately after breaking up of the hibernation colonies, but stay on in some Highveld caves even as late as November before migrating directly to the maternity caves (Van der Merwe, 1975). At this stage (November) many juveniles have already been born in the maternity caves (Van der Merwe, 1975). Many males also undertake this migration with the females. Some of them, however, migrate directly to the maternity caves to be there well in advance of the females. The vast majority of them do not stay long at the maternity caves, but start leaving them as there is an increased influx of pregnant females. Movements of the males after leaving the maternity caves are somewhat obscure. They may either stay in some other, still unknown, bushveld caves, or more possibly migrate directly back to the Highveld where they stay in bachelor groups, awaiting the return of the females towards late summer. Many males, however, do not seem to migrate long distances at all, but stay on in the Highveld caves after the hibernating colonies have broken up (Van der Merwe, 1975). The absence of males in the maternity caves is most probably to avoid any form of competition with the full-term or lactating females, especially competition for food, as these females have a very high metabolic rate during that stage. On the southern Transvaal Highveld local inter-cavern movements are common, even during winter when they are supposed to be in hibernation (Van der Merwe, 1973a; 1975). Hibernation has been found not to be a continuous period of torpidity, but is characterised by much activity such as intra and inter-cavern movements (Van der Merwe, 1973a; Norton & Van der Merwe, 1978). During periods of torpidity, however, these bats can be handled with ease for periods of more than 20 minutes before they will start waking up. Much of these activities during the hibernation period may be attributed to the relatively high cave temperatures prevailing in some of the Highveld caves during the winter months (Norton & Van der Merwe, 1978). In some of these Highveld caves their numbers vary from a few individuals up to an estimated number of 4 000 (Van der Merwe, 1973a). During the day they roost in caves and mine adits where total darkness prevails. They have been recorded from vertical mine shafts and in the roofs of houses (Smithers & Wilson, 1979). Roberts (1951) stated that they will also roost in crevices in rocks and trees. They hang from the ceilings of caves in tightly packed clusters, using their hind claws for anchorage. They may also use the walls and then the claws of both feet and thumbs are used to secure a hold. Cluster formation starts when a few individuals land on all fours on the ceiling, holding with the claws of both feet and thumbs. They clamber around until a suitable anchorage is found for the claws on the feet, then drop into the hanging position with the head down. New arrivals aggregate around them causing the cluster to expand rapidly. Late arrivals may land on the middle of the growing cluster in an attempt to secure a foothold on the ceiling. They use the claws of both feet and thumbs to cling to the heads and shoulders of other individuals in the clusters until a foothold is found, or they may crawl over the heads and shoulders of those already hanging towards the perimeter where they can secure a position. When landing on the walls, they do so with their heads upwards, then turn around until the head is down and crawl around until a secure hold is found. Occasionally individuals use crevices, crawling into them backwards, others following, until the crevice is full of bats (Van der Merwe, 1973b). In disbanding of the clusters the individuals on the outside tend to take to flight first, working towards the centre. In large clusters however, some of the individuals in the centre may wriggle loose, leaving a gap, from the edges of which individuals detach themselves. Schreibers' long-fingered bats are extremely fast fliers, their long, pointed wings adapted for this purpose. They tend to fly high, swooping in the air like swallows. When they hang in the roosting places, the long, tapering ends of the wings fold back, the tail and its interfemoral membrane rolling up against their bellies. In the caves they hang in closely packed clusters, estimated at densities of up to 2 800 individuals/m2 (Norton & Van der Merwe, 1978). The winters can be very cold on the Transvaal Highveld, and Twente (1955a) postulated that hibernating bats in temperate regions must roost in cool places in order to keep their body metabolism at a sufficiently low level to prevent over-utilisation of their fat resources before the onset of the next favourable feeding season. Conversely during their feeding season, when insect life is plentiful, as it is in the Transvaal during the summer months from about September to March, they must roost up in places warm enough for digestion and assimilation to proceed. Van der Merwe (1973c), after study of the temperature and humidity in a cave used for hibernation, came to the conclusion that low temperature was the deciding factor in its choice. Food Insectivorous. Reproduction In the Transvaal, Rautenbach (1982) found gravid females in October and November. Smithers & Wilson (1979) recorded them in September and October in Zimbabwe, and Herselman (1980) in the Cape Province in October. The females 93 normally produce one young at birth, but Herselman (1980) found one female, out of 150 examined, carrying twins. The pattern of reproduction is similar to that of temperate zone Miniopterus in that ovulation and fertilisation follow immediately upon copulation in the autumn, and neither sex stores sperm for prolonged periods (Van der Merwe, 1980). The females enter hibernation already pregnant, with the blastocyst lying unattached in the right uterine horn (Van der Merwe, 1980). On the Transvaal Highveld, Van der Merwe (1979) found that mating takes place in mixed clusters of males and females during late summer and autumn. Ovulations and conceptions peak towards the end of March and terminate towards the end of April. , , , Copulation and fertilisation are followed by a long gestation period of eight months, with the peak of births during the last half of November terminating towards mid December (Van der Merwe, 1979). After fertilisation the conceptus only develops as far as the bilaminar blastocyst stage and then shows no conspicuous development for four months while lying unattached in the right uterine horn (Van der Merwe, 1979, 1980). This period of delayed implantation overlaps to some extent with the winter hibernation period (Van der Merwe, 1980). Implantations occur during the winter with a peak towards the end of July when the hibernation period normally comes to an end (Van der Merwe, 1979). For the next four months embryonic and foetal development occur during which time the foetuses reach an average mass of 2,7 g at birth (Van der Merwe, 1979). , , . At birth the juveniles are pink, the only signs ot pigmentation being a faint coloration around the rostrum where there are a few faint vibrissae. The ears are fully erect and the eyes closed. The milk teeth are sharp and the thumbs and hind feet well developed. The feet and thumbs are very important as the females deposit their young on the roof only a few hours after being born and they are especially dependent on their hind feet for attachment (Van der Merwe, 1978). After birth the juveniles remain attached to their mothers for only a few hours, after which they are placed in a separate juvenile cluster. Normally a more secluded place within a maternity cave is selected to serve as the maternity chamber. This phenomenon is to promote high and stable temperatures within the chamber, which is important for maximum growth of the juveniles. In these chambers the adult females control the temperature by hanging at various distances from the juveniles. When the temperature drops too low, they will hang nearer to the juveniles, sometimes closely surrounding them, and when the temperature increases too much, they will move away. When the juveniles are older and more active they generate more heat themselves, which may result in the females hanging on the outside of the chamber to prevent it from overheating (Van der Merwe, 1978). Maternity colonies break up towards the end of the summer and the females and weaned young start migrating back to caves on the southern Transvaal Highveld where mating and hibernating colonies are formed (Van der Merwe, 1975). Subfamily VESPERTILIONINAE Key to the genera, after Meester et al. (1986) 1. Six upper, six lower cheekteeth on each side ... Myotis Fewer than six upper and six lower cheekteeth . . • 2 2. Five upper, five lower cheekteeth; ears not lengthened, two upper incisors on each side; wing with fifth finger not shortened; ears not joined at base ... Pipistrellus Four upper, five lower cheekteeth; ears notably lengthened only in exceptional cases ... J 3. Two upper incisors on each side One upper incisor on each side ... 6 4. Rostrum very short and broad; braincase very high, more than two-thirds of condylobasal length; lower lip with lobe at posterior angle; frequently with body pattern of spots and/or stripes, or conspicuous reticulation of wing membranes .. . Chalinolobus Rostrum not shortened; braincase not elevated; no body or wing pattern . . • v) 5. Ears 18 mm or more, about half length of forearm .. . Laephotis Ears less than 18 mm, much less than half length of forearm „ .. . Eptesicus 6 Size larger, forearm not less than 46 mm; first and second upper molars with W-pattern obsolescent; tragus long and tapering . . . bcotophilus Size smaller, forearm less than 46 mm; first and second upper molars with normal W-pattern; tragus either sickle-shaped or short and broad ^ 7. Skull with rostrum not broadened; palatal emargination narrow; anterior lower premolar about half the crown area of the next premolar; upper canines with rounded anterior surface; tragus halfmoon-shaped; penis not enlarged; forearm 29-33 mm ... Nycticeius Skull with rostrum notably broadened, particularly across lachrymals; palatal emargination broader; lower premolars equal in crown area; upper canines with broad, flat anterior face; tragus short and broad; penis greatly lengthened; forearm 28-38 mm . . . Scotoecus Genus Myotis Kaup, 1829 The generic name is derived from the Greek os otis an ear, and mys a mouse, referring to their large ears. Members of this genus can be separated from other vespertilionids by the elongated muzzle, the rich coppery-red colour of the body, the soft fur, which stands out from the body and the characteristic shape of the ear tragus. In the skull the braincase rises from the elongated muzzle in a rather even slope and they have three premolars in both the upper and lower jaws (Fig. 72.1). The dental formula is If C\ P| Mf = 38 the great number of teeth accounting for the elongated muzzle. The upper two incisors are bifid, the lower incisors trifid. The skull has no sagittal or supraoccipital crests. Key to the species after Hayman & Hill (1971) (Measurements in mm) 1. Size larger, forearm over 52 ... welwitschii Size smaller, forearm 52 or less ... 2 2. Wings with glands; first two upper premolars greatly reduced and closely crowded together laterally ... 3 Wings without glands; first two upper premolars not greatly reduced, not crowded together laterally (Fig. 72.1) 3. Size smaller, forearm 32,5-32,9; upper parts dull drab ... seabrai Size larger, forearm 34,5; upper parts honey-yellow ... lesueuri 94 4. Size larger, forearm 47-52 . . . tricolor Size smaller, forearm less than 47 . . . bocagei No. 68 Myotis welwitschii (Gray, 1866) Welwitsch's hairy bat Welwitsch se langhaarvlermuis Plate 5 Colloquial Name Named after Welwitsch, an Austrian botanist, who made extensive collections in Angola. Taxonomic Notes Meester et al. (1986) did not recognise any subspecies. Description Welwitsch's hairy bat is about 120 mm in total length, with a tail about 60 mm and a mass of 14,0 g (Table 68.1). It is the largest of the four species that occur in the Subregion. As in the other species of Myotis, the hair stands erect giving it a hairy look from which the colloquial name originates. The upper parts are a pale coppery-red, very similar to M. tricolor, but if anything, a shade darker. The individual hairs are black at the base and broadly tipped with copperyred. The top of the head and neck tend to be paler in colour than the remainder of the body. The under parts are whitish, tinged with coppery-red. The skin over the bones of the wings is light reddishbrown, the membranes dark, nearly black, and are reticulated with a profuse series of faint white lines, and with a few tiny yellow spots between the forearm and the fifth digit. The interfemoral membrane is coppery-red with a sparse coating of hair near the body and profusely covered with a series of small, irregularly shaped black dots. The ears are large with rounded points and are coppery-red in colour with narrow black edges. Table 68.1 Measurements (mm) and mass (g) of Welwitsch's hairy bat, M. welwitschii, from Zimbabwe (Smithers & Wilson, 1979) Irrespective of sex TL X n Range TL 119 7 105-127 T 58 7 52-63 Hf c/u 13 7 11-15 E 21 6 19-22 F/a 55 6 52-58 Mass 14,1 6 12,0-16,5 Distribution Records of this species are too few and scattered to allow an understanding of the limits within which they may occur. The original specimen from which the species was described is labelled "Angola", without further details of the locality from which it originated. It was taken, however, in this country by Hayman (1963). South of the Sahara, excluding the Southern African Subregion Recorded from the Ulurugu Mountains and from the east and southwest of Tanzania; from southeastern and northeastern Zaire; Kenya; Ethiopia; central Angola; northwestern Zambia and the Southern Province of Malawi. Southern African Subregion Recorded from Mashonaland and the Eastern Districts of Zimbabwe; parts of the central and eastern Transvaal; and the Orange Free State. Habitat A savanna woodland species. Habits A solitary species that has been taken roosting during the day hanging in bushes and trees (Rautenbach, 1982; Smithers & Wilson, 1979). They enter houses at night when foraging. Food Insectivorous. Reproduction No information available. No. 69 Myotis seabrai (Thomas, 1912) Angola hairy bat Angola-langhaarvlermuis Colloquial Name So called as the original specimen originated from Mossamedes on the southwestern coast of Angola. Taxonomic Notes No subspecies have been described. Description M. seabrai and M. lesueuri are very similar, M. seabrai, however, being slightly smaller with a forearm length of 32,5 to 32,9 mm as opposed to that of M. lesueuri at 34,5 mm. M. seabrai is, in addition, darker in colour, a dull drab with a tinge of yellow, the base of the hairs a pale slate-grey. Too few specimens of either species are available to properly 95 assess any variation in characters or factors which might assist in separating these two closely allied species. Distribution Angola Only known in Angola from the type locality Mossamedes in the southwest. Southern African Subregion Recorded from Berseba in Namibia and from three localities in the northwestern Cape Province, namely Goodhouse on the Orange River, Steyerskraal, Goegab and east and south of this (Herselman, 1980). Habitat Too little information is available to allow an assessment of the habitat requirements of this species. All the localities from which it has been taken lie in areas with a mean annual rainfall of less than 100 mm. Mossamedes is a coastal town lying in desert surroundings, Goodhouse a settlement on the Orange River with irrigated citrus orchards and riverine associations, and the other localities all lying in arid terrain. Habits Shortridge (1934) collected two at Berseba, Namibia, flying around a church, and he suspected that they roosted in the steeple during the day. Roberts (1951) stated that their flight was not so strong or erratic as the Cape serotine bat, E. capensis, and on emerging in the evening, they had a comparatively steady, direct flight. He observed that as dusk descended they circled low around trees and bushes, at Goodhouse around orange trees, snapping insects from the leaves. Food Insectivorous. Reproduction No information available. No. 70 Myotis lesueuri (Roberts, 1919) Lesueur's hairy bat Lesueur se langhaarvlermuis Colloquial Name Named after J.S. le Sueur of L'Ormarins in the Paarl District, Cape Province, who recovered the original specimen from his cat. Taxonomic Notes The two species M. lesueuri and M. seabrai are differentiated purely on a matter of size, M. lesueuri being the larger of the two, with a forearm length of 34,5 mm against that of M. seabrai at 32,5 to 32,9 mm. Hayman & Hill (1971) questioned the status of this species, noting that there has been some doubt as to whether it possesses wing glands. Herselman & Norton (1985) showed that some individuals had a gland on one or other of the wings, whereas in others it was absent. Hayman & Hill (1971) listed both species on the basis of difference in size. Description Lesueur's hairy bats have a total length of some 90 mm and tails 43 mm in length. The upper parts are honey-yellow in colour, the under parts much paler, a pale yellowish-white. The head is pale like the under parts, the face brown. The individual hairs on the body are black at the base. The wing and interfemoral membranes are dark brown. The two species M. lesueuri and M. seabrai are so alike that they can only be distinguished on size, the best criterion being the length of the forearm (see Taxonomic notes). Distribution Southern African Subregion Confined in their distribution to parts of the southwestern Cape Province where they occur from the vicinity of Franschhoek north to Citrusdal and northeast to Beaufort West and Mazelsfontein in the Great Karoo. Herselman (1980) in spite of the scarcity of this species, believed that in time they will be shown to occur more widely in the Cape Province. -r a / V N r -9 / U_ <*• ;? w ft Y r J ^ TTT V/M ) \/ \ TTTT v 9 _ \V M Ol/ / / StiSd^WfTll 96 Food Insectivorous. Habitat, Habits and Reproduction No information is available as the majority of the specimens have either been trapped or picked up dead. No. 71 Myotis tricolor (Temminck, 1832) Temminck's hairy bat Temminck se langhaarvlermuis Plate 4 Colloquial Name Named after C.J. Temminck who was the author of the Monographies de Mammalogie (1827) and whose name is perpetuated in that of the pangolin, Manis temminckii. Taxonomic Notes No subspecies are recognised. Description Temminck s hairy bats have a total length of about 100 mm with a tail about half this length. The upper parts are a pale coppery-brown, the individual hairs narrowly light grey at their bases, then broadly pale buff with coppery-brown tips. The under parts are lighter in colour, pale brown washed faintly with coppery-brown. The wing membranes are dark brown, the interfemoral membrane with a dense covering of coppery-brown hair, near the body naked towards the hind margin. The ears have rounded tips and are brown in colour. Distribution All records to date, with the exception of the single record from western Zaire (Hayman, Misonne & Verheyen, 1966) indicate an easterly distribution on the continent from western Ethiopia, from where there are a number of records, to the southwestern Cape Province where it has been taken in a number of localities. It is not, however, well represented from the intervening areas, records being few and scattered. South of the Sahara, excluding the Southern African Subregion They have been taken in western Ethiopia; in Kenya; Uganda; Zaire; Zambia; Malawi, but are not recorded from Tanzania or the northeastern parts of Mozambique, but probably do occur. Southern African Subregion They occur in eastern Zimbabwe and adjacent parts of Mozambique south of the Zambezi River, in parts of the Transvaal, Orange Free State, Natal and Transkei and in the southern and southwestern parts of the Cape Province. Not so far recorded from Lesotho, but may well occur. Habitat They occur predominantly in savanna woodland, but penetrate into drier, more open terrain in the southern parts of their range. Their occurrence is probably governed more by the availability of caves and mine adits, in which to roost during the day, than the vegetational associations in which they are found. Habits A gregarious species, occurring in colonies numbering dozens, roosting during the day in caves and mine adits. They appear to have a preference for those that are waterlogged thL6y 3re undisturbed (Roberts, 1951; Herselman, 1980). They hang from the ceilings or walls; in the former case Roberts (1951) stated that they clung either with the claws of the feet or with the aid of these as well as the claws on the thumbs. There is evidence of local movement, for Herselman (1980) reported that an individual ringed in De Hoop Cave, Cape Province, was recaptured in a cave at Montagu, a distance of 90 km away. In the roosting places they are often found in association with Schreibers' longfingered bats, Miniopterus schreibersii, and Cape horseshoe bats, Rhinolophus capensis. Food Insectivorous. Reproduction Bernard (1982b) found that copulation took place in midApril in Natal and ovulation and fertilisation in midSeptember. The gestation period was estimated to be 63 days, parturition occurring between mid-November to midDecember, followed by a six weeks period of lactation. He assumed that the females stored the sperm in their uterine horns between mid-April and mid-September, a reproductive strategy reported for north-temperate members of the genus. Herselman (1980) reported gravid females and juveniles in the Cape Province towards the end of October, females giving birth to their young from late October to the middle of November. No. 72 Myotis bocagei (Peters, 1870) Rufous hairy bat Rooi langhaarvlermuis Colloquial Name This species is often called Bocage's hairy bat or Bocage's banana bat. J.V. Barbosa du Bocage was an eminent Portuguese zoologist who worked during the 19th century, and who named material from the Congo and Angola collected by his Portuguese colleague Snr M. Jose d'Anchieta. 97 Taxonomic Notes Hayman & Hill (1971) listed two subspecies, M. b. cupreolus Thomas, 1904 from West Africa and M. b. bocagei from the remainder of the species' range. Description Rufous hairy bats have a total length of about 100 mm, with tails 40 mm in length and a mass of about 7,0 g. They are a very beautiful species with their coppery-red upper parts and almost black ears, wing and interfemoral membranes. The under parts are lighter in colour than the upper parts and are off-white washed with coppery-red. The hair on the body is narrowly black at the base with broad coppery-red tips. The inner margins of the ears are strongly convex, sweeping out to the broadly rounded tips, the outer edges strongly concave, giving the ear the appearance of a bluntpointed sickle. In members of the genus Myotis there are three premolars in each side both in the upper and lower jaw (Fig. 72.1). The ear tragus is long and has a rounded tip (Fig. 72.2). Angola border in the west and from the Southern Province of Malawi. Southern African Subregion Recorded from the Eastern Districts of Zimbabwe and Pafuri and Skukuza in the Kruger National Park, Transvaal. Habitat The West African subspecies, M. b. cupreolus, is a forest species; M. b. bocagei, the subspecies occurring in the Subregion, is associated predominantly with open savanna woodland. Habits Bocage's hairy bats occur solitarily or at most two occur together. They roost during the day in hollow trees, in the shelter of the leaves of Hyphaene palms, or in bunches of bananas, and have been taken from the sheath of an arum lily, and from damp undercover in thickly overgrown swamp (Rosevear, 1965). While there is no information on the roosting places in the Subregion as the specimens were netted, it is perhaps significant that in both localities bananas were being grown. In Zaire, Allen, Lang & Chapin (1917) recorded that the indigenous people know them as the "big red brother" of the banana bat, Pipistrellus nanus, as both occur together in bunches of bananas. They noted that they appear to avoid villages and therefore do not come to the notice of travellers. Food Insectivorous. Reproduction No information is available from the Subregion. Genus Pipistrellus Kaup, 1829 The name is derived from the Italian pipistrello from vispitrello, the diminutive form of the Latin vespertilio, a bat (Rosevear, 1965). Members of the genus are all small bats, difficult to distinguish from one another, the differences between them being slight. In this genus, as compared with Eptesicus, members have an anterior upper premolar (Fig. 75.1) which is generally lacking in Eptesicus. It is, however, sometimes absent in Pipistrellus, but for convenience the identity of the two genera is maintained. Members of the genus have two upper premolars, the dental formula being If C{ P| Mf = 34 Fig. 72.1. Skull of Myotis bocagei. 5sr x Fig. 72.2. Ear tragus of Myotis bocagei. Distribution Particularly in the eastern and southern parts of their distributional range Bocage's hairy bats are poorly represented in collections and it is not possible at the moment to show their limits of occurrence. In West Africa, Rosevear (1965) showed that M. b. cupreolus occurs from Liberia eastwards in the forest zone. Extralimital to the continent They occur in Aden in southwestern Arabia. South of the Sahara, excluding the southern African Subregion Recorded from Niger; Guinea and Liberia eastwards to southern Cameroun; Equatorial Guinea and Zaire. The original specimen was collected at Duque de Braganza, northern Angola, and remains the only record from that country. They are recorded from the Central African Republic; Uganda; Rwanda; Burundi; central Kenya and southwestern Tanzania; and in Zambia from Chingi on the 98 sometimes lying in the toothrow, in other specimens interna to the toothrow. The skull has no sagittal or supraoccipltal crest. Studies in progress by I.L. Rautenbach and R.L. Peterson suggest the presence of other species of Pipistrellus in southern Africa. Key to the subgenera 1. Inner upper incisors unicuspid or not very deeply bicuspid (Fig. 75.2); the under parts brown or grey .. . Pipistrellus Inner upper incisors deeply bifid; the under parts pure white . . . Vansoni Subgenus Pipistrellus Kaup, 1829 Key to the species after Meester et al. (1986) 1. Outer upper incisor more than half length of inner upper incisor; forehead strongly concave; upper canine and posterior upper premolar separated by a gap through which anterior upper premolar can be seen clearly rising above cingula of adjoining teeth; interdental' palate clearly longer than broad .. . nanus Outer upper incisor less than half length of inner upper incisor; forehead usually flat or weakly concave; upper canine and posterior upper premolar in contact or barely separated, anterior upper premolar seen from the side only with difficulty, usually not rising appreciably above cingula of adjoining teeth ... 2 2. Interdental palate as broad as long; forehead flat; total length of skull 12 mm or less .. . rusticus Interdental palate longer than broad; forehead at least slightly concave; total length of skull more than 12 mm ... 3 3. Posterior upper incisor barely extending beyond cingulum of anterior upper incisor; anterior upper premolar a pointed tooth rising above cingulum of canine but not above cingulum of posterior upper premolar; maxillary toothrow more than 4,5 mm . . . kuhlii Posterior upper incisor extending well beyond cingulum of anterior upper incisor; anterior upper premolar a flat-crowned tooth, completely below cingula of adjoining teeth; maxillary toothrow less than 4,5 mm .. . anchietai No. 73 Pipistrellus kuhlii (Kuhl, 1819) Kuhl's bat Kuhl se vlermuis Taxonomic Notes Hayman & Hill (1971) listed five subspecies from the continent, of which two occur in the Subregion, P. k. broomi Roberts 1948 from Natal and P. k. subtilis (Sundevall, 1846) trom other parts of the Subregion. Description Kuhl s bats are very similar in size and colour to rusty bats. I hey are only very slightly larger, with total lengths averaging about 76 mm in males to 80 mm in females and a mean mass of 4,6 g and 5,8 g respectively (Table 73.1). The colour ol the upper parts is light brown to russet, the under parts lighter coloured, in some buffy-white. Their ears are broader man in tne rusty bat, but similarly rounded at the tips. The wing and interfemoral membranes are dark, nearly black, the former with a narrow white border. The ear tragus is knife-shaped with a rounded point (Fig. 76.1.c). Table 73.1 Measurements (mm) and mass (g) of Kuhl's bats, P. kuhlii, from the Transvaal (Rautenbach, 1982) _ Males Females x n Range x n Range TL 76 16 63-80 80 5 76-84 T 29 16 20-33 30 5 26-32 Hf c/u 6 6 5-7 6 4 5-8 U 15 9-12 10 5 7-12 F/a 29 13 25-31 31 3 31 Mass 4,6 14 4,0-6,0 5,8 4 4,0-7,0 Distribution Extralimital to the continent Kuhl's bat originally was described from a specimen from Trieste at the north end of the Adriatic and has a wide distribution in Europe, the Middle East and eastwards to India. North of the Sahara They occur throughout Morocco; Tunisia; in Libya, being common in the coastal towns, and in Egypt to the shores of the Mediterranean. In Algeria they occur coastally and deep into the Sahara southward to the oasis of Messad and the Ahaggar Massif. South of the Sahara, excluding the Southern African Subregion Recorded from the northeastern and southeastern Sudan; Ethiopia; Cameroun; and in parts of Somalia and Kenya.' There is a single record from northeastern Tanzania and although they are at present only recorded from western and northeastern Zambia, Ansell (1978) believed that they are more widespread than present records indicate. They occur in southern Malawi with a single record from the extreme north of the country. At present there are no records from Mozambique, north of the Zambezi River. Southern African Subregion Recorded from central and eastern Zimbabwe and the Maputo District of Mozambique, south of the Zambezi River. In the Transvaal they occur across the middle of the province from near the Botswana border to the Mozambique border just north of Swaziland, and in the northeastern Orange Free State. They occur in Natal and coastally westwards in the Cape Province to about Knysna, being comparatively rare in this part of their range. Habitat It is difficult to assess the habitat requirements of a species that can utilise such diverse habitats as those found in Europe, in the deserts of Arabia or the Sahara and the forested terrain of the Knysna area on the south coast of the Cape Province. In Arabia, Harrison (1964) noted that although they foraged over the desert, they never moved far from the oasis with which they were associated. All the records from Zimbabwe were made in the vicinity of streams and rivers and their eastern distribution in the Subregion suggests that they prefer well-watered terrain. It seems likely that a water supply, the availability of suitable roosting places and a plentiful food supply are among their primary requirements. Habits Kuhl's bats are gregarious, occurring in small colonies in Zimbabwe numbering up to about 12. They are apparently far commoner in North Africa where Hufnagl (1972) stated that they are the commonest species in Libya, roosting in the roofs of bazaars and old houses. In the Cape Province on the other hand Herselman (1980) recorded them as being very rare. Judging from the number of specimens in collections in the Subregion, they are nowhere common. There is a scarcity of information on their daylight roosting places in the 99 Food Insectivorous. Reproduction In Zimbabwe gravid females were taken in January, the average number of foetuses being carried by females being 1,6 with a normal range of one or two. A female taken nearHarare carried three foetuses. In the Transvaal, Rautenbach (1982) recorded gravid females in October and November with one or two foetuses and a solitary female with three. He also took lactating females in September, November and December. In the northern parts of the Subregion these data indicate that the young are bom during the warm, wet summer months. Genus Scotophilus Leach, 1821 The ear tragus of members of this genus is characteristically long and fine-pointed, the outer margin convex with a lobe towards its base. In the posterior molars the usual W pattern on their crowns is reduced to two legs, only half the teeth being present. S. dinganii and S. borbonicus are both quite common in the Subregion, the latter tending to be confined to lower altitudes, the former common in the roofs of houses and recognisable from the yellow or yellowish-orange colour of the under parts. The giant yellow house bat, S. nignta, on the other hand is rare in the Subregion, only two specimens having been taken to date. The dental formula is I| CI P| M§ = 30 They have a single pair of upper incisors (Fig. 88.2), the anterior lower premolar much smaller than the posterior an tightly packed between it and the canine in the toothrow. Key to the species after Meester et al. (1986) (Measurements in mm) 1 Size large, forearm 75-80; total length skull over 30 ... nignta Size smaller, forearm 43-65; total length skull 16-23 ^ 2. Forearm 50-65; total length skull 20-23 ... dinganii Forearm 43-50; total length skull 16-19 ... borbonicus No. 87 Scotophilus nigritd (Schreber, 1774) Giant yellow house bat Groot geel dakvlermuis Colloquial Name This is the largest of the three yellow house bats which occur in the Subregion. Taxonomic Notes Robbins (1978) showed that Dobson (1875) when describing S. gigas was in fact redescribing S. nigrita and that since 1800 the name S. nigrita has been applied to the wrong taxon. As a result S. nigrita becomes the senior synonym of S. gigas; the next available name for the taxon previously known as S. nigrita is S. dinganii. This leaves three species which occur in the Subregion, S. nigrita, the largest, S. dinganii, a medium-sized species, and S. borbonicus, the smallest. Only the subspecies S. n. alvenslebeni Dalquest, 1965 occurs in the Subregion. Description As the name suggests this is a large Scotophilus, the tota length about 170 to 180 mm as against that of S dinganii at 130 mm and S. borbonicus at 110 to 120 mm (I able 87.1). In the Mozambique specimen the upper parts were dark sootybrown in colour, the ears and wing and interfemoral membranes black. There appears, however, to be considerable variation in colour, for Rosevear (1965) stated that three specimens from Malawi ranged from darkish red-brown to yellowish-greyish-brown on the upper parts, the under parts varying from pale straw colour to white. Among our insectivorous bats they rank second in size after Commerson s leaf-nosed bats, Hipposideros commersoni. The ear has a strongly rounded inner margin, the ear tragus typical of members of the genus with its evenly curved inner margin, long tapering shape ending in a fine point (Fig. 88.1.a). The sagittal crest on top of the skull is well developed and ends posteriorly in a rounded "helmet". Table 87.1 Measurements (mm) of giant yellow house bats, S. nigrita (Dalquest, 1965; Rosevear, 1965) Male from Zinave, Save River, Mozambique (Dalquest, 1965) TL T Hfc/u E F/a 175 77 14 20 77 Range of measurements as given by Rosevear (1965) HB T F/a 112-117 68-78 70-80 Distribution There are very few specimens of this, the largest of the members of the genus Scotophilus which occur on the continent. These were taken in a scatter of localities from Senegal to the Sudan and south to the Save River in Mozambique. It is impossible at the moment to attempt to assess their limits of distribution and only the localities from which they have been taken are mapped. South of the Sahara, excluding the Southern African Subregion Recorded from Senegal; Ghana; Nigeria; the Sudan; Kenya; eastern Zaire and southern Malawi. Some doubt remains as to the authenticity of the record from Ghana (Rosevear, 1965). 112 Southern African Subregion Represented by two specimens, one from Odzi, near Mutare, Zimbabwe, the other from the Zinave National Park on the Save River in Mozambique. Habitat Too little is known about giant yellow house bats to understand their habitat requirements. The type specimen came from rain forest in Nigeria, but another from the arid Sudan savanna (Rosevear, 1965). The two specimens taken in the Subregion were collected in the vicinity of major rivers lying within savanna woodland. Food Insectivorous. Habits and Reproduction No information available. No. 88 Scotophilus dinganii (A. Smith, 1833) Yellow house bat Geel dakvlermuis Plate 5 Colloquial Name This is the only member of the genus which occurs in the subregion in which the adults have yellow coloured under parts. The juveniles are not so clearly coloured, but usually show some suffusion of this colour. Taxonomic Notes Meester et al. (1986) recognised three subspecies as occurring in the Subregion: S. d. dinganii from Natal, Transvaal, Mozambique, Zimbabwe and eastern and northern Botswana; S. d. pondoensis Roberts, 1946 from the eastern Cape Province and Transkei, and S. d. herero Thomas, 1906 from northern Zambia. Description Yellow house bats are of medium size, the total length being about 130 mm, with tails of 50 mm long and a mass of about 27 g (Table 88.1). The colour of the upper parts is variable, ranging from a light olive-brown or a greyish-olive to a rich reddish-brown. The under parts may be bright yellow, ochre-yellow or much paler, almost off-white, with or without yellow on the flanks. The hair of the under parts continues broadly on to the wing membranes between the elbow and the thighs. The hair is soft and woolly with a distinct sheen. There is variation in the colour of both the upper and under parts even in a series from the same colony. The wing and interfemoral membranes are a dark translucent brown. The ears are relatively small, the inner edges strongly curved, the tips bluntly pointed. The ear tragus is characteristic in shape (Fig. 88.1.a). They have a single upper incisor on either side (Fig. 88.2). Table 88.1 Measurements (mm) and mass (g) of yellow house bats, S. dinganii, from Zimbabwe (Smithers & Wilson, 1979) TL T Hfc/u E F/a Mass 130 52 12 16 55 27,1 Males Females n Range X n Range 15 113-137 131 14 123-141 15 46-55 54 14 50-57 15 11-14 12 14 11-15 15 12-18 17 14 15-19 15 53-56 55 14 52-57 14 21,3-36,0 27,3 14 21,9-37,9 Distribution Extralimital to the continent Recorded from the extreme southwestern parts of Arabia and from the islands of Madagascar, Mauritius and Reunion, if S. robustus Milne-Edwards, 1881 is conspecific. South of the Sahara, excluding the Southern African Subregion Recorded from Senegal; Niger; Nigeria; Chad eastwards to Ethiopia and Somalia and southward in Uganda; Kenya; Tanzania; Zaire excluding the forest; Congo Republic; Angola; Zambia and Malawi. Not at present recorded from Mozambique, north of the Zambezi River, but likely to occur. Southern African Subregion Recorded from northern Namibia as far south as Windhoek; from northern and eastern Botswana; Zimbabwe; parts of the Transvaal; Mozambique, south of the Zambezi River; Natal and the eastern Cape Province where they occur along the south coast westwards to about 26 °E. There is an isolated record from the Kalahari Gemsbok National Park (Erasmus & Rautenbach, 1984) which suggests that they may have a wider distribution in this sector than formerly supposed. So far they have not been found in the northeastern Transvaal and adjacent parts of Zimbabwe and Mozambique. Fig. 88.1. Ear tragus: (a) Scotophilus dinganii (b) Nycticeius schlieffenii 113 Fig M.2 Front viow of tho skull of Scotophifus dinganii to show tho single uppor incisor on olther side and the canines. Habitat This is a savanna woodland species which is absent from desert areas and forest, although it occurs on forest fringes. Habits They are gregarious, but occur in small numbers of up to about a dozen together. They roost during the day tucked together into narrow crevices in the brickwork under the roofs of houses, or between the overlapping corrugated iron sheets. Numbers of these small clusters may be found under a single roof, the total numbers present amounting to dozens. They will also use hollow trees and have been taken tucked into crevices in the thatch inside disused rondavels. They have a tendency to be associated with built-up areas, occurring even in the peri-urban and urban surroundings of cities or in farm buildings. They quite often enter houses at night in foraging or fly around lights at night catching insects. They are fast fliers and are inclined to fly low over the ground. In Zimbabwe it was noticeable that they were absent from the plateau (1 200-1 500 m) during the colder months of the year from June to August, yet were present at lower altitudes (600 m) in the southeast of the country. Food Insectivorous. Reproduction In Zimbabwe gravid females were taken in September and October, and lariating and females with young clinging to them in November and December. Out of a sample of seven 1, 'TT 'hey ,were ""Ting twin foetuses in every nTTn ln«^n ^rrST RaLu,enbach H982) reported that out •t four gravid females taken in October, three carried twin dunnTMrl"6 ,nP Th,eS? observat'ons of parturition during early summer and the birth of twins have been (unp.Tl'f Pcnzhorn- ^"tenbach & van der Merwe, N.J. 1803) No 89 Scotophilus borbonicus (E. Ceoffroy, Lesser yellow house bat Klein geel dakvlermuis Colloquial Name T*J| fc smallest ye,,„K hoUM fa Taxonotnic Notes subsries as °™™8 —.su titrate 5 114 the remainder of the species range, S. b. borbonicus being extralimital. Description As the name implies the lesser yellow house bat is smaller than its near relative, the yellow house bat. The species has a total length of about 120 mm, with a tail of 46 mm and a mass of about 16 g (Table 89.1). The difference in size between the two species shows best in the length of the forearm which in this species ranges from 46-50 mm, in the yellow house bat from 52-57 mm. The colour of the upper parts varies from a light to a dark yellowish-brown, the under parts in the lighter coloured specimens being white, in the darker a greyish-white. In some specimens the under parts may be tinged yellow or there may be traces of yellow on the flanks. Because of the variation in colour the best criterion for distinguishing the two species is from their sizes. Table 89.1 Measurements (mm) and mass (g) of lesser yellow house bats, S. borbonicus, from Zimbabwe (Smithers & Wilson, 1979) Irrespective of sex X n Range TL 115 10 111-125 T 46 10 40-52 Hf c/u 11 10 10-15 E 14 10 10-16 F/a 48 6 46-50 Mass 15,7 7 13,0-18,0 Distribution The nominate S. b. borbonicus was described for Reunion Island and also occurs on Madagascar. South of the Sahara, excluding the Southern African Subregion Recorded from the Sudan; Kenya; southeastern Tanzania; Mozambique, north of the Zambezi River; Malawi and Zambia. While there are no records at the moment from ngola or Zaire, it seems likely, owing to their occurrence in adjacent parts of Zambia, that in time they will be found to occur. Southern African Subregion Recorded in northeastern Namibia; in Zimbabwe, excluding 71°Sk° • r.6 central P^teau; in Mozambique, south of the v17eZ' Rlv„er;. northern Botswana, in parts of the Transvaal and in Zululand. Plate 5 ivahlbergiS e^au^ettec^ b"uit bat, Epomophorus 49 St*!101*1"1" ' mauritianus 63. Egyptian free.tailed ^ (Tadarjda) aegyp] Egiptiese losstertvlermuis WtS' My°tis -elmtschii 77 Z^Sel-"„nuiS SS^aIino,obisi'~s ^el°dakvlermuis' Sc°tophiius dinSanii 98' Srn°ensSDt^ed b,at' Nycteris 100 HibS sPleetneusvlermuis brandti? horseshoe bat, Rhinolophus hild HO. Commbpandt.Se, saalneusvlermuis soni FSOn S af"nosed bat, Hipposideros comme Commersonsebladneusvlermuis PLATE 5 Habitat A savanna woodland species, in Zimbabwe restricted to altitudes of below about 1 000 m above sea level, although in the Transvaal this does not hold. They appear to be associated with riverine conditions or to areas with a mean annual rainfall in excess of 500 mm. Habits Little is known about the habits of this species except that they roost in small colonies in Zimbabwe in hollow trees. Cotterill & Giddings (1987) in netting at Tashinga in the Matusadona National Park, Zimbabwe, on the shores of Lake Kariba, found that they were active from 19h00 to 21h00, the peak of the male activity being 21h00 to 22h00, and of the females from 19h00 to 21h00. As the majority of the females were lactating, which involves added energy demands, and as insect abundance tends to be the greatest immediately after sundown, it was presumed that the females were benefitting by feeding early. In Zimbabwe, where they were using holes in trees, they changed roost trees from one day to the next and frequently moved between night and day roosts, usually staying, however, within a relatively small area of woodland (Fenton 1983; Fenton, Brigham, Mills & Rautenbach, 1985; Fenton & Rautenbach, 1986). Individuals feed from about an hour following sunset, after which they return to their day-time roosts. Very little activity occurs during the rest of the night. They commute over considerable distances to favourable feeding sites. Food Insectivorous. Reproduction Of 18 females collected in Zimbabwe in November, nine were gravid, each carrying two foetuses. Cotterill & Giddings (1987) recorded that a high percentage of 46 females taken at Lake Kariba were lactating in December. Van der Merwe, Rautenbach & Penzhorn (1988) recorded parturition during November/December, with a high incidence of twins delivered. Genus Nycticeius Rafinesque, 1819 No. 90 Nycticeius schlieffenii (Peters, 1859) Schlieffen's bat Schlieffen se vlermuis Plate 4 Colloquial Name Named after the collector Count Wilhelm von SchlieffenSchlieffenburg. Taxonomic Notes Meester et al. (1986) listed two subspecies from the Subregion, N. s. australis (Thomas & Wroughton, 1908) from Zululand; southern Mozambique; the eastern and northern Transvaal and Zimbabwe; and N. s. fitzsimonsi (Roberts, 1932) from northern Botswana and northern Namibia. Description Schlieffen's bats have a total length of about 75 mm, with tails 30 mm in length and a mean mass of 4,6 g (Table 90.1). The upper parts are light fawn, the individual hairs unicolour or only slightly lighter in shade at their bases, the under parts slightly paler. In contrast to the body colour, the wing and interfemoral membranes are dark brown and tend to show a series of dark, roughly parallel striations across them. The ears are strongly convex on their inner edges, concave on the outer, the tips rounded. The ear tragus is of characteristic shape (Fig. 88.1.b). The dental formula is If Cf Pf Mf = 30 The single premolar in the upper jaw is sharply pointed and about three-quarters the height of the canine. The anterior lower premolar is half the size of the posterior one and tightly packed between it and the canine. Table 90.1 Measurements (mm) and mass (g) of Schlieffen's bats, N. s. australis, from Zimbabwe (Smithers & Wilson, 1979), and N. s. fitzsimonsi from Botswana (Smithers, 1971) Irrespective of sex N. s. australis N. s. fitzsimonsi TL X n Range X n Range TL 75 12 69-80 71 10 64-78 T 30 12 28-31 29 10 24-30 Hf c/u 7 12 6-8 7 10 5-9 E 10 12 9-11 11 10 10-13 F/a 30 12 29-31 29 10 28-30 Mass 4.6 9 3,7-5,0 Distribution Schlieffen's bats have a wide distribution on the continent from Egypt southwards to Ethiopia and Somalia and westwards to Mali and south of this to northern Namibia and Zululand. Extralimital to the continent Recorded from the southwestern parts of Arabia. North of the Sahara Occurs along the western borders of the Red Sea and the Gulf of Suez to the delta of the Nile in Egypt. South of the Sahara, excluding the southern African Subregion Recorded from Mali; the northern parts of Ghana; southern Mauritania; northern Nigeria; Niger; Chad; northeastern and southern Sudan; Ethiopia; Somalia; parts of Uganda; Kenya; northeastern and southern Zaire; Tanzania; Angola; Zambia and Malawi. So far not recorded from the northeast117 ern parts of Mozambique, but little collecting has taken place there and they probably have been overlooked. Genus Scotoecus Thomas, 1901 Southern African Subregion They occur in northern Namibia; in the northern and northeastern parts of Botswana; in Zimbabwe and in Mozambique, south of the Zambezi River. In the Transvaal they occur in the north and east and southwards to the northern parts of Zululand. Habitat In West Africa they are absent from the High Forest Zone and parts of the Guinea savanna, tending to be associated with the more arid Sudan and Sahel savannas. In the more southerly parts of their distributional range they are associated with savanna woodland, but do not occur in the more arid open parts of the South West Arid Zone, nor locally in forest. In the eastern parts of the Subregion they are commoner at altitudes of less than 1 200 m. Habits A solitary species which roosts during the day in houses, huts and cellars (Rosevear, 1965) and has been taken in narrow crevices in branches of trees (Verschuren, 1957). Numbers occasionally congregate to forage. Roberts (1951) recorded that they emerge from their roosting places before dusk and have an erratic flight. They occasionally enter houses while foraging. Most of the material from the Subregion was netted or shot and therefore there is no record of their habits. Food Insectivorous. Reproduction Van der Merwe & Rautenbach (1986, 1987) recorded that, in the Transvaal, mating takes place during the winter months, predominantly in June. Spermatogenesis extends over a 10-month period with the first signs of spermatozoa in the epididymides by the end of April. Spermatozoa were present in the epididymides from the end of April until the beginning of September. Copulation begins during June (early winter) and the females have spermatozoa in the uterine horns from then until the end of August (late winter) when ovulation occurs. These bats are seasonally monoestrus with the great majority of births occurring during November. The number of conceptuses varied; a maximum of 5 pre-implanted embryos was recorded, but the maximum number of foetuses observed was 3. No. 91 Scotoecus albofuscus (Thomas, 1890) Thomas' house bat Thomas se vlermuis Colloquial Name Named after O. Thomas who described the species. Taxonomic Notes Meester et al. (1986) listed one subspecies, S. a. woodii Thomas, 1917 from the Subregion. The genus is subject to variability in its dental formula. Some individuals retain the tiny upper premolars that lie immediately behind the canines, in others they are missing. When they are present these tiny teeth are tucked away internal to the toothrow. In this species the broad rostrum, the shape of the ear tragus, and long bony penis, which reaches a length of a quarter of that of the head and body, mark them as clearly different from Schlieffen's bats, Nycticeius schlieffenii. S. a. albofuscus occurs in West Africa. Description Thomas' bats have a total length of some 70 mm, a mass of about 4,5 g and forearms of 30-31 mm. The colour of the upper parts is a light fawn-brown, the under parts paler and lighter still in colour on the chest. The wing membranes are white, brownish towards the edges, the interfemoral membrane brown. The ears are oval with broad rounded points, the tragus short and rounded. The dental formula is If Cf Pf Mf = 30 or when the anterior upper premolar is present If Cf Pf Mf = 32 Distribution At the moment there are only very few, scattered records from the continent and so little is known about this species that it is impossible to show its limits of distribution with accuracy. South of the Sahara, excluding the Southern African Subregion There are records from Senegal; Gambia; Nigeria; Kenya and the Southern Province of Malawi. Southern African Subregion So far recorded only from the Zinave National Park on the Save River in southern Mozambique. Habitat, Habits, Food and Reproduction Unknown. Subfamily KERIVOULINAE Genus Kerivoula Gray, 1842 All members of this genus are characterised by their long, soft, woolly fur which stands erect from the body, the tips of the individual hairs curled and often paler than the general colour, giving the body a grizzled appearance. In the skull the high, rounded braincase rises steeply from the long rostrum which is characteristic of members of this genus (Fig. 93.1). The dental formula is If Cf Pf Mf = 38 and is therefore the same as Myotis spp with three premolars on each side above and below. The two anterior upper 118 premolars are smaller than the third (Fig. 93.1) and the lower are the same size. The inner two lower incisors are trilobed, the outer conical. Key to the species, after Meester et al. (1986) (Measurements in mm) 1. Size larger, forearm 34-39; total length of skull 15 or more; colour of upper parts reddish-chestnut, under parts either whitish or white tinged buffy . . . argentata Size smaller, forearm 30-36; total length of skull 13,5 or less; colour brownish, greyish-brown, sometimes lightly grizzled with greyish-white or white, the under parts brown to greyish-brown to greyish-white to white .. . lanosa No. 92 Kerivoula argentata Tomes, i86i Damara woolly bat Damara-wolhaarvlermuis Colloquial Name Originally described from a specimen from Damaraland, Namibia; the soft, curly, woolly hair is characteristic of members of this genus. Taxonomic Notes Meester et al. (1986) listed three subspecies from the Subregion: K. a. argentata from northern Namibia, K. a. nidicola (Kirk, 1865) from central Mozambique and K. a. zuluensis Roberts, 1924 from northern Natal and Zululand. The paucity of material has rendered it difficult for authorities to determine the relationships of a number of species of this genus. Description The Damara woolly bat is the larger of the two species of Kerivoula that occurs in the Subregion. About 95 mm in total length, including the tail of about 47 mm in length, they have a mass of between 6,0 and 9,0 g (Table 92.1). The colour of the upper parts is a rich brown, many of the soft woolly hairs with silvery tips giving an overall grizzled appearance to the pelage. The individual hairs are dark grey at their bases, then broadly buffy with brown tips. The under parts are greyish-brown, tending to be lighter in colour towards the anus and on the sides of the belly. The wing and interfemoral membranes are brown, the latter lighter brown than the wing membranes. There is a sparse covering of hair on the interfemoral membranes, but a very distinct fringe of hair on their hind margins. The fringe near the point of the tail is comprised of stiff hair hooked inwards towards the tips. The ear tragus is long, thin and tapers to a point (Fig. 92.1.a). Table 92.1 Measurements (mm) and mass (g) of six Damara woolly bats, K. argentata, from Zimbabwe (Smithers & Wilson, 1979) Irrespective of sex X n Range TL 93 6 83-100 T 47 6 42-50 Hf c/u 10 6 10-11 E 14 6 13-15 F/a 37 6 36-41 Mass 7,6 6 6,0-9,0 (a) K. argentata (b) K. lanosa Distribution Throughout their distributional range the records are scattered which makes it difficult at the moment to assess the limits of their distribution. South of the Sahara, excluding the Southern African Subregion They are recorded from central Angola; parts of Zambia, and Malawi. There are only two records from Mozambique, north of the Zambezi River, these from the lower Zambezi River in the southern Zambezi District. Northwards there is a single record from central Tanzania and others from southern Kenya and southern Zaire. Southern African Subregion Originally the species was described from a specimen from Otjoro, Ovamboland, Namibia, and they occur in the eastern half of Zimbabwe and Mozambique, south of the Zambezi river, south to the Maputo District. They have been taken in Zululand and in Natal south to the White Umfolozi River, and in the Pafuri region of the Kruger National Park. Habitat A savanna woodland species with a tendency in the Subregion to be confined to well-watered areas or riverine associations in dry country. Habits Damara woolly bats are solitary or may occur in small groups numbering up to about six. Their habits have not been studied in detail and there are only casual observations on record. Shortridge (1934) stated that they roost singly or in pairs in exposed situations among clusters of dead leaves, on the rough bark of trees or in deserted birds nests. He recorded that in Zambia four were found clinging 119 so closely together under the eaves of a rondavel that they looked like the mud nest of a wasp. In Zimbabwe 12 of the total number of specimens numbering 15 were taken in the disused nests of masked weavers, Ploceus velatus, and Roberts (1951) recorded them in Zululand from the nests of spectacled weavers, Ploceus ocularis. In two cases in Zimbabwe there were three tightly packed in such a nest (Smithers & Wilson, 1979). Wright (in litt.) in Fort Victoria, Zimbabwe, collected five, all females, from a tight cluster on the outside wall of a rondavel, sheltered by the overhanging eaves. They have a slow fluttering flight and when foraging fly within two or three metres of the ground. Food Insectivorous. Reproduction There is no information available from the Subregion. No. 93 Kerivoula lanosa (A. Smith, 1847) Lesser woolly bat Klein wolhaarvlermuis Plate 4 Colloquial Name As can be seen from a comparison of the measurements and masses of K. argentata and this species, K. lanosa is all round smaller and less in mass. Taxonomic Notes Meester et al. (1986) recognised two subspecies in the Subregion: K. 1. lanosa from the southern and eastern coastal regions of the Cape Province extending eastwards to the Ciskei and K. 1. lucia Hinton, 1920 from northern Natal and Zululand, the Transvaal, eastern Zimbabwe and adjacent parts of Mozambique and northern Botswana. Description The lesser woolly bat is the smaller of the two species that occur in the Subregion. They have a total length of about 80 mm including a tail some 36 mm long and a mass of 6,0-8,0 g (Table 93.1). The upper parts are a dull huffy colour, the hair long, springy and curled which makes it stand out from the body. The bases of the individual hairs are dark in colour, the tips broadly paler and closely curled, giving the upper parts a grizzled appearance. The ears are very broad, the inner and outer edges curling inwards to give them a funnel shape, the ear tragus long and thin (Fig. 92.1 .b). The under parts are either white on the chest and belly or a lighter shade of buff, the throat the same colour as the upper parts. The wing and interfemoral membranes are brown, the curly hair extending sparsely along the forearm and fringes of the wing, on the tail and fringes of the interfemoral membrane. There is a very noticeable fringe of hairs on the hind edge of the interfemoral membrane, these hairs curving inwards like hooks. This fringe is a characteristic feature of bats of this genus. The ankle spur or calcaneum that supports the outer fringes of the interfemoral membrane is at least as long as the length of the tibia and foot, the membrane extending far beyond the feet. The shape of the skull is characteristic of members of this genus (Fig. 93.1). Fig. 93.1. Skull of Kerivoula lanosa Table 93.1 Measurements (mm) and mass (g) of lesser woolly bats, K. lanosa, from Botswana (Smithers, 1971) TL T Hfc/u E Length ear tragus Mass Male 77 37 7 13 9 6,0 Female 80 39 7 13 9 7,0 Female 78 37 8 14 9 6,0 Female 78 35 8 13 8 8,0 Distribution So few scattered records from the southern parts of the distributional range of the lesser woolly bat are available that it is impossible at this stage to present their limits of occurrence. Kingdon (1974) showed that they are distributed widely in East Africa, but Swynnerton & Hayman (1950) only listed K. africana Dobson, 1878 which occurs in the extreme northeast of Tanzania. South of the Sahara, excluding the Southern African Subregion Recorded from Liberia; Ivory Coast; Ghana; Gabon; Cameroun; Central African Republic; Ethiopia; Kenya; Tanzania; eastern and southern Zaire; southern Malawi and Zambia. Their occurrence on the Okavango River in northwestern Botswana suggests that in time they may be shown to occur in southeastern Angola in riverine situations. Southern African Subregion They occur in the riverine woodland of the Okavango River in the extreme northwestern parts of Botswana and have been taken at Sepopa on the western fringe of the Okavango Delta and near Molepolole in the southeast of the country. In Zimbabwe they occur on the Mozambique border east of Mutare. In the Transvaal recorded from the Njelele River, Soutpansberg as well as from Pafuri, in Zululand from 120 Ingwavuma and in the Cape Province from the Pirie Forest, King William's Town and Knysna. Habitat There is a distinct tendency for this species to be associated with riverine associations in dry country as well as occurring in well-watered areas. Roberts (1951) reported that a specimen from Zululand was taken in a forest, as was the King William's Town specimen which was collected in the Pirie Forest. Habits Little is known about the habits of this species. They have a slow fluttering flight like K. argentata and the same habit of using disused birds' nests to roost in during daylight hours. In northwestern Botswana a male and two females were taken from the nest of a masked weaver, Ploceus velatus, and a single female from the nest of a scarlet chested sunbird, Nectarinia senegalensis. In Zululand, Roberts (1951) collected specimens from the nests of spectacled weavers, Ploceus ocularis, three in one nest, two in the other. Food Insectivorous. Reproduction No information available from the Subregion. IX. Family NYCTERIDAE Slit-faced bats The Family gets its name from the Greek name for a bat nycteris. Members of the Family all have a slit from the nostrils to above the eyes, which overlies a concavity in the skull enclosing the complicated noseleaves which, unlike those in the Rhinolophidae, are only visible when the slit is open (Fig. IX.1). These noseleaves serve in echolocation, the emissions in members of this Family being through the nostrils and not through the mouth. A characteristic feature of all members is the large ears which make them easily identifiable (Fig. IX.1). In the common slit-faced bat, Nycteris thebaica, which is common throughout the Subregion, the ears reach a length of up to 37 mm. They have rounded ends and when the sides are bent forward the ears are nearly parallel-sided. The shape of the ear tragus is a useful character in the recognition of the species. They all have a small rounded protuberance on the lower lip, with a raised ridge on either side, the ridges converging but not meeting posteriorly. They have long tails, the whole of which is contained within the interfemoral membrane, the last vertebra bifurcating at its joint in a Y with broad open ends which help support the fringe of the membrane. This membrane is further supported by long calcanea arising from the ankles of the hind feet (Fig. IX.2). All members of the Family have long, soft fur, reddishbrown or greyish-brown in colour, and a bright orange phase is known to occur. The wings are broad and rounded at the ends. The skull is elongated and characterised by the deep concave depression in front which houses the noseleaves (Fig. IX.3). The dental formula is If Cj P| M§ = 32. The single upper premolar is well developed, the relative size of the posterior lower premolar in relation to the molar next to it is a useful character in separating the groups of species. Slit-faced bats are distributed widely throughout Africa and occur beyond the borders of the continent to Madagascar, Arabia, southeastern Asia and Malaya, being most abundant in the tropics. They are expert fliers and their powers of echolocation must be well developed as they are adept at avoiding being caught in mist nets, swinging away from them and over the top whereas other species fly directly into them. After having caught their prey they hang on temporary feeding perches where their prey is eaten, the unpalatable portions accumulating below them, forming useful indications of what they are eating. The Family is represented in the Subregion by a single genus and six species. Fig. IX.1. Head of a Nycteridae: Nycteris thebaica 121 Fig. IX.2. Tail of a Nycteris thebaica (a) calcaneum. Genus Nycteris G. Cuvier & E. Geoffroy, 1795 Key to the species after Meester et al. (1986) (Measurements in mm) 1. Upper incisors trifid (Fig. 94.1) ... 2 Upper incisors bifid ... 3 2. Forearm 36-45, ear 18-25; condylocanine length less than 18 ... hispida Forearm 57-66, ear 28-35; condylocanine length more than 18 .. . grandis 3. Ear tragus semi-lunate, base of its posterior margin not notched (aethiopica group) ... 4 Ear tragus pyriform, base of its posterior margin shallowly notched (thebaica group) ... 5 4. General colour greyish; forearm 37-42, ear 29-34 . .. woodi General colour brownish; forearm 45-50, ear over 21 ... macrotis 5. Size smaller, forearm, 42-52, ears 28-37; skull less massive, breadth of supraorbital flanges about 6,8 ... thebaica Size larger, forearm over 50, ears about 22; skull massive, rostrum 20% heavier than in N. thebaica, breadth of supraorbital flanges about 9,0 ... vinsoni Smithers (1983) noted that the ear length of N. vinsoni should be treated with caution as the ears of the type specimen were singed by fire (see Dalquest 1965) Hispida group No. 94 Nycteris hispida (Schreber, 1774) Hairy slit-faced bat Harige spleetneusvlermuis Colloquial Name This species is no more hairy than any other ol the slit-faced bats, the name having applied to it as a translation of the specific name hispida which is from the Latin hispidus, hairy or bristly, referring to the bristly hair lying around the facial slit. Taxonomic Notes The nominate N. h. hispida originally described from Senegal occurs throughout the northern parts of the species range, being replaced in the Subregion by N. h. villosa Peters, 1852. Description Hairy slit-faced bats have a total length of about 90 mm and a wingspan of about 280 mm (Table 94.1). The upper parts are sepia-brown, the under parts much paler than the upper parts and usually greyer. They and the large slit-faced bat, N. grandis, are the only two members of the Nycteridae in which the upper incisors have three lobes on their cutting edges (Fig. 94.1). The ear tragus, which is blunter than in the large slit-faced bat, is similar in that the outer edge is smooth. The ears and membranes on the wings are dark blackish-brown, the ears and tail long but not as long as in the common slit-faced bat, N. thebaica. Table 94.1 Measurements (mm) of hairy slit-faced bats, N. hispida (Rosevear, 1965) HB T E F/a 40-54 43-52 18-25 36-45 Distribution In the northern parts of their distributional range hairy slit-faced bats are one of the most commonly collected species and have one of the widest ranges. They occur from Senegal in West Africa eastwards to Ethiopia and south to the Subregion. Roberts (1951) listed two specimens from the "Cape of Good Hope" collected by Andrew Smith, but queried this locality as, at the time of publication of his work, there were no other records from southern Africa except those from Inhambane, Mozambique. He thought they might have been taken by A. Smith during his expedition to Zululand in 1831 for they do not occur anywhere 122 near the Cape. In the southern parts of their range they are not common. South of the Sahara, excluding the Southern African Subregion Hairy slit-faced bats occur from Senegal eastwards to Ethiopia and Somalia and southwards in Uganda; Kenya; Tanzania; Zaire; Cameroun; Gabon and in parts of Angola. They occur in the central and southwestern parts of Zambia and in the northwest and, while there are no records at the moment from the eastern parts, Ansell (1978) believed that they do occur, as they are found in the Southern Province of Malawi. They occur in the Tete District of Mozambique, north of the Zambezi River. Southern African Subregion While there are no records from Namibia, their occurrence in Angola as far southwest as Mossamedes suggests that in time they will be found to occur there. In Zimbabwe they are recorded from the Zambezi Valley in the northwest and in the Eastern Districts. They have been taken in Mozambique, south of the Zambezi River, from the Tete, Vila Pery and Inhambane districts. catch individuals in a room appreciated the agility with which they evaded capture in a butterfly net, calling it a "warm and exhilarating sport". Like the common slit-faced bat, N. thebaica, they are adept at avoiding capture in bird nets, swinging vertically up the face of the net at the last minute and escaping over the top. Although like other nycterids they are slow fliers, they appear to have very acutely tuned echolocation systems, and have the ability to jink and turn which enables them to penetrate deep into thick bush either in seeking roosting sites or in feeding in forest underbrush. Kingdon (1974) recorded that they become attracted to a roost and when disturbed from it return later to the exact same spot. They remain alert during the day and if disturbed, will flit around nearby bushes before returning to settle. Food Insectivorous. Reproduction There is no information from the Subregion. Habitat Hairy slit-faced bats are catholic in their habitat requirements, occurring in the High Forests of West Africa and Zaire as well as in relatively dry savanna woodland, as in Mozambique. Rosevear (1965) recorded their wide tolerance to a diverse range of vegetational associations and their habits show that they can use as diverse a range of shelters, not being restricted to any special type which in other species restricts their occurrence. Their distribution suggests that they avoid arid country. Habits Hairy slit-faced bats are predominantly solitary, but have been observed in small colonies numbering up to about 20 (Rosevear, 1965). They roost during the day in the shelter of dense, low bushes, hanging from the finer twigs, but are also found in thatched African huts, hanging from the thatch or thinner projections of the roofing timbers. Kingdon (1974) recorded that, in East Africa, they also use aardvark holes, holes in termitaria, hollow trees and papyrus crowns. In Mozambique two were taken in a cave in granite with a wet, muddy floor (Smithers & Wilson, 1979). They come to lights at night to hawk insects and frequently enter houses when foraging. Allen, Lang & Chapin (1917) in their endeavours to No. 95 Nycteris grandis Peters, 1865 Large slit-faced bat Groot spleetneusvlermuis Colloquial Name This is the largest member of the Family, all of which have a slit on top of the muzzle, from the nostrils to the base of the ears, under which lies the complicated noseleaves accommodated in a cup-like depression on the front of the skull. Taxonomic Notes The nominate N. g. grandis was described from Guinea, West Africa; only N. g. marica Kershaw, 1923 occurs in the southern parts of the species' range. Description The greater slit-faced bat is, as the name implies, the largest of the Nycteridae, adult males having a total length of about 160 mm and a mass of about 40 g (Table 95.1). They are light reddish-brown in colour, the under parts lighter in colour than the upper parts with a grey wash. The fur is long and soft, the wing membranes dark brown or nearly black and criss-crossed with fine parallel lines or dots. Their wings are broad and rounded at the ends, their ears broad and measuring up to 30 mm long. In common with N. hispida the upper incisors have three small lobes on their cutting edge (Fig. 94.1); the outer margins of the ear tragi are smooth. Although not so far seen in the Subregion, bright orange-coloured phases of this species are known. Table 95.1 Measurements (mm) and mass (g) of two male and one female large slit-faced bats, N. grandis, from Zimbabwe Males TL T Hfc/u E F/a Mass 164 84 15 31 64 35,7 157 70 15 31 64 42,5 Female 145 64 15 31 63 — Distribution In the southern parts of their range they probably occur more widely than present records indicate, as they probably do in Tanzania where at the moment there are no records between 123 the northwestern parts of the country, Dar-es-Salaam and the offshore islands. South of the Sahara, excluding the Southern African Subregion Recorded from Guinea eastward in most of the West African countries to Cameroun; Zaire, except in the southeast; Uganda; northeastern Tanzania and in the Dar-es-Salaam area and the offshore islands of Zanzibar and Pemba. In Zambia there is a record from the southern sector of the Luangwa Valley, and they occur in Malawi. Southern African Subregion In Mozambique, south of the Zambezi River, they are recorded from the western parts of the Vila Pery District adjacent to the border with Zimbabwe and may well occur in the Dondo and Amatongas forests in this sector. In Zimbabwe they occur in the Zambezi Valley east of Lake Kariba. Food Predominantly insectivorous, but with some carnivorous tendencies. In Zimbabwe, the discarded remains of food included much insect material, but also the remains of small fish and frogs Rana sp. It was thought that the fish were being taken from pools cut off from the river and then drying up. They used two types of perches, the hunting perch from which they took off to fly low over the shallow fringes of water, and the feeding perch to which they returned to consume the prey (Fenton et ah, 1987). Reproduction Fenton et al. (1987) recorded females each with a single clinging young in December in the Zambezi Valley at Mana Pools National Park, Zimbabwe. No. 96 Nycteris woodi K. Andersen, 1914 Wood's slit-faced bat Wood se spleetneusvlermuis Colloquial Name Named after Rodney C. Wood, a well-known naturalist and collector, who collected the original specimens at Chilanga, near Lusaka, Zambia. Taxonomic Notes Meester et al. (1986) listed two subspecies, N. w. woodi from Zambia and N. w. sabiensis Roberts, 1946 from Zimbabwe. Description A slightly smaller species than the common slit-faced bat, N. thebaica, this species is about 90 mm in total length with ears slightly shorter in series than N. thebaica (Table 96.1). The upper parts of the body are dark brown, the individual hairs grey towards their bases and about 10 mm long. The under parts are whitish tinged grey. The skull is typical of the genus with the concave depression in front which houses the noseleaves (Fig. IX.3). Habitat The greater slit-faced bat in the northern parts of its distributional range occurs in forest and the indications from the Mozambique specimens are that they are similarly associated. In Zimbabwe and Zambia, however, they are found in the well-developed evergreen riverine forests of the Luangwa and Zambezi rivers and should be looked for in other areas with this type of habitat in the southern parts of their range. Habits They occur in small colonies of up to about six individuals. Ansell (1978) mentioned that the Luangwa Valley specimens were taken from a colony, not recording the numbers. They roost during the day in hollow trees, one of the Zimbabwe specimens being taken from the hollow trunk ot a dead tree lying on the ground, a situation from which they have been collected in other parts of their range (Liberia; Allen & Coolidge, 1930). They have also been taken m culverts and shallow caverns in rocks (Rosevear, 1965). Fenton, Cumming, Hutton & Swanepoel (1987) recorded a colony of five roosting in a water tower in Mana Pools National Park, Zimbabwe. They return to the same feeding site to consume their food. At Mana Pools, Zimbabwe one regularly returned to hang up in the office of the Departmen of National Parks, scattering the discarded remains ot its food below the feeding site. Table 96.1 Measurements (mm) of Wood's slit-faced bats, N. woodi, from Zimbabwe (Smithers & Wilson, 1979) Males Females TL T Hf c/u E F/a 91 43 9 31 38 Range X n Range 88-93 91 5 88-96 41-46 46 5 44-49 8-10 9 5 8-10 29-33 33 5 31-35 37-39 39 5 38-40 Distribution Wood's slit-faced bats are confined in their distribution to central and eastern Zambia; to northeastern and southeastern Zimbabwe and to the Limpopo River valley and the northern parts of the Kruger National Park in the Transvaal. Habitat A savanna woodland species. Habits The Zimbabwe specimens were all netted and consequently there is no information on their daylight roosting places. In eastern Zambia they were roosting in rock fissures in a large rocky outcrop and one was taken in a building (Ansell, 1967a). 124 Food Insectivorous. Reproduction No information is available. No. 97 Nycteris macrotis Dobson, 1876 Greater slit-faced bat Groter spleetneusvlermuis Colloquial Name Although the specific name macrotis is derived from the Greek makros and otis meaning large eared, their ears are in fact smaller than in species such as N. thebaica. Taxonomic Notes The nominate N. m. macrotis was described from Guinea, West Africa; only N. m. oriana Kershaw, 1922 occurs in the Subregion and then only marginally in northeastern Zimbabwe. Description Greater slit-faced bats have a total length of about 100 mm (Table 97.1). The upper parts are warm reddish-sepia in colour, the under parts greyer. Like the common slit-faced bat N. thebaica the upper incisors have two lobes on their cutting edges, but the ear tragus is broader at the end and not so rounded and lacks the indentation on the lower part of the outer edge seen in N. thebaica. Table 97.1 Measurements (mm) of greater slit-faced bats, N. macrotis, from West Africa (Rosevear, 1965) HB 52-70 E 28-30 T 40-60 F/a 45-50 Distribution South of the Sahara, excluding the Southern African Subregion Recorded from Gambia eastwards to Nigeria; southern Sudan; western Ethiopia; Uganda; Kenya and Tanzania. They occur in northeastern Angola, in the central and parts of eastern Zambia and in Malawi. There are no records at the moment from northeastern Mozambique. Southern African Subregion So far recorded only from the Zambezi Valley in the northeast at the confluence of the Chewore and Zambezi Rivers and in the Matusadona National Park, Lake Kariba, in Zimbabwe. Habitat The specimens from the Subregion were netted in the riverine forest of the Zambezi River. In other parts of Africa they occur both in forest and in savanna woodland. Habits In other parts of Africa they have been taken roosting in caves, culverts and dark cellars. They will also hang up in rondavels. Rosevear (1965) believed that they may also roost hanging in bushes or under rocks like N. hispida. Food Insectivorous. Reproduction There is no information available from the southern parts of their distributional range. No. 98 Nycteris thebaica E. Geoffroy, 1813 Common slit-faced bat Gewone spleetneusvlermuis Plate 5 Colloquial Name This is one of the commonest and most widespread small insectivorous bats found in the Subregion. 125 Taxonomic Notes Numbers of subspecies have been described, but their validity remains uncertain. Meester et ai (1986) recognised two subspecies from the Subregion: N. t. capensis A. Smith, 1829 from the Cape Province eastwards to southern Mozambique, the Transvaal and Zimbabwe, and N. t. damarensis Peters, 1871 from the western parts of the species' range including northern Botswana. Description The conspicuous long ears make this species one of the more easily recognisable species that occur in the Subregion. Adult males have a total length of about 100 mm and a mean mass of 10,5 g, females, in series, slightly larger and heavier (Table 98.1). „ , . , , The upper parts of the body are buffy-brown, the base ot the hair slate-grey, the hairs on the sides of the neck and head with buffy bases. The under parts of the body are bufty or off-white. The long ears and the wing membranes are light brown. The hair is long and soft and on both the upper and under parts extends onto the bases of the forearms and on the under parts onto the wing membranes near the body. The light-coloured hair of the under parts extends upward onto the sides of the neck. Occasionally a rufous-coloured phase is encountered in the eastern parts of their distributional range. Specimens from Namibia are noticeably paler above and whiter below and have been considered as worthy ot subspecific rank as N. t. damarensis. Unlike N. grandis and N. hispida the upper incisors of N. thebaica have only two lobes on their cutting edge as against three in the other two species. The ear tragus is pear-shaped with an indentation on the lower part of its outer edge. Table 98.1 Measurements (mm) and mass (g) of the common slit-faced bat, N. thebaica, from the northern parts of the Subregion (Smithers & Wilson, 1979) TL T Hf c/u E Mass Distribution The common slit-faced bat has a wide distribution on the continent from parts of North Africa to the Cape Province. They also occur in Europe, having been recorded on the Island of Corfu, off the west coast of Greece, and in the Middle East. There are areas within the Subregion from which to date they have not been found to occur. As they range widely, this may be due to their having been overlooked; on the other hand they prefer woodland and some ot these areas such as southern Botswana and the grassland areas of the Transvaal may be too open for them. They occur throughout their range in association with buildings, yet examination of scattered stone buildings in southern Botswana did not reveal their presence. North Africa They occur coastally in Morocco; Algeria; Tunisia and Egypt and in Libya have been taken at Zuara in the Tibetsi Massif deep into the Sahara. South of the Sahara, excluding the Southern African Subregion They occur from Senegal eastwards to the Sudan; Ethiopia and parts of Somalia and northwards to Egypt and southwards in at least parts of all the countries to the borders ot the Subregion, excluding areas of tropical forest. Southern African Subregion Recorded as occurring widely in Namibia but in Botswana confined to the northern and eastern sectors and not so far recorded from the semi-desert central and southern parts of the country. Widespread in Zimbabwe; Mozambique, south of the Zambezi River, and Malawi. In the Transvaal not recorded to date from the grassland areas in the south but are otherwise widespread. Recorded from the Orange Free State; Natal and the Cape Province. Males Females X n Range X n Range 102 14 92-110 110 7 98-120 53 14 46-57 55 7 50-59 11 12 10-12 12 5 11-14 33 14 30-35 34 7 30-37 10,5 14 9,0-11,5 11,4 4 10,0-13,7 Habitat The common slit-faced bat is, throughout its range, associated with open savanna woodland although it has been taken on rare occasions in forests (Rosevear, 1965) and is found in oases in desert regions (Tibetsi, Libya; Hufnagl, 1972) and in isolated areas in the Sinai Peninsula and in Arabia, where there are scattered trees (Harrison, 1964). To some extent at least they have a wide habitat tolerance. In the Subregion they occur from sea level to over 1 500 m and in areas where the mean annual rainfall is as low as 100 mm and as high as 1 200 to 1 400 mm. Habits The common slit-faced bat is a gregarious species, occurring in small to medium-sized colonies of up to hundreds. They roost during the day in the substantial cover of caves, culverts under roads, in mine adits, in hollow trees (baobab, Adansonia digitata), in rock fissures and disused antbear holes (Temby, 1977) and are common occupants of the shelter under the roofs of houses or thatched rondavels. In these situations they hang from the ceilings, clinging on to ridge poles of huts or the thatch and, in situations such as culverts, to the small irregularities in the cement. They roost either in half light or in the darkest parts of the interior of these shelters, having a preference for the latter. They tend to roost in scattered groups, not closely packed together as do other species. In Drotsky's caves in Botswana, the colony was estimated to number 600, the individuals hanging from the 13 m high roof in total darkness, 270 m from the entrance. The atmosphere was humid with temperatures between 30 °C and 32 °C. They are late movers from their daylight roosting places, leaving well after sundown, usually about 20h30 to 21h00, and returning well before first light. On leaving they tend to scatter into small groups which break up, and the individuals forage separately. They are relatively slow fliers, but have great powers of jinking and twisting in flight. In foraging they may fly within a metre of the ground. Food Insectivorous, the food, as indicated by the discarded remains picked up under their feeding sites, includes a high proportion of moths and long-horned and short-horned grasshoppers. Felton (1956) recorded that in Namibia they 126 ate scorpions, including Opisthophthalmus wahlbergi and they will also take sun spiders, Solifugae. LaVal & LaVal (1980) provided a list of insects taken by the species in Natal, noting that many of the insects were taken while stationary rather than in flight. Once prey is secured they fly to established feeding sites, which they will use over prolonged periods, as is shown by the accumulations of discarded remains beneath them. Fenton (1975) showed, from the examination of the remains of insects collected at a feeding site, that Orthoptera remains accounted for 54% of the prey, Lepidoptera 45%. The Lepidoptera included 29 species of moths, one species, a sphingid, Polytychus compar, accounted for 32% of the remains, the next most common species 3,5%. This shows a remarkable selectivity on the part of the bat and compared with the details of the food of Lander's horseshoe bat, Rhinolophus landeri, suggests that there is a sharing of the food resources available between different species of bats. Reproduction In the Etosha National Park, Namibia it was found that after a brief period of courtship, copulation occurs in flight with both bats hovering. Copulation is of short duration and is repeated up to three times (Lindeque, 1987). In Natal, Bernard (1982a) found that ovarian and vaginal activity took place in two peaks, the first between April and early June, culminating in oestrus, the second between July and August during pregnancy. Copulation and fertilisation took place in early June, the young being born after a five month period of gestation in early November. Lactation lasted two months, followed by a period of anoestrus until the onset of pro-oestrus in April. In Zimbabwe, Smithers & Wilson (1979) found that they were seasonal breeders, gravid females commonly being taken during the months of August to October and not at other times of the year. Females with tiny young clinging to them were taken from October to December, the early part of the warm wet months of the year. A single young is produced at birth and is carried around by the mother attached to one of her nipples and clinging to her with the wings and feet. No. 99 Nycteris vinsoni Dalquest, 1965 Vinson's slit-faced bat Vinson se spleetneusvlermuis Colloquial Name Named after J. Vinson who sponsored an expedition to Mozambique during which the first specimens were taken (Dalquest, 1965). Taxonomic Notes Hayman & Hill (1971) stated that this species is more probably a member of the aethiopica group. Until its relationships are better known, however, it is considered to be a valid species. Only two specimens were collected; the species was described from an adult female. Description Dalquest (1965) in his description stated that it is a very large Nycteris, similar in proportions to N. thebaica but larger and stouter and with much shorter ears. One of the two specimens collected was in the orange phase, the other greyer like N. thebaica. N. vinsoni differs principally from N. thebaica in the features already mentioned and in its more massive skull and heavier teeth. The rostrum is 20% heavier than in N. thebaica and the supraorbital flanges overhang the eye orbits to a much greater degree, the breadth across the flanges being 6,8 mm in N. thebaica and 9,0 mm in N. vinsoni. The type specimen had the following measurements (mm): TL T Hfc/u E 125 55 13 22 The ear measurement given by Dalquest (1965) should be treated with caution as it had been singed in the fire lit to move them from the hollow tree—"although some allowance was made for this". Distribution So far only known from the south bank of the Save River in Mozambique at about 34 °E. Habitat Dry savanna woodland. Habits Two were taken from a small hollow at the base of a baobab, Adansonia digitata. Food Insectivorous. Reproduction Unknown. X. Family RHINOLOPHIDAE Horseshoe bats Members of this Family, all of which are of small to medium size, are found throughout the world, with the exception of the American Continent. It contains only one genus Rhinolophus, of which 30 species have been described from Africa, 10 occurring in the Subregion. They are called horseshoe bats from the shape of the anterior noseleaf, which is shaped like a horseshoe (Fig. 127 Fig. X.l. Head of a Rhinolophidae, Rhinolophus sp. The generic name Rhinolophus is derived from the Greek rhinos a nose and lophos a crest, referring to the crest of noseleaves. The noseleaf is a highly complicated organ and includes the nostrils which are situated centrally just behind the posterior fringe of the anterior noseleaf (Fig. X.2). The shape of the various parts of the noseleaves is useful in distinguishing between the species but they tend to distort in dried specimens and are appreciated more accurately in pickled specimens or better still in fresh material. Mohres (1953) found that the noseleaves in the genus act as transmitters of echolocation impulses, the impulses being emitted through the nostrils. It is the function of the noseleaves to channel and focus the emissions so that they achieve a maximum intensity at a point of focus ahead of the flying bat. They also shield the ears from the direct reception of the impulses, only the echoes being appreciated by them. which may serve as alternative clinging points for the young. In the skull the characteristic feature is the dome situated just above the nasal aperture (Fig. X.3). The two upper incisors are mounted in a projection of the palatine bone, which is partly cartilaginous and is often lost in preparation; these teeth are set well forward of the canines. On either side of this projection the skull is deeply emarginate. Fig. X.3. Skull of the Rhinolophidae, Rhinolophus fumigatus The dental formula is £ C± P M§ =28-32 The lower incisors are trifid. The anterior upper premolar is small and may be missing on both or either side. Its position, within or external to the toothrow, is important in the grouping of the species (Fig. X.4). Fig.X.2. Rhinolophidae noseleaf nomenclature: (a) nostril (b) sella (c) connecting process (d) posterior noseleaf (e) anterior noseleaf (f) secondary leaflet The ears are separated widely and are large and are capable of independent movement but lack ear tragi. The antitragi, however, are much enlarged and fold across the open base of the ears. The wings are short and rounded, the second finger consisting of the metacarpal only, without phalanges, the third, fourth and fifth fingers with two phalanges each which, when the bat is at rest, fold under the wing. The vertebrae of the tail end at the posterior fringe of the interfemoral membrane which is supported on either side by curved calcanea arising from the ankles. When at rest the tail and interfemoral membrane fold upwards. The fur is long and soft and while in general they are greyish-brown in colour, a reddish-brown colour phase is not uncommon. The females have a pair of false nipples situated low down on the abdomen just anterior to the genital orifice Genus Rhinolophus Lac<§p&de, 1799 Key to the species after Meester et al. (1986) 1. Face and/or lateral margins of sella liberally furnished with long hairs; connecting process low, rounded; greatest width of horseshoe over 9 mm ... 2 Sella naked; connecting process rounded or pointed; greatest width of horseshoe less than 9 mm ... 3 2. Forearm 62—67 mm .. . hildebrandtii Forearm 50—60 mm .. . fumigatus 3. Anterior upper premolar, when present, external to toothrow; canine and fourth upper premolar in contact; connecting process bluntly pointed ... 4 Anterior upper premolar in toothrow; canine and fourth upper premolar not in contact; connecting process blunt or sharply pointed ... 5 4. Forearm 50—57 mm ... clivosus Forearm 45-50 mm .. . darlingi 5. Connecting process rises to an erect point; interpterygoid groove usually shallow, not clearly defined by bordering ridges; anterior upper premolar usually not crowded between canine and fourth upper premolar, longer than wide ... 6 Connecting process with a low bluntly pointed tip; interpterygoid groove deep and clearly defined by bordering ridges; anterior upper premolar usually more or 128 less crowded between canine and fourth upper premolar, at least as wide as it is long ... 7 6. First phalanx of fourth finger notably shortened in relation to metacarpal length; connecting process pointed; molar width less than half width of palate between molars . . . landeri First phalanx of fourth finger not notably shortened in relation to metacarpal length; connecting process rises to a high narrow horn; molar width more than half width of palate between molars .. . blasii 7. Larger, forearm 47-51 mm, skull 20 mm or more ... capensis Smaller, forearm 40-46 mm, skull 18,7 mm or less ... 8 8. Sella broader; ears longer, 20-22 mm; condylocanine length more than 15,5 mm ... simulator Sella narrower; ears shorter, 18-20 mm; condylocanine length less than 15,5 mm ... 9 9. Front edge of connecting process almost concave; lancet with sides almost convex; fur whitish at base . . . denti Front edge of connecting process convex; lancet with sides concave, tip pointed; fur nearly unicoloured ... swinnyi Fig. X.4. Teeth (a) Rhinolophus darlingi, with the minute anterior premolar (c) situated external to toothrow (h) R. denti with the anterior upper premolar (d) situated in the toothrow. No. 100 Rhinolophus hildebrandtii Peters, 1878 Hildebrandt's horseshoe bat Hildebrandt se saalneusvlermuis Plate 5 Taxonomic Notes No subspecies are listed by Hayman & Hill (1971). Description Hildebrandt's horseshoe bats are the largest members of the genus that occur in the Subregion. Adult males have a total length of about 110 mm and a mass of about 27 g, the females slightly larger and heavier (Table 100.1). They are greyishbrown on the upper parts, the under parts about the same colour or very slightly lighter. The ears are conspicuously large and pointed at the tips, strongly convex on their inner edges and concave on the outer. The wing and interfemoral membranes are a translucent dark brown. The hair on the upper parts is unicoloured and is long and soft. Table 100.1 Measurements (mm) and mass (g) of Hildebrandt's horseshoe bats, R. hildebrandtii, from Zimbabwe (Smithers & Wilson, 1979) Males Females X n Range X n Range TL 114 15 108-121 116 15 109-125 T 38 15 32-40 39 15 31-43 Hf c/u 15 15 13-16 15 15 14-16 E 32 15 30-33 31 15 28-34 F/a 63 11 62-66 66 9 64-67 Mass 26,9 12 22,8-32,2 31,3 8 26,0-34,0 Distribution Confined to the eastern parts of the continent from the Transvaal north to southern Ethiopia and the Sudan. The most westerly records are from near Kabompo in Zambia. South of the Sahara, excluding the Southern African Subregion They occur in southern Ethiopia; Sudan; southern Somalia; in Kenya; Uganda and the extreme northeastern and southeastern parts of Zaire and in Rwanda. In Tanzania there are records from the eastern parts of the country, but their occurrence in Rwanda suggests that they have a wider distribution in this sector. There are records from Malawi, and in Zambia they are widespread except in the southwest. In Mozambique, north of the Zambezi River, they occur in the Tete District. While there are no records at the moment from the extreme northeastern parts of Mozambique, this area is little known and their occurrence in adjacent territories suggests that in time they will be shown to be present. Southern African Subregion They are widely distributed in Zimbabwe except in the northwest and occur across the border into Botswana in the Francistown district. They are recorded from Mozambique, south of the Zambezi River, as far south as the western parts 129 followed by a period of lactation and anoestrus of about 60 days. This is the first report of retarded embryonic development in the genus and only the third in the Chiroptera. In Zimbabwe (Smithers & Wilson, 1979) and the Transvaal (Rautenbach, 1982) gravid females have been taken in September and October, and Rautenbach (1982) recorded lactating females in the Transvaal in January. This suggests that the young are born early in the warm, wet summer months from about October to December. The females all carried one foetus. In the early part of their lives the young attach themselves firmly to the females' nipples and cling to them as they forage. The pair of functional nipples in the females is situated on the chest and in addition they have a pair of false nipples situated closely together, one on each side, just in front of the anus (Fig. 111.1). Genus Cloeotis Thomas, 1901 No. 112 Cloeotis percivali Thomas, 1901 Short-eared trident bat Drietand-bladneusvlermuis Plate 4 Colloquial Name So named from the three-pronged process at the top of the noseleaves and the noticeably short ears. Taxonomic Notes Meester et al. (1986) listed two subspecies, only one of which, C. p. australis Roberts, 1917, occurs in the Subregion; the other, C. p. per civali, occurs in East Africa. Description The smallest member of the Family Hipposideridae, the short-eared trident bat has a total length of about 70 mm, a short tail about 30 mm in length and a mean mass of about 4,0 g (Table 112.1). Only this species and the Persian leaf-nosed bat, Triaenops persicus, possess the three-pronged, trident-like process on top of the noseleaves. The colour of the upper parts is slate-grey, but some specimens from the Transvaal and in particular from Swaziland are buffy-brown. The hair is soft, silky and unicoloured and up to 9 mm long. The face is yellowish-white. The under parts are white tinged yellow, the individual hairs light slate-grey at their bases with yellowish-white tips, giving a general appearance of greyish under parts. The tiny ears are rounded, showing no sign of tips and lie close to the head, almost obscured by the long fur. The anterior upper premolar lies in the toothrow separating the canines and the posterior premolar, but lies to the external side of the mid-line of the toothrow; the anterior of the two premolars in the lower jaw is about half the height of the posterior. Table 112.1 Measurements (mm) and mass (g) of short-eared trident bats, C. per civali, from Zimbabwe (Smithers & Wilson, 1979) TL T Hf c/u E F/a Mass Distribution Too few records of this species are available to assess the limits of its distribution properly. Originally the species was described in 1901 from a specimen from Takaunga just north of Mombasa in coastal Kenya where it was taken from a coral Males Females X n Range X n Range 69 8 64-76 67 11 63-74 29 8 27-30 29 11 26-33 6 9 6-7 6 10 6-7 10 9 9-11 10 10 9-11 35 8 34-39 35 10 34-38 4,2 8 3,5-4.8 3,9 10 3,4-4,3 cave; Kingdon (1974) reported that since then only one specimen has been taken in the area. Roberts (1951) reported that in a cave near Pretoria, Transvaal, where he originallyhad found them to occur in large numbers, they were not found on a subsequent visit. South of the Sahara, excluding the Southern African Subregion In East Africa they are only known from coastal Kenya. Not recorded from Tanzania. Recorded from southeastern Zaire and although at present known only from two localities in Zambia, Ansell (1978) believed that they might be more widespread in the eastern parts of the country. In Mozambique, north of the Zambezi River, they are only known from Missale Mine in the northeastern Tete District. Southern African Subregion In Zimbabwe they are known from six localities from the Kariba Dam in the north to Mutare in the east and westwards to Gwanda. In the Transvaal they are recorded from three localities in the Pretoria, Rustenburg sector and at Komatipoort in the east; from Swaziland and Botswana in the Kanye and Molopolole areas in the southeast. Habitat There are insufficient data available at the moment to assess their habitat requirements. The presence of substantial shelter in the form of caves or mine adits appears to be an essential habitat requirement. Habits Short-eared trident bats are gregarious, occurring in colonies numbering hundreds. They roost during the day in caves and mine adits, hanging from the roof in tight clusters, far back into these shelters in total darkness. In Zimbabwe a very small colony of about 12 lived in an underground irrigation tunnel. Food Insectivorous. Reproduction In Zimbabwe gravid females each with a single foetus were taken in October, which suggests that the young are born during the warm, wet summer months from about October to December. 142 Genus Triaenops Dobson, i87i No. 113 Triaenops persicus Dobson, i87i Persian leaf-nosed bat Persiese bladneusvlermuis Colloquial Name Originally the species was described from a specimen from Shipaz, Persia (Iran) but subsequently has been found to occur in Arabia and on the continent of Africa. It is appropriately called the Persian trident bat because, like the short-eared trident bat, Cloeotis percivali, it has a threepronged trident at the top of the noseleaves. Taxonomic Notes Hayman & Hill (1971) listed two subspecies from the continent, T. p. majusculus Aellen & Brosset, 1968 from the Congo Republic and T. p. afer from the eastern part of the continent south to within the limits of the Subregion. Description The Persian leaf-nosed bat is about 140 mm in total length with a tail of about 40 mm long. The upper parts are yellowish-brown tinged with orange, the sides of the face yellow, the under parts lighter in colour than the upper parts, a shade browner on the throat. The hair is soft and silky, the bases of the individual hairs yellowish-brown, slightly browner towards the tips. The wing and interfemoral membranes are dark brown. The ears, which are characteristic in shape, are a translucent light brown. This bat is much larger in size than the short-eared trident bat, Cloeotis percivali, which is the only other species in the Subregion that possesses the three-pronged, trident-like, processes at the top of the noseleaves. The anterior upper premolar lies in the toothrow separating the canines and the posterior premolar, but lies to the external side of the mid-line of the toothrow; the anterior of the two premolars in the lower jaw is about half the height of the posterior. Distribution Extralimital to the continent Occurs in Iran, on the coast of the Persian gulf and in Arabia at an oasis in Oman in the east and Aden in the southwest, eastwards to Pakistan, and on Madagascar. South of the Sahara, excluding the Southern African Subregion Recorded from Uganda and a number of localities in Ethiopia, west in the valley of the Blue Nile to near the Sudan border; in western, northern and central Somalia; Ethiopia; Congo Republic; coastal Kenya and in two areas in the southeast of the country and coastal Tanzania. There are no records at the moment from the northeastern parts of Mozambique, but they occur in the Tete District, north of the Zambezi River. Southern African Subregion Recorded from the Mutare District in eastern Zimbabwe and in Mozambique, south of the Zambezi River, in the Tete and Inhambane districts. Habitat There are insufficient data available at the moment to assess their habitat requirements, but these include the availability of substantial shelter in the form of caves or mine adits. Hayman & Hill (1971) believed that they rarely occur inland, but since then they have been taken far inland including in Zimbabwe (Smithers & Wilson, 1979). In a coastal cave in the Inhambane District, Mozambique they occurred in hundreds, but only a single specimen was taken in a cave near Mutare, Zimbabwe. Habits Persian leaf-nosed bats are gregarious. Where they occur in large numbers they hang from the ceiling of the caves in large clusters, slightly separated from one another. Food Insectivorous. Reproduction No information available from the Subregion. 143