In the following accounts genera, species and subspecies are all arranged in alphabetical order. The number of species currently recognized in each genus is given at the beginning of each genus account. Each species account begins with the currently recognized scientific name and any vernacular names. Any appropriate information on the present or past taxonomic status of species/subspecies is given at this point. Each account is then divided into the following sections, which are only shown if information is available:
Priority Grade: Priority grades have been allocated to taxa to reflect their conservation status. These grades are used to prioritize recommended action (see Recommended Action section). The grades are as follows:
Grade Conservation status
1 Extinct
1 Extinct? (taxa thought to have become extinct recently)
1 Endangered (Limited Distribution)
2 Endangered
3 Vulnerable (Limited Distribution)
4 Vulnerable
5 Rare (Limited Distribution)
6 Rare
7 Indeterminate (Limited Distribution)
8 No Data (Limited Distribution)
9 Indeterminate
10 No Data
11 Not Threatened
Number of subspecies: If appropriate.
Distribution: Current known distribution. Localities are listed in alphabetical order. The distribution within countries or islands/island groups is given in parentheses. The geographical location of areas mentioned is shown in the accompanying locality maps (Figures 3 to 10).
Status: Information on current status if available. This section also includes any information on past status or declines and ends with information on any known threats.
Ecology: This includes information on the habitats used, on roost sites, feeding ecology and behavioural ecology. The food plants are listed in alphabetical order by family. Where known, the part of the plant that is utilized is listed thus:
FL Flowers
FR Fruits
LV Leaves
LB Leaf bracts
B Bark
S Seed Pods
C Cones
T Twigs
Population biology: Includes information on timing of copulation, length of lactation and gestation, the number of young born, the time to sexual maturity, longevity and survivorship.
Occurrence in protected areas: Information is listed in alphabetical order by country.
Summary of threats: A brief summary of the known major threats.
Recommended action: A list of suggested action that would assist in the conservation of this species/subspecies.
Principal authors: Where individuals other than the authors of this Action Plan have contributed specific information on species or subspecies, they have been credited.
Figure 3. Locality map for Africa. Political boundaries are indicated by heavy solid lines.
Figure 4. Locality map for the Indian Ocean.
Figure 5. Locality map for the Middle East and Asia. Political boundaries are indicated by heavy solid or dashed lines.
Figure 6. Locality map for South East Asia. Political boundaries are indicated by heavy solid or dashed lines.
Figure 7. Locality map for the Philippines.
Figure 8. Locality map for islands off eastern New Guinea. Political boundaries are indicated by heavy dashed lines.
Figure 9. Locality map for the western Pacific Ocean. Political boundaries are indicated by heavy dashed lines.
Figure 10. Locality map for Vanuatu.
Genus Acerodon (6 species)
All species were placed on Appendix II of CITES in 1989 in view of their similarity in appearance to Pteropus species in trade. The genera are distinguishable only by dental characters. Acerodon species have already appeared in the Guam trade in shipments from the Philippines.
Priority Grade: 10 (No Data).
Distribution: Salayar; Sangir Islands; Sula Islands (Sula Mangole); Sulawesi.
Status: Netted for food in northern Sulawesi (A. Marshall, pers. comm.).
Population biology: An adult female collected in February carried a large embryo (Bergmans and Rozendaal, 1988). Immature individuals collected in November had forearm lengths similar to those of adults while an immature collected in February was slightly smaller (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park.
Summary of threats:
Possibly hunting.
Recommended action:
Priority grade: 8 (No Data: Limited Distribution).
Distribution: Talaud Islands, north-east of Sulawesi.
Summary of threats:
Recommended action:
Priority Grade: 2 (Endangered).
Number of subspecies: 2.
A. j. jubatus
Priority Grade: 2 (Endangered).
Distribution: Philippines (Basilan, Biliran, Cebu, Dinagat, Leyte, Luzon, Mindoro, Negros, Panay, Samar, Sulu Archipelago [Bongao, Sanga Sanga, Sibutu, Tawitawi]) (J. Edwards Hill, pers. comm.).
Status: Populations have declined drastically in virtually all areas of the Philippines and this species is probably threatened with imminent extinction. Although widespread within its present distribution, it has vanished from a number of small and medium-sized islands and is also believed to have been completely hunted out or disappeared from larger islands, such as Cebu, which have been completely deforested. On Negros, because of heavy hunting pressure, and the great reduction of forest habitats, the current population is far below that present at the turn of the century. The small upland roosts of about 1000 individuals observed by Heideman and Heaney contrast markedly with the large roosts observed by Steere (quoted in Elliot, 1896) and Worcester (1899) in the late 1800s, by Taylor (1934) from the 1920s, and the roosts of thousands reported by Rabor (1977) in similar habitats in the 1950s and 1960s. There are also reports that in the 1920s joint camps of Pteropus vampyrus/A. jubatus contained up to 120,000 animals. The largest colonies now are about 5000 individuals, though most of them are only a few hundred - suggesting as much as a 96% decline in 60 years.
Despite these declines, small populations persist in many parts of the Philippines. However, the combination of habitat destruction and both subsistence and commercial hunting has resulted in a threat of extinction of A. jubatus on some islands in the next 10–20 years.
As the largest fruit bat in the Philippines and one which roosts in large colonies, this species is very easy to locate and thus exploit. Although it is much less common than Pteropus vampyrus, it is the preferred species for hunting (Bräutigam, 1989). It is hunted locally for subsistence purposes using fishing hooks on weighted lines hung from larger lines on trees and is believed to be taken in considerable numbers (Bräutigam, 1989). Many animals that are shot at the roost never fall from the trees, while others may glide out of sight after being shot. As a result, a hunter requiring 10 bats may kill 20–30 individuals. This species is sold in the markets in the Philippines and is commonly eaten, though considered a ‘poor man's food’ (Bräutigam, 1989).
The Philippines have been an important source of bats for the Guam market, with peak exports of 2471 in 1986 (Bräutigam and Elmqvist, 1990; Wiles, 1992). Bats have also been exported from the Philippines to the CNMI with a total of 2477 exported between 1986–89 (Stinson, et al., 1992). A. jubatus has been identified in a shipment confiscated in Guam that ostensibly originated in Saipan (Wiles, 1992).
Forest destruction has increased at a dramatic rate in the Philippines in recent years. As an example, primary forest cover on Negros was reduced from about 60% in 1945 to about 6% in 1987 (Heaney and Heideman, 1987). This has particularly serious implications for A. jubatus because of its reliance on primary forest.
Ecology: Taken from or near sea level to primary montane forest at 1100 m (Taylor, 1934 and authors' data). It is present only in or near areas with substantial tracts of forest, and is probably dependent on primary or old secondary forest. It has never been recorded foraging outside primary forest on Negros.
In the Balinsasayao region of Negros, A. jubatus feeds mainly on figs (Musaceae: including Ficus benjamina, F. chrysolepis, F. crassiramea and F. variegeta: Utzurrum (1984) and authors' data). It is certainly possible that this species may occasionally feed upon cultivated fruits, but current evidence suggests that the use of cultivated species is uncommon or rare.
Individuals probably fly long distances to feed. On Maripipi, bats were observed commuting at least 12 km to foraging sites on Biliran, and foraging distances could have been as high as 20 or 30 km each way. Those that fed at a submontane forest site on Negros probably travelled at least 5 km to feed.
Observations in the Balinsasayao region of Negros in 1981–84 suggested that average densities were one per 10–20 ha.
Population biology: Females probably produce no more than one young per year. Rabor (1977) reported that populations on Negros gave birth in April or May, and additional data from Negros and elsewhere in the Philippines suggest that young are born in approximately the same period on other islands.
Occurrence in protected areas: Certainly occurs within some designated reserves, but probably all individuals spend at least some time outside protected areas.
Summary of threats:
Recommended action:
A. j. mindanensis
Priority Grade: 2 (Endangered).
Distribution: Philippines (Mindanao).
Status: See A. j. jubatus.
Ecology: See A. j. jubatus.
Population biology: See A. j. jubatus.
Occurrence in protected areas: See A. j. jubatus.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
This species has recently been transferred from the genus Pteropus (Musser et al., 1982).
Priority Grade: 10 (No Data).
Number of subspecies: 2.
A. l. leucotis
Priority Grade: 10 (No Data).
Distribution: Philippines (Balabac, Calamian Group [Busuanga]).
Status: May be moderately common in good forest (Heaney et al., 1989). Given its restricted range and the current rate of deforestation in the Palawan chain, it seems likely that it is declining in abundance.
Summary of threats:
Recommended action:
A. l. obscurus
Priority Grade: 10 (No Data).
Distribution: Philippines (Palawan).
Status: See A. l. leucotis.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 1 (Extinct).
Distribution: Philippines (Panay).
Status: Almost certainly extinct as it is known only from a few specimens collected in 1888 and 1892.
A specimen (in the Field Museum of Natural History) from Concepcion, Panay, taken on 16th April 1892 by J. Steere was the last known individual. Steere (quoted in Elliot, 1896) noted that two males had come from a roost in some tall clumps of spiny bamboos (Gramineae: Bambusa arudinacea), among the rice fields and almost over some of the native houses. Hundreds of bats were thickly clustered in the tops of the bamboo. There appeared to be at least three species in the roost, although each species used separate trees.
The restricted range makes this species especially vulnerable to hunting and forest destruction. Deforestation has been as severe on Panay as on other islands in the central Philippines. The island retains about 10% of its forest cover, most of it on the highest mountains and ridges. Given the absence of specimens in the last 95 years and the level of deforestation on Panay, it is certain that this species is seriously threatened, if not extinct.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 5. With one exception (A. m. floresii) all subspecies are confined to single islands within the Lesser Sundas group. Goodwin (1979) noted that all taxa are based on slight differences in colour and/or size and doubted the necessity for recognizing them as subspecies. Sody (1936), in describing A. m. prajae, stated that it resembled floresii and Goodwin (1979) noted that the measurements given by Sody in his description fall within the range of the series from Timor.
A. m. alorensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Alor.
Summary of threats:
Recommended action:
A. m. floresii
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Flores; Sumbawa.
Summary of threats:
Recommended action:
A. m. gilvus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Sumba.
Summary of threats:
Recommended action:
A. m. macklotii
Priority Grade: 11 (Not Threatened).
Distribution: Timor.
Status: Common from sea level to an altitude of 450 m (Goodwin, 1979). Goodwin (1979) observed two colonies on Timor each with 300–500 individuals. At both sites the bats were spread out over a large portion of the crowns of large fig trees (Moraceae: Ficus sp.) on the edge of open forest; the roosting areas were largely defoliated.
Ecology: While no other species of bat roosted with Acerodon on Timor, Goodwin (1979) observed them feeding with Pteropus griseus, P. vampyrus and Dobsonia peronii; their staple food in March, April and May consisted of the fruit of at least two species of fig. The bats also fed upon some indeterminate part of the coconut palm (Palmae: Cocos nucifera). Most of their nutrition was probably obtained from the juice of the fruit. At dusk the bats flew in groups of two to six individuals and were most frequently observed flying into and out of the crowns of coconut palms (Goodwin, 1979).
Population biology: Pregnant females were seen on Timor in March and May (Goodwin, 1979).
Summary of threats:
Recommended action:
A. m. prajae
Priority Grade: 8 (No Data: Limited Distribution)
Distribution: Lombok.
Population biology: Kitchener et al. (1990) noted that of six females collected on Lombok in October 1987 two were lactating and three others appeared to be lactating. The appearance of a lactating female that was also pregnant demonstrates this subspecies is polyoestrous as a result of a post-partum oestrus.
Summary of threats:
Recommended action:
Genus Aethalops (1 species)
Priority Grade: 9 (Indeterminate).
Number of subspecies: 3. Recently, Kitchener et al. (1990) suggested, on the basis of examination of a number of specimens, that the subspecies A. a. aequalis should be considered as a separate species.
A. a. aequalis
Priority Grade: 9 (Indeterminate).
Distribution: Borneo (Brunei, Sabah, Sarawak).
Status: Probably fairly common in the hills of Borneo (C.M. Francis, pers. comm.). Threatened by deforestation (A. Suyanto and D. Kitchener, pers. comm.).
Ecology: All records are from mountains in the north-west including Gunung Kinabalu (up to 2700 m) and the Crocker Range in Sabah; Gunung Mulu and the Kelabit uplands in Sarawak (Payne et al., 1985). Apparently confined to montane forest above 1000 m (Payne et al., 1985).
Occurrence in protected areas:
Borneo
Brunei
Ulu Temburong National Park (proposed)
Sabah
Crocker Range National Park, Kinabalu Park
Sarawak
Gunung Mulu National Park
Summary of threats:
Recommended action:
A. a. alecto
Priority Grade: 9 (Indeterminate).
Distribution: Lombok; central Peninsular Malaysia; Sumatra.
Status: Lombok Kitchener et al. (1990) mist-netted a single adult male in low mossy forest at about 1700 m at Pos Tiga, Mt Rinjani.
Peninsular Malaysia Uncommon (Medway, 1978), known only from the forest above 900 m, on Maxwell's Hill, Gunung Benom and the main range from the Cameron Highlands to Selangor.
Threatened to a small degree by the so far limited disturbance of montane forest, but there are prospects of greater montane forest loss due to road building, expansion of temperate style agriculture, and building of montane holiday homes.
Sumatra The type locality is Indrapura Peak (2225 m), Sumatra.
Ecology: Recorded from montane forest over 900 m. Roosts singly or in small groups of two to three. Main food plants are unknown but presumably are the soft fruits of montane forest trees and climbers.
Population biology: In the Cameron Highlands in Peninsular Malaysia, pregnancies occur between April and June. A single female collected by Kitchener et al. (1990) on Lombok in early October 1987, had a large embryo but did not appear to be lactating.
Occurrence in protected areas:
Peninsular Malaysia
Cameron Highlands Wildlife Reserve
Fraser's Hill Wildlife Reserve
Krau Game Reserve (Gunung Benom)
Also along the main ridge of Peninsular Malaysia, some of which is Forest Reserve. It should be noted that Forest Reserves are ‘reserved for logging’, not totally protected. It might also occur in montane parts of Taman Negara, but little netting has been done there.
Sumatra
Bukit Barisan Selatan National Park
Gunung Leuser National Park
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
A. a. ocypete
Priority Grade: 6 (Rare).
Distribution: Jawa.
Status: Rare. Threatened by deforestation (A. Suyanto and D. Kitchener, pers. comm.)
Summary of threats:
Recommended action:
Genus Alionycteris (1 species)
Priority Grade: 5 (Rare: Limited Distribution).
Distribution: Philippines (Mindanao).
Status: Given its restricted range and the current extent of deforestation on Mindanao, it seems likely that it is declining in abundance and possibly threatened.
Ecology: Probably a primary forest species.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Genus Aproteles (1 species)
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: New Guinea (Western Province, Papua New Guinea).
Status: Rare. First described from 12,000-year-old fossils excavated from a rock shelter near Mount Elimbari, Chimbu Province (Menzies, 1977). It appears to have become extinct there about 9000 years ago. In 1975, it was discovered living in the Hindenberg Wall area of Western Province, Papua New Guinea where it inhabited a cave known as Luplupwintem, 2300 m above sea level. The cave is close to the edge of the Hindenberg Wall and difficult to reach, although it is close to the main track linking Tabubil and the Ok Tedi mine to Telefomin. The species was abundant in 1975, but on a return visit in 1977 only two bats were seen circling near the roost (Hyndman and Menzies, 1980). It is thought that increasing human population density in the Central Highlands may have led to the decline of the species there, and that it has survived in the remote western part of New Guinea where the human population density is low.
During a visit to the Ok Tedi mining area in April 1987, Flannery (1990) questioned informants who were familiar with this species in the 1970s, including one man who had accompanied Hyndman when he obtained animals in 1975. Informants made no distinction between this species and Dobsonia moluccense, except for noting a difference in size. They described how large numbers of Aproteles were obtained from Luplupwintem in the 1970s by use of a shotgun. From their accounts, it seems probable that disturbance to the Luplupwintem colony from shotgun hunting led to its decline. When the cave was checked on 11 and 12 April 1987, no bats were seen.
In 1991, a skull and skin of A. bulmerae were discovered at the Australian Museum. This animal had been collected in 1984 at Afektaman near Telefomin and confirmed that A. bulmerae had survived in this area until then. Funding is being sought for a project to search for and, if appropriate, undertake a study of this colony. In 1992, a colony of about 130 bats was discovered in Luplupwintem Cave (L. Seri, pers. comm.).
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Genus Balionycteris (1 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
B. m. maculata
Priority Grade: 11 (Not Threatened).
Distribution: Borneo (Brunei, West Kalimantan, Sabah, Sarawak).
Status: There are scattered records throughout the north and west including Kota Kinabalu, Sepilok, Madai and Tawau in Sabah; the upper Sungai Temburung and Tasek Merimbun in Brunei; Mulu, Niah, Gunung Dulit, and Kuching in Sarawak; Gunung Kenepai in West Kalimantan (Payne et al., 1985). Probably not threatened.
Ecology: Frequently netted in lowland dipterocarp forest.
Occurrence in protected areas:
Borneo
Probably found in all protected areas containing primary and lightly logged forest including:
Brunei
Ulu Temburong National Park (proposed)
Sabah
Danum Valley and Sepilok Reserve
Kinabalu Park
Balionycteris maculata (Photo by K-G. Heller)
Sarawak
Gunung Mulu National Park
Summary of threats:
Recommended action:
B. m. seimundi
Priority Grade: 9 (Indeterminate).
Distribution: Peninsular Malaysia; Riau Archipelago (Durian, Galang); Thailand.
Status: Peninsular Malaysia Widespread and locally common in forest at all elevations including Gunung Brinchang and Pahang (Medway, 1978). Probably commoner in lowland than montane forest.
Riau Archipelago No information, but a number of these islands are now heavily developed (e.g., Batam and Bintang).
Thailand Uncommon, known only from Khao Phu Pah in Trang Province (Lekagul and McNeely, 1977).
Large population reductions would result from forest loss. Currently, forest cover in Peninsular Malaysia is about 40%, with the greatest losses occurring in the lowlands.
Ecology: In Peninsular Malaysia, has been found roosting in small groups in crowns of palms and clumps of epiphytic ferns, rarely in caves (Payne et al., 1985).
Feeds on the fruits of understorey plants and small trees below the main canopy, does not congregate at large food sources as do some other species (e.g., Cynopterus).
Population biology: In Peninsular Malaysia, females produce a single young. Pregnancies or carried young have been recorded in nearly every month, even though Medway (1983) suggests a local breeding season at Cameron Highlands.
Occurrence in protected areas:
Peninsular Malaysia
Apparently all protected areas containing primary or lightly
logged forest contain this species, including:
Cameron Highlands Wildlife Reserve
Endau-Rompin Park (proposed)
Fraser's Hill Wildlife Reserve
Krau Game Reserve
Sungkai Game Reserve
Taman Negara
Also many reserves under the jurisdiction of the Forest
Department, Peninsular Malaysia.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
Genus Casinycteris (1 species)
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Southern Cameroun to eastern and central southern Zaire. A distribution map is given in Meirte (1984a).
Casinycteris argynnis (Photo K-G. Heller)
Status: Threatened by forest destruction.
Ecology: Occurs in lowland forest.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Chironax (1 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
C. m. melanocephalus
Priority Grade: 11 (Not Threatened).
Distribution: Borneo (Brunei, Sabah); Jawa; Nias; central Peninsular Malaysia; Sumatra; Thailand.
Status: Borneo Recorded from only two localities, Sepilok in Sabah and the upper Sungai Temburong in Brunei.
Jawa Rare (A. Suyanto and D. Kitchener, pers. comm.).
Peninsular Malaysia Locally common in the hills and foothills from the Cameron Highlands south to Selangor, mainly at elevations above 457 m (Medway, 1978). There is a potential for increased tourist and agricultural development of hilly and montane areas, causing loss of previously secure montane forest.
Thailand Recorded from the following provinces: Ranong; Surat Thani, Nakhon Si Thammarat and Yala (Yenbutra and Felten, 1986).
Ecology: In Borneo, several Sabah specimens were netted in the understorey of dipterocarp forest (Francis, 1989).
In Peninsular Malaysia, found roosting in small groups in tree ferns and in a shallow cave (Payne et al., 1985).
Information on food plants is scanty but presumably feeds on the fruits of understorey hill and montane forest trees. Known to feed on Ficus (Moraceae) (A. Marshall, pers. comm.).
Population biology: At the Cameron Highlands in Peninsular Malaysia, pregnancies in February and April may imply a local breeding season.
Occurrence in protected areas:
Borneo
Sabah
Sepilok Forest Reserve
Peninsular Malaysia
Cameron Highlands Wildlife Reserve
Fraser's Hill Wildlife Reserve
Krau Game Reserve (Gunung Benom)
Taman Negara (montane forest)
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
C. m. tumulus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: North and east-central Sulawesi.
Status: The type locality is the Dumoga-Bone National Park, north Sulawesi, where it is believed to be fairly common (W. Bergmans, pers. comm.).
Ecology: Apparently restricted to lowland forest (Bergmans and Rozendaal, 1988). Specimens have been taken in primary forest at altitudes of up to 960 m, in a number of cases near or over small streams.
Feeds on Ficus (Moraceae) (A. Marshall, pers. comm.).
Population biology: Pregnant females have been collected in March and April (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park.
Summary of threats:
Recommended action:
Genus Cynopterus (4 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 9. C. b. brachyotis includes the species C. archipelagus, which was described from a single immature specimen, now thought to be a variant of C. b. brachyotis (P. D. Heideman and L. R. Heaney, pers. comm.) and C. minor which was described from a single immature specimen (K. F. Koopman, pers. comm.).
C. b. altitudinis
Priority Grade: 10 (No Data).
Distribution: Highlands of Peninsular Malaysia, on the Main Ridge. Recorded from the Cameron Highlands south to Gunung Bunga Buah (Medway, 1978), but presumably also occurs on mountains north of the Cameron Highlands. Not found on isolated peaks away from the main ridge (e.g., Gunung Benom, Gunung Tahan).
Status: No information, but there is a proposed montane road from Cameron Highlands via Fraser's Hill south to the Genting Highlands. This would make large areas of montane forest susceptible to development and forest clearance.
Occurrence in protected areas:
Peninsular Malaysia
Cameron Highlands Wildlife Reserve
Fraser's Hill Wildlife Reserve
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
C. b. brachyotis
Priority Grade: 11 (Not Threatened).
Distribution: Bangka; Belitung; Borneo (Brunei, Central, East, South and West Kalimantan, Sabah, Sarawak); Lombok; Nicobar Islands; Peninsular Malaysia (including islands of Babi, Great Redang, Langkawi, Pangkor, Penang, Pisang [off Johore], Sri Buat, Tinggi, Tioman and Tulai); Philippines (Balabac, Calamian Group [Busuanga], Culion, Luzon, Maripipi, Negros, Palawan, Polillo); Riau Archipelago; Singapore; Sulawesi; Sumatra; Talaud Islands and adjacent small islands; Thailand.
Cynopterus brachyotis (Photo by K-G. Heller)
Status: Borneo Probably one of the commonest mammals of lowlands and hills. Recorded from all parts of the island including Gunung Penriven, the Kelabit Uplands and Gunung Kinabalu up to 1676 m. Also found on islands of: Ereban, Kalimantan; Satang, Sarawak; and Balambangen, Sabah (Medway, 1978; Payne et al., 1985). Occurs in most habitats including lower montane forest, dipterocarp forest, gardens, and mangrove and strand vegetation.
Lombok Kitchener et al. (1990) mist-netted this bat in a wide range of habitats, both disturbed and ‘natural’. It was most common, however, near Desa Kuta where 20 were collected along a dry watercourse fringed with banana plantations (Musaceae: Musa spp.) and village gardens. It appeared equally abundant from sea level to about 200 m, but only one animal was collected at Batu Koq (at 400 m).
Peninsular Malaysia Common everywhere in the lowlands and in montane forest on peaks away from the Main Ridge. Mainly in disturbed forest and cultivation, and only occasionally in primary forest.
Philippines Ranges from sea level to at least 1500 m, although rarely found above 1000 m. Common in vegetated urban areas and in agricultural areas, especially in gardens and orchards, where 7% to 44% of all captures were made (Guerrero and Alcala, 1973; Heaney et al., 1989). Rare in primary forest, but more abundant in clearings and patches of secondary growth forest (6–20% of captures; Heaney et al., 1989; Heideman and Heaney, 1989). In general, therefore most common in agricultural and disturbed habitats and uncommon in undisturbed primary forest. An exception is Maripipi, where it is most abundant in primary forest; this may be related to the absence of Haplonycteris fischeri and Ptenochirus major which are similar in size and potential competitors (E. A. Rickart, pers. comm.).
In lowland orchards, capture rates (and probably density) were high (44% of total captures; Guerrero and Alcala, 1973; 9% of captures and an average of 1 individual per net-night; Heaney et al., 1989 and unpublished data). In upland primary forest on Negros, capture rates were only 1 per 10 net-nights, and these were almost exclusively in clearings (Heideman and Heaney, 1989), and still lower on Leyte (3 to 5 per 100 net-nights; Heaney et al., 1989).
Because of its preference for disturbed habitats and success in agricultural habitats, it is not currently at risk in the Philippines. Its small size and choice of inconspicuous roost sites reduces its vulnerability to hunters.
Sulawesi Has been collected in primary forest, disturbed primary forest, partly disturbed mangrove forest, over brackish water less than 100 m from the sea, coastal alang-alang secondary forest mosaic, plantations/secondary forest mosaic, in a solitary fig tree (Moraceae: Ficus sp.) in a deforested village area, in a grass plain with scattered trees, in a limestone area and among fruit trees in a garden. Altitudes vary from sea level to about 1100 m (Bergmans and Rozendaal, 1988).
Thailand Recorded from Chiang Mai south to Pattani and east to Udon Thani and Chanthaburi (Lekagul and McNeely, 1977).
Ecology: Normally roosts in pairs or small groups in trees, especially under dead palm leaves and among stems and leaves of palms (Taylor, 1934; Lawrence, 1939; Lim, 1966; Lekagul and McNeely, 1977; Payne et al., 1985). It is suggested that it modifies leaves to form a tent (Phillips, 1924; Lekagul and McNeely, 1977), but this is disputed by Rickart et al. (1989). Also roosts in the threshold zone of caves.
Feeds primarily upon fruits, but also takes nectar and pollen (Gould, 1978). Diet includes the following plants (Lim, 1966; Guerrero and Alcala, 1973; Gould, 1978, Medway, 1978 and author's data): Anacardiaceae: Mangifera indica (FR); Bombacaceae: Durio zibethinus (FL); Melastomataceae: Melastoma malabathricum (FR); Moraceae: Ficus spp. (FR); Musaceae: Musa spp. (FL,FR); Myrtaceae: Psidium guajava (FR); Palmae: Cocosnucifera (FL); Piperaceae: Piper aduncum (FR); Sapotaceae: Chrysophyllum cainito (FR), Palaquium sp. (FR), Pouteria sapota (FR); Sonneratiaceae: Sonneratia spp. (FL).
Marshall (1985) listed 27 plant genera fed upon by members of the genus Cynopterus. Faecal analysis suggests that Ficus accounts for a large part of the diet.
At one site on Negros, nine animals moved an average of 650 m between their first and second capture (Heideman and Heaney, 1989). Apparently does not forage in flocks, and it is rare to capture more than one or two at any time.
Animals captured in primary forest at Sepilok, Sabah, were significantly smaller than those in secondary forest, but it is not known whether this reflects dietary differences or taxonomic differentiation (Francis, 1989).
Population biology: In the Philippines, most populations probably produce two young per year (Heideman, 1987). Pregnant females may be found in all or almost all months, although births are restricted to two three-month periods. Females may only produce a single young annually in some populations.
On Negros, most females become pregnant at about 6–8 months of age, while males become sexually mature at approximately 1 year (Heideman, 1987; pers. obs.). Gestation is approximately 3.5–4 months; lactation about 6–8 weeks (Heideman, 1987; pers. obs.). On Lombok, in September and October 1987, Kitchener et al. (1990) noted that of 17 females examined, 13 were pregnant, each with a single embryo, and one was lactating. Three of the pregnant females were also lactating, suggesting that it may at least be seasonally polyoestrus. In May 1988, of three adult females collected, one was lactating and two appeared to have been pregnant recently. In Sulawesi, five females collected in January and March had one embryo each. Immature individuals have been collected in January, February and March and in October and November (Bergmans and Rozendaal, 1988). The greatest documented longevity in the wild is 4.5 years (Heideman and Heaney, 1989). Annual survivorship of yearlings/adults in the Balinsasayao region on Negros was estimated to be 70%, implying that about 5% of yearlings may live to 6 years of age (Heideman and Heaney, 1989).
Occurrence in protected areas:
Borneo
Brunei
Temburong National Park (proposed)
Peninsular Malaysia
Taman Negara
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Principal authors for this subspecies: P. D. Heideman, L. R. Heaney.
C. b. brachysoma
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Andaman Islands.
Summary of threats:
Recommended action:
C. b. ceylonensis
Priority Grade: 10 (No Data).
Distribution: Sri Lanka.
Summary of threats:
Recommended action:
C. b. concolor
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Enggano.
Summary of threats:
Recommended action:
C. b. hoffeti
Priority Grade: 11 (Not Threatened).
Distribution: Vietnam.
Status: Appears to be widespread. Recorded from 15 localities in 11 provinces throughout southern Vietnam (Van Peenen, 1969).
Summary of threats:
Recommended action:
C. b. insularum
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Kangean Islands; Laut Kecil Islands.
Summary of threats:
Recommended action:
C. b. javanicus
Priority Grade: 10 (No Data).
Distribution: Bali; Jawa; Madura; Penida.
Status: Jawa Collected in a garden in Jakarta and in the Botanical Garden in Bogor in 1981, where it was common (W. Bergmans, pers. comm.).
Summary of threats:
Recommended action:
C. b. minutus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Nias.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 4.
C. h. harpax
Priority Grade: 11 (Not Threatened).
Distribution: Peninsular Malaysia (Penang); Sumatra; Thailand.
Status: Peninsular Malaysia Widespread throughout the mainland and also occurs on the island of Penang (Medway, 1978).
Thailand Rare, with specimens from Trang Tala and Tak (Lekagul and McNeely, 1977).
Ecology: Occurs in the lowlands in Peninsular Malaysia, but its upper altitudinal limit is not known. It is more common in disturbed habitats, such as gardens, orchards, rubber estates and wasteland, than in forest. However, nearly everywhere it is less abundant than C. brachyotis and generally its distribution is patchier.
Congregates at large food sources such as mango trees (Anacardiaceae: Mangifera indica). Pakarnseree (1986) lists the following as food plants of Cynopterus (species unidentified) in Thailand: Bignoniaceae: Dolichandrone spp., Markhamia spp., Oroxylum indicum; Bombacaceae: Bombax spp.; Compositae: genus unidentified; Leguminosae: genus unidentified; Musaceae: Musa spp.; Myrtaceae: Eugenia spp.; Sapotaceae: Palaquium spp.; Sonneratiaceae: Sonneratia caseolaris; Ulmaceae: Trema spp.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
C. h. horsfieldii
Priority Grade: 11 (Not Threatened).
Distribution: Jawa; Lombok.
Status: Lombok Kitchener et al. (1990) mist-netted it in most habitats, particularly disturbed situations, but it was most common at Pelangan and Batu Koq.
Population biology: Kithener et al. 1990 noted that all 13 females examined from Lombok in October 1987 showed some reproductive activity. Six were pregnant and three of these were also lactating, suggesting polyoestry. All of the remaining females showed evidence of recent parturition or early pregnancy, four of these were also lactating. A single female caught on Lombok in May 1988 was recently pregnant.
Summary of threats:
Recommended action:
C. h. persimilis
Priority Grade: 9 (Indeterminate).
Distribution: Borneo (Brunei, Central, South and West Kalimantan, Sabah, Sarawak).
Status: Reported from scattered localities in all districts except East Kalimantan, with cave roosts known from Gunung Kinabalu and Gomantong in Sabah, and near Kuching in Sarawak.
Ecology: Often roosts in rock shelters or caves, usually near the entrance. Occasionally found in trees or palms (Palmae) (Payne et al., 1985).
Summary of threats:
Recommended action:
C. h. princeps
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Nias.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 7.
C. s. angulatus
Priority Grade: 11 (Not Threatened).
Distribution: Borneo (Central Kalimantan); south China; Peninsular Malaysia; Sumatra; Thailand.
Status: Borneo Only record is from Central Kalimantan (Payne et al., 1985).
Peninsular Malaysia Known from Selangor northwards (Medway, 1978). Appears to be commoner in the northern states (Kedah, Perlis, Kelantan) and declines in abundance southwards. Occurrence is patchy but it is common where it is found (G. W. H. Davison, pers. comm.).
Thailand Recorded from Chiengmai, Bangkok, Siracha, Trang, and the area of the Malay peninsula (Lekagul and McNeely, 1977).
Ecology: The information provided by Pakarnseree (1986) (see C. h. harpax) for specimens from Thailand applies equally to this subspecies, though G. W. H. Davison (pers. comm.) mentions it also feeds on figs (Moraceae: Ficus spp.).
Summary of threats:
Recommended action:
C. s. babi
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Babi Island, near Simalue Island, west Sumatra.
Summary of threats:
Recommended action:
C. s. gangeticus
Priority Grade: 11 (Not Threatened).
Distribution: Central Provinces and Palnpur, north-west India.
Status: Prater (1971) described it as common, although he did not distinguish between the subspecies that occur in India.
Ecology: Roosts singly or in small groups among palm leaves, aerial roots of banyan (Moraceae: Ficus benghalensis), tree hollows, and similar situations, rarely being found in ruins or caves (Prater, 1971).
Leaves roost early in the evening to forage. Feeds on fruit and nectar. Often flies off with ripe fruit, which is eaten elsewhere (Prater, 1971).
Population biology: Pregnant females have been obtained in February and young probably born in March and September have been collected, suggesting two birth peaks (Prater, 1971). Young carried by the mother have been seen in September and it is thought that the gestation is 115–125 days (Prater, 1971).
Summary of threats:
Recommended action:
C. s. pagensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Mentawai Islands (Pagai Islands [North Pagai], Siberut and Sipura).
Summary of threats:
Recommended action:
C. s. scherzeri
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Nicobar Islands (Car Nicobar and ?Great Nicobar).
Summary of threats:
Recommended action:
C. s. serasani
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Natuna Islands (Serasan).
Summary of threats:
Recommended action:
C. s. sphinx
Priority Grade: 11 (Not Threatened).
Distribution: Bangladesh; Burma (Shan States); India (Assam, Bengal, Bhutan Duars, Bombay, Calcutta, Chin Hills, Kumaon, Madras, Sikkim); Sri Lanka; north Thailand.
Status: Bangladesh Seems to be common and widespread. Often found in Dhaka city being mobbed by crows (Khan, 1985).
India See C. s. gangeticus.
Sri Lanka Locally common, throughout almost the whole of the cultivated areas in the lowlands and also throughout the higher hills of the central massif up to 1828 m. Generally more abundant in the dry zones than in the wet (Phillips, 1980).
Ecology: In Sri Lanka, roosts in colonies of 6–12 inside seed clusters in kitul palms (Palmae: Caryota urens) and in talipot palms (Corypha umbraculifera). Sometimes roosts in hollow trees (Phillips, 1980). In Bangladesh, lives singly or in small numbers among dead palm leaves and in tree hollows (Khan, 1985). Younger males roost with females, while older males usually roost alone (Phillips, 1980). Adult males can be wanderers, spending their day many miles from the main colony (Phillips, 1980).
In Sri Lanka, feeds on fruits of soursop (Annonaceae: Annona muricata), guava (Myrtaceae: Psidium guajava) and mango (Anacardiaceae: Mangifera indica) and the flowers and fruits of palms and plantains (Musaceae: Musa spp.) (Phillips, 1980).
Population biology: In Sri Lanka, females with young are found in most months, thus breeding is probably intermittent throughout the year. Females have one young per year (Phillips, 1980).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
C. t. major
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Nias.
Summary of threats:
Recommended action:
C. t. terminus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Timor.
Status: Common in 1968 (Goodwin, 1979), but no recent information on status.
Ecology: At low elevations, roosts in palms and modifies the leaves to make tents. At high elevations, roosts in hollow trees (Goodwin, 1979).
Summary of threats:
Recommended action:
C. t. titthaecheilus
Priority Grade: 11 (Not Threatened).
Distribution: Bali; Jawa; Krakatau Islands; Lombok; Sebesi; Sumatra.
Status: Jawa Very common in disturbed situations from low altitudes (Bogor Botanical Gardens) to 1600 m (Cibodas Botanical Gardens) (Kitchener et al., 1990).
Krakatau Islands One of the most common bats in the Krakatau Islands (Tidemann et al., in press).
Lombok Kitchener et al. (1990) mist-netted these bats in October 1987 on Lombok, finding them from sea level to 400 m in all habitats, particularly disturbed situations, but most commonly at low altitudes, particularly Pelangan and Kuta. Also collected by A. Suyanto and NAMRU II (Navy Army Medical Research Unit) at Sewela and Kelayan, Lombok Timur, respectively.
Ecology: On Anak Krakatau probably roosts in trees because there are no caves. Circumstantial evidence from the Krakatau Islands suggests that this bat moves between islands (Kitchener et al., 1990).
Population biology: Kitchener et al. (1990) suggested possible asynchrony in the reproductive cycle in different areas on Lombok. Of 12 females from Kuta, two had not bred, five were apparently primiparous, four were pregnant but not lactating and one was pregnant and lactating. At Pelangan, Suranadi and Batu Koq all but two females were lactating. These observations were taken to indicate that in the moister areas (which have streams with running water or pooled water) females had generally bred once and were pregnant again, while in the very dry areas of Des Kuta on the central south coast (no potable water) several females had not bred and most were only recently pregnant.
Summary of threats:
Recommended action:
Genus Dobsonia (11 species)
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Biak; Owii; and Waigeo off north-west New Guinea.
Status: Waigeo Known from one cave (Nja-njef).
Population biology: The holotype was lactating at time of capture, on 25 December 1909. All other six females collected at this time contained embryos. There might be a single breeding season with births taking place by the end of December and in January (Bergmans, 1975b).
Summary of threats:
Recommended action:
Priority Grade: 1 (Extinct?).
Distribution: Philippines (Cebu, Negros).
Status: Presumed to have been extinct since about 1970 (Heaney and Heideman, 1987). All known specimens, except one from Cebu, were taken from Negros between 1949 and 1964. No specimens have been taken since, despite extensive efforts in its former habitats and collection sites.
Two series are in museums. The first was taken in 1949 at 200 m elevation in Mambaho Cave on Negros, which at the time was surrounded by large tracts of primary dipterocarp forest (Rabor, 1952). The cave was apparently the largest of a series in a ridge of soft limestone that parallels the Bagtik River, a tributary of the Ilog. Rabor estimated about 300 bats in groups of 4–12, with one group of 30, in a poorly lit portion of the cave. All bats taken were subadult males, indicating that this was a bachelor colony. The second series was taken at Lobogon Barrio, near Basay, Negros, in 1964. The series contained adult males and females and was probably from a breeding colony.
In 1945, about 60% of Negros was forested, but a boom in logging and an expansion of sugar plantations reduced this to 12% by 1975 (Alcala, 1976). Almost no forest remained below 800 m, the recorded upper elevational limit of this species. Between 1981 and 1987, Heaney and Heideman (1987) surveyed 10 large caves and numerous smaller caves on southern Negros (including several near the type locality). They found only a single fragment of an apparently old mandible of this species in guano deposits on the floor of one cave. Extensive netting in the area where Rabor collected his series also failed to produce any specimens. It seems likely that destruction offorests, disturbance of cave roosts by guano mining, and hunting of bats for meat probably all played a role in the demise of this species (Heaney and Heideman, 1987).
Ecology: All specimens from Negros were taken at elevations of between sea level and 800 m. Most appear to have been taken within or on the edge of primary forest.
A female shot in a coconut tree (Palmae: Cocos nucifera) in 1948 had been feeding on betel nut (Areca catechu) fruits, an introduced palm.
Population biology: Births occur in May or June and young fly by August and September. A subadult collected in December 1964 was probably weaned.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Biak;?Numfoor; and Owii off north-west New Guinea.
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Distribution: Sulawesi.
Status: Hunted for food in northern Sulawesi (Bergmans and Rozendaal, 1988)
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
D. i. inermis
Priority Grade: 10 (No Data).
Distribution: Bougainville; New Guinea; Solomon Islands (Alu, Fauro, Guadalcanal, Malaita, New Georgia, Nggela Sule, San Cristobal, Shortland, Uki Ni Masi, Vella Lavella).
Summary of threats:
Recommended action:
D. i. minimus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Choiseul, Santa Isabel).
Summary of threats:
Recommended action:
Priority Grade: 6 (Rare).
Distribution: Bagabag; north and west New Guinea; Sulawesi; Yapen.
Status: Uncommon (Flannery, 1990).
Ecology: Encountered in both primary and secondary rain forest (Flannery, 1990), and two were obtained hanging by day from the branch of a broad-leaved shrub (McKean, 1972). No records of roosting in caves, which contrasts greatly with the habits of the larger Dobsonia moluccense. Mist-netted in open areas 1 m from the ground (Greig-Smith, 1975) but is usually caught in nets set over small, overgrown streams or paths (Flannery, 1990). Flannery (1990) never encountered this species at altitudes in excess of 250 m, although Ziegler (1982) suggests the altitudinal limit may be as high as 600 m.
Population biology: A female collected on 18 January and two females caught in late May and early June had single embryos (McKean, 1972). In addition, four females captured in the Yapsiei area in January 1984 were all pregnant (Flannery, 1990).
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2. The species D. magna is now considered to be a subspecies of D. moluccense.
D. m. magna
Priority Grade: 11 (Not Threatened).
Distribution: Australia (New South Wales, Queensland); New Guinea and islands off west coast (Batanta, Misool, Waigeo).
Status: Australia Rare (Strahan, 1983).
New Guinea Common (Flannery, 1990). Abundant in most habitats up to 2700 m in altitude, but may be uncommon or absent in areas of the savannah woodland in the south, as Waithman (1979) reports that it was only captured along the coastal strip during his survey of the Morehead region (Flannery, 1990).
Ecology: Usually roosts in the twilight zone of caves and sometimes forms immense colonies; a cave in the Nong River Valley near Telefomin, New Guinea, is home to a colony of many thousands. Australian groups are much smaller. Colonies of 100 or so have been seen scattered in dark areas under piles of large boulders. Roosts have also been found in old mines, abandoned houses, and dense vegetation; usually in near-darkness. It is the only Australian megachiropteran to roost in caves (Strahan, 1983). Sometimes roosts singly and Flannery (1990) records seeing solitary bats roosting in small or easily accessible caves, or even under shallow overhangs in the Telefomin area. Vocalization occurs almost continuously in colonies, but the bats forage alone and are silent at night (Dwyer, 1975). Occasionally, individuals also roost in tree hollows and even in the crowns of trees. (Hyde et al., 1984). Brass (1964) even records a group of adults and young taken from a hollow nursery tree in the Markham Valley, New Guinea, in November 1959.
Is a strong flier and Dwyer (1975) reports seeing bats fold their wings and plummet 300 m almost vertically to reach the entrance of their roost cave. Wilson (1985) and Richards (1986b) report that D. m. magna manoeuvres well and can fly slowly; it is even capable of flying backwards. This allows it to forage below the forest canopy and reach roosts and food unavailable to Pteropus species. Specimens are occasionally caught in deadfall traps set for bandicoots on Mount Karimui, which suggests that it may land on the ground to forage (Hyde et al., 1984). Food taken includes cultivated bananas (Musaceae: Musa spp.), papaya (Caricaceae: Carica papaya), and wild Ficus (Moraceae) (Hyde et al., 1984). In Australia, the bloodwood (Myrtaceae: Eucalyptus sp.) is one of its favourite food trees (Strahan, 1983).
Population biology: Breeding appears to be seasonal, with most births occurring September-October. The young do not become sexually mature until they are 2 years old, and are presumably nursed for 5–6 months. The overall pattern of reproduction in the Mount Elimbari area (Chimbu Province, New Guinea) seems to be: copulation at the close of the wet season (April–June); birth at the close of the dry season (August–November); weaning at the height of the wet season (February–April). It is not known whether this reproductive pattern is seen in areas of New Guinea with a less seasonal climate than Chimbu Province, but evidence from the Yapsiei area of west Sepik Province, where seasons are not clearly demarcated, suggests that it is the case (Flannery, 1990). In Australia, males mature at about 2 years of age and copulation occurs at the end of the wet season in May and June. Between September and November, each female gives birth to a single young, which is carried for about 1 month and nursed for a further 4–5 months (Strahan, 1983).
Summary of threats:
Recommended action:
D. m. moluccense
Priority Grade: 10 (No Data).
Distribution: Ambon; Aru Islands; Buru; Halmahera; Kai Islands; Seram.
Occurrence in protected areas:
Seram
Manusela National Park
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Number of subspecies: 3. The subspecies D. p. anderseni and D. p. pannietensis were formerly included as subspecies of D. moluccense.
D. p. anderseni
Priority Grade: 10 (No Data).
Distribution: Bismarck Archipelago (Admiralty Islands [Manus], Boang, Duke of York, Emirau, Lihir, New Britain, Tabar); Umboi.
Summary of threats:
Recommended action:
D. p. pannietensis
Priority Grade: 10 (No Data).
Distribution: D'Entrecasteaux Islands (Fergusson Island, Goodenough Island, Normanby Island); Louisiade Archipelago (Misima Island, Panaeati Island, Rossel Island, Sudest Island);Muyua.
Summary of threats:
Recommended action:
D. p. remota
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Trobriand Islands (Kiriwina).
Summary of threats:
Recommended action:
Priority Grade: 9 (Indeterminate).
Number of subspecies: 4.
D. p. grandis
Priority Grade: 9 (Indeterminate).
Distribution: Komodo; Lombok; Penida; Sumbawa.
Status: Bergmans (1978c) notes that from their scarcity in collections one might infer that in the Lesser Sundas, Dobsonia is either not very common or mostly restricted to less accessible habitats. As it has been found in caves and hollow trees, near the coast and inland, and from sea level up to an altitude of considerably more than 880 m (on Timor), a relatively low density seems rather more likely than inaccessible habitats. The vertical range for lowland animals from the Lesser Sundas is 0–1000 m, and locally even 0–1200 m (Rensch, 1936). A plausible explanation for a low density could be an equally low occurrence of suitable habitats, since man has destroyed more than 50% of the forest on these islands (Rensch, 1936) and this must have seriously affected bat populations.
Lombok Kitchener et al. (1990) collected two adult males and two adult females in 1987 at Pelangan. These were netted using oval fishing nets by village people among coconuts (Palmae: Cocos nucifera) in the early evening. Another specimen was collected by NAMRU II (Navy Army Medical Research Unit) in a garden at Bilekedit, west Lombok on 24 March 1979.
Ecology: A large series was collected from a hollow tree on Sumbawa (Mertens, 1936). Another specimen, from Komodo, was taken from a colony of about 25 animals hanging in an open cave (Bergmans, 1978c).
Population biology: The two females collected by Kitchener et al. (1990) in October 1987 were lactating.
Summary of threats:
Recommended action:
D. p. peronii
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Timor.
Status: See D. p. grandis.
Summary of threats:
Recommended action:
D. p. sumbana
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Sumba.
Status: See D. p. grandis.
Summary of threats:
Recommended action:
D. p. ssp. incertae sedis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Alor; Babar; Flores; Wetar.
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Distribution: Bismarck Archipelago (Duke of York, New Britain, New Ireland).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
D. v. crenulata
Priority Grade: 10 (No Data).
Distribution: Bacan; Banggai Islands; Halmahera; Morotai; Rau; Sulawesi; Ternate; Togian Islands; Waigeo.
Ecology: Two bats were caught by Bergmans and Rozendaal (1988) over brackish water in partially disturbed, tall mangrove forest, within 100 m of the sea on the Banggai Islands.
Population biology: One of the males caught by Bergmans and Rozendaal (1988) on the Banggai Islands in November 1981 was sexually mature.
Summary of threats:
Recommended action:
D. v. viridis
Priority Grade: 10 (No Data).
Distribution: Ambon; Banda Islands; Buru; Kai Islands; Misool; Sangir Islands; Seram.
Summary of threats:
Recommended action:
Genus Dyacopterus (1 species)
Priority Grade: 6 (Rare).
Number of subspecies: 2. Specimens from the Philippines have not been identified subspecifically.
D. s. brooksi
Priority Grade: 10 (No Data).
Distribution: Sumatra.
Summary of threats:
Recommended action:
D. s. spadiceus
Priority Grade: 6 (Rare).
Distribution: Borneo (Brunei, Sabah, Sarawak); central Peninsular Malaysia.
Status: Borneo Recorded from Sepilok and Baturong Caves in Sabah; Baram district, Niah, and near Sibu in Sarawak (Payne et al., 1985), and from Brunei.
Peninsular Malaysia Rare, recorded only in Selangor and Negri Sembilan (Medway, 1978).
Presumably threatened with forest loss; known from both lowland and montane forest and has also been recorded in open country. It is rarely netted, and the reasons for its apparent rarity are unknown.
Ecology: In Borneo, it has been caught near caves, but its roosting sites are unknown. In Peninsular Malaysia it has been found roosting in a tree trunk (Payne et al., 1985) and has recently been netted in forest canopy in the Krau Game Reserve (C. M. Francis, pers. comm.).
One flock was seen feeding in a fig tree (Moraceae: Ficus sp.) with Horsfield's fruit bat (Cynopterus horsfieldii) (Payne et al., 1985).
Occurrence in protected areas:
Borneo
Brunei
Ulu Temburong National Park (proposed)
Sabah
Sepilok Forest Reserve
Not known but likely to occur in large forest areas such as Kinabalu Park.
Peninsular Malaysia
Krau Game Reserve
Not known but likely to occur in large forest areas such as Taman Negara.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
D. spadiceus (subspecies unknown)
Priority Grade: 10 (No Data).
Distribution: Philippines (Luzon, Mindanao).
Ecology: Probably a primary forest species.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Genus Eidolon (2 species)
Priority Grade: 11 (Not Threatened). This species was formerly considered a subspecies of E. helvum but has recently been elevated to a full species (Bergmans, 1990).
Distribution: Madagascar and offshore islets.
Status: Rather widespread and at least locally common (M. Nicoll, pers. comm.).
It is hunted for food, at least locally (Wilson, 1987). As it depends on trees for food, the deforestation in Madagascar threatens its continued existence in many places. All fruit bat colonies at Ankarana Special Reserve are in inaccessible places because local people use the bats as food (J. Wilson, pers. comm.).
Ecology: Wilson (1987) reports on a colony at Ankarana Special Reserve subsisting apparently almost entirely on the fruit of ebony (Ebenaceae: Diospyros sp.).
Occurrence in protected areas:
Madagascar
Ankarana Special Reserve (but protection apparently not implemented).
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
A full review of this species is given by DeFrees and Wilson (1988).
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
E. h. helvum
Priority Grade: 11 (Not Threatened).
Distribution: Angola; Benin; Burkina Faso; Burundi; Cameroun; Central African Republic; Chad; Congo; Equatorial Guinea (Bioko, Mbini, Pagalu); Ethiopia; Gabon; Gambia; Ghana; Guinea; Guinea-Bissau; Ivory Coast; Kenya; Liberia; Malawi; Mali; Mozambique; Namibia; Niger; Nigeria; Rwanda; São Tomé and Príncipe (Príncipe, São Tomé); Senegal; Sierra Leone;?Somalia; South Africa; Sudan; Tanzania (Mainland, Mafia, Pemba, Zanzibar); Togo; Uganda; Zaire; Zambia; Zimbabwe.
Moves from its prime habitat, the tropical forests of central Africa, where a variety of fruits is available throughout most of the year, to areas both north and south of this optimal territory. The availability of food determines occurrence elsewhere. Areas where it may be considered a year-round resident include parts of north-eastern Zambia, Malawi, and possibly north-eastern Mozambique and north-eastern Natal (Skinner and Smithers, 1990). Records are infrequent from 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 south-west province of Transvaal these bats are considered ‘wanderers’. In addition, has been taken in the Orange Free State and in the eastern Cape Province as far west as the Bredasdorp district (Skinner and Smithers, 1990). Rosevear (1965) reported it had been taken at sea 250 km from the nearest land. Has been recorded from several islands off Cameroun (Ambas and Bimbia) and Sierra Leone, and on islands in the Congo River and in Lake Kivu (Zaire).
Status: In general common throughout its range. Rosevear (1965) noted that in West Africa it was exceedingly common and very widespread. Sometimes occurs in enormous colonies of 100,000–1,000,000 individuals of both sexes. However, it is locally threatened.
M. D. Tuttle (pers. comm.) believes hunting in West Africa is sufficient to seriously threaten populations. Is eaten in all communities in south-western Nigeria; hunters are often hired to shoot bats for hotels and restaurants. Most people eat whole bats including the bones (Funmilayo, 1978). Although the subspecies as a whole is not threatened, with the human population in Africa rapidly increasing, and with progressive forest destruction, this situation may change sooner than expected, because the large aggregations in high trees are easy prey for hunters. On the other hand, it is quite adaptable. Colonies are known from the centres of large, busy cities (Lagos, Libreville, Kampala) and from the campus of the University of Ife, Nigeria, where it was hunted both for food and for dissection by zoology students on a sustainable basis (Halstead and Segun, 1975). In other parts, tradition ensures the protection of colonies and consequently, unmolested colonies may exist in the centres of villages and towns. Occasionally the skulls and smoke-dried bodies of this bat are found in ju-ju stalls because it is believed that they improve fertility in women (Shoga, 1974).
Could become a pest in fruit orchards and plantations, eating mango (Anacardiaceae: Mangifera indica), guava (Myrtaceae: Psidium guajava), papaya (Caricaceae: Carica papaya) and avocado pear (Lauraceae: Persea americana) (Funmilayo, 1976). These bats are certainly fond of bananas and plantains (Musaceae: Musa spp.) and can become a nuisance in these plantations. In Nigeria, it is killed for meat and because it is regarded locally as a pest because of mechanical damage to tree crops and competition with man for fruit (Funmilayo, 1979). Happold (1987) believes that there is little evidence that these bats damage commercial fruit. In the Ivory Coast it is a threat to non-native pine plantations because it gnaws the bark, wood and leaves killing the trees (Malagnoux and Gautun, 1976). May also eat and destroy dates (Palmae: Phoenix dactylifera) to such a degree that protective measures are required (Nowak, 1991). Guano creates problems in some villages and possibly poses a threat to the health of villagers (Funmilayo, 1979). For example, leaves used in cooking are often contaminated by faeces (Funmilayo, 1976). However, present control methods are wasteful and need regulation (Funmilayo, 1978).
Has also been shown to be a vector of Lagos bat rabies virus (Boulger and Porterfield, 1958) and as such may be in danger from attempts at extermination. Its large roost size and habit of roosting in areas of high human population, make it particularly susceptible.
Ecology: Inhabits forest and savannah, and is found up to an elevation of 2000 m in the Ruwenzori Mountains (Kingdon, 1974).
Is gregarious and prefers to roost in tall trees by day, but has also been found in lofts and in caves in rocks (Nowak, 1991). In Nigeria, selects trees of particular species for roosting (Okon, 1974) and some tree species, even though common, are not used. Common trees for roosts include Eucalyptus saligna (Myrtaceae), Cocos nucifera (Palmae), Elaeis guineensis (Palmae), and three species of Ficus (Moraceae) (Jones, 1972). Prefers dead trees that have bare branches; if living trees are used, the leaves are soon broken and the branches become bare. Trees used as day roosts are large with spreading branches, commonly found in dense groves with thick undercover. At night, roosts are chosen according to food availability. Trees are of various heights and sizes, some in groups, others widespread (Okon, 1974). Colonies can number up to 1,000,000; sleeping groups about 100 (Kulzer, 1969; Nowak, 1991). Roosting clusters are located 6–20 m above ground on sturdy branches (Jones, 1972). During periods of migration, colonies disperse into small groups and form temporary roosts from which they eventually form ‘regular’ roosts (Mutere, 1980). Baranga and Kiregyera (1982) reported a colony of 70,388 bats in Uganda. The average number of bats per tree was 310; the average number of clusters per branch 4; and the average cluster size 8. Most colonies use the same roosts for many years, but because of local fluctuations in food, some colonies make regular seasonal migrations returning after a few months to their former roosting sites (Happold, 1987). For example, in the Ivory Coast, has been observed to migrate, from the tropical forest zone where it stays between June and December, to the Niger basin in the interior where it appears in January and stays until May (Huggel-Wolf and Huggel-Wolf, 1965).
Feeds on the following fruits (both wild and cultivated) and on some flowers (Allen, 1939; Booth, 1959; Harris and Baker, 1959; Osmaston, 1965; Rosevear, 1965; Mutere, 1966, 1967; Jones, 1972; Ayensu, 1974; Kingdon, 1974; Funmilayo, 1979; Thomas, 1982; Dobat and Piekert-Holle, 1985; Fujita and Tuttle, 1991): Anacardiaceae: Anacardium occidentale (FR, FL), Mangifera indica (FR), Pseudospondias sp. (FR), Spondias mombin (FR); Annonaceae: Annona sp. (FR); Bignoniaceae: Kigelia aethiopica (FL), K. pinnata (FL); Bombacaceae: Adansonia digitata (FL), Bombax buonopozense (FL), Ceiba pentandra (FL), Ochroma pyramidale (FL); Caricaceae: Carica papaya (FR); Cecropiaceae: Musanga cecropioides (FR); Chrysobalanaceae: Parinari excelsa (FR); Combretaceae: Terminalia sp. (FR); Convolvulaceae: Ipomoea albivenia (FL); Cunoniaceae: Codia sp.; Euphorbiaceae: Bridelia ferruginum (FR), Sapium ellipticum (FR); Lauraceae: Persea americana (FR); Leguminosae: Albizia sp. (L), Erythrina sp. (L), Parkia clappertoniana (FL), P. filicoidea (FL), P. roxburghii (FL); Loganiaceae: Anthocleista spp.; Meliaceae: Azadirachta indica (FR,FL); Moraceae: Antiaris africana (FR), Artocarpus sp. (FR), Chlorophora sp. (L), C. excelsa (FR), Ficus exasperata (FR), F. leprieuri (FR), F. mucosa (FR), F. natalensis (FR), F. thonningii (FR), F. umbellata (FR), F. vogelii (FR); Morinaceae: Morina lucinda (FR); Musaceae: Musa sapietum (FR), M. s. paradisica (FR); Myristicaceae: Pycnathus angolensis (FR); Myrtaceae: Eucalyptus sp. (FL);Psidium guajava (FR), Syzygium sp.; Palmae: Borassus aethiopum (FR), Elaeis sp., Phoenix dactylifera (FR); Passifloraceae: Adenia cissampeloides (FR), Passiflora sp.; Rhamnaceae: Maesopsis sp.; Rosaceae: Eriobotrya sp.; Sapotaceae: Vitellaria paradoxum (FR,FL); Solanaceae: Solanum anomalum (FR); Sterculiaceae: Cola sp., Theobroma cacao (L); Ulmaceae: Celtis sp.; Verbenaceae: Vitex sp. (FR).
In Nigeria, feeds almost exclusively at night, visiting only trees that have food resources, whereas trees visited during the day are only for roosting (Okon, 1974). At night, small groups of bats fly to foraging areas in straight lines. On many occasions, near Ibadan, 1000 bats per minute have been counted flying along a fairly narrow flight path. The foraging area is not known, but the powerful flight suggests that these bats utilize food sources many kilometres from their roosts (Happold, 1987). Nowak (1991) suggested that foraging range may be at least 30 km for some of the larger colonies. They may assist in the pollination of the flowers of some trees (Baker and Harris, 1959) but probably not to the same extent as some of the smaller fruit bats (Happold, 1987). Vast quantities of fruits must be required to sustain large colonies; the daily foraging flights, and local seasonal migrations, are clearly related to the availability and abundance of food, the fruiting times of different tree species, and the size of the colony. Colonies do not appear to break up into smaller sub-colonies in times of food shortage, although individuals scatter and forage in smaller groups each night. The gregarious habits of these bats probably evolved in conjunction with their ability to forage on many types of food resource; obviously a species that feeds on only one or two food items could not be sustained in large numbers in one place for more than a short time (Happold, 1987).
Although predation is infrequent and seemingly poses little threat to populations, several animals eat this species; spotted eagle owl (Bubo africanus), crows, steppe buzzards, black kite (Milvus migrans; Kingdon, 1974), snakes, palm civets, genets and hawks (Funmilayo, 1979), pottos (Perodicticus potto; Jones, 1972), Ayer's hawk eagle (Hieraaetus dubius; Wolf, 1984), and African hawk eagle (Hieraaetus spilogaster; Louette, 1975). Kingdon (1974) recorded E. h. helvum attacking a pied crow.
Population biology: Essentially monoestrus, giving birth to one young annually. Two reproductive strategies are reported; one involving a gestation offour months with conceptions in the autumn and births in the spring (Andersen, 1912), the other involving an extended gestation of at least nine months, incorporating a period of delayed implantation which results in births timed to coincide with the onset of the rainy season (Mutere, 1967; Fayenuwo and Halstead, 1974; Funmilayo, 1979). Anciaux de Faveaux (1978) recorded two colonies of E. helvum both breeding at 2° of latitude S, as having different conception times. One colony followed a boreal pattern of conceptions in September/October and births four months later, while the other followed an austral pattern with conceptions in June/July and births four months later. Anciaux de Faveaux (1978) suggested that this might be a direct result of the migratory nature of the species, the colony with the boreal pattern migrating from the north and the other with the austral pattern colony coming from the south. Wolton et al. (1982) suggested that breeding takes place in the dry season in West Africa. In Cameroun, Eisentraut (1941) noted that copulation took place at the end of the rainy season (October and November) with births occurring in February and March. No records of females in breeding condition exist in southern Africa suggesting the improbability of breeding there due to low population density of these migratory animals (Mutere, 1967). The only known longevity record for E. h. helvum is 21 years 10 months (Nowak, 1991).
Occurrence in protected areas:
Ghana
Mole National Park
Ivory Coast
Tai National Park
Malawi
Kusungu National Park
Nigeria
Borgu Game Reserve
Pandam Wildlife Park
Summary of threats:
Recommended action:
E. h. sabaeum
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Saudi Arabia (Nader, 1985b); Yemen.
Ecology: In Taizz, Yemen, a specimen was collected from a papaya tree (Caricaceae: Carica papaya) (it is not known whether it was feeding). In Al Shugayri, Saudi Arabia, individuals were seen feeding in a Ficus tree (Moraceae)(Nader, 1985b).
In Yemen, Allen (1939) recorded bats eating ripe dates (Palmae: Phoenix dactylifera) in the middle of August, and the fruits of a Borassus palm.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Eonycteris (2 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
E. m. major
Priority Grade: 11 (Not Threatened).
Distribution: Borneo (Brunei, East and South Kalimantan, Sabah, Sarawak)
Status: Scattered records from most areas, except Central and West Kalimantan, including Kota Kinabalu and Ranau in Sabah, Gunung Dulit and Kuching in Sarawak, Kutai in East Kalimantan, and upper Sungai Tengah in South Kalimantan. Cave roosts have been found in Sarawak along the Sungai Tinjar and in the Bintulu district.
Ecology: Roosts in caves and hollow trees (Payne et al., 1985).
Summary of threats:
Recommended action:
E. m. robusta
Philippine nectar-feeding fruit bat
Priority Grade: 6 (Rare).
Distribution: Philippines (Biliran, Leyte, Lubang, Luzon, Maripipi, Mindanao, Negros, Siargao).
Status: Rare wherever it occurs (e.g., only 5 of 2122 bats trapped and less than one capture per 100 net-nights [authors' data]; none in over 300 net-nights in primary forest on Leyte [ Heaney et al., 1989]).
This bat is very vulnerable to capture at its roost caves, which it shares with E. spelaea and probably Rousettus (Rousettus) amplexicaudatus. The latter species are gathered at caves for food throughout the Philippines, with hunters routinely removing ‘sacks-full’ of individuals. Undoubtedly these sacks sometimes include E. m. robusta.
The authors captured no E. m. robusta in caves on Negros, despite extensive sampling between 1981 and 1989. Its current rarity in caves and its strong association with primary forest suggests that it has declined sharply in parallel with the deforestation of lowland Negros. May well be threatened throughout its range, but is apparently so rare that the seriousness of the threat is difficult to assess. Even without protection, will probably persist, albeit in very low numbers, as long as undisturbed cave roosts are available in areas near primary, or perhaps secondary, forest.
Ecology: Taken from near sea level to 1100 m. Some Philippine records are from caves (Taylor, 1934; Lawrence, 1939). All records with associated habitat information indicate that the specimens were taken in or near primary forest, and this bat is probably dependent on these forests. Available evidence suggests that this bat roosts primarily in caves, and may be dependent upon them for successful reproduction.
Undoubtedly feeds on nectar and pollen, as do all other macroglossines (Marshall, 1985).
Population biology: An adult female captured on 28 April 1984 on Biliran was carrying a 13 g juvenile (E. A. Rickart, pers. comm.).
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
E. s. glandifera
Priority Grade: 4 (Vulnerable).
Distribution: Bali; Borneo (East and South Kalimantan, Sabah, Sarawak); east Jawa; Lombok; Muna off south-east Sulawesi; Philippines (Carabao, Cebu, Marinduque, Mindanao, Negros, Siargao, Siquijor, Sulu Archipelago [Sanga Sanga], Tablas [J. Edwards Hill, pers. comm.]); south-east Sulawesi; Sumba; Timor.
Status: Borneo Cave roosts have been found at Gomantong, Sukau, Sungai Segama and Madai in Sabah; Niah, Sungai Tinjar, and near Kuching in Sarawak. Also recorded from the upper Sungai Tengah in South Kalimantan and Kutai in East Kalimantan (Payne et al., 1985).
Lesser Sundas (Bali, Lombok, Muna, Sumba, Timor) Considered endangered because of habitat destruction, cave disturbance and severe hunting pressure, throughout Jawa and the Lesser Sundas (A. Suyanto and D. Kitchener, pers. comm.). It plays an important role in the pollination of durian (Bombacaceae: Durio zibethinus). Kitchener investigated the relationships between the species, its roosting caves and the durian crop and found serious conservation problems with Indonesian caves being heavily exploited for limestone, guano and bats for food. Fifteen Lombok caves have good bat populations but all are affected by human pressures, particularly fires and removal of bats for food by knocking them down with sticks. This situation will be exacerbated as the human population continues to migrate eastwards along the Lesser Sundas. The human population is increasing and so is its longevity. There is an increasing demand for protein, and bats are a good, often easily accessible food source (A. Suyanto and D. Kitchener, pers. comm.).
Philippines Capture rates of E. s. spelaea were 0.2 per net-night in urban orchards (Heaney et al., 1989), and 0.1 per net-night in forest clearings in the Balinsasayao region of Negros, but these bats were very rare within primary forest (Heaney et al., 1981; Heideman and Heaney, 1989). Eonycteris made up 2% of total captures in an orchard site on Negros (Heaney et al., 1989), as it did in a similar orchard sampled nearby by Guererro and Alcala (1973).
In the Philippines, tolerant offorest clearance, and able to use the nectar and pollen of many trees planted by humans in rural and urban areas. However, because it roosts almost exclusively in caves, usually within 10–20 m of the entrances, it is extremely vulnerable to hunting and disturbance. In the Philippines, many populations have been much reduced by hunters, who capture the bats in their roosts. Some caves with colonies of up to 500 bats in the 1970s and/or early 1980s, held from none to fewer than 50 animals in the late 1980s. At some caves, the only signs of bats were heaps of decomposing skins, sometimes mixed with charred bones.
As long as some roost caves are inaccessible to hunters, viable, if small populations, should remain. It rarely, if ever, attacks fruit, but is an important pollinator of many commercially important species including durian, kapok (Bombacaceae: Ceiba pentandra), abaca (Musaceae: Musa textilis), wild banana (Musa spp.), mangroves and many commercially valuable forest trees (Start and Marshall, 1976; Gould, 1978).
Ecology: Mist-netted on Lombok from sea-level in all habitats but most commonly in banana plantations, near mango trees (Anacardiaceae: Mangifera indica) and over watercourses, full or dry. Not recorded in caves (Kitchener et al., 1990).
On Timor recorded from solution caves and cave-like crevices near the coast; in one cave roosted only 5 m above high tide level (Goodwin, 1979).
In the Philippines, ranges from sea-level to at least 1100 m. Most abundant in agricultural habitats, especially orchards, or naturally disturbed forest habitats, particularly those with species of bananas, and is uncommon or rare in submontane primary forest and absent from mossy forest (Heaney et al., 1981, 1989).
Usually roosts in large colonies in caves. In the Philippines, generally found in chambers to which some light penetrates (R. C. B. Utzurrum, pers. comm.). Colonies may be large (e.g., tens of thousands [Start and Marshall, 1976]; thousands [authors' data]) but most Philippine colonies are well under 500. These smaller colonies may be the result of the hunting of bats at roosts or may reflect a greater abundance of suitable cave roosts. There is some segregation by sex and age at roosts, with individual clusters composed of males, females or subadults (Bhat et al., 1980 and authors' data).
A pattern of sexual segregation by elevation suggests that the maternity roost requirements (caves) tie most reproductive females to suitable roosts in the lowlands in the Philippines, where caves tend to be more common than in the volcanic uplands. Start and Marshall (1976) noted that adult males preferentially foraged in mangrove forests and females preferred inland forest, while subadult bats stayed closer to the roost cave than adults.
In the Philippines, commuting distances of at least 10–15 km are likely in many areas; individuals are commonly captured at these distances from known cave sites. Has been reported to forage in flocks, usually of 5–20 but occasionally as many as 50 (Start and Marshall, 1976) and observations in the Philippines provide some evidence for small flocks. Start (1974) found that marked individuals returned to the same site for weeks or months after marking, despite the extensive foraging range he documented (Start, 1974; Start and Marshall, 1976).
Population biology: Observations by Kitchener et al. (1990) on Lombok suggest that by October, prior to the onset of the wet season, the period of births had all but finished, or at least was going through a temporary lull, and that most young had assumed a degree of independence. One-third of females of adult size had not bred. There was no indication of asynchronous breeding at Batu Koq, Pelangan, Suranadi and Kuta. At the end of the wet season there were few juveniles captured but all adult females were in active breeding condition. Goodwin (1979) reported a lactating female on Timor in March, but located no nursery roosts during March, April and May, nor were juveniles collected during that period.
In the Philippines, most populations apparently reproduce asynchronously, either entirely aseasonally or with only slight seasonal variation (Heideman, 1987).
Most adult females produce two young per year in the Philippines (Heideman, 1987). Healthy females could possibly produce three young per year, allowing about 4 months per cycle.
Females appear to become fertile at about 6 months, with males maturing later, possibly as late as 1 year or more.
Occurrence in protected areas:
Borneo
Sabah
Gomantong Caves
Sarawak
Niah Caves
Summary of threats:
Recommended action:
Principal authors for this subspecies: P. D. Heideman, L. R. Heaney.
E. s. rosenbergii
Sulawesi dawn bat
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: North Sulawesi.
Status: Hunted for food (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
E. s. spelaea
Dawn bat, Cave fruit bat
Common nectar-feeding fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: Andaman Islands; Burma; south China; north-east and north-west India; west Jawa; Peninsular Malaysia (including islands of Langkawi and Tioman); Singapore; Sumatra; Thailand; Vietnam.
Status: Peninsular Malaysia Widespread and locally common on the mainland. Beck and Lim (1973) reported colonies of tens of thousands in large limestone massifs in Peninsular Malaysia.
Thailand Rather common throughout Thailand (Lekagul and McNeely, 1977).
Vietnam Recorded from Quang Triand Tuyen Doc Provinces (Van Peenen, 1969).
In Jawa, considered endangered; threatened by habitat destruction, cave disturbance and severe hunting pressure (A. Suyanto and D. Kitchener. pers. comm: see E. s. glandifera).
In Peninsular Malaysia, threatened by forest clearance in general, and of mangrove in particular. Also threatened by blasting, guano digging and disturbance of caves.
Ecology: In Peninsular Malaysia, is a generalist utilizing both agricultural land and primary forest.
In Peninsular Malaysia, feeds on the following plants (Start and Marshall, 1976 and authors' data): Bombacaceae: Bombax spp. (FL), Durio zibethinus (FL), Durio spp. (wild varieties) (FL); Lecythidaceae: Barringtonia spp. (FL); Leguminosae: Parkia spp. (FL); Moraceae: Artocarpus spp. (FL); Myrtaceae: Eugenia malaccensis (FL); Sonneratiaceae: Duabanga grandiflora (FL), Sonneratia spp. (FL); Verbenaceae: Avicennia spp. (FL).
In Peninsular Malaysia, may travel up to 38 km to feed (Start and Marshall, 1976).
Population biology: Bhat et al. (1980) found that an Indian colony reproduced aseasonally and asynchronously, as did the population studied in Peninsular Malaysia by Start (1974) and, probably, that studied by Beck and Lim (1973) in the same country. In contrast, Lim (1973) reported seasonal reproduction in yet another Peninsular Malaysian population.
Beck and Lim (1973) estimated a gestation of about 6 months, but the accuracy of their estimate was questioned by Start (1974); Bhat et al. (1980) estimated a gestation of 3–4 months and a lactation of 5–8 weeks.
In Peninsular Malaysia, a marked animal was at least 5 years old at the time of recapture (Start, 1974).
Occurrence in protected areas:
Peninsular Malaysia
Batu Caves (protected from quarrying since 1980)
Krau Game Reserve, Pahang
Kuala Selangor Nature Park, Selangor (only seen feeding)
Taman Negara National Park
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
Genus Epomophorus (6 species)
Priority Grade: 6 (Rare).
Distribution: Angola; Namibia.
Status: Restricted distribution, occurring only in woodland areas in western Angola and the extreme north of Namibia, but is locally common (Bergmans, 1988; Crawford-Cabral, 1989) Must, however, be susceptible to habitat destruction.
Feiler (1988) described a trip to the National Park of Kangandala in 1982, and gave data on the massive decline of large mammals because of the civil war. Feiler's conclusion was that only peace, together with well-organized habitat and species protection supported by international help, could save the large mammals from extinction. It is not likely that small mammals such as Epomophorus have suffered, but habitat protection is necessary for all species.
Ecology: As a fruit-eating species, probably confined to areas of riverine and other types of evergreen forest where there are fruit-bearing trees (Smithers, 1986).
Shortridge (1934) recorded that in Ovamboland bats hung singly from the bare branches of large Acacia (Leguminosae) trees near the Cunene River.
Eats cultivated tree fruits (Monard, 1935).
Population biology: Shortridge (1934) recorded newly-born young clinging to their mothers in September and October in Ovamboland.
Occurrence in protected areas:
Angola
Parc Nacional da Mupa
Parc Nacional do Bikuar
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
Epomophorus gambianus (Photo by A. M. Hutson)
E. g. crypturus
Peter's epauletted fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: Angola; Botswana; Malawi; Mozambique; Namibia; South Africa; Tanzania; south-east Zaire; Zambia; Zimbabwe.
Status: Probably not threatened.
Ecology: Predominantly associated with evergreen forests in the higher rainfall areas but occurs in evergreen riverine forests in otherwise dry and unsuitable terrain (Smithers, 1986). Requirements are exemplified in northern Botswana, where it is common throughout the Okavango delta, with its riverine vegetation. It is not found in the drier associations of mopane, (Leguminosae: Colophospermum mopane), Acacia spp. (Leguminosae) or Terminalia spp. (Combretaceae) that surround it and offer insufficient cover and food. It has, however been recorded from mopane woodland in Zimbabwe (A. M. Hutson, pers. obs.). Where there are isolated areas of slightly higher rainfall within otherwise drier country, such as in the Fort Victoria or Matopos Hills areas in Zimbabwe, with pockets of evergreen vegetation supported by run-off from granite bosses, it settles in the moister woodlands and moves into the dry surrounding country to feeding sites. In some areas, man has provided additional suitable conditions in orchard developments, where fruits such as guavas (Myrtaceae: Psidium gaujava), plums (Rosaceae: Prunus domestica), mangoes (Anacardiaceae: Mangifera indica) and other soft fleshy fruits provide it with food and where surrounding exotic trees such as Cyperus sp (Cyperaceae). provide the dense foliage shelter required for roosting (Smithers, 1986).
Occurs in colonies up to hundreds of individuals. In Zimbabwe, commonly uses trees such as the wild fig (Moraceae: Ficus spp.) or the sausage tree (Bignoniaceae: Kigelia pinnata), but will use any evergreen tree provided it has dense foliage and twigs thin enough to allow the bats to cling on to them (Smithers, 1986). In Botswana, used 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 stems were weighed down to within 3 m of the ground (Smithers, 1986).
Feeds on the following plants (Smithers and Wilson, 1979; Pineaar et al., 1980; Smithers, 1986): Anacardiaceae: Mangifera indica (FR), Sclerocarya birrea caffra (FR); Apocynaceae: Rauvolfia caffra (FR); Bignoniaceae: Kigelia pinnata (FL); Bombacaceae: Adansonia digitata (FL); Caricaceae: Carica papaya (FR); Chrysobalanaceae: Parinari curatellifolia (FR); Ebenaceae: Diospyros mespiliformis (FR), D. senensis (FR); Euphorbiaceae: Uapaca kirkiana (FR); Leguminosae: Xanthoceras zambesiaca; Moraceae: Ficus spp. (FR); Myrtaceae: Psidium guajava (FR); Rhamnaceae: Berchemia discolor (FR); Rosaceae: Cydonia sp. (FR), Eriobotrya japonica (FR), Prunus spp. (FR); Sapindaceae: Litchi chinensis (FR); Sapotaceae: Mimusops zeyheri (FR).
Was observed feeding with Epomops dobsonii and foraging on the same Diospyros senensis shrubs as Rousettus (Rousettus) aegyptiacus (Ansell, 1960; Thomas and Fenton, 1978).
Smithers (1986) noted that it eats orchard and garden fruits such as guava, apricots (Prunus armeniaca), peaches (Prunus persica) and loquats (Eriobotrya japonica) and if papaya (Carica papaya) are left to ripen on the tree, damages these by biting into them and scratching them as they cling. Apples (Rosaceae: Malus spp.), pears (Rosaceae: Pyrus spp.) and other hard fruits remain untouched.
Population biology: Females normally bear a single young annually between September and February (Smithers and Wilson, 1979; Pienaar et al., 1980; Rautenbach, 1982). Twins are known (Smithers, 1986).
Occurrence in protected areas:
Angola
Parc Nacional de Cameia
Botswana
Moremi Wildlife Reserve (Okavango)
South Africa
Kruger National Park
Zaire
Parc National de l'Upemba
Zambia
Luangwa Valley
Nyika National Park
Zimbabwe
Charara Safari Area
Chizarira National Park
Gonarezhou National Park
Hwange National Park
Inyanga National Park
Matopos National Park
Selukwe Reserve
Zambezi National Park
Summary of threats:
Recommended action:
E. g. gambianus
Gambian epauletted fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: Benin; Burkina Faso; Cameroun; Central African Republic, Chad; Ethiopia; Gambia; Ghana; Guinea; Guinea-Bissau; Ivory Coast;?Liberia; Mali; Niger; Nigeria; Senegal; Sierra Leone; Sudan; Togo.
Status: Considered ‘widespread and common’ in West Africa (Rosevear, 1965). Happold (1987) considered it to be the commonest of the non-colonial species of fruit bats in Nigeria. Probably not threatened.
Hunted and eaten locally but this is probably not a serious threat yet.
Burkina Faso partially protects this species under wildlife regulations.
Ecology: Common throughout savannahs and woodlands of western Africa. The Sahel Acacia-wooded grassland and deciduous bushland forms the northern limit. Is a woodland species that is found along the forest edge (Bergmans, 1988). Specimens have been collected in the closed rain forest south-west of Kumasi, Ghana (Rosevear, 1965).
Roosts in large hollow trees, dense foliage, accumulated roots along stream banks, and below the thatch of open sheds. Often roosts where there is considerable light, and sometimes groups of about six hang from the midribs of palm fronds (Palmae) in plain sight (Nowak, 1991). Roosts in a variety of trees, including Khaya senegalensis (Meliaceae), Kigelia pinnata (Bignoniaceae), Azadirachta indica (Meliaceae), Cola sp. (Sterculiaceae), Mangifera sp. (Anacardiaceae), Trichilia emetica (Meliaceae), and Ficus sp. (Moraceae) (Baker and Harris, 1957; Rosevear; 1965; Ayensu, 1974; Poche, 1975; Thomas and Fenton, 1978; Marshall and McWilliam, 1982). Bats roosting in a neem tree (Meliaceae: Azadirachta indica), did not feed on its fruit (Ayensu, 1974). The African Mammal Project of the National Museum of Natural History, Washington DC, observed roosting with Micropteropus pusillus in a neem tree in Ghana.
Feeds on the following plants (Baker and Harris, 1957; Booth, 1959; Harris and Baker, 1959; Rosevear, 1965; Ayensu, 1974; Dobat and Peikert-Holle, 1985; Fujita and Tuttle, 1991 and authors' data): Anacardiaceae: Anacardium occidentale (FR), Mangifera indica (FR); Bignoniaceae: Kigelia pinnata (FL); Bombacaceae: Adansonia digitata (FL), Bombax spp. (FL), Ceiba pentandra (FL); Caricaceae: Carica papaya (FR); Chrysobalanaceae: Maranthes polyandra (FL); Ebenaceae: Diospyros mespiliformis (FR), D. senensis (FR); Leguminosae: Parkia clappertoniana (FL), P. filicoidea (FL); Meliaceae: Azadirachta indica (FL), Trichilia roka (FR); Moraceae: Ficus umbellata (FR); Musaceae: Musa spp. (FL); Myrtaceae: Psidium guajava (FR); Sapotaceae: Vitellaria parkii (FR).
Rather nomadic depending on the location of suitable food. At Ibadan in Nigeria, for example, it was particularly common when mango (Mangifera indica) and Parkia trees were in flower, and when mangoes were fruiting, but at other times was seen less frequently (Happold and Happold, 1978) Many individuals congregated to feed at a fruiting tree, while at other times bats probably fed singly or in small groups (Happold, 1987).
Probably pollinates some trees, especially Parkia clappertoniana and the silk cotton tree (Ceiba pentandra) (Baker and Harris, 1957; Harris and Baker, 1959). These trees have large spherical inflorescences, which hang downwards and are supported on thick stems. When the bats come to feed on the nectar, they land on the inflorescences and push their noses into the cluster of flowers, flapping their wings to maintain balance. Pollen adheres to the pelage of the neck and ventral surface, and is later transferred to other inflorescences on the same or different trees. They usually arrive at the flowering trees at dusk, feeding for only 20–30 minutes and leave due to the arrival of the smaller, but dominant, Veldkamp's dwarf fruit bats (Nanonycteris veldkampii). Thus both species utilize the same food resource, but at different times (Happold, 1987). E. gambianus generally feeds singly (Poche, 1975; Thomas and Fenton, 1978), but was observed feeding on Adansonia digitata in mixed-species flocks with Micropteropus pusillus and N. veldkampii. In these observations, none of the bat flocks stayed more than 45 minutes; arrival and departure of the flocks was not synchronized. This species may be a dispersal agent for guava (Psidium guajava), cashew (Anacardium occidentale) and neem. It has been observed leaving neem trees carrying the fruits. However, the relationship between this species and mango trees is a one-sided interaction; the bats were observed destroying as much as one-third of the fruit while foraging, but did not seem to pollinate the flowers or disperse the fruit (Ayensu, 1974).
There are a few records of predation. Pied crows (Corvus albus) have been observed to prey upon roosting bats during the day but only when the crows were raising young (Smalley, 1984). A specimen in the British Museum is recorded as having being killed by a crow (Rosevear, 1965).
Population biology: Females collected in western Niger during the peak of the dry season (February–May) were either pregnant or lactating. Pregnant females were collected in February, March and May. Lactating females were taken in February. Weaning probably occurs when abundant food resources are available during the period of maximum vegetative growth in the summer (June–August) rainy season (Poche, 1975). This limited information, probably also applicable to northern Nigeria, suggests that breeding occurs during the dry season (Happold, 1987).
In Burkina Faso, pregnant females were captured in March, May, and September (Koopman et al., 1978).
Marshall and McWilliam (1982), working in Ghana, concluded that this species gave birth at the beginning of the rainy season (April), but that it may be polyoestrus. Thomas and Marshall (1984), classed it as bimodally polyoestrus in the Ivory Coast.
This species may thus be classified as bimodally polyoestrus with two parturition periods, followed by post-partum oestrus, during the rainy season. The timing of the breeding seasons may be determined by the lactating female's need for food rather than the needs of the young after weaning. The first period of birth occurred in April, coinciding with the onset of the rainy season, and the second in October, the last rainy month. The first lactation occurred during peak fruiting; the second followed 5–6 months later, the length of gestation, regardless of available resources (Thomas and Marshall, 1984).
Occurrence in protected areas:
Burkina Faso
Po National Park
Central African Republic
Bamingwi-Bangoran National Park
Ghana
Mole Game Reserve
Shai Hills Game Reserve
Niger
Park W
Nigeria
Borgu Game Reserve
Darazo Forest Reserve
Kainiji Lake National Park
Nagaruta Forest Reserve
Pandam Wildlife Park
Upper Ogun Game Reserve
Yankari Game Reserve
Summary of threats:
Recommended action:
E. g. pousarguesi
De Pousargue's epauletted fruit bat
Priority Grade: 6 (Rare).
Distribution: Central African Republic.
Status: Known only from the type locality (between Mokorou and Yabanda) and from Bangui, both in the Central African Republic (Bergmans, 1978a). Probably not threatened.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 5 (Rare: Limited Distribution)
Distribution: Angola; Congo.
Status: Known only from the type locality (Dundo) in Angola and Pointe Noire in Congo.
Population biology: A male caught by W. Bergmans in December 1972 at Pointe Noire was a very young adult. The female holotype collected in September 1948 was carrying a juvenile male with milk dentition.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened)
Distribution: Burundi; Chad; Congo; Ethiopia; Kenya; Malawi; Nigeria; Rwanda; Sudan; Tanzania; Uganda; north-east Zaire.
Status: Probably not threatened.
Ecology: Occurs in bushlands and woodlands.
Feeds on the following plants (Verschuren, 1957; Verschuren, 1965; Okia, 1974a; Dobat and Peikert-Holle, 1985; Fujita and Tuttle, 1991):
Anarcardiaceae: Mangifera indica (FR); Bignoniaceae: Kigelia aethiopica (FL), K. pinnata (FL); Moraceae: Ficus natalensis (FR), F. vallis-choudae (FR); Simaroubaceae: Irvingia smithii (FR).
Population biology: Seasonally polyoestrus with a bimodal pattern, conceptions occurring twice a year in late March/early April and late September/early October, with births in September and March after a 5–6 month gestation.
Occurrence in protected areas:
Malawi
Kasungu National Park
Rwanda
Parc National de Kagera
Sudan
Nimule National Park
Uganda
Murchison Falls National Park
Queen Elizabeth National Park
Zaire
Parc National de Virunga
Parc National de la Garamba
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened)
Distribution: Ethiopia; Kenya; Malawi; Rwanda; Somalia; Sudan; Tanzania (mainland, Zanzibar); Uganda; south-east Zaire; Zambia.
Status: Probably not threatened.
Ecology: Found in woodlands, savannah areas with forest patches, bushlands, and even drier vegetation types.
Feeds on the following plants: Combretaceae: ?Terminalia catappa (FR); Moraceae: Ficus (FR); Salvadoraceae: Salvadora persica (FR).
Population biology: Very little information available on population biology specifically for E. minor because in the past it has been regarded as, or confused with, E. labiatus. The data available suggests that the breeding cycle may be similar to E. labiatus, with births in March followed immediately by a second pregnancy.
Occurrence in protected areas:
Kenya
Meru Game Reserve; species not identified
Malawi
Kasungu National Park
Tanzania
Lake Manyara National Park
Zambia
Luangwa Valley; species not identified
Lukusuzi National Park
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
On the basis of differences in dimensions many authors, e.g. Hayman and Hill (1971), have long recognized two subspecies, wahlbergi and haldemani, but Bergmans (1988) has shown that this is not tenable.
Distribution: Angola; Burundi; Cameroun; Congo; Equatorial Guinea (Mbini);?Ethiopia; Gabon; Kenya; Malawi; Mozambique; Namibia; Somalia; South Africa; Swaziland; Tanzania (mainland, Pemba, Zanzibar); Uganda; south-east and south-west Zaire; Zambia; Zimbabwe.
Status: Not threatened.
Readily visible and easily shot. Known to cause damage to litchi orchards (Sapindaceae: Litchi chinensis) in South Africa, although Rousettus (Rousettus) aegyptiacus is more important in this respect.
In 1980, rabies serotype 2 (Lagos Bat Virus) was isolated from specimens captured in and around Durban, South Africa. The infected bats were known, in at least one case, to have flown at (attacked?) a human. Lagos Bat Virus, however, is not known to affect humans. This incident, nevertheless, precipitated local bat extermination programmes and if the problem arose again, could seriously threaten insectivorous bat populations as well as fruit bats.
The male bats, with their repetitive bell-like call, are known to keep people awake all night. This, coupled with the problem of faeces splashed over white walls, causes roosts to be sought out and the bats shot.
Dried bodies of this species are used to a limited extent by Zulu witchdocters as ‘muti’, but probably not enough to constitute a threat.
Ecology: Occurs in tropical forest and evergreen riverine forests where there are fruit-bearing trees. While its occurrence lies largely within areas with a mean annual rainfall in excess of 700 mm, it penetrates up river valleys carrying evergreen forest into otherwise much drier country with a minimum annual rainfall as low as 250 mm (Limpopo Valley) (Smithers, 1986).
Roosts in the canopies of dense evergreen trees, sometimes in colonies numbering dozens of individuals and often in association with E. gambianus crypturus, which is always in much greater numbers (Smithers, 1986). In coastal cities in Mozambique it roosts in evergreen trees in city parks, and even in trees along busy streets (Smithers, 1986).
Feeds on the following plants (Kingdon, 1974; Wickler and Seibt, 1976; Sowler, 1984; Dobat and Peikert-Holle, 1985; Smithers, 1986; Fujita and Tuttle, 1991). Anacardiaceae: Anacardium occidentale (FR), Mangifera indica (FR); Bombacaceae: Adansonia digitata (FR); Caricaceae: Carica papaya (FR); Chrysobalanaceae: Parinari curatellifolia (FR); Combretaceae: Terminalia catappa (FR); Ebenaceae: Diospyros squarossa (FR); Euphorbiaceae: Uapaca kirkiana (FR), U. sansibarica (FR); Moraceae: Ficus sycamorus (FR), F. natalensis (FR), F. polita (FR), F. vogelii (FR); Musaceae: Musa spp. (FR); Myrtaceae: Psidium guajava (FR), Syzygium cordatum (FR); Rosaceae: Prunus armeniaca (FR), P. domestica (FR); Sapindaceae: Litchi chinensis (FR); Zygophyllaceae: Balanites wilsoniana (L).
Moves considerable distances in its search for food. At preferred feeding sites it may settle in the cover of evergreen trees during the day and then quite suddenly, when the supply is exhausted, move off elsewhere (Smithers, 1986).
Population biology: Conceptions occur from May to December, the peak months being May, June and July. Births occur from October to June with the peak birth season in November and December. The majority of females undergo one pregnancy per year terminating in November/December with a small percentage terminating in April (Sowler, 1984). In Kenya, Wickler and Seibt (1976) noted births occurring in January, while O'Shea and Vaughan (1980) found volant young, pregnant and lactating females continuously from November to May. In Zaire, Anciaux de Faveaux (1972) described a biannual cycle with births in March and in October and November, but did not exclude the possibility of continuous polyoestrus reproduction. Reproductive information extracted from specimens collected in Congo, Zaire, Kenya, Tanzania and Zambia support Anciaux de Faveaux's (1972) suggestion.
Occurrence in protected areas:
Angola
Reserva do Luando
Kenya
Amboseli Game Reserve
Masai Mara Reserve
Serengeti National Park
Tsavo National Park
Mozambique
Parque National de Gorongosa
South Africa
Addo Elephant Park
Empisini Nature Reserve (Umkomaas)
Enseleni Nature Reserve
Hans Merensky Nature Reserve
Harold Johnson Nature Reserve
Hluhluwe Game Reserve
Inanda Game Reserve
Itala Nature Reserve
Kenneth Stainbank Reserve
Keurbooms Reserve
Kosi Bay Nature Reserve
Kruger National Park (and adjacent private game reserves on western boundary)
Loskop Dam Nature Reserve
Mkuzi Game Reserve
Ndumu Game Reserve
Ngoye Forest Reserve
Oribi Gorge Nature Reserve
St Lucia Lakes Game Reserve
Sodwana Nature Reserve
Umfolozi Game Reserve
Umgeni Valley Nature Reserve
Umlalazi Nature Reserve
Umtamvuna Nature Reserve
Vernon Crookes Nature Reserve
Wolkberg Wilderness Area
Swaziland
Hlane Game Reserve
Malolotja Nature Reserve
Mlilwane Wildlife Sanctuary
Tanzania
Manyara National Park
Mikumi National Park
Selous Game Reserve
Serengeti National Park
Zaire
Parc National de Virunga
Zimbabwe
Chizarira National Park
Gonarezhou Game Reserve
Inyanga National Park
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Epomops (3 species)
Priority Grade: 4 (Vulnerable)
Distribution: Ghana; Guinea; Ivory Coast; Liberia; Nigeria; Sierra Leone.
Status: As a forest species, will suffer from the severe forest destruction in West Africa and should be considered vulnerable, or possibly endangered.
Ecology: An inhabitant of the wetter and drier types of the Guineo-Congolean rain forests of the western Upper Guinea forest block and the adjoining mosaic of lowland rain forest and secondary grassland, with some possibly isolated occurrences in the eastern Upper Guinea region (the Nigerian localities). In Liberia is reported as favouring areas of secondary bush or cultivated land in preference to primary rain forests (Wolton et al., 1982).
Feeds on the following plants (Rosevear, 1965; Wolton et al., 1982; Dobat and Peikert-Holle, 1985; Fujita and Tuttle, 1991) Bignoniaceae: Spathodea campanulata (FL); Bombacaceae: Ceiba pentandra (FL); Combretaceae: ?Teminalia catappa (FR); Ebenaceae: Diospyros mespiliformis (FR); Leguminosae: Parkia roxburghii (FL); Moraceae: Chlorophora excelsa (FR), Ficus vallis-choudae (FR); Musaceae: Musa spp. (FL?); Myrtaceae: Psidium guajava (FR); Passifloraceae: Adenia cissampeloides (FR), Smeathmannia pubescens (FR); Solanaceae: Solanum erianthum (FR), S. torvum (FR).
Population biology: Bergmans (1975a) suggested two breeding cycles per year for Ivory Coast populations. More extensive research showed that in the Ivory Coast there are two parturition periods (March and September) occurring during the two rainy seasons, each followed by apparently immediate embryonic development (Thomas and Marshall, 1978). On this basis it could be classified as bimodally polyoestrus. In Liberia, Wolton et al. (1982) also found two breeding peaks per year, births occurring just before the main rainfall peak in July/August and again at the end of the rains in late November/December. It seems to compress its breeding seasons more closely than E. franqueti, resulting in a gestation of 3–4 months.
Occurrence in protected areas:
Ghana
Bimpang Forest Reserve
Ivory Coast
Tai National Park
Liberia
Mount Nimba
Nigeria
Pandam Wildlife Park
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened)
Distribution: Angola; Botswana; Malawi; Rwanda; Tanzania; south-east Zaire; Zambia; Zimbabwe.
Status: Probably not threatened.
Ecology: Largely restricted to woodland vegetation, dominated by Brachystegia, Julbernadia and Isoberlinia (Leguminosae). Is restricted by shrubland belts and desert vegetation of the Atlantic coast, by the Lower Guinea rain forest block in the north and by the drier vegetation of the Zambezian deciduous forest and secondary grassland to the south, south-east and east.
Population biology: Females caught at Mount Soque in Angola at the end of August were either pregnant or lactating, which suggests that births occur in August/September. There is no information to suggest bimodality.
Occurrence in protected areas:
Botswana
Chobe National Park
Malawi
Kasungu National Park
Zaire
Parc National de l'Upemba
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
Two subspecies have been described, E. f. franqueti (Tomes) and E. f. strepitans Andersen. Bergmans (1989) found these to be poorly differentiated, and synonymized them.
Distribution: Angola; Benin; Cameroun; Central African Republic; Congo; Equatorial Guinea (Mbini); Gabon; Ghana; Ivory Coast; Nigeria; Sudan; Tanzania; Togo; Uganda; Zaire; Zambia.
Status: Unknown, but probably on the whole not threatened. Happold (1987) noted that it did not seem to be as common in the western area of Nigeria as Epomophorus gambianus, Micropteropus pusillus and Nanonycteris veldkampii but was very numerous in some parts of its geographical range such as Gabon (Brosset, 1966b).
Ecology: Essentially a species of the Guineo-Congolian lowland rain forest, occurring in both wetter and drier types. Ranges from eastern Upper Guinea to the whole of Lower Guinea. Within the forest belt it probably does not enter the large area of swamp forest in north-east Congo and adjacent Zaire. On the continental side of the rain forests, occurs in various mosaics of rain forest with woodland and grassland (Bergmans, 1989).
During the day roosts, usually in groups, suspended from small branches, 4–6 m above the ground and often close to water (Brosset, 1966b; Jones, 1972).
Feeds on the following plants (Jones, 1972; Bradbury, 1981; Fujita and Tuttle, 1991): Anacardiaceae: Mangifera sp. (FR); Annonaceae: Annona sp. (FR); Combretaceae: Terminalia catappa (FR); Lauraceae: Persea americana; Moraceae: Artocarpus sp. (FR), Ficus sp. (FR); Myrtaceae: Psidium guajava (FR); Solanaceae: Solanum torvum (FR).
Happold (1987) noted that in Nigeria, it was not known to assist in the pollination of trees. Often congregates to feed on fruiting trees, and, therefore, may be locally common while adequate food is available. Little is known about seasonal movements and flight; tends to fly more frequently at 4–6 m above the ground than at 0–2 m (Jones, 1972), but on occasions will feed on fruit on the ground (Kingdon, 1974).
Rosevear (1965) noted that, along with Hypsignathus monstrosus, it is considered a great nocturnal pest of fruit orchards.
Population biology: Working in central Africa, Anciaux de Faveaux (1972) concluded a continuous polyoestrus breeding pattern. Okia (1974b) working in Uganda and Bergmans (1979) in Congo both concluded a bimodally polyoestrus pattern with births timed to occur at the beginning of the two rainy seasons. Okia (1974b) reported a 5–6 month gestation, with copulation in April and early September and births in early September and early February.
Data from museum specimens suggest births in August/September/October in Nigeria, but this may be only a partial picture and the true pattern could be either bimodal or continuous reproduction. Museum data for Cameroun point to bimodality, with births in January/February and in June/July.
Occurrence in protected areas:
Congo
Parc National d'Odzala
Nigeria
Mamu River Forest Reserve
Olokomeji Forest Reserve
Omo Forest Reserve
Uganda
Murchison Falls National Park
Zaire
Parc National de Virunga
Parc National de la Garamba
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Haplonycteris (1 species)
Priority Grade: 4 (Vulnerable).
Distribution: Philippines (Dinagat, Leyte, Luzon, Mindanao, Mindoro, Negros). A record from Palawan (Kock, 1969) is probably erroneous.
Status: Rare even in areas of mixed agricultural land, secondary growth, and patches of secondary forest near to continuous primary forest (Heaney et al., 1989 and unpublished data). Even in areas of continuous primary forest, it was about eight times more numerous within forest as within small clearings in forest (Heideman and Heaney, 1989), in contrast to the pattern for some other small fruit bats (e.g., Cynopterus brachyotis, Macroglossus minimus and Ptenochirus jagorii).
The most abundant bat in primary forest on Negros (55% of captures between 800 m and 1000 m and 59% of captures between 1250 m and 1300 m elevation). About 4 individuals were recorded per hectare in the Balinsasayao region forest on Negros (Heideman and Heaney, 1989).
No evidence of self-sustaining populations in non-forested areas. Too small and inconspicuous to be threatened by hunting, but is seriously threatened by habitat destruction. The forest habitats of this species have been reduced from about 75% of the land area of the Philippines to perhaps 10 or 20% in the last 100 years, and populations have probably declined similarly. Is threatened by forest destruction throughout its range, but there is some evidence that small populations can persist in secondary forest.
Ecology: Has been taken at elevations from below 100 m to 1550 m. One of the smallest (typically 18–20 g) primary and old secondary forest fruit bats in the Philippines (Heaney et al., 1981, 1989; Heideman and Heaney, 1989).
Feeds on several species of figs (Moraceae: Ficus spp.), including Ficus chrysolepis. Captives readily ate fruits of Musa sp. (Musaceae), and it is likely that they also feed on fruits of at least one species of Piper (Piperaceae). Numbers seem to be positively correlated with the abundance of two species of Piper vines in the Balinsasayao region of Negros, but it is not clear to what extent the species relies on Piper fruits or vice versa.
Flies and forages largely in the understorey and in the subcanopy, but also feeds in the canopy. Captured in small clearings (< 1 ha) and occasionally along edges of larger clearings, but rarely more than 10 m from the forest edge. Feeding roost sites usually were branches less than 5 m high on a sapling, often out of sight of possible fruit source trees. Individuals in the Balinsasayao region of Negros moved, on average, 300 m between net captures (Heideman and Heaney, 1989).
Heideman et al. (1987) found teeth from a single H. fischeri in one of 23 carnivore scats from the Balinsasayao region that contained small mammals. The teeth were far more worn (to the roots) than any other of this species seen by Heideman and Heaney, suggesting that the bat had been very old when eaten. Although mammalian carnivores may take H. fischeri opportunistically, the bats' primary predators are probably owls and arboreal snakes.
Population biology: Heideman (1988, 1989) describes in detail the reproductive pattern in the Balinsasayao region. Females produced one young per year; births were highly synchronized in late May, June and early July, the earliest part of the wet season. The duration of lactation was about 3 months, ending in September or early October. The timing of birth periods was different on some other islands (Heideman, 1988). Females reached maturity at about 3–5 months while males were not mature until 10–11 months.
Individuals have been documented to live at least 4 years in the wild; their apparently high annual survivorship rate (about 80%) suggests that about 5–10% of yearlings reach an age of about 10 years (Heideman and Heaney, 1989).
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Genus Harpyionycteris (1 species)
For a review of the genus, see Tate (1951).
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
H. w. celebensis
Priority Grade: 10 (No Data).
Distribution: Sulawesi.
Status: Hunted for food in north Sulawesi (Bergmans and Rozendaal, 1988).
Population biology: Females collected in January and September carried embryos. A female caught in January 1983 was carrying a young. Immature but rather large specimens were obtained in January, May, June and July while two nearly full-grown females were obtained in June (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
H. w. negrosensis
Priority Grade: 4 (Vulnerable).
Distribution: Philippines (Negros).
Status: Typical capture rates in habitats where it was most common were about one animal for every 10 net-nights (Heaney et al., 1989; Heideman and Heaney, 1989). Heideman and Heaney (1989) estimated a density of about one individual for every 1.4 ha in the Balinsasayao submontane forest region.
Uses clearings as flyways, and tolerates low levels of disturbance. However, it seems to be intolerant of complete forest removal, and does not persist in agricultural or urban areas. It is not threatened by hunting, but by habitat destruction.
Ecology: Occurs at maximum density in lower montane forest (Heaney et al., 1989). Is relatively rare or absent from upper mossy forest, and requires primary forest or lightly disturbed forest in good condition to maintain populations (Heaney et al., 1989; Heideman and Heaney, 1989). Will venture short distances across cleared areas to feed, but has never been taken more than a few km from good forest.
Feeds most heavily on the bright red fruits of the vine-forming pandans (Pandanaceae: Freycinetia spp.) that are present in virtually all primary forest, and are most abundant in montane forest (Heaney et al., 1989). The diet also includes species of Ficus (Moraceae).
Probably forages primarily in the canopy and upper portion of the subcanopy. Many of the captures were in open areas on ridgetops or along roads, although some were captured on densely vegetated ridgetops in primary forest. Evidence suggests that it has a moderately large home range (Heideman and Heaney, 1989).
Population biology: There appear to be two synchronous birth periods, one in January and early February (mid-rainy season) and the second in July and early August (early rainy season; Heideman, 1987). Gestation is about 4–5 months and lactation 3–4 months. Most females apparently have two young per year, but at least a few females fail to produce young during the January/February birth period. Some females give birth for the first time at the age of one year (Heideman, 1987).
Summary of threats:
Recommended action:
Principal authors for this subspecies: P. D. Heideman, L. R. Heaney.
H. w. whiteheadi
Priority Grade: 10 (No Data).
Distribution: Philippines (Biliran, Camiguin, Leyte, Maripipi, Mindanao, Mindoro).
Status: On Leyte, occurs at maximum density in lower montane forest (Heaney et al., 1989). Is relatively rare or absent from upper mossy forest, and requires primary forest or lightly disturbed forest in good condition to maintain populations (Heaney et al., 1989; Heideman and Heaney, 1989).
For threats see H. w. negrosensis.
Ecology: See H. w. negrosensis.
Population biology: See H. w. negrosensis.
Summary of threats:
Recommended action:
Principal authors for this subspecies: P. D. Heideman, L. R. Heaney.
Genus Hypsignathus (1 species)
Priority Grade: 11 (Not Threatened).
Distribution: Angola; Burkina Faso; Cameroun; Central African Republic; Congo; Equatorial Guinea (Bioko, Mbini); Gabon;?Gambia; Ghana; Ivory Coast; Kenya; Liberia; Nigeria; Sierra Leone; Sudan; Togo; Uganda; Zaire.
Status: Unknown, but said to be common and widespread throughout forested Africa (Rosevear, 1965). It will be threatened by deforestation.
Ecology: Is a species of moist forest but has occasionally been found outside the original west and central African rain forest blocks. Occurs in both wetter and drier types of the Guineo-Congolian lowland rain forest, in the mosaic of both, in the swamp forest and in the mosaic of swamp forest with lowland rain forest (Bergmans, 1989).
Feeds on the following plants (Lang and Chapin, 1917; Rosevear, 1965; Brosset, 1966a; Kingdon, 1974; Bradbury, 1977; Thomas, 1982; Happold, 1987; Fujita and Tuttle, 1991): Anacardiaceae: Mangifera indica (FR); Annonaceae: Annona spp. (FR); Bombacaceae: Ceiba sp.; Cecropiaceae: Musanga cecropioides (FR); Loganiaceae: Anthocleista sp. (FR); Moraceae: Chlorophora sp., Ficus lyrata (FR), F. ovata (FR), F. scott-ellioti (FR); Musaceae: Musa spp. (FL); Myrtaceae: Psidium gujava (FR); Passifloraceae: Adenia cissampeloides (FR); Solanaceae: Solanum spp. (FR).
Rosevear (1965) noted that this species, along with Franquet's epauletted bat (Epomops franqueti), is a considered great nocturnal pest.
Population biology: Most available data (from Zaire, Uganda, Cameroun, Liberia and Gabon) points to a bimodally polyoestrus breeding pattern. The data from Ghana is inconclusive, but suggests births taking place from July to September.
Bradbury (1977), working in Gabon, found two copulation periods, one in July to August, and the other December to February with births occurring in December to February and July to August. Exhibits a classical lek mating system, with traditional arena locations, highly ritualized male displays, extreme skew in mating success among males and the existence of mating centres on lek sites.
In Liberia, Wolton et al. (1982), inferred from a small sample size that there are two birth peaks, a pronounced one in August/September (mid-rains) and possibly a second peak at the end of the rains (October/December). This differs from Bradbury's (1977) findings that the two birth peaks were timed to coincide with the two dry seasons in Gabon. Growth of the young is fairly slow, with males reaching adult body size at 12–18 months and females at 6–7 months (Happold, 1987).
Occurrence in protected areas:
Congo
Parc National d'Odzala
Ivory Coast
Tai National Park
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Latidens (1 species)
Priority Grade: 5 (Rare: Limited Distribution).
Distribution: India (Madurai District).
Status: Has not been recorded since its description by Thonglongya (1972). The type specimen was collected in 1948, but was incorrectly identified as Cynopterus sphinx.
Summary of threats:
Recommended action:
Genus Macroglossus (2 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 5.
M. m. lagochilus
Dagger-toothed flower bat
Priority Grade: 11 (Not Threatened).
Distribution: Ambon; Banda Islands; Borneo (Brunei, East, South and West Kalimantan, Sabah, Sarawak); Buru; Natuna Islands (?Bunguran Besar, Serasan); Nias; Peleng; Peninsular Malaysia (including islands of Langkawi, Sri Buat and Tioman); Philippines (Cagayan Sulu, Cuyo, Mindanao, Negros, Panay, Samar, Tablas); Sangir Islands (Sangir); Seram; Singapore; Sulawesi; south Thailand; Timor; Vietnam.
Status: Borneo Recorded from most areas including Kota Kinabalu, the Witti Range, Sepilok, Sukau and Tawau in Sabah; near Bandar Seri Begawan in Brunei; the Kelabit Uplands, Niah and Bako in Sarawak; Gunung Kenepai in West Kalimantan; near Kutai in East Kalimantan; upper Sungai Tengah in South Kalimnatan (Payne et al., 1985).
Peninsular Malaysia Widespread in the coastal lowlands and mangrove. Also recorded from the islands of Sri Buat (Johore), Tioman and Langkawi (Medway, 1978).
Philippines Most commonly recorded from habitats that have suffered disturbance either through natural causes (e.g., landslides, treefalls, river courses) or human activities (e.g., orchards and gardens). Sometimes the most regularly recorded bat in such sites (Heaney et al., 1981, 1989; Heideman and Heaney, 1989). Uncommon in other habitats. Disturbed sites are colonized by wild banana (Musaceae: Musa sapientum) and abaca (Musa textilis), which presumably provide abundant food. In disturbed habitats it typically made up 5–20% of total captures (Guererro and Alcala, 1973; Heaney et al., 1989; Heideman and Heaney, 1989).
Heideman and Heaney (1989) estimated a density of about 2 bats per ha in the Balinsasayao region of Negros. This area had been about 15–30% deforested at the time of the study, and held abundant wild bananas and planted abaca in the small clearings.
Thailand Only recorded from Chantaburi Province in the south-east (Yenbutra and Felten, 1986).
Vietnam Recorded from Con Son and the Provinces of Gia Dinh, Long Kranh and Tuyen Doc (Van Peenen, 1969).
Not threatened by habitat destruction. Although it cannot use areas planted with most field crops, it can utilize small areas that have been planted with trees that produce nectar or pollen on which it can feed. It is too small and inconspicuous to be attractive to hunters.
Ecology: In the Philippines, occurs from sea-level to at least 1500 m. It uses a wider range of habitats than any other fruit bat species in the Philippines, from urban orchards at sea-level to mossy forest (Heaney et al., 1989).
In the Philippines, can very occasionally be found roosting beneath the dead leaves of banana. Usually only a single individual or a mother with a single young are found at the roost site. Start (1974) and Start and Marshall (1976) reported that Peninsular Malaysian animals roosted either alone or in small groups in palms, bananas or other trees in the vicinity of their major food source, such as Sonneratia spp. (Sonneratiaceae).
Feeds on the flowers of the following plants (Start and Marshall, 1976; Fujita and Tuttle, 1991): Anacardiaceae: Mangifera sp.; Bombacaceae: Ceiba pentandra, Durio zibethinus; Flacourtiaceae: Muntingia sp.; Musaceae: Musa spp.; Myrtaceae: Eugenia malaccensis, Syzigium malaccense; Palmae: Cocos nucifera; Rhizophoraceae: Rhizophora spp.; Sonneratiaceae: Sonneratia alba, S. caseolaris, S. ovata.
Of these, the most important are cultivated bananas, coconut (Cocos nucifera) and two species of mangroves (Sonneratia alba, and S. caseolaris). In Peninsular Malaysia, is most abundant in mangrove areas, in which it feeds upon all chiropterophilous plants present (Start and Marshall, 1976); in the wider range of habitats used in the Philippines, the diet composition is somewhat broader. In disturbed forest habitats on Negros, it almost certainly feeds heavily on chiropterophilous Musa (those with pendulous inflorescences; Start and Marshall, 1976).
Feeds in much the same way as Eonycteris spelaea, landing on inflorescences and using its long tongue to pick up nectar and pollen, and to groom pollen from its fur (Start and Marshall, 1976; Gould, 1978). Flies in the subcanopy and readily crosses open areas.
In the Balinsasyao region of Negros, typical movements between captures were about 250 m with a maximum of 750 m (Heideman and Heaney, 1989), although sample sizes were small and re-captures rare. An upland population in mixed agricultural land and patches of riverine forest in Peninsular Malaysia foraged solitarily and travelled up to 2 km to feed (Start and Marshall, 1976).
Population biology: In the Philippines, apparently reproduces aseasonally (Heideman, 1987). Evidence suggests that females produce a second young very soon after weaning the current one as a result of a post-partum oestrus. Most females probably produce two young per year in the Philippines, very rarely only one, and a few probably manage three births within 12 months in some years.
Start (1974) found that one of the two populations he studied in Malaysia reproduced aseasonally, but data for the second population strongly suggested seasonal reproduction. He estimated a gestation of 120 +/- 10 days, a lactation of 60–70 days, and an interval between successive pregnancies of 140–160 days.
Low long-term recapture rates in the Balinsasayao forests on Negros (Heideman and Heaney, 1989) imply relatively low survivorship (perhaps 40–50% annual survivorship for adults). However, the low recapture rates could be due to the disappearance of bats from the study area due to migration rather than mortality.
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Principal authors for this subspecies: P. D. Heideman, L. R. Heaney.
M. m. microtus
Priority Grade: 10 (No Data).
Distribution: Bougainville; the D'Entrecasteaux Islands (Fergusson Island); Solomon Islands (Choiseul, Fauro, Guadalcanal, Kolombangara, Malaita, Nggela Sule, San Cristobal, Santa Isabel, Vella Lavella).
Ecology: Collected on Bougainville in both primary and secondary rain forest up to an altitude of 1155 m (Strahan, 1983).
Summary of threats:
Recommended action:
M. m. minimus
Common long-tongued fruit bat
Priority Grade: 10 (No Data).
Distribution: Bali; Jawa; Kangean Islands; Lombok; Madura.
Ecology: Kitchener et al. (1990) mist-netted 29 individuals on Lombok in October 1987. Roughly equal numbers were found at locations ranging from 50–400 m altitude. Most were captured among banana (Musaceae: Musa spp.) plantations but three were captured along a track in dense primary rain forest at Suranadi. Others were mist-netted over water courses at Pelangan and Batu Koq, but none was collected at a coastal site at Desa Kuta.
Population biology: Kitchener et al. (1990) reported that only three of the 13 females, all apparently adult, collected on Lombok in late September and October, 1987 showed any indication of reproductive activity. The single female collected on Lombok in May 1988 was pregnant but not lactating.
Summary of threats:
Recommended action:
M. m. nanus
Northern blossom bat
Priority Grade: 11 (Not Threatened).
Distribution: Aru Islands; Bismarck Archipelago (Admiralty Islands, New Britain); Kai Islands; Misool; New Guinea.
Status: New Guinea Common (Flannery, 1990). Widespread in lowland New Guinea. At some localities it is more common than Syconycteris australis, although it is usually much rarer. With increasing altitude it becomes increasingly rare relative to S. australis. The exception to this is in the Munbil area of the Star Mountains, where M. m. nanus was not much rarer than S. australis at 1000 m.
Ecology: In New Guinea, occurs in a variety of forested areas including rain forest, sago palm swamp (Palmae: Metroxylon spp.), mangroves and paperbark woodland (Myrtaceae: Melaleuca spp.) at altitudes ranging from sea-level to 260 m.
Roosts singly or in small groups in the canopy of large-leafed palms and trees and in the roofs of disused buildings (McKean, 1983).
Undoubtedly a nectar feeder; specimens taken from New Guinea lacked fruit in the stomach (McKean, 1983) and captive individuals refuse to eat fruit. Stomach contents contained pollen grains and occasional (probably accidentally ingested) insect remains (McKean, 1983).
Population biology: In New Guinea it is likely that births may occur at any time of year (Strahan, 1983). Hood and Smith (1989) report uterine storage of sperm in bats on New Guinea.
Summary of threats:
Recommended action:
M. m. pygmaeus
Priority Grade: 11 (Not Threatened).
Distribution: Australia (Northern Territory, Queensland [Murray Island], Western Australia).
Status: Common, although it has a limited distribution (Strahan, 1983). This species is unprotected in Western Australia.
Ecology: In northern Australia has been recorded from lowland rain forest, paperbark swamps, bamboo thickets and monsoon scrub along watercourses, and in banana plantations (Musaceae: Musa spp.) (Strahan, 1983).
Known roost sites include bamboo thickets and the rolled-up young leaves of banana plants.
Feeds actively upon two introduced plants, the sausage tree (Bignoniaceae: Kigelia pinnata), and the century plant (Agavaceae: Agave americana), both of which are pollinated by bats in their countries of origin (Strahan, 1983).
Is an important pollinator of paperbarks, particularly the night-flowering Melaleuca cajuputi (Myrtaceae) (Strahan, 1983).
Population biology: In Northern Territory, births may occur during the dry season in August and September.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
M. s. fraternus
Priority Grade: 8 (No Data: Limited Distribution). Distribution: Mentawai Islands (Siberut, Sipura).
Summary of threats:
Recommended action:
M. s. sobrinus
Priority Grade: 11 (Not Threatened).
Distribution: Burma; Jawa; Krakatau Islands (Krakatau); Nias; Peninsular Malaysia; Sumatra; Thailand.
Status: Jawa Very common in a garden in Jakarta in 1981 (W. Bergmans, pers. comm.).
Peninsular Malaysia Occurs from sea-level up to at least 1800 m. Is rarely recorded in mangroves but is present regularly in lowland and montane forest. Lim (1966) collected most specimens in montane forest, but found it could be locally common in lowlands, old fruit orchards and villages with bamboo.
Thailand Recorded throughout with records from the following provinces: Mae Hong So, Chiang Mai, Chiang Rai, Phetchabun, Nakhon Ratchasima, Uthai Thani, Chanthaburi, Ranong, Surat Thani, Trang, Satun, Songkhla, Pattani and Yala (Lekagul and McNeely, 1977; Yenbutra and Felten, 1986).
Macroglossus sobrinus (Photo by K-G. Heller)
Ecology: In Peninsular Malaysia, roosts singly or in small groups of up to 10 under branches, under roofing near forest, in palm tree crowns, and occasionally in rolled young banana leaves (Musaceae: Musa spp.) where it competes for sites with Myotis spp.
In Peninsular Malaysia, feeds on the pollen and nectar of three species of wild bananas, and some soft fruit. Fujita and Tuttle (pers. comm.) noted that it had also been recorded feeding on the flowers of Duabanga grandiflora (Sonneratiaceae) in Malaya.
Start (1974) estimated that an adult M. sobrinus could survive on the nectar and pollen output of about two or three flowering Musa spp., each of which may flower for up to 5 months. This suggested that home ranges could be very small and/or population density very high where Musa spp. are dense. At a site in Peninsular Malaysia where Musa spp. were scattered in small clusters, Start (1974) found typical movements of about 1–2 km. It appeared that individuals foraged singly or in pairs of a mother and her young (Start and Marshall, 1976). Individuals may be trapline foragers (Gould, 1978), following set routes from plant to plant on successive nights.
Population biology: In Thailand, probably breeds throughout the year (Lekagul and McNeely, 1977).
Occurrence in protected areas:
Peninsular Malaysia
Pasoh Forest Reserve
Taman Negara
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
Genus Megaerops (4 species)
Priority Grade: 11 (Not Threatened).
Distribution: Borneo (Brunei, East and West Kalimantan, Sabah, Sarawak); Peninsular Malaysia; west Sumatra; Thailand; Vietnam.
Status: Borneo Has been netted mainly in tall forest, and recorded from Gunung Kinabalu (up to 1500 m), Danum, Tawau and Tenom in Sabah; upper Sungai Temburong and Tasik Merimbun in Brunei; the Kelabit Uplands in Sarawak; Sungai Kapuas in West Kalimantan; and the Kutai district in East Kalimantan (Payne et al., 1985).
Peninsular Malaysia Occurs from sea-level up to the mountain tops at about 1800 m. Found in forest or dense vegetation and open country (Medway, 1978). Generally found singly and is widespread, although the population density is low. Recent studies in Kuala Lumpur suggest this species may be fairly common in the canopy of lowland forest (C. M. Francis, pers. comm.).
Thailand Found throughout Thailand with records from the following provinces: Nakhon Ratchisma; Ranong; Surat Thani; Satun; Songkhla and Yala (Lekagul and McNeely, 1977; Yenbutra and Felten, 1986).
Vietnam Recorded from the provinces of Khanh Hoa, Kontum, Long Khanh and Ninh Thuan (Van Peenen, 1969).
Ecology: Feeds on figs (Moraceae: Ficus spp.).
Population biology: There is an extended breeding season in the early months of the year (Medway, 1983), with about 50% of females pregnant in February and other pregnancies recorded in June, at least in the Cameron Highlands in Peninsular Malaysia.
Occurrence in protected areas:
Borneo
Brunei
Temburong National Park
Sabah
Kinabalu Park
Sepilok Forest Reserve
Peninsular Malaysia
Krau Game Reserve
Taman Negara
Summary of threats:
Recommended action:
Principal author for this species: G. W. H. Davison.
Priority Grade: 6 (Rare).
Distribution: Jawa.
Status: Rare (A. Suyanto and D. Kitchener, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Distribution: North-east India; Thailand; and Vietnam.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
M. w. albicollis
Priority Grade: 9 (Indeterminate).
Distribution: Borneo (Brunei); south Peninsular Malaysia.
Status: Borneo Records are from the extreme lowlands. This habitat is under greatest threat from development.
Peninsular Malaysia The first record was from Pasoh Forest Reserve, an area of primary lowland dipterocarp forest in the extreme lowlands. Several further individuals have been netted in the canopy of lowland forest in the Krau Game Reserve (C. M. Francis, pers. comm.).
Occurrence in protected areas:
Peninsular Malaysia
Krau Game Reserve
Pasoh Forest Reserve
Summary of threats:
Recommended action:
M. w. wetmorei
Priority Grade: 10 (No Data).
Distribution: Philippines (Mindanao).
Ecology: Probably limited to primary forest (Heaney et al., 1989).
Summary of threats:
Recommended action:
Genus Megaloglossus (1 species)
Priority Grade: 11 (Not Threatened).
The subspecies prigoginei has been proposed (Hayman (1966) in Hayman et al., 1966) to separate specimens from north-east Zaire and ‘West Africa’, on the basis of size. Bergmans and Van Bree (1972) found indications of a clinal size variation and rejected subspecific divisions. This conclusion had been anticipated by Hayman and Hill (1971).
Distribution: Angola; Cameroun; Central African Republic; Congo; Equatorial Guinea (Bioko, Mbini); Gabon; Ghana; Guinea; Ivory Coast; Liberia; Nigeria; Togo; Uganda; Zaire.
Status: Not known. Rosevear (1965) believed that it was probably not very common. Threatened by deforestation.
Ecology: Is a lowland forest species. In Nigeria, most of the specimens were taken in the lower strata of the rain forest or in forest clearings (Happold, 1987). In other parts of Africa, recorded from banana plantations, close to cultivated areas, and over water (Brosset, 1966b; Jones, 1971).
Appears to be solitary, but may congregate at suitable flowering trees. In Gabon (Brosset, 1966b) and Ghana (Jeffrey, 1975) groups of males alone have been recorded, and at other times and places, males and females have been found together; this seems to suggest a rather well defined social organization with segregation of the sexes at certain times of year (Happold, 1987).
There is very little direct evidence that this species is predominantly a nectar feeder, as some have thought. It has been suggested that it feeds on the flowers of the following plants (Kock, 1972; Jeffrey, 1975; Dobat and Peikert-Holle, 1985): Bignoniaceae: Crescentia cujete, Kigelia sp.; Caricaceae: Carica papaya; Convolvulaceae: Ipomoea albivenia; Leguminosae: Mucuna flagellipes, Parkia bicolor; Musaceae: Musa sp.
In Nigeria, is found in the rain forest where flowers are present throughout most of the year; even so, may be nomadic or even make seasonal migrations to find flowering trees (Happold, 1987).
Population biology: From specimens caught in Rio Muni (Equatorial Guinea) and Ghana (Jones, 1971) and in Congo (Adam and Le Pont, 1974; Bergmans, 1979) the reproductive pattern is either seasonally polyoestrus with two breeding seasons a year, or aseasonally polyoestrus. If the former is true, births occur in March/April and September/October. However, the sample size is only seven. In Liberia, Wolton et al. (1982), also from a small sample, suggested that the species would appear not to have a well defined breeding season.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Melonycteris (3 species)
An undescribed and highly endangered Melonycteris species occurs on Malaita and San Cristobal in the Solomon Islands (T. Flannery, pers. comm.).
T. Flannery (pers. comm.) considers M. aurantius to be a subspecies of M. woodfordi.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Choiseul, Nggela Sule).
Status: Probably rare (T. Flannery and L. Seri, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: Bismarck Archipelago (Duke of York, New Britain, New Ireland);?New Guinea.
Status: Probably common (T. Flannery and L. Seri, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: Bougainville; Solomon Islands (Alu, Choiseul, Fauro, Guadalcanal, Russell Islands [Pavuvu]).
Status: Abundant (T. Flannery and L. Seri, pers. comm.).
Ecology: Kress (1985) recorded feeding at flowers of Heliconia solomonensis (Heliconiaceae).
Summary of threats:
Recommended action:
Genus Micropteropus (2 species)
Priority Grade: 6 (Rare).
Distribution: Angola; south-west Zaire.
Status: Known from only four specimens; one from Dundo, Angola, one from Luluabourg, Zaire, one from Banana/Netonna, and one from Thysville, Zaire (Bergmans, 1989).
Population biology: The type specimen from Dundo was pregnant at the time of capture (June/July) (Bergmans, 1989).
Occurrence in protected areas: Not known to occur in any protected areas.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
Distribution: Angola; Benin; Burkina Faso; Burundi; Cameroun; Central African Republic; Chad; Congo; Equatorial Guinea (Mbini); Ethiopia; Gabon; Gambia; Ghana; Guinea; Guinea-Bissau; Ivory Coast; Kenya; Liberia; Mali; Nigeria; Senegal; Sierra Leone; Sudan; Tanzania; Togo; Uganda; north-east, south-east and south-west Zaire; Zambia.
Status: General status unknown, but locally common. In Nigeria, common, often occurring in large numbers at fruiting trees (Happold, 1987).
Ecology: In the north remains within the northern limit of the east-west belt of Sudanian and Ethiopian woodlands. In West Africa some localities approach the latitude of 14°N, but in Sudan and Ethiopia it is not known from north of 11°N. The eastern limit of the range appears to be set by the Ethiopian highlands. These are only penetrated by low-lying river valleys on the western side and by the central rift valley in the south, where the only known collecting locality is also the easternmost point of the distribution. In the southern part of the range it remains within the North Zambesian woodland in north-west Angola, central and southern Zaire and on the north-west Zambian border (Bergmans, 1989).
Feeds on the following plants (Harris and Baker, 1959; Green, 1983; Dobat and Peikert-Holle, 1985; Happold, 1987; Fujita and Tuttle, 1991): Anacardiaceae: Anacardium occidentale (FL); Annonaceae: Annona chrysophila (FR), A. senegalensis (FR); Bignoniaceae: Kigelia pinnata (FL), Spathodea campanulata (FL); Bombacaceae: Adansonia digtata (FL), Ceiba pentandra (FL); Chrysobalanaceae: Maranthes polyandra (FL), Parinari polyandra (FL); Leguminosae: Parkia clappertoniana (FL), P. roxburghii (FL); Moraceae: Ficus capensis (FR); Myrtaceae: Psidium guajava (FR); Sapotaceae: Vitellaria parkii (FR).
In Nigeria, occurs in widely scattered localities, and may be very abundant when it congregates to feed on flowering and fruiting trees (Happold, 1987). Nomadic, and in any particular locality numbers fluctuate depending on the availability of food. Feeds on smaller, and perhaps softer, fruits than do the large fruit bats. In addition, also feeds on nectar, and, like Epomophorus gambianus, pollinates the flowers of sausage trees (Kigelia pinnata) (Baker and Harris, 1957; Harris and Baker, 1958). Sausage tree flowers hang on thick stalks and have deep, wide corollas; when a bat alights on the lip of the flower, it pushes its head into the corolla to reach the nectar, and, in doing so, covers the top of its head and shoulders with pollen (Baker and Harris, 1957; Harris and Baker, 1958).
Population biology: Verschuren (1957) working in north-east Zaire, suggested a bimodal pattern of births during November/December and in late February. In Rio Muni (Equatorial Guinea), Jones (1972) suggested breeding occurs throughout the year but mostly in March/April and again in November. Bergmans (1979) reported that in Congo a bimodal pattern occurred with births in September and January/February. In Ghana, births occurred at the beginning and end of the rains in April and the beginning of September (Marshall and McWilliam, 1982). A similar pattern was recorded in the Ivory Coast with births at the beginning and end of the rainy season in April and October (Thomas and Marshall, 1984). It may therefore be concluded that M. pusillus is bimodally polyoestrus with a 5–6 month gestation.
Occurrence in protected areas:
Angola
Reserva do Luando
Burkina Faso
Po National Park
Ghana
Mole National Park
Nigeria
Kainji Lake National Park
Olokomeji Forest Reserve
Pandam Wildlife Park
Zaire
Parc National de Virunga
Parc National de la Garamba
Parc National de l'Upemba
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Myonycteris (3 species)
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: São Tomé and Principe (São Tomé).
Status: A forest species threatened by habitat destruction. Little forest seems to be left on São Tomé (A. Feiler, pers. comm.). Untouched rain forests remain at high altitudes in the centre and south-west of the island (Imboden, 1987; Anon., 1988a). New specimens have been collected from more than one locality on the island by a Spanish expedition though data from these are not yet published.
Occurrence in protected areas: There are no protected areas on São Tomé.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Kenya; Tanzania.
Status: Its known range is the coastal Shimba Forest of south-east Kenya and east Tanzania in Pangani, Bagamoya, Kisarawe and Rufiji districts south-westwards to the east slope of the Uzungwa mountains (K. M. Howell and D. Kock, pers. comm.). It may occur in other forested areas in East Africa (Bergmans, 1980; Schlitter and McClaren, 1981). Status is unknown but as the pressure on the remaining forests in East Africa is very high, should probably be regarded as seriously threatened.
Threatened by habitat degradation, even in the protected Shimba Hills National Reserve, and also by habitat destruction, as in large parts of the Usambaras.
Ecology: Essentially a lowland forest inhabitant, known to occur from below 200 m (the type locality) up to about 900 m (Nguru Mountains). Most lowland fruit bats may still be found at 1500 m, and some even higher.
Occurrence in protected areas:
Kenya
Shimba Hills National Reserve
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
Distribution: Angola; Cameroun; Central African Republic; Congo; Equatorial Guinea (Bioko, Mbini); Gabon; Ghana; Guinea; Ivory Coast; Liberia; Nigeria; Sierra Leone; Togo; Uganda; north-east Zaire; Zambia.
Status: In Nigeria, considered to be the rarest fruit bat, known from only two localities (Happold, 1987). Apparently not uncommon locally, but as a forest species is certainly threatened by habitat destruction and degradation.
Ecology: Recorded from moist forest at low and medium altitudes and surrounding forest-savannah mosaics and woodland-savannah mosaics. In Nigeria, appears to be solitary, living mostly in the rain forest zone or near the forest-savannah boundary (Happold, 1987).
Probably roosts in trees, although one individual in Gabon (Brosset, 1966b) roosted in gaps between the leaves of a banana plant (Musaceae: Musa sp.) about 5 m above the ground.
Observed eating bananas (Brosset, 1966b). Has been caught near these and near guavas (Myrtaceae: Psidium guajava) and mangoes (Anacardiaceae: Mangifera indica). Dobat and Peikert-Holle (1985) report on possible visiting of flowers of Spathodea campanulata (Bignoniaceae), Ceiba pentandra (Bombacaceae) and Parkia roxburghii (Leguminosae).
Population biology: In Liberia, Wolton et al. (1982) reported an extended breeding season, beginning in July and continuing well into the dry season (at least until March), with most females giving birth in September at the peak of the rains. Thomas (1983) working in the Ivory Coast, recorded two birth periods: one in February/March and the other in September/October. Brosset (1966b) reported that in Gabon, births occur twice a year, in June and in December/January, in captivity. In the wild, he caught either pregnant or lactating females from November to March, but sexually inactive females in June and July. Anciaux de Faveaux (1972) postulated a reproductive cycle extending from September to May for Cameroun to Congo, with copulations around September, births in December/January and lactation to May.
Myonycteris torquata (Photo by K-G. Heller)
Occurrence in protected areas:
Ghana
Mole National Park
Congo
Parc National d'Odzala
Zaire
Parc National de la Garamba
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Nanonycteris (1 species)
Priority Grade: 11 (Not Threatened).
Distribution: Cameroun; Central African Republic; Ghana; Guinea; Ivory Coast; Liberia; Nigeria; Sierra Leone; Togo. Previous records from Zaire or Congo are incorrect (Bergmans, 1989).
Status: Apparently not uncommon locally, but as a predominantly West African forest species is no doubt suffering from large scale deforestation. In Ghana is abundant in the forest zone in the dry season and migrates into the southern Sudan savannah in the rainy season (Thomas, 1983). Even if part of the year is spent outside the forest zone, as in Ghana (Thomas, 1983) and very likely also elsewhere (Bergmans, 1989), this habitat is essential for this species.
Ecology: A forest and savannah species. Feeds on the following plants (Harris and Baker, 1959; Ayensu, 1974; Dobat and Peikert-Holle, 1985; Fedden and Macleod, 1986; Fujita and Tuttle, 1991): Bignoniaceae: Kigelia sp. (FL); Bombacaceae: Adansonia digitata (FL), Ceiba pentandra (FL); Caricaceae: Carica papaya (FR); Chrysobalanaceae: Parinari polyandra (FL); Leguminosae: Eperua falcata, Mucuna flagellipes (FL), Parkia clappertoniana (FL), P. roxburghii (FL); Moraceae: Ficus umbellata (FR); Myrtaceae: Psidium guajava (FR); Proteaceae: Protea ellioti (FL); Sapotaceae: Vitellaria parkii (FR).
Population biology: Marshall and McWilliam (1982), working in Ghana, postulated polyoestry, with an extended period of parturition synchronized with the early rains, probably in May and June. Data collected in the Mount Nimba region in Liberia by J. Verschuren and M. J. Coe (in: Wolton et al., 1982) appear to support this, as do data from Cameroun.
Occurrence in protected areas:
Ghana
Mole National Park
Nigeria
Igangan National Park
Pandam Wildlife Park
Sapoba Forest Reserve
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Neopteryx (1 species)
Priority Grade: 5 (Rare: Limited Distribution).
Distribution: North and west Sulawesi.
Status: Hunted for food in north Sulawesi (Bergmans and Rozendaal, 1988).
Ecology: Specimens were caught simultaneously in three mist-nets set over Sungai Tumpah, which runs through lowland primary rain forest (Bergmans and Rozendaal, 1988). Population biology: A female caught in March was carrying an embryo (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Genus Notopteris (1 species)
Priority Grade: 9 (Indeterminate).
Number of subspecies: 2. There are two recognized subspecies, N. m. macdonaldii in Fiji and Vanuatu and the smaller N. m. neocaledonica in New Caledonia (Sanborn and Nicholson, 1950). Some specimens bridge the size gap, but populations in different island groups are presumably isolated. Andersen (1912) noted specimens of N. m. macdonaldii reported to be from Pohnpei, Federated States of Micronesia, by Jentink, but questions the locality. As noted by Medway and Marshall (1975), given the lack of additional specimens from later work on Pohnpei, and the enormous range extension involved, this record is almost certainly erroneous.
N. m. macdonaldii
Priority Grade: 4 (Vulnerable).
Distribution: Fiji (Viti Levu); Vanuatu.
Status: Medway and Marshall (1975) recorded it from all six islands sampled in the New Hebrides in Vanuatu, from Santo in the north to Anatom in the south. All caves were close to sea-level, but animals were netted at all elevations sampled up to 1100 m. The colonies are referred to as large, but no estimates of numbers are given. Some respondents to a recent questionnaire on fruit bats in Vanuatu mentioned this bat, but it is apparently not exploited (Chambers and Esrom, 1988). Both islands (Pentecost Islands and Vanua Lava in the Banks Islands) for which Chambers and Esrom (1988) obtained local names for Notopteris lacked previous literature records. Both Andersen (1912) and Sanborn (1931) report specimens taken over 80 years from what is probably the same locality, Colombo Cave, near Suva, Viti Levu, Fiji. Pernetta and Watling (1978) list this bat as abundant on the large islands of Fiji.
Insufficient recent data are available to assess trends or threats. If experience with New World phyllostomid generalist nectarivores can be used as a guide, the mosaic of regenerating habitats created by periodic storm disturbance or moderate human intervention in rain-forest habitats may favour this bat. However, because it roosts in traditional well-known cave sites, it is highly vulnerable to direct exploitation for food, vandalism, repeated disturbance by guano mining or curious visitors and incidental destruction in the course of quarrying. Evidence of rapid decimation of cave-dwelling pteropodid populations is unfortunately increasingly common (Heaney and Heideman, 1987; D.J. Kitchener, pers. comm.). T. Flannery (pers. comm.) has expressed serious concern about the status of Notopteris in Vanuatu and New Caledonia. Recent faunal surveys, which included visits to known cave roosts, yielded no observations of Notopteris.
International trade in this species is regulated under wildlife laws.
Ecology: Favours lowland forest and patchy secondary forest in Fiji, rather than montane or swamp forest (Pernetta and Watling, 1978). However, D. Watling (pers. comm.) suggests tentatively that Notopteris is more closely associated with intact forest than disturbed habitats. Roosts in caves (see above).
Medway and Marshall (1975) noted that roosts in Vanuatu were dimly illuminated, but from Sanborn's (1931) report on shooting bats in Colombo Cave on Vitu Levu, Fiji, total darkness could be inferred. G. Graham (pers. comm.) reports roosting Notopteris from dark portions of a cave on Vitu Levu.
Morphology (reduced dentition, elongate brush-tipped tongue) argues strongly for a nectar and pollen specialist, but data are scant. Medway and Marshall (1975) frequently netted Notopteris near bananas (Musaceae: Musa spp.); scratches on the bracts of bananas suggested the flowers are an important food source.
Summary of threats:
Recommended action:
N. m. neocaledonica
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: New Caledonia.
Status: Occurs in caves on New Caledonia. Sanborn and Nicholson (1950) found 200 bats at Hienghene and 300 at Poya. They were told that it occupied only one of several hundred caves at Poya and that it sometimes roosts in hollow trees. There is no recent information on status.
The threats this bat faces would be similar to those discussed under N. m. macdonaldii.
The hunting of this species is regulated under wildlife laws.
Ecology: See N. m. macdonaldii.
Sanborn and Nicholson (1950) observed that roosting animals on cave walls in New Caledonia were in clusters of 5–25, resembling some Microchiroptera. Males as well as females with young were taken from single clusters, so there is apparently no sexual segregation while the young are prevolant.
Population biology: Sanborn and Nicholson (1950) collected pregnant animals and young in August, suggesting a birth peak slightly earlier than Pteropus in New Caledonia. They reported one young per female.
Summary of threats:
Recommended action:
Principal author for this species: W. E. Rainey.
Genus Nyctimene (14 species)
Priority Grade: 6 (Rare).
Distribution: Misool; New Guinea.
Status: New Guinea Uncommon (Flannery, 1990). Widely distributed from Geelvink Bay in the west (Thomas, 1922) to Milne Bay in the east (Thomas, 1900). Also known from the Torricelli Mountains.
Ecology: Of the specimens reported upon by McKean (1972), all were collected in primary rain forest, except one from secondary forest and one from a swamp. The altitude ranged from sea-level to 260 m. The single specimen obtained at 990 m in the Torricelli Mountains was captured by a local hunter.
Has been observed feeding on figs (Moraceae: Ficus spp.) (Flannery, 1990).
Population biology: McKean (1972) collected three pregnant females in January and one in February. Greig-Smith (1975) noted that the four specimens collected by him were netted in male-female pairs.
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
N. a. albiventer
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Halmahera; Morotai; Ternate.
Summary of threats:
Recommended action:
N. a. papuanus
Priority Grade: 11 (Not Threatened).
Distribution: Bismarck Archipelago (Admiralty Islands, New Britain); Kai Islands; New Guinea.
Status: Extremely widespread throughout New Guinea and surrounding islands and is the most commonly caught Nyctimene when mist-netting at low altitudes (Flannery, 1990).
Ecology: Commonly caught in both primary and secondary rain forest (Flannery, 1990). A single specimen was taken in Melaleuca (Myrtaceae) savannah, and another in monsoon forest in the Morehead region of New Guinea (Waithman, 1979). Seems to have a greater altitudinal range than most other species in New Guinea, being found as high as 1650 m (McKean, 1972). Most stomach contents examined have included pulped vegetable remains, but one contained beetles and ants, and a second had traces of moths (Vestjens and Hall, 1977). Greig-Smith (1975) recorded that it was caught only at dawn, and all individuals were netted at heights of from 0.5–2 m above the ground.
Population biology: Two females caught in January, three in July and one in August were pregnant with a single embryo (McKean, 1972).
Summary of threats:
Nyctimene albiventer (Photo by B. H. Gaskell)
Recommended action:
Principal author for this subspecies: T. Flannery.
This species was formerly included as a subspecies of N. aello.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Western New Guinea. A record from Halmahera is based on a juvenile N. albiventer in the British Museum (Natural History) (J. Edwards Hill, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: Ambon; Australia (Queensland); Buru; Numfoor; New Guinea; Seram; Sulawesi; Tenimber Islands; Timor; Umboi.
Status: New Guinea Rare (Flannery, 1990). Has only previously been recorded on the mainland of New Guinea at Mari and Bulla, in the Morehead region of southern Papua New Guinea (Waithman, 1979). However, a previously unreported male specimen is held in the Australian Museum; it was collected during the 1885 Royal Geographical Society Expedition to New Guinea, probably along the lower Strickland River.
Sulawesi Hunted for food in north Sulawesi (Bergmans and Rozendaal, 1988).
Ecology: The only comment on diet is the observation by Muller (1841) that the stomach of a specimen contained only fragments of Coleoptera and Diptera.
Population biology: In Sulawesi, females collected in October and January were pregnant (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Seram
Manusela National Park
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Priority Grade: 9 (Indeterminate).
Number of subspecies: 2. The subspecies certans may well represent a distinct species (Flannery, 1990).
N. c. certans
Priority Grade: 6 (Rare).
Distribution: Bismarck Archipelago (New Britain); central and eastern New Guinea.
Status: New Guinea Rare (Flannery, 1990).
Ecology: A female was netted on the margin of a patch of primary rain forest at 830 m at Lake Katubu, Southern Highlands Province, New Guinea (McKean, 1972). Flannery (1990) obtained a single specimen at Telefomin in March 1986 during 8 days mist-netting at the site. It was caught in a mist-net set in a taro garden at 2300 m in an area that also yielded Syconycteris hobbit and S. australis. The folk ‘taxon’Brulim, tentatively identified with this species, is held in awe by the Telefol, who believe it is extremely bad luck to kill one (Flannery, 1990).
Summary of threats:
Recommended action:
N. c. cyclotis
Priority Grade: 10 (No Data).
Distribution: Western New Guinea.
Status: Recorded only from Vogelkop.
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Priority Grade: 6 (Rare).
This was formerly included as a subspecies of N. albiventer.
Distribution: New Guinea.
Status: Rare (Flannery, 1990). Has been recorded previously only on two occasions; once from the Lorentz River (type locality) and once from the upper Fly River (Koopman, 1982). The record from Rauit Village, West Sepik Province (Greig-Smith, 1975) is erroneous, the specimens now being identified as Paranyctimene raptor (Flannery, 1990). Flannery (1990) obtained two specimens in a mist-net at Kiunga, apparently referable to this species.
All of these records are from west of 144°E. Given the difficulty of distinguishing this species from N. albiventer it may well be that it is more abundant and widespread than the above data suggest.
Ecology: The Kiunga specimens collected by Flannery (1990) were caught in a mist-net set next to an oxbow of the Fly River, in a small forest clearing.
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 4.
N. m. geminus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: D'Entrecasteaux Islands (Fergusson Island, Goodenough Island); Louisiade Archipelago (Logeia Island); Trobriand Islands (Kiriwina).
Summary of threats:
Recommended action:
N. m. lullulae
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Muyua.
Summary of threats:
Recommended action:
N. m. major
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Bismarck Archipelago (Duke of York, New Britain, New Ireland).
Summary of threats:
Recommended action:
N. m. scitulus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Choiseul, Guadalcanal, Malaita, New Georgia, Nggela Sule, Shortland).
Summary of threats:
Recommended action:
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Malaita).
Summary of threats:
Recommended action:
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Bismarck Archipelago (New Ireland).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
N. m. minutus
Priority Grade: 10 (No Data).
Distribution: Sulawesi.
Summary of threats:
Recommended action:
N. m. varius
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Buru.
Summary of threats:
Recommended action:
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Philippines (Negros).
Status: Recorded from 200 m to 1300 m, but probably rarely occurs below 600 m. Very rare outside of primary or good secondary forest, and not found in agricultural or urban habitats (Heaney et al., 1989). Does not tolerate disturbance well. Heaney et al. (1989) captured only one at a netting site at 600 m in agricultural land and secondary growth 2 km from continuous primary forest, and suggested that this young animal was dispersing from a narrow strip of forest on a ridge 200 m from the capture point. Made up about 4% of total captures in the Balinsasayao forest region, but was only 1% of total captures at 1280 m, and was not taken at higher elevations. Density at Lake Balinsasayao was about one per 3 ha (Heideman and Heaney, 1989).
Seriously threatened by habitat destruction, but not by hunting. Known breeding populations persist only in a narrow elevational band of forest near the tops of high ridges and on the sides of tall mountains. The upper limit of their distribution is about 1300 m. The current lower limit is about 800–900 m, which is rising rapidly as the forest is cleared. At current rates of deforestation, little habitat for N. rabori will remain in 10 years. Might persist in forest patches for some time, but its low density suggests that small forest patches will not support viable populations. As it is known only from southern Negros, its extinction within the next 20 years is probable unless special measures are taken.
Ecology: Often captured near fruiting figs (Moraceae: Ficus spp.), including Ficus chrysolepis.
Three animals moved between 320 m and 1260 m between first and second captures (Heideman and Heaney, 1989). It is unclear whether these are typical movements for the species, or whether individuals may range over somewhat larger areas.
Population biology: Females produce only one young per year, in April or early May, after a gestation of 4.5–5 months. The duration of lactation is about 3–4 months (Heideman, 1987). Young females become pregnant at about 7–8 months of age, and produce their first young at an age of 1 year. Evidence suggests that males reached sexual maturity at about 1 year.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 11 (Not Threatened).
Distribution: Tropical and subtropical eastern Australia (New South Wales, Queensland) (Richards, 1983).
Nyctimene robinsoni (Photo by W. E. Rainey)
Status: Common (Strahan, 1983).
Ecology: Permanent roosts per se are not utilized, and individuals appear to roost in the vicinity of the tree on which they are feeding at the time (Spencer and Fleming, 1989; C. R. Tidemann, J. R. Nelson and G. C. Richards, pers. comm.).
Based on capture locations and some feeding observations, Richards (1986a) showed that there was a preference for understorey fruits of canopy trees in rain forest. Fruits known to be eaten include: Moraceae: Ficus copiosa (FR), F. nodosa (FR), Ficus sp. (?variegata) (FR); Myrtaceae: Eugenia erythrocalyx (FR), Psidium guajava (FR), Syzygium cormiflorum (FR); Rubiaceae: Randia sessilis (FR).
Population biology: Females give birth to one young between October and December.
Occurrence in protected areas: Common in national parks with rain forest.
Summary of threats:
Recommended action:
Principal author for this species: G. C. Richards.
Priority Grade: 1 (Extinct).
Distribution: Solomon Islands (Santa Cruz Islands).
Status: Probably extinct (T. Flannery, pers. comm.). Last seen in 1907 on Nendo in the Santa Cruz Islands. A subsequent expedition to find this bat was fruitless. As it is a large and distinctive species it is now presumed extinct. Disturbance has probably caused its disappearance (T. Flannery and L. Seri, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
The subspecies N. v. bougainville was formerly considered a subspecies of N. aello and N. v. vizcaccia a subspecies of N. cephalotes.
N. v. bougainville
Priority Grade: 10 (No Data).
Distribution: Bougainville; Solomon Islands (Choiseul, Fauro, Guadalacanal, Kolombangara, Santa Isabel).
Summary of threats:
Recommended action:
N. v. vizcaccia
Priority Grade: 10 (No Data).
Distribution: Australia (Queensland [Moa Island, Torres Strait]); Bismarck Archipelago (Admiralty Islands);?north-east New Guinea; Umboi.
Summary of threats:
Recommended action:
Genus Otopteropus (1 species)
Priority Grade: 9 (Indeterminate).
Distribution: Philippines (Luzon).
Status: Moderately common at middle elevations in primary lowland forest on southern Luzon, and present but uncommon in montane and mossy forest. Given the restricted range and the extent of deforestation on Luzon, it may be threatened.
Ecology: Taken from 200 m to 1750 m at localities scattered throughout the mountains of Luzon. Never been recorded outside of primary or good secondary forest (unpublished data and specimens at University of the Philippines at Los Banos and Philippine National Museum).
Fig (Moraceae: Ficus sp.) seeds were found in the faeces of many individuals.
Population biology: May produce either one or, possibly, two young per year on southern Luzon. Pregnancy may include a period of post-implantation developmental delay. Births occur synchronously within the population.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Genus Paranyctimene (1 species)
Priority Grade: 6 (Rare).
Distribution: New Guinea.
Status: Uncommon (Flannery, 1990). Recorded as far east as Morwuna, Milne Bay Province (Koopman, 1982), and as far west as the upper Fly River (type locality) and the northern foothills of the Star Mountains. Probably widespread in New Guinea, with the lack of records in Irian Jaya almost certainly due to a lack of collecting there (Flannery, 1990).
Ecology: Specimens from Rauit Village (originally referred to Nyctimene draconilla; see Hill, 1983) were taken at 525 m above sea-level on one night in a single mist-net, and seemed to form a group that consisted of three males and one female (Greig-Smith, 1975).
The dentition is most unusual. The canines are extremely fine and the cheekteeth are also long and pointed. While this suggests an unusual diet, there are as yet no studies to indicate what this may be (Flannery, 1990).
Population biology: Females pregnant with a single embryo have been collected in January, February and May (McKean, 1972). Flannery (1990) netted two lactating females in May 1986 near Yapsiei, West Sepik Province. A female collected by Greig-Smith (1975) in July–August had a single young.
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Genus Penthetor (1 species)
Priority Grade: 11 (Not Threatened).
Distribution: Borneo (Central, East, South and West Kalimantan, Sabah, Sarawak); Peninsular Malaysia; Riau Archipelago; Singapore; Sumatra.
Status: Borneo Apparently widespread. There are specimens from: the foot of Gunong Kenepai, Sungai Kapuas, Puruk Tjahu, Barito and Perbuwa in Kalimantan; Sebuyau and Gunong at 610 m in Sarawak; and Tenom, Sabah. It is also found from cave roosts at Sikadar cave and Bidi near Bau, I Division, Lobang Pedang Salai, Sungei Baram, and Gua Lobang Sungei Tinjar, all in Sarawak (Medway, 1977).
Peninsular Malaysia Irregularly distributed mainly in the lowlands but up to 1828 m at the Cameron Highlands (Medway, 1978; Medway, 1983).
Threatened generally by forest destruction.
Ecology: Roosts in limestone caves in quite light conditions near the entrances, in rock clefts and between big river boulders. Medway (1983) suggests that this may limit distribution. May also roost in small clusters or singly in hollow trees.
Eats a wide variety of hard fruits and seeds of forest trees, both of the canopy and understorey layer.
Population biology: Medway (1983) reported that in Selangor there were many pregnancies in September, few in June, none in January, February, March or July. Gives birth to one young each.
Occurrence in protected areas:
Borneo
Occurs patchily in lowland forest and at low densities in most forest reserves and other protected areas, such as:
Brunei
Ulu Temburong National Park (proposed)
Sabah
Kinabalu National Park
Sarawak
Gunung Mulu National Park
Niah Caves
Peninsular Malaysia
Cameron Highlands Wildlife Refuge
Endau-Rompin Park
Fraser's Hill
Krau Game Reserve
Taman Negara
Sumatra
Gunung Leuser National Park (W. Bergmans, pers. comm.)
Summary of threats:
Recommended action:
Principal author for this species: G. W. H. Davison.
Genus Plerotes (1 species)
Priority Grade: 6 (Rare).
Distribution: Western central Angola (on the Angola plateau; Crawford-Cabral, 1989); south-east Zaire; north Zambia (Bergmans, 1989).
Status: Unknown. Threatened by loss of woodland habitat. Appears to be highly specialized and as such will be very vulnerable to destruction of food plants.
Ecology: All localities are within the wetter Zambezian miombo woodland, with the possible exception of Kasama in Zambia, which is in the mosaic of Zambezian dry evergreen forest and wetter miombo woodland. All collecting localities are at altitudes of at least 1000 m.
Nothing is known of feeding habits. From its weak jaws and dentition, a diet of pollen and nectar may be expected (Allen, 1939).
Occurrence in protected areas:
Zaire
Parc National de l'Upemba
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Ptenochirus (2 species)
Priority Grade: 11 (Not Threatened).
Distribution: Philippines (Biliran, Bohol, Camiguin, Carabao, Dinagat, Leyte, Luzon, Maripipi, Mindanao, Mindoro, Negros, Siargao, Sulu Archipelago [Sanga Sanga], Tablas).
Status: Second most abundant species in the submontane forest in the Balinsasayao region (830–1000 m) on Negros but was extremely rare at 1300 m, and not present at higher elevations (Heaney et al., 1989). Comprised about 30% of all captures. Density estimates on Negros ranged from 1.1 to 3.1 individuals per ha. (Heideman and Heaney, 1989).
Occasionally occurs outside primary or good secondary forest (Guerrero and Alcala, 1973 and authors' data), but does not persist outside forest. Forages in urban or agricultural areas near forests, and it is possible that it is limited by roost sites rather than by an inability to use urban/agricultural habitats. Although the evidence suggests that most individuals roost in tree hollows, it has been observed in the entrances of caves in agricultural areas near secondary forests on Leyte and Bohol, but always in locations that make it very vulnerable to hunters. Thus, the combination of habitat destruction and hunting may be acting synergistically to prevent its persistence outside forest.
Suitable habitat (primary or good secondary forest) has declined in the Philippines from about 80% to less than 10% of total land area; population levels probably have declined commensurately. Not seriously threatened at present, as it is probably able to survive at reduced density in secondary forest.
Ecology: Taken from sea level to 1300 m in appropriate habitat. Most abundant in primary forest or good secondary forest at lower or middle elevations (Heaney et al., 1981; Heaney et al., 1989).
Apparently roosts singly and in groups of up to 10 or more. In forest, roosts are probably usually in tree hollows, and perhaps in foliage, but individuals may be found roosting in cave entrances in localities where caves are abundant (E. A. Rickart, pers. comm.). An apparent maternity roost was in a standing hollow dead tree with an opening 12 m above ground in primary forest. Rabor (1977) noted that it can be found ‘…singly or in small numbers, suspended on the walls of shallow cliff caves formed by overhanging rocks…’ and reported capturing a single male in an earthen bank. Lawrence (1939) reported collecting a single male from the underside of a coconut frond on Mindoro.
On Negros, feeds heavily on figs (Moraceae: Ficus spp.) including Ficus chrysolepis, fruits of at least one unidentified plant (Lauraceae), and cultivated or wild bananas (Musaceae: Musa spp.). Figs appear to comprise the major part of its diet. Many were captured beneath a pair of fruiting coffee trees (Rubiaceae: Coffea arabica), a dozen coconut palms (Palmae: Cocos nucifera) and several dozen abaca plants (Musa textilis) in the middle of a clearing about 50 m from the forest edge. The bats were apparently feeding on the sweet coffee fruits; they may also have been feeding on abaca flowers or fruit or coconut flowers. Sanborn (1952) reported Rabor's capture (Rabor, 1977) of two individuals apparently feeding on young fruits or flowers of kapok (Bombacaceae: Ceiba pentandra).
Average movements for adult males were 320 m at an upland site on Negros; this was significantly less than the mean (690 m) for adult females and young of either sex (Heideman and Heaney, 1989). This may indicate fidelity to a resource that males defend (for example, a roost) and that males may remain near a roost site to increase their contact with females (Heithaus and Fleming, 1978), or that males may move shorter distances because they use temporary roosts near food sources while females prefer scarce tree cavities (Morrison, 1978).
P. jagorii flying near and feeding in a 14-m canopy Ficus chrysolepis tree flew along the edge of the crown, only occasionally landing in the tree, and then apparently only briefly, to remove a fruit. The bats flew elsewhere with the fig fruits. Ejecta were uncommon under the crown, but there were clusters of 5–50 ejecta at three localities, presumably feeding roosts, 10–50 m from the tree. Feeding roosts have been found with 100 or more ejecta, indicating the consumption of perhaps 10 or more figs. These limited observations suggest that bats may return to a given feeding roost at least several times (or, perhaps, feed in small groups), but they sometimes use a given feeding roost only once.
Population biology; In the central Philippines, probably undergoes two synchronized birth periods each year, separated by 4 months, with some variation in timing from island to island (Heideman, 1987). Unusual in occasionally producing twins (less than 1% of pregnancies). The duration of gestation is about 4 months and that of lactation about 3 months (Heideman, 1987). On Negros, females undergoing their first pregnancy give birth midway between the two birth peaks of parous females (Heideman, 1987).
Longevities of 5 years documented in the wild; individuals of between 4–5 years of age were healthy and still reproducing (Heideman and Heaney, 1989). The teeth of these animals were appreciably worn, however, and other signs of age were apparent. Estimated annual survivorship was approximately 70% for marked animals, suggesting that about 5 out of 100 yearlings would reach an age of 8 years.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 11 (Not Threatened).
Distribution: Philippines (Biliran, Dinagat, Leyte, Mindanao). A record from Palawan is probably erroneous.
Status: Abundant within primary lowland forest, common in montane and mossy forest (up to 1250 m), uncommon in secondary forest, and absent outside of forest (Heaney et al., 1989; E. A. Rickart, pers. comm.). On Leyte, most frequently netted in primary forest (Heaney et al., 1989).
Currently, large populations are present in forest. The principal habitat of this bat is threatened by deforestation, but it would probably persist in good secondary forest.
Ecology: A single male was observed roosting among the leaves of a two metre high understorey shrub in primary forest on Leyte.
Fig seeds (Moraceae: Ficus spp.) were often present in the faeces.
Population biology: Females on Leyte and Biliran produce one young synchronously in July/August; they may or may not give birth again between November and January, but data are lacking for these months.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Genus Pteralopex (3 species)
T. Flannery (pers. comm.) notes that a very distinctive, as yet undescribed, new Pteralopex species has recently been found on Guadalcanal in the Solomon Islands.
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Fiji (Taveuni). Only two specimens are known, captured in montane forest on Taveuni (Hill and Beckon, 1978).
Status: Endangered or extinct. Not found on most recent expedition (T. Flannery and L. Seri, pers. comm.).
Ecology: There is no direct information on feeding ecology, but the highly cuspate teeth and well-developed jaw musculature of the genus suggests a diet of fruits and other plant material that is tougher than typical for Pteropus.
Population Biology: There is no direct information on population biology, but the lack of observations by local residents and visiting scientists suggests that this species lives solitarily or in small groups in the montane forest.
Summary of threats:
Recommended action:
Principal author for this species: W. E. Rainey.
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Bougainville; Solomon Islands(Choiseul).
Status: Endangered or extinct. Six months of recent fieldwork failed to locate a single specimen (T. Flannery and L. Seri, pers. comm.). Probable reason for decline is disturbance from increasing human population. The islands in the north-west Solomons are the most heavily populated and have the greatest disturbance factors (T. Flannery and L. Seri, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Solomon Islands (Guadalcanal, Santa Isabel).
Status: Endangered or extinct (T. Flannery and L. Seri, pers. comm.). See also P. anceps.
Summary of threats:
Recommended action:
Genus Pteropus (57 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 4.
P. a. admiralitatum
Priority Grade: 10 (No Data).
Distribution: Bismarck Archipelago (Admiralty Islands, Emirau, New Britain, Tabar).
Summary of threats:
Recommended action:
P. a. colonus
Priority Grade: 10 (No Data).
Distribution: Solomon Islands (Choiseul, Mono, Shortland).
Summary of threats:
Recommended action:
P. a. goweri
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (N'Dai).
Summary of threats:
Recommended action:
P. a. solomonis
Priority Grade: 10 (No Data).
Distribution: Solomon Islands (Guadalcanal, Ghizo, Kolombangara, Malaita, Ranonggo, Russell Islands [Mbanika], Simbo, Vella Lavella).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 4.
P. a. alecto
Priority Grade: 10 (No Data).
Distribution: Lombok; Salayar; Sulawesi.
Status: Sulawesi Hunted for food in north Sulawesi. In January 1982 about 100 live P. alecto were offered for sale in Unjung Padang market. They were virtually undamaged and evidently had been mistnetted (Bergmans and Rozendaal, 1988).
Population biology: A pregnant female was caught in Sulawesi in January (Bergmans and Rozendaal, 1988).
Summary of threats:
Recommended action:
P. a. aterrimus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Bawean; Kangean Islands.
Summary of threats:
Recommended action:
P. a. gouldi
Gould's fruit bat, black or blackish fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: South New Guinea and the north and east coast of Australia (New South Wales, Northern Territory, Queensland, Western Australia).
Status: Australia Common (Strahan, 1983). Occurs from Shark Bay in the west to northern New South Wales in the east. Mangrove islands in the estuaries of most northern rivers usually contain camps, often consisting of hundreds or thousands of individuals. Smaller camps of several hundred to several thousand may be found in mangroves, paperbark swamps (Myrtaceae: Melaleuca spp.) or occasionally in patches of rain forest (Strahan, 1983).
This species is unprotected in Queensland and Western Australia and listed as ‘noxious’ in Northern Territory.
New Guinea Rare (Flannery, 1990). Previously reported from mainland New Guinea on only two occasions, once at Bugi, Western Province in 1937 (Tate, 1942), and more recently by Waithman (1979), also in Western Province. Waithman (1979) found the species widespread, although scattered, in the Morehead area in June and September 1973. The Australian Museum holds a large series collected in the 19th century on Yule Island, Central Province (Flannery, 1990).
Ecology: In New Guinea occurs from sea level to 60 m (Waithman, 1979).
Is a high-roosting species and seeks dense leaf-cover. The perimeter of the camp is guarded by old males, which observe intruders and, if they come too close, fly into the camp to raise an alarm. Also roosts on sheltered vertical rock faces in arid areas of Queensland, Australia (A. M. Hutson, pers. comm.).
In Australia, preferred food includes the blossoms of eucalypts (Myrtaceae: Eucalyptus spp.), paperbarks and turpentines. Groups are known to travel as far as 50 km from their camps to feed. Other native and introduced blossoms and fruits, particularly mangoes (Anacardiaceae: Mangifera indica), are also eaten (Strahan, 1983).
Fujita and Tuttle (1991) summarized information on food plants collected by Australian researchers as follows: Anacardiaceae: Mangifera indica (FR); Leguminosae: Castanospermum australe (FL); Loranthaceae: Loranthus sp. (FL); Moraceae: Ficus macrophylla (FR); Myrtaceae: Angophora costata (FL), A. lanceolata (FL), A. subvelutina (FL), Eucalyptus acmenoides (FL), E. alba (FL), E. baileyana (FL), E. camaldulensis (FL), E. citroidea (FL), E. cloeziana (FL), E. corymbosa (FL), E. crebra (FL), E. grandis (FL), E. gummifera (FL), E. hemiploia (FL), E. intermedia (FL), E. longifolia (FL), E. maculata (FL), E. miniata (FL), E. paniculata (FL), E. pilularis (FL), E. polycarpa (FL), E. propinqua (FL), E. ptychocarpa (FL), E. saligna (FL), E. tereticornis (FL), E. tesselaris (FL), E. tetrodonta (FL), E. torelliana (FL), E. umbra (FL), Lophostemon suaveolus (FL), L. confert (FL), Melaleuca dealbata (FL), M. leucadendron (FL), M. quinquenervia (FL), M. viridiflora (FL), Syncarpia laurifolia (FL); Proteaceae: Banksia aemula (FL), B. integrifolia (FL), B. robur (FL), B. serrata (FL), Grevillea robusta (FL); Rubiaceae: Nauclea orientalis (FL); Rutaceae: Citrus sp. (FR).
Population biology: Copulations reported from March to April. Single young born from August to the end of November with peak in October. Young do not leave the camp until they are about 3 months old.
Summary of threats:
Recommended action:
Principal author for this subspecies: T. Flannery.
P. a. morio
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Savu; Sumba.
Summary of threats:
Recommended action:
Priority Grade: 9 (Indeterminate).
Number of subspecies: 7.
Endemic to Vanuatu. Felten and Kock (1972) recognized seven subspecies. As noted by Medway and Marshall (1975), the significant morphological divergence among adjacent islands and the apparent absence of the species from Tanna Island, suggests very limited dispersal. Chambers and Esrom (1988) list local names for P. anetianus on islands from which it has not previously been reported, emphasizing that its distribution and systematics are incompletely known.
Not protected by law in Vanuatu.
P. a. anetianus
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Vanuatu (Anatom, Erromango).
Status: Unknown.
An important food item for humans on Vanuatu. Chambers and Esrom (1988) reported that most people hunted bats for their own consumption, but some sold them in the market and bats were on menus at some tourist hotels. They also noted that, although local customs restricted the take of fruit bats in only a few areas of the country, there was no evidence that current hunting levels were causing declines in populations. The only threat to bat populations they identified was the greatly increased killing of fruit bats by villagers immediately following typhoons.
From the evidence available, it seems that neither direct exploitation nor habitat destruction through logging or agricultural conversion have yet seriously affected populations, offering time to establish a long-term management policy.
Ecology: Neither Baker and Baker (1936) nor Medway and Marshall (1975) observed conspicuous diurnal aggregations in trees comparable with those of Pteropus tonganus geddiei. Chambers and Esrom (1988) noted that colonies were much smaller and quieter than those of P. t. geddiei.
Observed feeding with P. t. geddiei on fruits of Ficus (Moraceae) on Erromango (Medway and Marshall, 1975), although it was reported to be more frequently a blossom feeder than P. t. geddiei. Sanborn and Nicholson (1950) noted that both species fed in coconut groves, but P. anetianus arrived earlier in the day.
Based on the abundance of bat remains in peregrine falcon (Falco peregrinus) eyries in Fiji (White et al., 1988), it seems likely that this bird, which is widely distributed in Vanuatu (Medway and Marshall, 1975), is an important natural predator.
Summary of threats:
Recommended action:
P. a. aorensis
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Vanuatu (Aore, Espiritu Santo).
Status: Baker and Baker (1936) reported that it was far less common than P. t. geddiei near Hog Harbour, Espiritu Santo. Sanborn and Nicholson (1950) reported it as numerous on Espiritu Santo, but less common than P. t. geddiei.
A summary of the threats is given under P. a. anetianus.
Ecology: See P. a. anetianus.
Population biology: Males become most active sexually between October and January. Females probably have a birth peak in August to September, concurrent with P. t. geddiei (Baker and Baker, 1936).
Summary of threats:
Recommended action:
P. a. bakeri
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Vanuatu (Efate, Emao, Nguna).
Status: Unknown except for general comments under P. a. anetianus which includes information on threats.
Ecology: Medway and Marshall (1975) observed it feeding with P. t. geddiei on fruits of Syzygium (Myrtaceae). See also general comments under P. a. anetianus.
Summary of threats:
Recommended action:
P. a. banksiana
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Vanuatu (Banks Islands [Ureparapara, Vanua Lava]).
Status: Unknown except for general comments under P. a. anetianus, which includes information on threats.
Ecology: See P. a. anetianus.
Summary of threats:
Recommended action:
P. a. eotinus
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Vanuatu (Aoba, Maewo, Malakula, Malo, Pentecost).
Status: Unknown except for general comments under P. a. anetianus, which also includes information on threats.
Ecology: See P. a. anetianus.
Summary of threats:
Recommended action:
P. a. motalavae
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Vanuatu (Banks Islands [Mota Lava]).
Status: Unknown except for general comments under P. a. anetianus, which also includes information on threats.
Ecology: See P. a. anetianus.
Summary of threats:
Recommended action:
P. a. pastoris
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Vanuatu (Emae, Tongoa).
Status: Unknown except for general comments under P. a. anetianus, which also includes information on threats.
Ecology: See P. a. anetianus.
Summary of threats:
Recommended action:
Principal authors for this species: W. E. Rainey, E. D. Pierson.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Type locality is possibly Ambon, but there is doubt about the origin of this specimen. Specimens from Sulawesi have been transferred to Acerodon celebensis (Musser et al., 1982).
Summary of threats:
Recommended action:
Priority Grade: 1 (Extinct?).
Distribution: Percy Island, Queensland, Australia.
Status: Almost certainly ‘extinct’, although considerable doubt exists as to the validity of this species. (K. Koopman, pers. comm.). Corbet and Hill (1991) thought that the record of this species may refer to a vagrant Pteropus hypomelanus.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
P. c. caniceps
Priority Grade: 10 (No Data).
Distribution: Bacan; Halmahera; Morotai; Sula Islands (Sulabesi); Ternate.
Summary of threats:
Recommended action:
P. c. dobsoni
Priority Grade: 10 (No Data).
Distribution: Sulawesi.
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Distribution: Ambon; Asilulu; Buru; Gorong Islands (Panjang); Keffing; Seram; Watubela Islands.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
P. c. chrysauchen
Priority Grade: 11 (Not Threatened).
Distribution: Bacan; Gebe; Misool; Morotai; north-west New Guinea; Obi; Salawati; Schouten Islands; Ternate.
Status: New Guinea Uncommon (Flannery, 1990). Records from mainland New Guinea are few, it being restricted to the north-west.
Summary of threats:
Recommended action:
P. c. conspicillatus
Priority Grade: 11 (Not Threatened).
Distribution: Alcester Island; Australia (north-east Queensland [Fitzroy Island]); D'Entrecasteaux Islands; Louisiade Archipelago; Muyua; south-east New Guinea; Trobriand Islands (Kiriwina).
Status: Australia Common in its limited range (Strahan, 1983), although new information indicates a decline in the Cairns-Atherton-Innisfail region. Currently has unprotected status in Queensland.
New Guinea Uncommon. Records from mainland New Guinea are few and this species is confined to the south-east.
Ecology: In Australia, camps may be located in patches of mangroves but are more frequently in tall trees in swamps and rain forest (Strahan, 1983). Usually located in the vicinity of rain forest (Richards 1987; Richards, 1990a).
In Australia, favourite foods are fruit, particularly light-coloured rain-forest fruit that has a high visibility at night, and the ratio of fruit:flowers is approximately 4:1 (Richards, 1990b). G. C. Richards showed that feeding territoriality contributed to dispersal of seeds over long distances. When individuals raided a tree in which others had established territories, they were typically evicted but did not leave without carrying a fruit away. The author lists the following food plants: Anacardiaceae: Anacardium occidentale (FR), Mangifera indica (FR); Annonaceae: Annona muricata (FR), A. squamosa (FR), Polyalthia micheali (FR), Rollinia deliciosa (FR); Burseraceae: Canarium muelleri (FR); Caricaceae: Carica papaya (FR); Chrysobalanaceae: Parinari nonda; Combretaceae: Terminalia arenicola (FR), T. catappa (FR), T. sericocarpa (FR); Elaeocarpaceae: Elaeocarpus angustifolius (FR), E. bancroftii (FR); Lauraceae: Neolitsea dealbata (FL); Leguminosae: Albizia procera (L), Castanospermum australe (FL), Tamarindus indica (FR); Meliaceae: Melia azedarach var. australasica (FR); Moraceae: Ficus crassipes (FR), F. fraseri (FR);F. pleurocarpa (FR), F. triradiata (FR), F. virens var. sublanceolata (FR); Musaceae: Musa sp. (FR); Myrtaceae: Decaspermum humile (FR), Eucalyptus citriodora (FL), E. cloeziana (FL), E. intermedia (FL), E. polycarpa (FL), E. tereticornis (FL), Psidium guajava (FR), Syncarpia sp. (?glomulifera) (FL), Syzygium dictyophlebium (FR), S. forte (FR,FL), S. kuranda (FR), S. wesa (FR); Palmae: Archontophoenix alexandrae (FR), Arecastrum romanzoffianum (FR); Proteaceae: Grevillea robusta (FL); Rubiaceae: Nauclea orientalis (FR); Rutaceae: Acronychia acidula (FR), Citrus sinensis (FR), C. reticulata (FR); Sapindaceae: Litchi chinensis (FR); Sapotaceae: Manilkara kauki (FR); Verbenaceae: Faradaya splendida (FR).
Fujita and Tuttle (1991) additionally listed the following food plants: Myrtaceae: Eucalyptus torelliana (FL), Melaleuca leucadendron (FL).
Can drink sea-water and crocodiles have been known to catch them as they do so (Flannery, 1990).
Population biology: In Australia, copulation occurs from March to May and one young is born between October and early December (Strahan, 1983). Occurrence in protected areas: In Australia, most colony sites are not protected but during seasonal movements some sites in national parks are occupied.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. C. Richards.
Priority Grade: 1 (Endangered: Limited Distribution).
Number of subspecies: 5.
P. d. daitoensis
Daito fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Ryukyu Archipelago (Kita-daitojima, Minami-daitojima). Originally assumed to be confined to Minami-daitojima (Kuroda, 1921; Kuroda, 1933,; Kuroda, 1940) but recently confirmed also to occur on Kita-daitojima (Ikehara, 1973).
Status: Ikehara (1973) counted two and nine animals simultaneously on Kita and Minami-daitojima, respectively, in October 1972. Takara (1976) estimated the population size to be at least 50 on each island.
Probable that destruction of natural environments caused by human exploitation and by introduced animals and plants is causing reduction in available food plants and roosting sites. Shimojana (1978) and Maruyama (1982) noted that several inhabitants pointed out the recent reduction in numbers on Minami-daitojima, although there are at present no scientific data to confirm this. It has been reported that several animals were recently electrocuted on power cables (M. Toyama, pers. comm.).
Ecology: Has been recorded as using the following sites as roosts: in a branch of Casuarina equisetifolia (Casuarinaceae) in the coastal bush; in a bamboo grove; in the top of Livistona chinensis (Palmae) about 13 m above ground (Ikehara, 1973); in bushes of Casuarina equisetifolia and Pandanus tectorius (Pandanaceae) near the coast; in a Casuarina equisetifolia bush on an islet in an inland lake (Maruyama, 1982).
Feeds on the following plants (Ikehara, 1973; Takara, 1976; Shimojana, 1978; Maruyama, 1982): Guttiferae: Garcinia subelliptica (FR); Moraceae: Ficusmicrocarpa (FR), F. superba var. japonica (FR), Morus australis (FR); Palmae: Cocos nucifera (FR), Livistona chinensis (FL).
Occurrence in protected areas: Although it is protected as a national monument by the Japanese Government, no special areas are designated as sanctuaries.
Summary of threats:
Recommended action:
P. d. dasymallus
Erabu fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Ryukyu Archipelago (Oosumi Group: Kuchinoerabujima, Tokara Group: Akusekijima (Yamagata, 1929; Funakoshi, 1990), Nakanoshima (Funakoshi, 1990), Tairajima (Funakoshi, 1990), Takarajima (Nagai, 1929; Funakoshi, 1990)). Funakoshi (1990) also lists records, based on information from inhabitants, for Kuchinoshima and Suwanosejima in the Tokara Group.
Earlier records for Okierabujima (Matsuura, 1920) and south Japan (Uchida, 1963) are considered unreliable because they were based on scant or misleading information. Kuroda (1933) reviews other misleading locality records provided by previous authors (e.g., Siebold, 1824; Temminck, 1825; Dobson, 1878; Andersen, 1912).
Status: Shimojana (1978) observed no individuals on Kuchierabujima in August, 1977. He also reported that the inhabitants talked of recent reduction in populations of this bat. Kunisaki and Kamishikirei (1988) reported that the maximum number of bats observed simultaneously on Kuchierabujima was 9 during the survey in 1986.
Protected as a national monument by the Japanese Government.
Ecology: Funakoshi (1989) listed the following food plants: Ebenaceae: Diospyros sp. (FR,FL); Elaegnaceae: Elaegnus sp. (FR,L); Elaeocarpaceae: Elaeocarpus decipiens (FR); Gingkoaceae: Gingko sp. (L); Moraceae: Ficus erecto (FR), F. microcarpa (FR,B,L), F. pumilo (FR), F. superba (FR,L), Morus australis (FR); Musaceae: Musa sp. (FR); Myrsinaceae: Ardisia crenata (FR); Palmae: Livistona chinensis (FL); Rhamnaceae: Rhamnus liukiuensis (FR); Rosaceae: Rhaphiolepis indica (FL); Theaceae: Eurya emarginata (FR).
Occurrence in protected areas: There are no protected areas within the range of distribution.
Summary of threats:
Recommended action:
P. d. formosus
Taiwanese fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Taiwan (Kashoto[=Lutao] (Kishida, 1924; Chen, 1984), ?Kotosho [=Lanyu] (Kano, 1934; Chen, 1984), ?Main Island: Takoo [=Takao or Kaohsung] (Andersen, 1912; Chen, 1984), Karenko [=Hualien] (Horikawa, 1932; Chen, 1984)). Recent intensive surveys have confirmed its occurrence only on Kashoto.
Status: On Kashoto, where it has been hunted for food, population has been remarkably reduced during the last few decades (information from inhabitants). Handling is currently regulated by the Government of the Republic of China under the law validated in June, 1989.
Occurrence in protected areas: There are no protected areas within the range of distribution.
Summary of threats:
Recommended action:
P. d. inopinatus
Orii's fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Ryukyu Archipelago (Okinawajima). Has occasionally been observed on Minnajima, an islet off-shore of the Motobu Peninsula of Okinawajima (Hinoue, pers. comm.). However, because this animal is not observed on the latter island throughout the year, it is highly probable that the above observations indicate a temporary migration from Okinawajima.
Status: Seems to be abundant in the southern and central parts of Okinawajima (Maruyama, 1980; Shiroma et al., 1981). There is no information concerning its status in the Motobu Peninsula or the northern part of the island.
Ecology: Feeds on the following plants (Maruyama, 1980; Ikehara et al., 1981; Shiroma et al., 1981): Actinidiaceae: Actinidia rufa (FR); Araliaceae: Schefflera actophylla (T); Bromeliaceae: Ananas comosus (FR,T); Elaeocarpaceae: Elaeocarpus decipiens (T), E. japonicus (T); Ericaceae: Pieris japonica (FR); Euphorbiaceae: Bischofia javanica (FR,T); Fagaceae: Castanopsis sieboldii (LB); Guttiferae: Garcinia subelliptica (FR); Moraceae: Ficus microcarpa (FR,T), F. stipulata (FR), F. septica (FR), Morus australis (T,L); Musaceae: Musa paradisiaca (FR); Rutaceae: Citrus spp. (FR); Smilacaceae: Smilax spp. (T); Theaceae: Schima wallichii (T); Zingiberaceae: Alpinia intermedia (FR,T).
Population biology: Maruyama (1980) observed a ‘possible copulation’ on 15 September. Shiroma et al. (1981) reported that one juvenile was born on 13 May in captivity.
Occurrence in protected areas: There are no protected areas within the range of distribution.
Summary of threats:
Recommended action:
P. d. yayeyamae
Yaeyama fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Ryukyu Archipelago (Miyako Group: Taramajima (Ikehara and Shimojana, 1975); Yaeyama Group: Haterumajima, Hatomajima, Iromotejima, Ishigakijima, Kohamajima, Kuroshima, Taketomijima, Yonagunijima [Ikehara et al., 1984]).
Status: Population sizes seem relatively large on Ishigakijima, Iriomotejima and Yonagunijima. However, populations have been remarkably reduced during the last decade on Haterumajima and Kohmajima. In 1981, H. Ota recorded 127 bats on Haterumajima. In 1985, he counted only 33 animals and found that most of the forest they used for daytime roosting had been turned into sugar cane fields. He found no bats on a visit in 1988. It seems likely, therefore, that it is seriously threatened.
Occurrence in protected areas: Exploitation of native vegetation is strictly regulated by the Japanese Government in nearly half the forest on Iriomotejima where this bat seems abundant. However, there is no regulation specific to handling bats.
Summary of threats:
Recommended action:
Principal author for this species: H. Ota.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Nicobar Islands (Car Nicobar).
Summary of threats:
Recommended action:
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Vanuatu (Banks Islands [Mota Lava]).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 4.
P. g. ariel
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Maldives.
Status: Dolbeer et al. (1988) reported on methods used to control fruit bats on the Maldives. This action was taken because of supposed damage to fruit crops (almonds (Rosaceae: Prunus dulcis), guavas (MyrtaceaePsidium guajava and mangoes (Anacardiaceae: Mangifera indica). They reported that bats were common, but not overly abundant, on most islands visited in 1986–87. Population densities ranged from 0.6 to 2.1 bats/ha on 5 islands that were censused. Thoddoo Island, with at least 150 bats, had the largest bat population. Populations on other islands were as follows: Feridhoo (82 bats), Alifushi (79 bats), Kuramathi/Rasdhoo (50 bats) and Mualku (40 bats). Banyan trees (Moraceae: Ficus benghalensis) were by far the most common daytime roosting sites (Dolbeer, 1987). To reduce the size of bat populations, mist-nets were erected, and the bats caught were killed with chloroform-soaked rags. Mist-netting allowed populations to be very significantly reduced, in one case by almost 80%. In this case the cull left only an estimated 16 bats on the island of Alifushi. Estimated reductions of 55% and 67% were achieved on Thoddoo and Feridhoo respectively. Dolbeer et al. (1988) estimated that bat populations would increase at a rate of 49% a year, and further hypothesized that a bat population would take at least 3.5 years to recover to its previous level following culling. On this basis, they recommended reducing the population by 75% every 3 to 4 years. They further recommended that culling should take place in March and April, when pregnant females would be present. They stated that bat populations should never be reduced to below 0.25 bats/ha, which translates to 10 to 20 bats for a typical 40 to 80 ha island.
Pteropus giganteus (Photo by T. H. Kunz)
It is clear that under the regime described above, populations must be considered seriously threatened. The suggested culling rates have been put forward using the assumption that the population size is known and that all other mortality factors are taken into account. At best this is a very dangerous suggestion. With the small population sizes involved in the Maldives, there is a real danger of precipitating the extinction of many populations. The importance of bats as potential ‘keystone’ species in such island situations is not discussed by Dolbeer et al. (1988).
Ecology: Dolbeer et al. (1988) stated that bats were known to feed on commercial crops of almond, guava, and mango and were considered to be serious pests, although no evidence to support this claim was put forward.
Population biology: Dolbeer et al. (1988) obtained information on reproductive status from bats that were killed during culling. Pregnant females were encountered only during April 1987. Litter size was always one and most foetuses were at a similar stage of development indicating that the reproductive period was synchronized within the population and that parturition would take place from mid-April to early May. Immature bats were not collected during April but were collected at other times of the year. In 1987, parturition coincided with the average onset of the rainy season in the Maldives.
Summary of threats:
Recommended action:
P. g. chinghaiensis
Priority Grade: 10 (No Data).
Distribution: North-west China (Chin Hai province). This record is well outside the currently known distribution of P. giganteus and it is possible that this subspecies is not endemic to this region of China.
Summary of threats:
Recommended action:
P. g. giganteus
Priority Grade: 11 (Not Threatened).
Distribution: Andaman Islands; Bangladesh; Burma; India (north to Punjab and eastwards to Sikkim, Bhutan Duars and Pegu); Pakistan;Sri Lanka.
Status: Bangladesh Widely distributed (Khan, 1985). The largest colony seen by Khan was at Barisal town in 1978, where he counted over 2500 bats on 11 trees of Ficus religiosa (Moraceae), Albizia richardiana (Leguminosae), Enterolobium saman (Leguminosae) and Anthocephalus indica (Rubiaceae). By 1981 the size went down to a little over 1000 bats because some of the trees were cut down for use in the construction of buildings (Khan, 1985).
India Thought not to be seriously threatened at present. However, bats are killed for food, because they are thought to be of medicinal value and also because of damage to fruit orchards. Are included under the Wildlife Protection Act as vermin. In a number of areas, they are considered sacred. For instance, a colony of about 500 bats in a banyan tree (Ficus benghalensis) complex in a small village east of Madurai in southern India is considered sacred and is treated with special care. It is thought that people who disturb the colony will be punished by the god Muni.
Sri Lanka Plentiful over most of the cultivated and jungle areas of the whole island with the exception of the highlands. Often particularly numerous along the densely populated coastal belt from north of Colombo south to Matarut appears to be absent, during the greater part of the year, from the Hambantota district. In the hills it is resident up to around 915 m, being found in great numbers around Kandy; quite recently, a colony has been established in the Lower Dikoya Valley (1067 m) but it does not appear to be resident at high altitudes. However, many will make nightly raids into the higher hill country, when certain trees are in flower. Thus, in the Passara District where the nearest colony is said to be over 32 km distant, numbers appear during September in areas above 915 m to feed on flowers of red gum trees (Myrtaceae: Eucalyptus robusta) (Phillips, 1980).
In Pakistan this species is specifically exempted from protection under wildlife regulations.
Ecology: Roosts are very conspicuous and often in large trees such as banyans or tamarinds, generally colse to water. Colonies may number many hundreds (Phillips, 1980). In Bangladesh, live in large colonies ranging from several hundred to a few thousand. The most preferred roosting trees in Bangladesh are Ficus benghalensis, F. religiosa, Dendrocalamus strictus (Graminae), Enterolobium saman, Albizia spp., Borassus flabellifer (Palmae), Cocos nucifera (Palmae), Phoenix sylvestris (Palmae), Eucalyptus globosus (Myrtaceae), Mangifera indica (Anacardiaceae), Tamarindus indica (Leguminosae), Artocarpus lakoocha (Moraceae) and Casuarina equiestifolia (Casuarinaceae) (Khan, 1985).
Feeds on the following plants (Phillips, 1980; Fujita and Tuttle, 1991; author's data): Anarcardiaceae: Anacardium occidentale (FR), Mangifera indica (FL); Annonaceae: Annona muricata (FR), Polyalthia longifolia (FR); Bombacaceae: Ceiba pentandra (FL); Caricaceae: Carica papaya (FR); Combretaceae: Terminalia catappa (FR); Ebenaceae: Diospyros embryopteris (FR), D. melanoxylon (FR); Euphorbiaceae: Hevea brasiliensis (FL); Guttiferae: Calophyllum inophyllum (FR); Leguminosae: Albizia falcata (FL), Erythrina lithosperma (FL); Meliaceae: Melia azedarach (FR); Moraceae: Artocarpus integra (FR), Ficus benghalensis (FR), F. benjamina (FR), F. infectoria (FR), F. microcarpa (FR), F. mysorensis (FR), F. racemosa (FR), F. religiosa (FR); Musaceae: Musa paradisiaca (FR); Myrtaceae: Eucalyptus robusta (FL), Psidium guajava (FR), Syzygium cumini (FR), S. jambos (FR); Palmae: Cocos nucifera (FL); Proteacae: Grevillea robusta (FL); Rhamnaceae: Ziziphus sp. (FR); Rubiaceae: Anthocephalus cadamba (FR); Sapotaceae: Madhuca indica (FR).
Can be destructive in orchards and various methods have been tried to eradicate them, although these have met with little success (Prater, 1971).
Population biology: In India, produces a single young that is born in February or March after a gestation of 140–150 days (Neuweiler, 1969; Prater, 1971). Neuweiler (1969) studied a colony of 800–1000 bats in India and discovered a vertical rank order among males, each having a particular roosting area in the tree. Copulation took place from July to October and births took place mostly in March. When the young became independent they separated from the colony and gathered in a neighbouring tree.
In Sri Lanka, copulation was observed during March with the single young born from the end of May to early July (Bhat, 1942). Both sexes shared the same camps.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. Marimuthu.
P. g. leucocephalus
Priority Grade: 10 (No Data).
Distribution: North-east India (Assam, Manipur); Nepal.
Status: In Nepal this species is partially protected under wildlife regulations.
Summary of threats:
Recommended action:
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Bismarck Archipelago (New Britain).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
P. g. griseus
Priority Grade: 10 (No Data).
Distribution: Bonerate; Jampea; Semau; Timor.
Summary of threats:
Recommended action:
P. g. mimus
Priority Grade: 10 (No Data).
Distribution:?Philippines (Luzon); Salayar; Sulawesi.
Summary of threats:
Recommended action:
P. g. pallidus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Banda Islands.
Summary of threats:
Recommended action:
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Ontong Java Atoll).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 16.
P. h. annectens
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Natuna Islands (Serasan, Subi Besar).
Summary of threats:
Recommended action:
P. h. cagayanus
Island flying-fox
Priority Grade: 11 (Not Threatened).
Distribution: Philippines (Cagayan Sulu, Dinagat, Guimaras, Leyte, Luzon, Maripipi, Mindanao, Negros, Panay).
Status: Taken from sea-level to over 900 m. Rarely netted or observed in primary forest (Heaney et al., 1981, 1989; Heideman and Heaney, 1989). Most common in disturbed habitats, and frequently observed in coconut (Palmae: Cocos nucifera) groves and orchards, either at roosts or during foraging at night. Occasional individuals were found entangled in high electrical wires or tower support cables near urban orchards.
Relative abundance is difficult to assess, because individuals normally fly well above the heights at which bat nets are usually set.
Quite tolerant of disturbance, and persists in some agricultural habitats, although nowhere in the Philippines does it seem to be particularly abundant, perhaps because it is often hunted. Roosts in noisy and highly visible colonies of ten to several hundred individuals, and is vulnerable to hunters. It is probable that hunters at times exterminate local populations, especially given its low reproductive rate.
Ecology: Roosts in tall trees or coconut palms in groups of a few to several hundred (Lim, 1966 and authors' data). When very tall emergent trees were available, the bats usually choose one of these.
Some bats have been observed landing on coconut palms, and farmers indicate they feed upon the coconut flowers. Also been observed feeding on mango (Anacardiaceae: Mangifera indica) and chico (Sapotaceae: Pouteria sapota); undoubtedly also eats fruit and possibly nectar and pollen of some native species (e.g., Ficus spp. (Moraceae)).
Probably flies fairly long distances to forage at times, but those captured were never more than about 6–8 km from a known roost. A number of individuals may feed simultaneously in the same tree.
Population biology: The very limited data available suggest that females give birth in April and May on Negros and Maripipi (Heideman, 1987). Lactation lasts at least 3 months, and reproductive maturity in both sexes occurs at an age of more than 1 year (Heideman, 1987).
Summary of threats:
Recommended action:
Principal authors for this subspecies: P. D. Heideman, L. R. Heaney.
P. h. canus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Natuna Islands (Laut, Pandak, Panjang).
Summary of threats:
Recommended action:
P. h. condorensis
Priority Grade: 11 (Not Threatened).
Distribution: Cambodia; Thailand; Vietnam (Con Son).
Status: Thailand Lekagul and McNeely (1977) record it as being found off the south-east coast, while Yenbutra and Felten (1986) record it from the Provinces of Rayong and Surat Thani.
Vietnam Van Peenen (1969) recorded only one locality (Con Son) in southern Vietnam.
Summary of threats:
Recommended action:
P. h. enganus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Enggano; Mentawai Islands (Siberut, Sipura).
Summary of threats:
Recommended action:
P. h. fretensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Peninsular Malaysia (Jerak, west of the Sembilan Islands); Berhala (west of Jerak in the Straits of Malacca).
Status: No information.
These islands are increasingly visited by holidaymakers so there is an increased risk of shooting and disturbance. This bat is seldom seen as a pest by island residents, but may be shot by mainland villagers when it flies to feed on the adjacent mainland.
In Malaysia this species is partially protected under wildlife regulations.
Ecology: Recorded as eating ripe mango (Anacardiaceae: Mangifera indica). Apart from this, diet is unknown, but presumably includes forest fruits.
Occurrence in protected areas: Not in a protected area.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
P. h. geminorum
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Burma (Mergui Archipelago [South Twin Island]);Peninsular Malaysia (Paya Island southeast of Langkawi off Kedah); Thailand (coastal islands of Ko Samui and Ko Tao).
As noted above (P. h. fretensis) these islands are increasingly being disturbed by holidaymakers. On the islands off Thailand, bats are eaten, usually being killed at the roost site with a long stick or, increasingly, by shooting. The bats are claimed to cure asthma. The roost sites in coconut palms (Palmae: Cocos nucifera) are susceptible: they are rather low and exposed, and easy to find because the bats are noisy.
Occurrence in protected areas: Paya Island is a marine reserve. This protection refers only to the waters, not to the land itself, but a management plan is being prepared by the Fisheries Department, Peninsular Malaysia, to conserve the land resources.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
P. h. hypomelanus
Priority Grade: 10 (No Data).
Distribution: Halmahera; Ternate.
Summary of threats:
Recommended action:
P. h. lepidus
Priority Grade: 10 (No Data).
Distribution: Anamba Islands (Jimaja); Peninsular Malaysia (islands of Aur, Great Redang, Lang Tengah, Pemanggil, Perhentians, Tioman); Tambelan Islands (Saddle Island).
Pteropus hypomelanus (Photo by T. H. Kunz)
Ecology: In Tioman Island Wildlife Reserve in Peninsular Malaysia, roosts were found in palm trees every few hundred metres along the coast, but not inland.
Occurrence in protected areas:
Peninsular Malaysia
Tioman Island Wildlife Reserve
Shooting occurs in this reserve. Bats were common here in 1981, but less so in 1989 (G. W. H. Davison, pers. comm.).
Summary of threats:
Recommended action:
P. h. luteus
Priority Grade; 11 (Not Threatened).
Distribution: Bismarck Archipelago (New Britain); D'Entrecasteaux Islands; Louisiade Archipelago; New Guinea; Solomon Islands (Russell Islands); Trobriand Islands (Kiriwina).
Status: New Guinea Rare (Flannery, 1990). Only recorded once, at Marienberg, 61 km inland from the mouth of the Sepik River (Sanborn, 1931).
Summary of threats:
Recommended action:
P. h. macassaricus
Priority Grade: 10 (No Data).
Distribution: Sangir Islands; Sulawesi; Talaud Islands.
Summary of threats:
Recommended action:
P. h. maris
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Maldives.
Status: Unknown but probably threatened by culling (see above under Pteropus giganteus ariel).
Summary of threats:
Recommended action:
P. h. robinsoni
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Peninsular Malaysia (Sembilan Islands [Lallang, Rumbia]).
Status: Unknown. These islands are increasingly visited by holidaymakers and the bats are shot when they feed on the mainland (see P. h. fretensis).
Ecology: See P. h. fretensis.
Occurrence in protected areas: Does not occur in any protected areas.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
P. h. simalurus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Simalue Islands (Tapeh Islet).
Summary of threats:
Recommended action:
P. h. tomesi
Priority Grade: 10 (No Data).
Distribution: Borneo (Coastal islands: East Kalimanatan [Bilang-bilang]; Sabah [Mantananai, Mengalum]); Philippines (Sulu Archipelago [Sibutu]).
Status: Unknown.
Ecology: Roosts in the fronds of coconut palms (Palmae: Cocos nucifera) or in the branches of small trees (Payne et al., 1985). Sometimes travels to the mainland to feed, but no information on diet (Payne et al., 1985).
Occurrence in protected areas:
Borneo
Sabah
Mantanai Islands Wildlife Reserve
Summary of threats:
Recommended action:
P. h. vulcanius
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Manam (= Vulcan) off north-east New Guinea.
Summary of threats:
Recommended action:
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Federated States of Micronesia (Chuuk).
Status: Does not receive local protection from the Federated States of Micronesia or the Chuuk State Government.
Little information available on abundance. Bruner and Pratt (1979) saw groups of three or four bats flying on Moen in 1976 and 1978 and noted the bats were not wary of humans. In 1983, J. Engbring (pers. comm.) observed colonies estimated to contain about 1000 bats at the summit of Tol and about 100 bats at the top of Dublon. Bats were commonly seen in 1984 on Moen, Dublon, Fefan and Tol (J. Engbring inWiles and Payne, 1986). The population was estimated at 5628 in 1986 (Engbring, 1986). Numbers have almost certainly declined substantially since then, because imports into Guam from Chuuk have increased dramatically since that date. Recently, large numbers of bats were being shipped to Guam from islands within the Chuuk lagoon. There has been a noticeable decline recently in bat numbers on Moen. During a 4-day visit to Chuuk in February 1989, G. Wiles observed moderate numbers near the summit of the mountain on Tol. However, a colony of 200 observed by Engbring in 1984 appears to have disappeared. On Moen, Wiles saw very few bats and was told by residents of the island that numbers had declined significantly during the past few years. Rainey (1990) also reported low numbers during a brief visit to several islands in 1989.
A large commercial trade has existed in Micronesia during the last 20 years, with large numbers of bats captured on a number of islands and exported to the Mariana Islands for use as food (Wiles and Payne, 1986). Records reveal that 5795 P. insularis were exported from Chuuk to Guam between 1978 and 1989 (Wiles, 1992), with more than half of this amount, 3723 animals, exported in 1988 and 1989. The residents of Chuuk are not known to hunt this species for local use.
Chuuk's dense human population has destroyed much of the native forest on the islands in the Chuuk Lagoon. Remnants of these forests occur on the mountain tops on Moen, Dublon, Fefan, Tol and other small islands. Most of the lower slopes of these islands have been converted to coconut (Palmae: Cocos nucifera) and breadfruit (Moraceae: Artocarpus altilis) forest, with mangoes (Anacardiaceae: Mangifera indica), bananas (Musaceae: Musa spp.) and other food crops also present. This type of habitat change has probably affected fruit bats by altering their food sources and reducing colonial roosting sites.
Ecology: Observed to feed on the flowers of coconut.
Occurrence in protected areas: No protected areas have been established within the geographic range of this species.
Summary of threats:
Recommended action:
Principal author for this species: G. Wiles.
Priority Grade: 4 (Vulnerable).
Distribution: Philippines (Catanduanes, Dinagat, Luzon).
Status: Unknown. Rare and very probably endangered by deforestation on Luzon. Known predominantly from ‘politically sensitive’ areas, which may preclude the initiation of wide ranging status surveys (R. Wirth and W. L. R. Oliver, pers. comm.). The status on Luzon and Dinagat could be investigated as part of the cloud rat project on these islands (R. Wirth and W. L. R. Oliver, pers. comm.).
Ecology: Has been taken only on ridgetops near primary forests. Very little is known about its habitat. The majority of specimens have been taken from cloud forests and storm forests (the latter heavily damaged by typhoons at irregular intervals), (Heaney et al., 1987; R. Wirth and W. L. R. Oliver, pers. comm.).
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Comoros (Anjouan, Moheli).
Status: Anjouan According to Humboldt (Milne-Edwards and Oustalet, 1888) was abundant in the great forests of Anjouan's peaks in the late 19th Century. However, by 1977 Cheke and Dahl (1981) reported it to be extremely rare and threatened by active and progressive deforestation. Sightings of ‘a number’ of bats had been reported near Dzialandze in 1977, and over 60 were seen there in 1988. In 1989, the University of East Anglia Comoro Islands Expedition reported this population to be much reduced, with only a handful of sightings. Census work showed the presence of a group of 30–40 in primary forest about 3 km south of Dzialandze. The bats appeared to have moved to this new site due to disturbance from people and as a result of Cyclone Calasanji (January 1989). The forest in this area is rapidly being underplanted and cleared and there is a prospect of a highland road being built. The estimated total population was about 60 individuals. In July 1990, J. B. Carroll (pers. comm.) located a single roost estimated to contain between 60 and 120 bats. A single bat was seen flying at another locality on the islands.
Moheli Reports of a few individuals seen in 1975 and 1977. The UEA expedition in 1988 found no evidence of this species, although they did not rule out the possibility of relict populations being present.
Grand Comore The UEA expedition in 1988 found no evidence of this species on Grand Comore. They did not rule out the possibility of relict populations in the Karthala Forest, although these should have been noticed by the many collectors who have visited the site since the mid-1800s. Is now extremely rare and threatened by a number of factors.
Deforestation is a serious problem in the Comoros, but there are difficulties in quantifying and monitoring the destruction. The deforestation is a gradual process of underplanting with fruit, coconuts (Palmae: Cocos nucifera), manioc (Euphorbiaceae: Manihot esculenta), maize (Graminae: Zea mays), peas (Leguminosae: Pisum sativum) and sweet potatoes (Convolvulaceae: Ipomoea batatas). This type of agriculture necessitates the removal of scrub and herbaceous layers and some large trees. This, coupled with grazing, severely affects regeneration. It also appears that in this new type of forest, P. seychellensis comorensis (the common Comorian fruit bat) outcompetes P. livingstonii. Wheras P. s. comorensis was formerly confined to areas below 400 m, roosts were recorded at 610 m and 1000 m in 1989 (Thorpe, 1989). P. livingstonii is confined to diminishing patches of primary forest. On Anjouan, virtually all of the remaining primary forest is on uncultivatable slopes, while on Moheli some is still accessible but is declining rapidly. Grande Comore fares the best but this is due to the large size of the Karthala Forest. The annual loss of forest on Grande Comore is alarming. Other causes of deforestation include losses due to the cultivation of cash crops such as cloves (Myrtaceae: Syzygium aromaticum), particularly at the farmland-forest interface. Some wood is cut for fuel. Commercial exploitation is low but can be very damaging to non-target species. Anjouan is more densely populated than any African country and pressure for land is intense.
Cyclones are common in the Indian Ocean, although few affect the Comoros. A serious cyclone (winds in excess of 64 knots) appears to hit the Comoros every 10 or so years. It seems likely that the Moheli population was wiped out as a result of two major cyclones: Elinah in January 1983 and Kaimsey in April 1984. Kaimsey, with winds up to 100 knots, destroyed 80% 0f the food crops on Moheli. The low relief and small area of Moheli makes it more susceptible than the larger Anjouan with its complex topography. The recent incidence of major cyclones is unprecedented in climatic records for the Comoros. This could represent a shift in the storm belt of the western Indian Ocean. Whereas the Comoros were once on the edge of the cyclone belt, they are now within it (I. Thorpe, pers. comm.).
The 1989 UEA Expedition noted that part of the population had moved from Dzialandze to Hasiaka. At Dzialandze local people saw them as guardians of the lake and protected them. This belief is not well known at Hasiaka and an influx of refugees from Madagascar (who eat bats) has further diluted the belief. Young boys now have no qualms about catapaulting bats for sport or food (Thorpe, 1989).
There is no legal protection of endangered species on the Comoros.
In 1990 the Jersey Wildlife Preservation Trust was planning to take 10 bats into captivity for breeding at Jersey Zoo as a safeguard against extinction (I. Thorpe, pers. comm.). J. B. Carroll (pers. comm.) reports that this expedition was unsuccessful at obtaining any bats for captive breeding, but it is hoped to mount another expedition in the future.
Ecology: Was seen feeding on the fruits of Ficus lutea (Moraceae) (I. Thorpe, pers. comm.).
Population biology: Voeltzkow (1904) saw numerous adult females with almost fully grown adult young in the second week of October.
Occurrence in protected areas: No protected areas have been established in the Comoros.
Summary of threats:
Recommended action:
Principal author for this species: J. B. Carroll.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
P. l. lombocensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Flores; Lombok.
Summary of threats:
Recommended action:
P. l. solitarius
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Alor.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: Cambodia; Thailand; Vietnam.
Status: Thailand Lekagul and McNeely (1977) reported it being found along the coastal area of the Gulf of Thailand, south to Phetburi, north to Ayutthaya and east to Cambodia.
Vietnam Recorded only from Saigon (Van Peenen, 1969).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2.
P. m. epularius
Priority Grade: 11 (Not Threatened)
Distribution: Australia (Queensland: Boigu Islands); New Guinea.
Status: Australia Rare. Only recorded from the Boigu Islands, 10 km south of New Guinea in the Torres Strait (G. C. Richards, pers. comm.).
New Guinea Common (Flannery, 1990). Has been recorded at various localities in southern New Guinea from the Oriomo Plateau (Western Province) in the west to Milne Bay (Milne Bay Province) in the east. Koopman (1982) records that it was also found in several localities in northern New Guinea. Thus it is probably widespread throughout the lowlands.
Ecology: Possibly forages in drier areas than other Pteropus species (Ziegler, 1982), and McKean (1972) noted that of the four specimens reported by him, one was captured in a mist-net in dryish monsoon forest and that three were shot at night while feeding on the flowers of coconut palms (Palmae: Cocos nucifera) in the same general environment.
Summary of threats:
Recommended action:
Principal author for this subspecies: T. Flannery.
P. m. macrotis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Aru Islands (Wokam).
Summary of threats:
Recommended action:
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Bougainville; Solomon Islands (Santa Isabel).
Status: Rare and possibly endangered (T. Flannery, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade; 1 (Endangered: Limited Distribution).
Number of subspecies: 7. Yoshiyuki (1989) considered P. m. loochoensis to be a full species.
Populations of P. mariannus on a number of the Mariana Islands have never been identified to subspecies level. Recently, Wiles and Glass (1990) questioned the validity of distinguishing two subspecies of P. mariannus in the Mariana Islands. This was based on the evidence of movements of P. mariannus between islands, which would facilitate gene flow through the chain.
P. m. loochoensis
Greater loochoo fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Ryukyu Archipelago (Okinawajima).
Status: Imaizumi (1970) reported that only two specimens are known to have been collected. Kuroda (1932) remarked that it ‘seemed to be very rare’. Current information on status is unavailable.
May be threatened by typhoons. These are rare but can cause serious problems. Strong typhoons (windspeed in excess of 130 knots) occur once every 10–15 years. They can denude large areas of foliage and fruit and there have been reports from Indian Ocean islands of dead bats being found under trees after storms.
There are problems of habitat destruction in the Ryukyus. In some areas, most of the forest that bats use for daytime roosting has been turned over to sugar cane fields (Graminae: Saccharum officinarum). It is not known how this may have affected bats.
Legal status in the Ryukyus unknown.
Ecology: No specific information available but there is extensive data available for P. mariannus in the Marianas (see below).
Summary of threats:
Recommended action:
Principal authors for this subspecies: H. Ota, G. Wiles.
P. m. mariannus
Marianas fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Commonwealth of the Northern Mariana Islands (Aguijan, Rota, Saipan, Tinian); Guam.
Status: The Guam population of P. m. mariannus is listed as endangered by the US Fish and Wildlife Service and the Government of Guam. It is locally protected on the remaining Mariana Islands by the Commonwealth of the Northern Mariana Islands. Populations on Saipan, Tinian and Aguijan are listed as Category 1 Endangered Species by the US Fish and Wildlife Service, meaning that the agency supports full endangered species status for these populations but does not have sufficient time or resources to finalize the listing. Populations on Rota are Category 2 Candidate Endangered Species, meaning that the US Fish and Wildlife Service has some evidence of vulnerability but not enough to justify an endangered listing at present.
Aguijan Intensive surveys in 1983 and 1984 (Wiles et al., 1989) indicated that there were fewer than 10 bats on Aguijan. However, approximately 300 bats were seen in 1988 (Wiles and Glass, 1990). The population declined to about 50 bats in 1989 (Stinson et al., 1992).
Guam Full details of the history of the decline of bat populations on Guam is given by Wiles (1986, 1987a, 1992). Crampton (1921) reported bats as ‘not an uncommon sight’ in 1920. By 1931 (Coultas, 1931) they were considered uncommon on the island, the introduction of firearms had led to their destruction. In 1945, Baker found bats uncommon in the northern half of Guam and confined primarily to the forested clifflines. He failed to locate a single colony in one year's work. In 1958, Woodside (in Wiles, 1987a) reported a maximum population of 3000, a ‘greatly reduced’ number. Between 1963 and 1968 monthly counts showed the island's population dropping. The last large roost in south Guam was of 150 bats at Orote Point Island in 1971. In 1972, a colony on the USAF base numbered 500–600 and made up the bulk of the island's population (estimated at less than 1000). This colony disappeared soon after and from 1974–5 the island's population was estimated at less than 100 bats. In 1978, a survey by Wheeler and Aguon (1978) estimated the population to be fewer than 50 bats. In 1980, there was a dramatic increase with a new colony at Pati Point numbering several hundred bats. In total, there were 240 bats in early April and 508 bats in mid-May. The sudden increase was probably due to immigration from Rota (Wiles and Glass, 1990). In 1981–82, the colony increased from an estimated 600 at the end of 1981 to 780–850 in 1982. In 1983, there was a decline to an estimated 500–600 bats. There were 50–100 in the forest within 1.5 km of the north cliffline from Bijia Point to Lates Point, with most animals between Achae Point and Tarague point. There were also 25–50 bats in south Guam in Nav May, Talofofo and Malojloj regions. The total island-wide estimates were: for 1981, 650–750; for 1982, 850–100; and for 1983, 600–775. In 1984, between February and April, bats were located at 11 sites, all on USAF-controlled land in northern Guam (Wiles, 1987a). The estimated total was 450–500 bats between Achae and Lafac Point in northern Guam. No bats were recorded from southern Guam or from the north-east and north-west coasts, although up to 50 bats may still inhabit these areas. A roost at Mergagan Point lost up to 40–50 bats to poachers in 1984, giving a new overall estimate of 425–500 bats (Wiles, 1987a). In 1985 the population was estimated at 695–800, although this was reduced to 500–600 by 1987. In 1988 the population increased to 600–650 due to immigration from Rota. There is currently no recruitment of young into the population due to predation by brown tree snakes (Boiga irregularis) (Wiles, 1987b).
Rota Wheeler (1980) believed that only 200–400 bats were left on Rota in 1979, but his estimate was probably far too low. Wiles et al. (1989) estimated the population at 1500–2000 bats in 1983 and 800–1000 bats in 1984. The latter figure was also probably an underestimate. More detailed and reliable surveys of the population were conducted between August 1986 and January 1988, with estimates of 2000–2500 bats obtained (Stinson et al., 1992). A severe typhoon struck the island in January 1988. Four surveys conducted between April 1988 and July 1989 indicate that population has declined to an estimated 1000–1400 bats, with the reduction caused by increases in poaching rather than storm-related factors (Stinson et al., 1992).
Saipan Saipan may well have had several thousand bats until as recently as the early 1970s (Wiles et al., 1989). Wheeler (1980) failed to observe any bats on the island. More intensive surveys in 1983 and 1984 (Wiles et al., 1989) indicated that there were fewer than 50 bats on Saipan. Numbers on Saipan increased to about 75–100 animals in 1985 and 1986 (P. O. Glass, pers. comm.).
Tinian Wheeler (1980) estimated that there were 25–100 bats on Tinian. More intensive surveys in 1983 and 1984 (Wiles et al., 1989) indicated that there were fewer than 25 bats on Tinian.
P. m. mariannus faces a number of threats:
Hunting: On Guam bats have never been viewed as agricultural pests and have been taken solely for human consumption. Overhunting has been the most important cause of decline (Wiles, 1987a). P. m. mariannus was probably greatly reduced in numbers in the early 1900s. The expanding population and the use of firearms has accentuated the decline. Bats were removed from the list of Unprotected Wildlife in 1965 and in 1966 a 10-week hunting season was imposed. In successive years the hunting seasons became shorter and the catches more restrictive, culminating in a hunting ban in 1973 (Wiles, 1987a). In 1981 and 1984 there was additional protection when both species on Guam (P. mariannus and P. tokudae) were included in the Guam and US Endangered Species Lists respectively. But illegal hunting continues. Between 1981–84 there were 8 reported cases of hunting at colonies and 7 reported cases of hunting along flyways or at feeding sites. Colony hunting is very destructive and 50+ can be caught in one night (Wiles, 1987a). Night hunting is now opportunistic. Between 1981 and 1990 there was only one arrest and conviction, resulting in a $100 fine.
Micronesian islanders hunt P. mariannus as a local food source throughout its range, except on Kosrae. Hunting for this purpose has been most intensive in the Marianas. Numerous observers on Guam, beginning with Coultas (1931), have implicated overhunting in the decline of P. m. mariannus on the island. Illegal hunting was responsible for the drastic decline in fruit bats on Rota in 1988 (Stinson et al., 1992), with most of the bats there killed for local consumption. Hunting for local use is also presumed to have caused severe declines in the bat populations on Saipan, Tinian and Aguijan in the 1960s and 1970s, although some of the harvested bats were exported to Guam (Wiles and Payne, 1986).
A large commercial trade involving at least four subspecies of P. mariannus has existed during the last 20 years, with large numbers of bats captured and exported to the Mariana Islands for use as food (Wiles and Payne, 1986; Wiles, 1992). Most of the bats were shipped to Guam but smaller numbers have also entered Saipan, Tinian and Rota. Records indicate that between 1975 and 1989, Guam imported an estimated 16,324 P. m. mariannus from neighbouring islands in the south Marianas (Wiles and Payne, 1986; Wiles, 1992).
Forest destruction: Deforestation is not currently a major conservation problem on the Micronesian islands inhabited by P. mariannus. Rapid modern development has occurred since 1985 on Guam and Saipan, but the resulting losses of forest have generally not taken place in areas used by bats. In the 1920s and 1930s much of the forest on Saipan, Tinian, Aguijan and Rota was cleared for agriculture by the Japanese. During and after World War II, mailitary invasions and subsequent construction of military bases on Guam, Saipan and Tinian also resulted in some forest loss. However, secondary forests have regrown in many areas, particularly on Guam. Many of these forests are substantially modified, but contain a combination of introduced and native plants used by fruit bats.
Predation by brown tree snakes (Boiga irregularis): On Guam, predation on young P. m. mariannus by brown tree snakes has become a greater threat to the long-term survival of the island's population than illegal hunting. The snake was introduced to Guam shortly after the end of World War II and became abundant, particularly in southern Guam. Large numbers of snakes did not occur in the north of the island, where most bats remained, until the early 1980s (Wiles, 1987b). Observations of bat colonies from 1984 to 1988 indicated that virtually all juvenile fruit bats did not survive beyond an estimated age of 1–2 months, almost certainly because they were being eaten by snakes (Wiles, 1987b). B. irregularis is nocturnal and highly arboreal, and is already responsible for the near total disappearance of the island's forest-dwelling avifauna (Savidge, 1987).
Typhoons: These are a rare but serious threat. Strong typhoons (wind speed in excess of 130 knots) occur about once every 10–15 years. There is no evidence that typhoons have caused major declines of bats on Guam (Wiles, 1987a) but under present conditions, with most animals in a single colony, a typhoon directly striking the roosting area could have disastrous results.
Ecology: All Micronesian subspecies of P. mariannus congregate in colonies in native forest, coastal strand, and mangroves (Perez, 1968; Wiles, 1987a, 1987c; Falanruw, 1988a; Wiles et al. 1989, 1991; J. Engbring, pers. comm.). The populations include a small percentage of animals that roost alone or in small aggregations. Within colonies, bats gather in harems, which contain a male and 2–15 females, and bachelor groups, or roost as individuals on the periphery of these groups (Wiles, 1987b; Wiles et al., 1991)
On Guam and Rota, colonies normally occur at isolated locations along cliffs, where human disturbance is minimal and the surrounding terrain offers some protection from winds.
The known food plants of P. mariannus in the Marianas include the following species (data from Wiles, 1987b; P. O. Glass, pers. comm.). Agavaceae: Agave rigida (FL); Anacardiaceae: Mangifera indica (FR); Annonaceae: Annona reticulata (FR); Apocynaceae: Neisosperma oppositifolia (FR), Ochrosia mariannensis (FR); Bombacaceae: Ceiba pentandra (FL); Caricaceae: Carica papaya (FR); Casuarinaceae: Casuarina equisetifolia (FL), C. litorea (FL); Combretaceae: Terminalia catappa (FR,FL); Cycadaceae: Cycas circinalis (FR); Elaeocarpaceae: Elaeocarpus joga (FR,FL); Guttiferae: Calophyluum inophyllum (FR), Mammea odorata (FR,FL); Hernandiaceae: Hernandia sonora (FR); Lauraceae: Persea americana (FL); Lecythidaceae: Barringtonia asiatica (FL); Leguminosae: Cynometra ramiflora (L), Erythrina variegata (FL), Mucuna gigantea (FL); Loganiaceae: Fagraea bateriana (FL); Meliaceae: Aglaia mariannensis (FR); Moraceae: Artocarpus altilis (FR,L), A. mariannensis (FR,L,T), Ficus prolixa (FR), F. tinctoria (FR); Musaceae: Musa sp. (FR); Myrtaceae: Psidium guajava (FR); Palmae: Cocos nucifera (FL); Pandanaceae: Freycinetia reineckei (FR,FL), Pandanus fragrans (FL), P. tectorius (FR,FL); Passifloraceae: Passiflora suberosa (FR); Rubiaceae: Guettarda speciosa (FL); Sapindaceae: Tristiropsis obtusangula (FR); Solanaceae: Cestrum diurnum (FR); Urticaceae: Dendrocnidne latifolia (FR,FL), Pipturus argenteus (FR); Verbenaceae: Premna obtusifolia (FR).
Movements of P. mariannus are poorly known. Bats from Guam's only remaining colony are suspected to forage up to 10–12 km from their roosting site. Wiles and Glass (1990) provide evidence of inter-island movements. On rare occasions, groups of 50–300 bats fly 60–80 km between Rota and the neighbouring islands of Guam and Aguijan. Hunting and other forms of human disturbance are believed to cause most of these flights. Additional movements between other islands by solitary bats and groups of several individuals have also been noted.
Population biology: On Guam, copulation and lactation have been observed throughout the year (Wiles, 1987b). Approximately 7–20% of all harem females possess small to medium-sized young each month with no consistent peak in births noted between years.
Occurrence in protected areas: The Government of Guam has designated four conservation reserves that total approximately 1700 ha, but none contain significant numbers of fruit bats. However, most of the island's bats roost and feed on Andersen Air Force Base (6250 ha), with small numbers also present on the US Naval Facility (134 ha), US Naval Communications Area Master Station (1196 ha) and US Naval Magazine (3578 ha). Military bases on Guam have functioned as partial refuges for fruit bats and other wildlife in the past 35 years. By limiting access to civilians and clearing few additional tracts of native forest, the military has unintentionally reduced illegal hunting and maintained habitat.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. Wiles
P. m. paganensis
Pagan Island fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Commonwealth of the Northern Mariana Islands (Almagan, Pagan).
Status: Locally protected in the CNMI. Populations on islands north of Saipan are Category 2 candidate endangered species, meaning that the US Fish and Wildlife Service has some evidence of vulnerability but not enough to justify an endangered listing at present.
Alamagan Wiles et al. (1989) surveyed the island in 1983, though only two counts were made. No bats were seen on either occasion. A recent trip to Alamagan in March 1988 discovered moderate numbers of fruit bats in many of the island's forested ravines (P. O. Glass, pers. comm.). This suggests that numbers have perhaps increased since 1983.
Pagan Wiles et al. (1989) surveyed the island in 1983. They noted one colony believed to contain 2000 bats (this was probably the largest colony in the Marianas). They estimated 2500 bats on the island as a whole.
Threatened by hunting and by the effects of typhoons (see P. m. mariannus).
Ecology: See P. m. mariannus.
Population biology: See P. m. mariannus.
Occurrence in protected areas: There are no protected areas within the geographic range.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. Wiles.
P. m. pelewensis
Belau fruit bat
Priority Grade; 1 (Endangered: Limited Distribution).
Distribution; Belau.
Status: Extensive population surveys have never been made. Owen (1977) described fruit bats as common but declining, yet visitors to Belau from 1983 to 1985 found bats to be common in many areas (Wiles and Payne, 1986). In 1988, Wiles and Conry (1990) estimated 130–170 bats in the Ngerukewid Islands Wildlife Preserve, a somewhat isolated group of islands in south-central Belau. Fruit bat abundance appeared to be much lower on several other islands in central and southern Belau at the same time.
Threatened by hunting and the effects of infrequent typhoons (see P. m. mariannus). Guam imported an estimated 112,184 bats from Belau between 1975 and 1989 (Wiles, 1992).
Not protected on Belau. The recently amended CITES regulations do not cover trade between Belau and Guam because this is deemed not to be international.
Ecology: The diet probably contains an even greater variety of foods than that described above for P. mariannus in the Mariana Islands, reflecting the fact that Belau's forests are the most diverse in Micronesia, but only a few food plants have thus far been reported (Perez, 1968; Wiles and Conry, 1990): Anacardiaceae: Mangifera indica (FR); Apocynaceae: Neisosperma oppositifolia (FR); Combretaceae: Terminalia catappa (FR); Cycadaceae: Cycas circinalis (FR); Leguminosae: Intsia bijuga (FL); Moraceae: Ficus spp. (FR); Myrtaceae: Eugenia malaccensis (FR); Palmae: Gulubia palauensis (FL); Pandanaceae: Pandanus tectorius (FR).
Population biology: Females have been observed flying with young in January (Wiles and Conry, 1990), but not in April (Perez, 1968).
Occurrence in protected areas: Occurs in the small Ngerukewid Islands Wildlife Preserve, which is the only nature reserve in the Caroline Islands.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. Wiles.
P. m. ualanus
Ualan fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Federated States of Micronesia (Kosrae).
Status: Coultas (1931) observed only a few during 5 months of collecting birds on Kosrae for the Whitney South Sea Expedition in 1931. He reported bats to be nearly extinct and attributed their rarity to a disease that reportedly killed ‘thousands’ of animals in about 1927. In 1983 J. Engbring (pers. comm.) observed several colonies, each with about 100–400 bats, high in the interior of the island. No current population estimates are available. Kosrean government officials believe that the island's population has declined significantly in recent years (G. Jackson, pers. comm.). A brief survey in 1989 revealed that bats were not common (Rainey, 1990).
Has been threatened by hunting for export to the Marianas. Government officials in Kosrae stated in 1989 that significant numbers have been exported to the Marianas in recent years (W. E. Rainey, pers. comm.), although the Guam Department of Agriculture does not have documentation to support this. Recently amended CITES regulations have made this trade illegal.
Also threatened by the effects of infrequent typhoons (see P. m. mariannus).
Not locally protected on Kosrae.
Ecology: There is no specific information on this subspecies, although feeding habits are probably similar to those of P. mariannus in the Marianas.
Population biology: See P. m. mariannus.
Occurrence in protected areas: There are no protected areas on Kosrae.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. Wiles.
P. m. ulthiensis
Ulithi fruit bat
Priority Grade; 1 (Endangered: Limited Distribution).
Distribution: Federated States of Micronesia (Yap [Ulithi Atoll]). Enquiries to islanders on Ulithi resulted in reports of bats occurring on several other atolls where they had previously been unreported. They were reported to occur on Fais and Satawal, single islands located 80 km east and 800 km south-east of Ulithi, respectively, and on Woleai, 500 km south-east of Ulithi. These are presumably P. mariannus, possibly P. m. ulthiensis, or perhaps P. insularis, known from Chuuk (Wiles et al., 1991).
Status: In March 1986, Wiles et al. (1991) surveyed 14 islands within Ulithi Atoll. A total of 715 bats were observed during the census and the overall population was estimated at 895–1060 bats at an overall density of 0.43–0.51 bats per ha. Fruit bats were most numerous on Sorlen, Fossarai (including Lolang) and Asor, each with a population of at least 120 animals. The largest number and highest density was on Sorlen, where a colony of 146 bats resided. Most other islands apart from the above three had estimated populations of 40–70 bats. Eight other islands in the atoll had suitable habitat but were not surveyed. If the densities of bats were the same as elsewhere, this would increase the overall estimated population to 1128–1333 bats. The bats do not avoid islands with people, but did avoid villages by day. There was some evidence of deliberate disturbance by Ulithians, with small numbers occasionally shot by hunters. The estimated densities for fruit bats are the highest recorded for Pteropus.
Although hunting does take place, it is on a relatively small scale and has probably had little effect on the overall population (Wiles et al., 1991). Fruit bats have been protected from hunting since 1981 (Falanruw, 1988a, 1988b). In November 1988, a month-long hunting season was introduced. Methods of harvest other than air guns and traditional methods were prohibited. The number of animals killed during the season is unknown, but two shipments totalling 37 bats were exported to Guam (Wiles et al., 1991)
Ulithi was struck by two severe typhoons in December 1986 and January 1987. It is not known what effect the storms had on the bat population.
Ecology: Most day roosting is colonial, with 68% of bats observed during a survey in March 1986 roosting in colonies of > 20 bats (Wiles et al., 1991). Colonies were present on most islands, typically in the upper halves of emergent trees with crowns several metres above the surrounding forest canopy. Colonies were composed mainly of harem groups of one male and several females and usually contained some solitary individuals, mostly males (Wiles et al., 1991).
Feeds on the following plants (Wiles et al., 1991): Apocynaceae: Neisosperma oppositifolia (FR); Guttiferae: Calophyllum inophyllum (FR); Moraceae: Artocarpus altilis (FR,L), Ficus prolixa (FR); Musaceae: Musa sp. (FL); Myrtaceae: Eugenia javanica (FR); Palmae: Cocos nucifera (sap); Pandanaceae: Pandanus tectorius (FR); Rubiaceae: Guettarda speciosa (FR,L).
In Ulithi, fruit bats undoubtedly play a major role in dispersing fruits of trees that have seeds >1 cm in diameter, such as Pandanus, Guettarda, Neisosperma, Eugenia, and Calophyllum. Micronesian starlings (Aplonis opaca) are the only other frugivorous vertebrates on the atoll, but they feed primarily on smaller fruit and fruit with smaller seeds (Baker, 1951; Jenkins, 1983). Coconut crabs (Birgus latro) feed on fallen fruit (Reyne, 1939; Holthuis, 1963) and may rival Pteropus as seed dispersal agents for some island tree species, but this aspect of crab biology is poorly understood (Wiles et al., 1991).
Population biology: Four of 19 females in harems were noted with small and medium-sized young in March (Wiles et al., 1991). See also general comments under P. m. mariannus.
Occurrence in protected areas: There are no protected areas on Ulithi Atoll.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. Wiles.
P. m. yapensis
Yap fruit bat
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Federated States of Micronesia (Yap).
Status: Abundance declined sharply because of overhunting in the late 1970s (Falanruw, 1988a, 1988b). Observations in 1976 and 1978 indicated that fruit bats were less common on Yap than on other islands in the Carolines (Bruner and Pratt, 1979). By 1981, just 1000 animals were believed to be left in the island group (M. V. C. Falnaruw inWiles and Payne, 1986) causing the Yap Government to ban all hunting. Surveys in 1984 and 1986 revealed that numbers had increased to an estimated 2500–5000 bats (Engbring, 1985, 1986).
Has been threatened by hunting both for local use and for export. Guam imported an estimated 23,410 bats from Yap between 1975 and 1989 (Wiles, 1992). The recent ban on hunting has lessened this threat for the time being, although some illegal hunting continues (M. V. C. Falanruw, pers. comm.). The recently amended CITES regulations have made the trade in fruit bats to Guam illegal. Also threatened by typhoons (see P. m. mariannus).
Locally protected on Yap.
Ecology: Falanruw (1988a) listed the following food plants: Anacardiaceae: Campnosperma brevipetiolata (FR), Mangifera indica (FR), Semecarpus veneosus (FR); Annonaceae: Annona muricata (FR); Bombacaceae: Ceiba pentandra (FR); Caricaceae: Carica papaya (FR); Chrysobalanaceae: Parinari spp. (FL); Combretaceae: Lumnitzera littorea (FL), Terminalia catappa (FR); Euphorbiaceae: Glochidion sp. (FL); Guttiferae: Calophyllum inophyllum (FR,FL); Leguminosae: Inocarpus fagifer (FR); Moraceae: Artocarpus altilis (FR), A. heterophyllus (FR), Ficus prolixa (FR); Musaceae: Musa spp. (FR); Myrtaceae: Eugenia spp. (FR); Palmae: Cocos nucifera (FR); Pandanaceae: Freycinetia sp. (FL), Pandanus tectorius (FR); Rutaceae: Citrus spp. (FR); Sonneratiaceae: Sonneratia alba (FR,FL,L).
Population biology: Reproduces throughout the year, but with more births possibly occurring from February to March and August to October (Falanruw, 1988a). See also P. m. mariannus.
Occurrence in protected areas: There are no protected areas on Yap.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. Wiles
P. mariannus (subspecies unknown)
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Commonwealth of the Northern Mariana Islands (CNMI) [Agrihan, Anatahan, Ascuncion, Farallon de Medinella, Guguan, Maug, Sarigan].
Status: Locally protected in the CNMI, although a limited special hunt was allowed on Anatahan in 1988. Populations on islands north of Saipan are Category 2 candidate endangered species, meaning that the US Fish and Wildlife Service has some evidence of vulnerability but not enough to justify an endangered listing at present.
Agrihan Wiles et al. (1989) made counts of two separate colonies in 1983. They estimated Agrihan's bat population to be about 1000 animals. This was considered conservative and as no surveys were made of the island's upper hillsides and crater, which are partly forested, it is likely that several thousand bats could be present on this large island.
Anatahan Wiles et al. (1989) report that circumnavigation of the island in 1983 revealed two bat colonies on the lower slopes of the east and north-east coasts. Counts of 628 and 576 bats were obtained. Their survey did not include the central crater. They suspected that similar numbers of animals dispersed along the southern coast and into the island's craters, and estimated that about 2500 bats occurred in these two colonies. T. O. Lemke climbed to the southern rim of the large central crater in 1984, and noted that fruit bats were common inside the crater but did not make counts. The overall estimate for the island was 3000 bats.
Ascuncion Wiles et al. (1989) made one count in 1983, sighting 394 bats. In 1984 a high level of bat activity was seen. Adequate forest cover and food are available on the island to support a relatively small but dense population of fruit bats. Wiles et al. (1989) estimated 400 fruit bats to be on Ascuncion.
Farallon de Medinilla Modest numbers of bats were reported in 1971. Since then, the US military has used the island as a bombardment range. Wiles et al. (1989) report on a visit in 1984 during which no bats were seen.
Farallon de Parajos Wiles et al. (1989) made 1-day surveys in 1983 and 1984 but located no fruit bats on the island and found that suitable habitat for bats was absent.
Guguan Although fisherman have observed fruit bat colonies in the past in Guguan's eastern ravines (J. Eads, pers. comm.), none was located by Wiles et al. (1989) during a survey in 1983. However, they estimated that southern Guguan supported a dense population of fruit bats: of a total of 400, approximately 150 bats occurred in the centre of the island and 250 on the southern plateau and surrounding cliffs.
Maug Eldredge et al. (1977) and visiting fisherman previously noted small numbers of fruit bats on Maug. Wiles et al. (1989) reported on a visit in 1984, which located no bats. A typhoon in 1981 destroyed much of the existing forest and may have killed most of the bats. They estimated that Maug supported a small population of fewer than 25 bats.
Sarigan Wiles et al. (1989) surveyed the island in 1983 and found no colonies. They estimated the total population to be 125 animals. A second visit in 1983 located a small colony of 30 bats in a Ficus (Moraceae) tree in an isolated valley not previously visited.
Threatened by hunting and the effects of typhoons (see P. m. mariannus).
Ecology: See P. m. mariannus.
Population biology: See P. m. mariannus.
Occurrence in protected areas: The CNMI has declared the islands of Ascuncion, Farallon de Parajos and Maug as wildlife sanctuaries.
Summary of threats:
Recommended action: See P. m. mariannus.
Principal author for these populations: G. Wiles.
Priority Grade: 8 (No Data: Limited Distribution).
It has been suggested by K. Koopman (pers. comm.) that P. mearnsi may be synonymous with Pteropus speciosus.
Distribution: Philippines (Basilan, Mindanao [only on the Zamboanga Peninsula]).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
P. m. aruensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Aru Islands.
Summary of threats:
Recommended action:
P. m. keyensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Kai Islands (Little Kai).
Summary of threats:
Recommended action:
P. m. melanopogon
Priority Grade: 10 (No Data).
Distribution: Ambon; Banda Islands; Boano Buru; Gorong Islands (Manawoka); Saparua; Seram; Tenimber Islands.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 6.
P. m. melanotus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Nicobar Islands (Car Nicobar).
Summary of threats:
Recommended action:
P. m. modiglianii
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Enggano.
Summary of threats:
Recommended action:
P. m. natalis
Priority Grade: 11 (Not Threatened).
Distribution: Christmas Island off the south coast of Jawa.
Status: Christmas Island remained unsettled by humans until 1888, when it was colonized by a small group to exploit phosphate. Andrews (1900, 1909) provided comments on the status of fruit bats at the time of first human settlement of the island and 20 years after settlement in 1908. Unlike several other indigenous mammals on the island, which seemingly had become extinct by 1908, fruit bats were still present in good numbers. Visits in 1932 and 1947 showed that the fruit bats were still present in some abundance (Gibson-Hill, 1947). In 1984, about 3500 bats were estimated to be using six identified camps and perhaps the same number were roosting singly or in small groups dispersed through the forest. Two other camps have since been located (J. Tranter and H. Yorkston, pers. comm.). The maximum size of the population seems unlikely to exceed 10,000. The camps appear to be important as maternity sites, and perhaps as assemblage areas for copulation.
Fruit bats are preyed on by feral cats;Pteropus constituted 21% by weight of the gut contents of 95 feral cats and are thus an important source of food for this animal. However, this situation has probably existed since the turn of the century and appears stable (Tidemann, 1989). Introduced house rats (Rattus rattus) also form an important part of the diet of feral cats (31% by weight) and probably for this reason, feral cats are commonest in areas of seral vegetation (non-climax vegetation, which occurs on ground that has been mined for phosphate) which sustain large numbers of rats. In seral vegetation, Pteropus feed close to the ground on Muntingia (Flacourtiaceae) fruit and it is likely that most fall prey to cats there.
Until recently, a workforce of about 3000 people was stationed on the island in connection with phosphate mining. A number hunted fruit bats for consumption and quite large catches were made, in some cases 200 at a time. Hunting is a simple matter because these bats to a large extent are diurnal and evince little fear of humans. Information on when most catches were made was difficult to obtain, but it seems likely that most hunting was done at about the same time as the peak of births (Tidemann, 1985, 1987). Phosphate mining ceased in 1987 because it was no longer profitable and the human population of the island has since become much reduced. Hunting does not now seem to be a serious threat to survival. A much reduced mining venture is to be started up soon, but it will be restricted to areas that have already been cleared of natural vegetation.
Probably of far greater concern in the longer term is the fact that in mid-1987 the rat snake (Family Colubridae: Lycodon aulicus capucinus) was accidentally introduced to the island from Asia (probably in building materials) and since then about 30 individuals have been found (Smith, 1988; D. Phillips, pers. comm.). Given the threat posed to P. mariannus on Guam by the introduced colubrid snake, Boiga irregularis, it seems likely that Lycodon could also become a serious problem on Christmas Island.
Ecology: Utilizes most vegetation types present, including seral vegetation, which occurs on ground that has been mined for phosphate.
Camps were located in Pisonia (Nyctaginaceae), Barringtonia (Lecythidaceae) and Ficus (Moraceae) canopy emergents, but dispersed individuals were present in many vegetation associations. An important feature of camp locations seems to be the ease with which departing animals can gain access to wind currents, which they use extensively to commute to feeding locations.
The following food plants were noted either through direct observation or analysis of gut contents. Pollen from many other taxa were also isolated from gut contents, but were unidentifiable due to a lack of reference material. Anacardiaceae: Mangifera indica (FR); Caricaceae: Carica papaya (FR); Combretaceae: Terminalia catappa (FR,FL); Euphorbiaceae: Macaranga tanarius (FL); Flacourtiaceae: Muntingia calabura (FR); Lecythidaceae: Barringtonia asiatica (FL), B. racemosa (FL); Meliaceae: Dysoxylum gaudichaudianum (FL), Melia azedarach (FL); Moraceae: Ficus spp. (FR,L); Myrtaceae: Eugenia grandis (FR,FL), Psidium guajava (FR); Palmae: Arenga listeri (FL), Cocos nucifera (FL); Rubiaceae: Morinda citrifolia (FL); Sapindaceae: Tristiropsis acutangula (FR,FL); Sapotaceae: Plachonella nitida (FR,FL); Solanaceae: Physalis sp. (FL); Ulmaceae: Celtis cinnamomea (FR); Urticaceae: Dendrocnide sp. (FL), Pipturus argenteus (FL).
The fruit of the introduced plant Muntingia calabura provides an important food source, particularly in areas that have been cleared for phosphate mining. Muntingia is a colonist of disturbed areas and its seeds are dispersed by P. melanotus.
A revegetation programme is also about to be initiated by the Australian National Parks and Wildlife Service. This should ensure an expansion of the feeding resources available.
Population biology: There is a single breeding season with a peak period of births in February. One young is produced after a gestation of about 5 months. Females mature rapidly and can become pregnant when they are only 6 months old. By contrast, males take about 18 months to reach maturity. The result of these differences in maturation rates is a highly skewed population structure in which mature females outnumber mature males by about 3:1 which is reflected in the social structure of camps and a consequently polygamous mating system.
Occurrence in protected areas: Christmas Island National Park is under the control of the Australian National Parks and Wildlife Service. The National Park has recently been extended substantially and now covers 63% of the island. This action ensures protection of the major camps and feeding areas.
Summary of threats:
Recommended action:
Principal author for this subspecies: C. R. Tidemann.
P. m. niadicus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Nias.
Summary of threats:
Recommended action:
P. m. satyrus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: North Andaman Islands.
Summary of threats:
Recommended action:
P. m. tytleri
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: South Andaman Islands.
Summary of threats:
Recommended action:
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Federated States of Micronesia (?Chuuk [Mortlock Islands], Pohnpei [Ant Atoll, Pakin Atoll]).
Status: Does not receive local protection from the Federated States of Micronesia or the Pohnpei State Government.
Chuuk Reported to occur in the Mortlock Islands (Thomas, 1882), but its status and distribution there is unknown. Because of the small area of these islands, which totals only 12 sq km, the overall population probably does not contain more than several thousand individuals.
W. E. Rainey, based on an examination of the relevant specimens, but without actual fieldwork, believes that this locality record is erroneous and that there are no occurrences outside Pohnpei, Ant and Pakin.
Pohnpei Described as common at all elevations in the 1950s (Jackson, 1962). Bruner and Pratt (1979) observed ‘many’ bats throughout the island during visits in 1976 and 1978. Locally common in the northern half of Pohnpei in July 1981 (Wiles and Payne, 1986). Evidence gathered in 1989 indicated that significant declines in the populations have occurred recently (Rainey, 1990).
Ant Atoll: Common in 1957 on this atoll which lies 15 km west of Pohnpei (Jackson, 1962).
Pakin Atoll: Reported on this atoll (approximately 30 km west of Pohnpei) by Pohnpeian government staff (W. E. Rainey, pers. comm.).
Has been threatened by the large commercial trade that has developed in the past 20 years. Records show that 15,223 bats were exported from Pohnpei to Guam between 1979 and 1989 (Wiles, 1992). Shipments to Guam increased significantly in 1989, with 6478 animals exported (Wiles, 1992). Fruit bats from the Mortlock Islands have also been shipped to Guam in the past few years, but the volume of this trade and the species composition is unknown. Hunting for local use is virtually nonexistent because bats are viewed as unacceptable food by Pohnpeians. The recently amended CITES regulations have made this trade to Guam illegal.
Many areas of native forest at lower elevations on Pohnpei have been converted to agroforest, which contains breadfruit (Moraceae: Artocarpus altilis), coconut trees (Palmae: Cocos nucifera), mangoes (Anacardiaceae: Mangifera indica), bananas (Musaceae: Musa spp.), and other food crops, or secondary forest. This activity probably affects fruit bats by altering food sources and reducing colonial roosting sites.
Ecology: Jackson (1962) and Bruner and Pratt (1979) saw many individuals and pairs of bats around the island, an indication that a large proportion may roost outside colonies. G. Wiles observed a colony of about 200–300 individuals in forest on top of Sokeh's Rock in 1981.
Has been observed to eat the fruits of Clinostigma (Palmae) and the flowers of Ceiba pentandra (Bombacaceae) and Cocos nucifera (Jackson, 1962; author's data).
Population biology: Jackson (1962) collected unweaned young on Pohnpei in February, September and November.
Occurrence in protected areas: No protected areas have been established within the range of P. molossinus.
Summary of threats:
Recommended action:
Principal author for this species: G. Wiles.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 2
P. n. hilli
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Bismarck Archipelago (Admiralty Islands).
Summary of threats:
Recommended action:
P. n. neohibernicus
Priority Grade: 11 (Not Threatened).
Distribution: Bismarck Archipelago (Duke of York, Mioko [near Duke of York], New Britain, New Ireland); Gebe; Karkar Island;Misool; New Guinea; Sakar; Umboi.
Status: New Guinea Common (Flannery, 1990). Seems to be widespread on the mainland, being recorded from the Morehead region of the trans-Fly (Waithman, 1979), Mount Dayman (Brass, 1956), the Torricelli Mountains and Andei (=Manokwari) and Mansinam, Vogelkop Peninsula (Andersen, 1912). Indeed, it is probably present throughout most of the country below 1000 m. Can clearly survive on small islands, as demonstrated by its presence on Karkar and Sakar. Why it has not spread to other larger islands in the region, such as the Louisiade Archipelago and the D'Entrecasteaux Islands, remains mysterious.
Ecology: In New Guinea, forms massive camps in swamp forest in areas such as the upper Sepik, from which it flies into foothill forest and surrounding floodplain forest each night to feed on fruiting trees. The purple fruits of a sapotaceous tree are reported to be particularly attractive (Brass, 1964). One colony at Madang is composed of thousands of individuals. The clatter of noise of this colony can be heard for 0.5 km or so. A sweet odour characterizes the roosting site (Smith and Hood, 1981). On one night at about 21.30 h in January, Flannery (1990) observed hundreds of animals in the Yapsiei area, flying at considerable altitude, in an easterly direction from swamps along the Sepik into foothill forest. The wingbeats of this bat are extremely loud when they are circling overhead looking for food or about to alight in a tree. Smith and Hood (1981) report that on New Ireland this species stripped a Ficus tree (Moraceae) of fruit within 3 nights, and that bat activity around the tree dwindled to nothing after 5 nights. This species has a distinctive smell that it is difficult to describe, but it is nevertheless avidly sought as a food item in most of New Guinea.
Dillon (1960) reports on a very curious occurrence involving large pteropodids, probably P. neohibernicus, on the north coast of Irian Jaya at dawn. The bats were attempting to pluck floating fruit out of the sea and occasionally dipped into the water too far and splashed in. When this happened, they came to rest quietly on the surface with the wings well spread and gradually rode in on the waves of the rising tide. On reaching the beach they tumbled out of the light surf, and dragged themselves up the beach to the dunes above the high-water mark. From here they climbed the trunks of saplings to 2.4 or 3 m, and launched themselves into the air.
Population biology: Two out of four females taken in the Yapsiei area in January 1984 were pregnant. Smith and Hood (1981) reported finding individuals in breeding condition in June–August 1979 on New Ireland, and many females carrying half-grown young were observed. Hyde et al. (1984) believed that populations may separate into male and female camps at certain times of year but could not confirm this.
Summary of threats:
Recommended action:
Principal author for this subspecies: T. Flannery.
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Réunion (now extinct); Mauritius.
Status: Mauritius Locally common (Cheke and Dahl, 1981). There were some camps of a few hundred but most bats were in groups of 1–15. At night they ranged 16–24 km from the roost. All the camps were in hilly wooded country. Camps were found from Montagne Fayence in the east, south through the Bamboa Mountains, the southern foothills of the central Plateau, Black River gorges and west to Montagne du Rempart. They were absent from the Moka Range and Nicoliere in the north-west.
The population was estimated at 10,000 in 1974 but recent evidence suggests it may be higher (Cheke and Dahl, 1981).
Réunion It was originally described from here by Brisson (Andersen, 1912), but has long since been extinct.
Rodrigues A specimen of P. rodricensis found by J. Anderson in 1881 was re-identified as P. niger by Andersen (1912). If this species were ever established on Rodrigues, it colonized from Mauritius.
It is shot for food and sport. In 1974, the annual bag was 1000 individuals, although recent evidence suggests it may be higher. Large numbers of bats are shot at but few are killed, so many are probably wounded. Hunting is traditional and therefore difficult to ban, but it should be possible to reduce the wastage. Legislation is before parliament to omit bats from the list of game species permissible to hunt without giving them special status.
Cyclones can be a serious problem. Cyclone Carol in 1960 was very severe. ‘Gervaise’ in 1975 was the most severe since ‘Carol’ and ‘Claudette’ in 1979 also caused havoc. The effects of cyclones are that all the fruit is stripped off the trees, bats are swept from their roosts and there is a small amount of food available for the survivors
On Réunion this species is fully protected under wildlife regulations.
Ecology: Known to feed on mango (Anacardiaceae: Mangifera indica), litchi (Sapindaceae: Litchi chinensis), longan (Sapindaceae: Dimocarpus longan), jackfruit (Moraceae: Artocarpus heterophyllus), fruits of badamier (Combretaceae: Terminalia catappa), flowers of traveller's palm (Streliziaceae: Ravenala spp.), kapok (Bombacaceae: Ceiba pentandra), Mauritius hemp (Agavaceae: Furcraea gigantea) and fruit of native trees in the indigenous forest (Cheke and Dahl, 1981).
Population biology: Cheke and Dahl (1981) reported that females with large, almost fully independent young were seen in December. Copulation was observed in April. Copulating pairs separated from the flock to copulate elsewhere.
Summary of threats:
Recommended action:
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Santa Cruz Islands [Nendo]).
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Distribution: Buru; Seram.
Summary of threats:
Recommended action:
Priority Grade: 9 (Indeterminate).
Number of subspecies: 2.
P. o. auratus
Priority Grade: 8 (No Date: Limited Distribution).
Distribution: New Caledonia (Loyalty Islands [Lifou, Mare]).
Status: See P. o. ornatus.
Ecology: See P. o. ornatus
Population biology: see P. o. ornatus.
Summary of threats:
Recommended action:
P. o. ornatus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: New Caledonia (New Caledonia).
Status: Sanborn and Nicholson (1950) reported that ‘P. ornatus makes up the bulk of the fruit bat population in New Caledonia’ and that they were less numerous than in the New Hebrides or Solomon Islands. They observed only two large fruit bat camps, of 4000 and 1500, with all other aggregations averaging around 300 animals per site.
Current camp sizes or recent population estimates are unavailable, although recent commercial imports to New Caledonia of P. tonganus geddiei from Vanuatu (E. Banei, pers. comm.) suggest reduced local availability or restrictions on commercial harvest in New Caledonia. Hunting of fruit bats is controlled by local wildlife laws and commercial sale is prohibited.
Insufficient recent data are available to assess status or identify threats. Fruit bats remain a favoured food item in New Caledonia and are available in restaurants by arrangement (A. M. Bauer, pers. comm.).
Ecology: Sanborn and Nicholson (1950) noted that bats rested during the day in aggregations covering several acres which were typically located at the upper end of dense rain forest on mountain slopes. Aleurites moluccana (Euphorbiaceae), the tallest tree in these patches, was favoured for roosting. Both obvious damage to roost trees from prolonged site use and reluctance to abandon a site in response to shooting or other disturbance indicate substantial site fidelity. However, they also noted that site occupancy varied seasonally in response to a north-south gradient in fruiting and flowering of fruit trees. Few bats were observed roosting alone. When P. o. ornatus and P. tonganus were found in the same camp, they typically did not roost in the same tree.
Sanborn and Nicholson (1950) suggested that P. o. ornatus ranged from sea level to at least 1066 m, but they failed to observe fruit bats in ‘the dense forest commonly found on the tops of mountains…’ above that elevation.
Sanborn and Nicholson (1950) observed that this species foraged predominantly nocturnally, leaving the roost an hour before sunset. However, small numbers of animals were observed foraging after 14.00 h. Feeds on the following plants (Sanborn and Nicholson, 1950): Anacardiaceae: Mangifera indica (FR); Caricaceae: Carica papaya (FR); Elaeocarpaceae: Elaeocarpus percisifolius (FR); Juglandaceae: Carya spp. (FL); Musaceae: Musa spp. (FR); Myrtaceae: Melaleuca viridiflora (FL), Psidium guajava (FR), Syzygium spp. (FR); Palmae: Cocos nucifera (FL); Passifloraceae: Passiflora spp. (FR).
Population biology: Data suggest that there is a six month gestation and a birth peak from late August to October (Sanborn and Nicholson, 1950). Sanborn and Nicholson (1950) concluded animals did not breed until their second year and observed no instances of more than one embryo or young per female.
Summary of threats:
Recommended action:
Principal author for this species: W. E. Rainey.
Priority Grade: 10 (No Data).
Distribution: Bacan; Halmahera; Morotai;?Sulawesi; Ternate.
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Federated States of Micronesia (Chuuk [Mortlock Islands]).
Status: Does not receive local protection from the Federated States of Micronesia or the Chuuk State Government.
Mortlock Islands The Mortlock Islands are considered to include all of the islands south-east of the Chuuk Lagoon, from Nama Island to Satawan atoll. These islands are sometimes distinguished as the Upper Mortlocks (Nama Island and Losap atoll) and Lower Mortlocks (Namoluk atoll to Satawan atoll). Status and exact distribution are unknown. Thomas (1882) reported two species of small Pteropus, P. phaeocephalus and P. molossinus, to be present in the Mortlocks but did not provide specific island locations for his specimens. Because older references often list the Mortlocks as including only the atolls of Etal, Lukonor and Satawan, it is possible that Thomas's (1882) specimens originated from one of these three atolls.
Girschner (1912) and Marshall (1975) recorded Pteropus on all islets in Namoluk atoll, but did not identify which species was present or comment on general abundance.
Government officials in Chuuk report that fruit bats also occur on Losap atoll. Because the atoll is located about 80 km south-east of the Chuuk Lagoon, where P. insularis occurs, it is possible that this is the species present rather than P. phaeocephalus or P. molossinus.
No information is available on population size or trends. However, because of the small land area in the Mortlock Islands, which totals only about 12 sq km, the overall population probably does not contain more than several thousand animals.
Girschner (1912) reported that fruit bats are eaten by the residents of Namoluk atoll. Recent information from the Mortlocks suggested that bats were being harvested for commercial export to the Mariana Islands. In February 1989 G. Wiles learned from government officials in Chuuk that several people from the atolls of Losap, Etal and Satawan were sending regular shipments of bats to Moen, Chuuk, for export to the Marianas. Regular overnight boat traffic between the Mortlocks and Moen made the shipment of bats possible without spoilage. The amendment of the CITES regulations in 1989 has made this trade illegal.
Summary of threats:
Recommended action:
Priority Grade: 1 (Extinct).
Distribution: Belau.
Status: The government of Belau does not give this species any legal protection.
Presumed to be extinct. Known only from two specimens (both in the British Museum [Natural History]), which were collected prior to 1874 (K. Koopman, pers. comm.). Coultas (1931) spent 3 months in Belau in 1931 as part of the Whitney South Sea Expedition, but did not find P. pilosus. There have also been no observations of this bat in more recent studies (Perez, 1968; Owen, 1977; Wiles and Conry, 1990). In addition, none has been seen amongst shipments of P. mariannus pelewensis, which were confiscated and examined on Guam between 1984 and 1989 (Wiles, 1992).
Causes for the probable extinction are unknown. The bat's disappearance was perhaps related to the subsistence hunting of fruit bats by Belauans. P. pilosus is considerably larger than P. m. pelewensis and may have been selected for by hunters. It is possible that any animals that may have survived into the 1960s or 1970s were commercially harvested along with large numbers of P. m. pelewensis for export to the Marianas.
Summary of threats:
Recommended action:
Principal author for this species: G. Wiles.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Yapen off north New Guinea.
Status: Last seen in 1932, but there has been no recent expedition to investigate (T. Flannery and L. Seri, pers. comm.).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: Eastern Australia (New South Wales, Queensland, Victoria).
One of the few species in the genus that is confined to a continental land mass. It extends to higher latitudes than any other pteropodid. The most southerly breeding camp is near Ulladulla at a latitude of 35°16′ S. Endemic to the south-eastern forested areas of Australia, principally east of the Great Dividing Range. In the north its range overlaps with P. alecto and over most except the southern part it is seasonally sympatric with P. scapulatus (Hall, 1987). Camps containing all three species may be seen in the vicinity of the New South Wales/Queensland border. The distribution may be extending southwards (Strahan, 1983; Menkhorst and Dixon, 1985; Aston, 1987). Camps are scattered at more or less regular intervals down the latitudinal range, but tend to be occupied on a seasonal basis.
Pteropus poliocephalus (Photo by T. H. Kunz)
Status: There seems little doubt that numbers have declined substantially since the 1930s when the first estimate of abundance was carried out (Ratcliffe, 1931, 1932), and although an overall assessment of the population has yet to be carried out, it appears to be still relatively common. The major cause of the reduction in numbers appears to be the clearing of native vegetation for agriculture. Eucalypt forest has been extensively cleared over much of its range, and rain forest over the same area is now reduced to remnants, many of which support fruit bat camps. Forays from these and camps in other vegetation types to surrounding fruit crops cause significant economic damage (Fleming and Robinson, 1987; Jamieson, 1988). Shooting of fruit bats on crops is widely carried out as a control method, although the total number killed is likely to be insignificant at population level. Of much greater concern is the shooting of animals in camps, as is still practised in Queensland, which causes considerable disruption, often when pregnant or lactating females are present. The only completely effective method of crop protection available to fruit-growers at present is exclusion netting, which is costly to erect and maintain and is impracticable to use in some situations.
This species is protected in Victoria, but is currently unprotected in Queensland. It is protected in New South Wales, but permits may be obtained to destroy animals on crops.
Ecology: Numbers of animals in camps are linked to the reproductive cycle as well as food availability. Pregnant females congregate in the maternity camps some time before giving birth and numbers of males do not peak until later during lactation. Copulation occurs in these camps. Camps are usually in rain forests and less frequently in mangroves. In some cases they may be in very small isolates of rain forest, less than 2 ha, which are very prone to edge effects. In the south of its range camps are usually in gullies supporting small patches of forest within eucalypt forest.
The social behaviour associated with camping and breeding follows a fixed pattern. Females in advanced pregnancy segregate from the males and each bears a single young. Females carry their young for about 4–5 weeks. During the nursing period, males rejoin the females. Competition between males and courtship of females lead to pair-formation and the establishment of territories. The camp then divides into family units, groups of unattached adults, and juvenile packs, with guard groups of unattached old males on the periphery (Strahan, 1983).
Long distance movements between camps, both north and south, have been demonstrated through tracking of radio-tagged individuals (Eby, in press) and banded animals. There seems little doubt that the population is continuous with unrestricted gene flow.
In some years the rate of successful pregnancies is low and this is often coincident with high mortality rates of unweaned young. Significant adult mortality sometimes occurs over wide areas, probably through ingestion of plants containing toxic substances (L. S. Hall, pers. comm.). These events appear to coincide with poor flowering of various food plants.
Utilizes a wide range of flowers and fruits of both native and introduced plants, but main food source is the blossoms of various species of Eucalyptus and other species in the family Myrtaceae. Seasonal and spatial changes in the availability of this resource are the probable cause of long-distance movements (Ratcliffe, 1931, 1932; Nelson, 1965a, 1965b; Eby, in press). Animals present at one site over an extended period also show seasonal changes in diet reflecting locally changing food availability (Parry-Jones and Augee, in press). Individuals may forage up to 50 km away from a camp (Eby, in press). It is regarded as a pest of cultivated fruit over much of its range (Hall, 1987).
The following is a list of known food plants. Most cultivated fruits that are eaten have not been included; for a detailed discussion of these see Jamieson (1988). Sources are Ratcliffe (1931, 1932), Nelson (1965a, 1965b), Hall and Richards (1979), McWilliam (1985–86), Parry-Jones and Augee (in press). Apocynaceae: Melodinus australis (FR); Cunoniaceae: Schizomeria ovata (FR); Ebenaceae: Diospyros sp. (FR); Leguminosae: Bauhinia hookeri F. Mueller (FL);Castanospermum australe Cunn. (FL); Loranthaceae: Amyema spp. (FL); Moraceae: Ficus sp. (FR), Cudrania cochinchinensis (FR), Morus sp. (FR); Musaceae: Musa sp. (FR); Myrtaceae: Angophora costata (FL), A. floribunda (FL), A. subvelutina (FL), Callistemon spp. (FL), Eucalyptus acmenioides (FL), E. alba (FL), E. beyeri (FL), E. citriodora (FL), E. cloeziana (FL), E. crebra (FL), E. fibrosa (FL), E. grandis (FL), E. gummifera (FL), E. haemastoma (FL), E. intermedia (FL), E. longifolia (FL), E. maculata (FL), E. microcorys (FL), E. moluccana (FL), E. paniculata (FL), E. pilularis (FL), E. punctata (FL), E. racemosa (FL), E. resinifera (FL), E. robusta (FL), E. siderophloia (FL), E. tereticornis (FL), E. tessellaris (FL), E. tindaliae (FL), Leptospermum spp. (FL), Melaleuca quinquenervia (FL), Melaleuca sp. (FL), Psidium guajava (FR), Syncarpia glomulifera (FL), S. hillii (FL); Palmae: Archontophoenix cunninghamiana (FR) Livistona australis (R. Br.) Mart. (FL), Phoenix canariensis (FR); Pittosporaceae: Pittosporum undulatum (FR); Proteaceae: Banksia aemula (B. serratifolia) (FL), B. integrifolia (FL), B. serrata (FL), Grevillea robusta (FL); Rubiaceae: Morinda jasminoides (FR); Salicaceae: Populus sp. (L); Solanaceae: Solanum mauritianum (FR); Verbenaceae: Avicennia marina (L); Vitaceae: Cissus sp. (FR); Xanthorrhoeaceae: Xanthorrhoea spp. (FL).
Fujita and Tuttle (1991) also list the following food plants: Moraceae: Ficus macrophylla (FR); Myrtaceae: Angophora lanceolata (FL), Eucalyptus baileyana (FL), E. corymbosa (FL), E. fastigata (FL), E. globoidea (FL), E. hemiploia (FL), E. intermedia (FL), E. obliqua (FL), E. planchoniana (FL), E. propinqua (FL), E. radiata (FL), E. reanans (FL), E. saligna (FL), E. sieberi (FL), E. umbra (FL), E. viminalis (FL), Lophostemon confert (FL), Melaleuca leucadendron (FL), Syncarpia laurifolia (FL); Proteaceae: Banksia robur (FL); Sapindaceae: Diploglottis cunninghamii (FL).
Population biology: Reproductive biology has been summarized in Martin et al. (1987). Copulation peaks in March–April and most births occur in September–October. Both sexes appear to require 18 months or so to reach sexual maturity (Nelson, 1965b). Lactation may last for 5–6 months (Martin et al., 1987), although the young can fly when about 8–10 weeks old (Strahan, 1983).
Occurrence in protected areas: Colonies occur in many national parks and other protected areas, which contain large patches of eucalypt and some rain forest, which provide food. Because of the wide-ranging movements, and the incomplete nature of information about these and overall resource requirements, the degree of protection afforded is somewhat uncertain.
Summary of threats:
Recommended action:
Principal author for this species : C. R. Tidemann.
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Japan (Kazan-retto [Kita-iwojima, Iwojima, Minami-iwojima], Ogasawara-shato [Chichijima, Hahajima]).
Status: Kuroda (1930) noted that this species was abundant in Kazan-retto, but had been considerably reduced in the Ogasawara-shato. Ogasawara-shato and Kazan-retto were removed from Japan and assigned to the US Government from 1945 to 1967. During that period, Hahajima in Ogasawara-shato remained uninhabited, and several hundred bats were seen there just after the island was re-assigned to Japan in 1968, although a large number of animals seemed to have been regularly captured each year (Hasuo, 1969, 1970a; Nakane et al., 1970). The number of bats seems to have fallen rapidly since 1968, and Matsumoto (1978) reported that only three animals were simultaneously observed in the summer of 1978 when he and his colleagues surveyed the island. He also noted that this species had already been extirpated on Chichijima (Matsumoto, 1978, 1980).
An intensive survey was conducted on Minami-iwojima, an uninhabited island in Kazan-retto, in the middle of June 1982 recording a maximum of 79 animals observed simultaneously (Ishii, 1983). On the basis of these results, Ishii (1983) estimated a population size of ‘a hundred to several hundreds’. There is no information available for other islands.
Matsumoto (1978, 1980) assumed the reduction of available food plants to be the primary factor causing the rapid decline on Hahajima.
Designated as a natural monument by the Japanese Government in 1968, thus regulating its handling.
Ecology: Feeds on the following plants (Layard, 1829; Hasuo, 1969, 1970b; Imaizumi, 1970; Matsumoto, 1978, 1980): Agavaceae: Agave americana. (FL); Caricaceae: Carica papaya (FR); Lauraceae: Persea kobu (FR); Moraceae: Morus spp. (FR); Musaceae: Musa paradisiaca (FR); Pandanaceae: Freycinetia boninensis (FR, FL), Pandanus boninensis (FR); Sapotaceae: Pouteria obovata (FR).
Population biology: Ishii (1983) reported that two females captured on Minami-iwojima in the middle of June were pregnant, each with one embryo.
Occurrence in protected areas: Most of the range of distribution is within protected areas.
Summary of threats:
Recommended action:
Principal author for this species: H. Ota.
Priority Grade: 4 (Vulnerable).
Distribution: Miangas (formerly in the Philippines, now in Indonesia); Philippines (Balut, Camiguin, Leyte, Maripipi, Mindoro, Negros, Sarangani Islands, Sulu Archipelago [Bongao, Sanga Sanga, Tawitawi], Tablas).
Status: Philippines Rare at all elevations on most larger islands (Negros and Leyte; Heaney et al., 1989; Luzon; unpubl. data). Common on the very small island of Maripipi, but none was captured on adjacent Biliran (E. A. Rickart, pers. comm.). Many of the areas where it occurs are ‘politically sensitive’.
None of 27 bats marked and released in primary forest at Lake Balinsasayao on Negros was recaptured (Heideman and Heaney, 1989). It made up less than 0.5 to 3% of captures in forest and forest clearings, respectively, on Negros (Heaney et al., 1981; Heideman and Heaney, 1989). As it tends to forage and fly in the canopy, and hence is not commonly netted except on ridgetops, it may be more abundant than capture rates imply. However, it is possible that individuals range over large areas, and average population densities may be quite low.
Tolerates only a small amount of disturbance. Has been seen foraging in clearings within and near forest. It may fly at least 2 km from forest, but has never been taken at greater distances. Records prior to 1970 document presence at lowland sites on Negros where it was absent in sampling between 1981 and 1989. Not hunted systematically, as its roosts are apparently not conspicuous and its body size is relatively small for the genus. Threatened mainly by habitat destruction, and extirpation is likely on some islands within the next 20 years if forest clearance continues at the current rate.
Ecology: A bat of primary and good secondary forest at elevations from 200 m to at least 1000 m; probably occurs down to sea-level in suitable forest habitat. Absent from purely agricultural habitats (Heaney et al., 1989), but has been taken in mixed secondary growth, young secondary forest, and agricultural land within 2 km of forest (Heaney et al., 1989).
Apparently, a roosting bat has been observed only once. This was a single individual roosting from a frond of a 5 m canopy tree fern (Cyatheaceae: Cyathea) in secondary forest at 600 m elevation on Maripipi (E. A. Rickart pers. comm.) Probably roosts either in small groups or alone in inconspicuous sites in forest vegetation.
Lawrence (1939) reported capturing individuals feeding on a partially ripened fruit in a kapok tree (Bombacaceae: Ceiba pentandra) and another in an ‘alemandras tree’. Also feed on figs (Moraceae: Ficus spp.).
Aggregates in small numbers at fruiting trees, but is usually silent while feeding. In a fruiting tree in a clearing on Maripipi 15–20 individuals made occasional relatively quiet calls (E. A. Rickart, pers. comm.). They remained within the fruiting tree to feed; it is not clear how often they use feeding roosts away from fruiting trees. Six to twelve Acerodon jubatus and about the same number of smaller bats (probably Cynopterus brachyotis and/or Ptenochirus jagorii) fed simultaneously in the same tree.
Population biology: The few data available (Heideman, 1987) do not provide a clear indication of the timing of reproduction. On some islands there is a suggestion of asynchrony, while on others females appear to be fairly tightly synchronized. The occasional capture of females simultaneously lactating and in early pregnancy suggests that a post-partum oestrus occurs and that some or most females give birth twice a year.
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 7.
P. r. cognatus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (San Cristobal, Uki Ni Masi).
Summary of threats:
Recommended action:
P. r. grandis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Bougainville; Buka; Solomon Islands (Alu, Choiseul, Santa Isabel, Shortland).
Summary of threats:
Recommended action:
P. r. lavellanus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Ghizo, Ranonggo, Vella Lavella).
Summary of threats:
Recommended action:
P. r. monoensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Mono).
Summary of threats:
Recommended action:
P. r. rayneri
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Guadalcanal, Malaita).
Summary of threats:
Recommended action:
P. r. rennelli
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Rennell).
Summary of threats:
Recommended action:
P. r. rubianus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Kolombangara, New Georgia, Simbo).
Summary of threats:
Recommended action:
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Mauritius (extinct), Rodrigues.
Status: Mauritius A skull found on Round Island off Mauritius (Mason, 1907) was P. rodricensis. It would have been expected to have been found in dry woodland characteristic of north-west Mauritius. The last relict of this habitat type remained on Round Island long after it was destroyed on the mainland.
Rodrigues Cheke and Dahl (1981) documented the decline on Rodrigues. In the 18th and 19th centuries and until 1916 it was reported as abundant. In 1955, large numbers (about 500) roosted in tamarinds (Leguminosae: Tamarindus indica) on the east flank of Baie aux Huitres valley below Jardin Mamzelle. There were 100 or so at Cascade Pigeon, and ‘plenty’ in Anse Mourouk and elsewhere. In total there were about 1000 individuals. In 1965, there were fewer bats but they were still common. There were 200 at Cascade Pigeon and good numbers at Jardin Mamzelle. Between 1967 and 1971 Elysee noted a marked decline in numbers. In 1971–72 Ales noted no more than 10 bats at Jardin Mamzelle. In 1974 very low numbers were reported (a maximum of 69+ in July). In 1975, Gardner noted 70+ at Cascade Pigeon. In 1976, Durrell counted up to 130 individuals. He caught 25, 18 for captive breeding. Eight (3 males and 5 females) went to Mauritius and 10 (3 males and 7 females) went to Jersey Zoo. In September 1978, Hartley and Cheke counted 151. In 1979, Cyclone Celine II reduced the population to about 70. By 1981, numbers had risen to at least 200 and more probably 250 (Carroll and Mace, 1988). In 1988, a single count of bats leaving the roost gave a figure of 412 (Carroll and Mace, 1988). At the end of February 1990 the population was estimated to be greater than 1000, based on an actual count of 650. A cyclone hit the island in February 1991 and a count immediately after the cyclone suggested little change in the population size. However, in the summer of 1991, an expedition from the University of Dublin counted a maximum of 350 bats (J. B. Carroll, pers. comm.).
Deforestation has been a serious threat. Between 1955 and 1968, there was clear felling of a large stand (100++) of old mature tamarinds at a place still known as Tamarins, on the west slope of the Baie aux Huitres valley. Ripe tamarind pods are a favourite food source. There have also been losses of other food and shelter trees at other sites.
Cyclones destroy remnant forest, and with it feeding areas. Both ‘Monique’ in 1968 and ‘Fabienne’ in 1972 probably killed many bats. This bat is known to be a poor flier in moderate wind.
Administrators and other Mauritians on duty on the island, in the habit of hunting fruit bats in Mauritius, have also done so on Rodrigues, being no doubt unaware the species was in danger. After a quiet period from 1972 to 1977, Mauritian visitors have again been shooting bats.
At the end of 1991 a total of 9 zoos held colonies of captive-bred P. rodricensis: Mauritius Black River Zoo; Jersey Zoo; Paignton Zoo; Chester Zoo; London Zoo; Bronx Zoo, New York; Brookfield Zoo, Chicago; Folsome Childrens Zoo, Nebraska; and the Lubee Foundation, Florida. The total captive population was greater than 250 animals (J. B. Carroll, pers. comm.).
Because of the success of the captive-breeding programme plans are being formulated to introduce the species to an Indian Ocean island outside the cyclone belt.
In Mauritius this species is fully protected under wildlife regulations.
Ecology: Tamarinds are very important food plants. Also feeds on the rose apple (Myrtaceae: Eugenia jambos), mango (Anacardiaceae: Mangifera indica), native Pandanus spp. (Pandanaceae), palms (Palmae: Hyophorbe verschafeltii and possibly Latania verschafeltii), Pyrostria trilocularis (Rubiaceae) (said to be abundant by Balfour, (1879), but now common only in Cascade Pigeon and Cascade St Louis), Terminalia catappa (Combretaceae) (formerly more common) and figs (Moraceae: Ficus spp.) (now very rare).
Population biology: There is an annual reproductive cycle. Dependent young are present from late August through to early February. Births in captivity are from October to April. Young are normally weaned at 10–11 weeks but occasionally suckle at 6 months and associate with their mother for 1 year.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Andersen (1908) proposed to separate populations from southern Madagascar as P. rufus princeps (type locality: Fort Dauphin). This was based on a difference in size and on the implicit assumption that the southern populations would be somehow geographically isolated from the typical northern and central ones. Later collecting and observations have shown the species's distribution to be continuous, and subspecific divisions can probably not be maintained.
Distribution: Madagascar.
Status: Mostly confined to coastal areas, offshore islands and low-lying plains towards the interior. Is entirely absent from the central high plateau, where Eidolon is the only fruit bat. Most common on the west and east coasts and less common on the arid south coast. Most roosts occur within 100 km of the coast, the only exception is one 80 km south of Ihosy in the central south.
Widely hunted for food, but there is no indication that the population is in decline. The effects of habitat destruction are unknown, but the bats will roost in primary forest, secondary forest and domestic fruit trees.
This species is listed as ‘noxious’ under wildlife regulations.
Ecology: Reported to visit flowers of Ceiba sp. (Bombacaceae) (Dobat and Peikert-Holle, 1985). B. Stequert reported to the Zoological Museum in Amsterdam that the species was observed to fly from the island Nosy Tany Kely to Nosi Be to feed on papaya (Caricaceae: Carica papaya), guava (Myrtaceae: Psidium guajava) and mango (Anacardiaceae: Mangifera indica) every evening (July, 1975).
Occurrence in protected areas: Roosts occur in the following protected areas:
Berenty private reserve
Mananara MAB Biosphere Reserve
RNI 9 Bemahara
RNI 12 Marojejy
The following protected areas are at least visited by Pteropus:
Analabe private reserve, Morondava
Andranonema special reserve
Montagne d'Ambre National Park
RNI 7 Ankarafantsika
Swiss cooperation forest, Morondava
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, M. Nicoll, S. Sowler.
Priority Grade: 4 (Vulnerable).
Number of subspecies: 2. The subspecies P. s. nawaiensis was considered by Andersen (1912) to be a distinct species, but was considered a subspecies of P. samoensis by Wodzicki and Felten (1975).
Is a Category 2 Candidate Endangered Species under the US Endangered Species Act (US Fish and Wildlife Service, 1988) meaning that the US Fish and Wildlife Service has some evidence of vulnerability but not enough to justify an endangered listing at present.
P. s. nawaiensis
Priority Grade: 9 (Indeterminate).
Distribution: Fiji (Nayau, Ovalau, Taveuni, Vanua Levu, Viti Levu).
Status: Noted as ‘abundant’ on Viti Levu and Vanua Levu by Pernetta and Watling (1978). A 1-week survey by the US Fish and Wildlife Service and Bat Conservation International in the vicinity of Suva, Viti Levu, in July 1989, located some individuals (G. Graham, pers. comm.), but current status, distribution, and population trends elsewhere in Fiji are unknown.
On Fiji, pteropodids are an important food item for peregrine falcons (Falco peregrinus). In rain-forest habitat, 56% of all prey-remains collected in eyries contained bats (White et al., 1988). There are four pteropodid species on Fiji, and it is not known what proportion of the bats taken are P. s. nawaiensis.
Fiji also lies within the typhoon belt. The effects of typhoons on populations in Samoa have been well documented (see P. s. samoensis).
Ecology: Does not form large colonies, and is most often found roosting alone (Pernetta and Watling, 1978).
According to Pernetta and Watling (1978) feeds in open areas on Fiji. D. Wilson (pers. comm.) observed one feeding on nectar or leaf buds of a Pandanus sp. (Panadanaceae) on Fiji.
Observations in 1989 on Viti Levu, Fiji, suggest P. s. nawaiensis is less diurnal than P. s. samoensis. P. s. nawaiensis was never observed foraging before 16.00h, did not exhibit soaring behaviour, and was seen flying only below the forest canopy (G. Graham, pers. comm.).
Summary of threats:
Recommended action:
P. s. samoensis
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: American Samoa (Tutuila); Western Samoa.
Status: In 1986 American Samoa passed legislation restricting local hunting, and prohibiting export of fruit bats. Similar regulations were instituted in Western Samoa in 1989.
Cox (1983, 1984b) observed indications of drastic declines on ‘Upolu, Western Samoa, in the 1980s due to commercial hunting and forest clearance. Subsequent work by the authors suggest generally low densities in Samoa, with probably no more than 500 animals left on Tutuila, the largest island on American Samoa, and populations seriously depleted on ‘Upolu. In 1984, Cox petitioned the US Fish and Wildlife Service (USFWS) to list P. s. samoensis as endangered under the US Endangered Species Act (US Fish and Wildlife Service, 1985). In 1988, based on a 1986 USFWS survey, the Service issued a finding of ‘not warranted’ but retained the species as a candidate, category 2 (US Fish and Wildlife Service, 1988). A 1989 USFWS survey concluded the species to be in ‘no immediate danger of extinction’ (Wilson, 1990), although reporting a 27% decline in total number of observations, and a 70% decline in number of animals observed on the large island of ‘Upolu between 1986 and 1989. The 1989 survey noted 176 animals at 49 survey sites. Anecdotal data after an intense typhoon in February 1990 indicate significant mortality, particularly from the increased post-storm hunting.
The primary cause of population declines in Samoa in recent years has been the commercial exportation of fruit bats for a luxury food market in Guam. From 1982–1986, Western Samoa was the primary supplier (Wiles and Payne, 1986; Wiles, 1992), shipping between 2700 and 8350 animals per year. Since export documents did not distinguish between the two Samoan species (P. s. samoensis and P. tonganus tonganus), it is impossible to assess what proportion of these shipments consisted of the endemic P. s. samoensis. Based on relative abundance, the expectation would be that most animals in any shipment would be P. t. tonganus. Chiefs on Savai'i, however, report instances of buyers on Guam seeking exclusively P. s. samoensis. There is also evidence that P. s. samoensis is behaviourally more vulnerable to hunting than P. t. tonganus, because it is a relatively slow, with poor manoeuvring ability, is primarily diurnal, and shows little inclination to escape when approached at a roost or shot at on the wing (authors' pers. obs.; L. Manuele, pers. comm.). The amendment of the CITES regulations in 1989 has made the international trade in bats illegal.
Although commercial hunting poses the greatest threat, chiefs in Western Samoa report that bat populations began to show a decline with the introduction of firearms (La Mosetele, pers. comm.). Harvesting of bats for local consumption may pose a threat to populations, especially those that have already been placed under stress by commercial harvest and deforestation. There are several examples of non-commercial hunting causing or contributing to pteropodid extinctions (e.g. P. tokudae on Guam [Wiles, 1987a];Dobsonia chapmani in the Philippines [Heideman and Heaney, 1988]).
Loss of habitat, due to logging and agricultural conversion, poses the second most significant threat to populations in Samoa. In Western Samoa, commercial logging is an important source of revenue (Engbring, 1989), and in 1985 only 38% of land on ‘Upolu, and 59% on Savai'i was classified as rain forest (Iakopo, 1985). Knowles (1988) estimated that 1–2% of rainforest habitat is being lost annually in American Samoa, mainly for agricultural purposes. The extremely steep (>30%) slope of much American Samoan land provides some limit on development (Engbring, 1989), but extensive loss of lowland forest can reduce wildlife populations relying on altitudinal variation in seasonal fruiting patterns (Cox et al., 1992).
The range of P. samoensis is within the typhoon belt. Samoa experienced at least 39 typhoons between 1831 and 1926, and there have been three very severe storms in the past 24 years, in 1966, 1987 and 1990 (Amerson et al., 1982; Knowles, 1988; authors' pers. obs.). Typhoon Tusi devastated the Manua Islands of American Samoa in January, 1987, and the more severe Typhoon Ofa hit all the islands of both Western and American Samoa in February, 1990, but damaged Savai'i most extensively. Post-storm surveys of the Manua Islands in 1987 suggested severe depletion of bat populations (Cox, 1987; Knowles, 1988), but considerable recovery of the forest and the bats by 1988 (Knowles, 1988). Reports from American and Western Samoa (Daschbach, 1990; B. Landin, pers. comm.) in February, 1990 indicated intense post-storm hunting pressure on surviving bat populations in the weeks following Typhoon Ofa.
Ecology: Does not form large colonies, and is most often found either alone, or in small family groups (adult male, adult female and a single young) in emergent canopy trees near the tops of ridges (Cox, 1983; authors' pers. obs.). Occasionally, aggregations of between 9 and 40 have been observed (I. Gurr, pers. comm; Wilson, 1990; authors' pers. obs.). On rare occasions, individuals have been found roosting in association with a P. tonganus tonganus colony (I. Gurr, pers. comm.). The male and female of individual pairs often roost in separate trees, but will usually fly out together, and return to the roost at approximately the same time (authors' pers. obs.).
Feeds primarily on the fruits and flowers of forest trees (Cox, 1983, 1984a), and is found in agroforest and villages much less frequently than P. t. tonganus. Although it has been observed feeding in coconut (Palmae: Cocos nucifera) plantations, in breadfruit (Moraceae: Artocarpus altilis) trees, and along the forest edge (Knowles, 1988; Engbring, 1989; authors' pers. obs.), it appears to prefer primary forest.
Serves as a major pollinator for the nectarless, dioecious liana, Freycinetia reinecki (Pandanaceae), by feeding on its fleshy bracts (Cox, 1984a). Also feeds on the following plants (Cox, 1983, 1984a; Engbring, 1989; Cox et al., 1992; D. Wilson, pers. comm.): Anacardiaceae: Mangifera indica (FR);Rhus taitensis (FL); Annonaceae: Cananga odorata (FL); Bombacaceae: Ceiba pentandra (FL); Caricaceae;Carica papaya (FR); Lecythidaceae: Barringtonia asiatica (FL); Leguminosae: Erythrina sp. (FL); Liliaceae: Collospermum samoense (FR); Loganiaceae: Fagraea beretiana (FR); Meliaceae: Dysoxylum samoense (FR), D. moata (FR); Moraceae: Artocarpus altilis (FR), Ficus graeffi (FR); Pandanaceae: Freycinetia reineckei (FL); Sapindaceae: Cupaniopsis samoensis (FR); Sapotaceae: Planchonella torricellensis (FR)
Primarily diurnal, and can be seen soaring on thermals (Cox, 1983), often spiralling upwards in pairs. Although its activity patterns differ seasonally, it most frequently feeds in the early morning and late afternoon (authors' pers. obs.). Observations at roosts and feeding trees, using night vision equipment, strongly suggest that animals return to a home roost close to dark, and do not leave the roost until shortly before dawn (authors' pers. obs.).
Individual animals, recognizable by colour patterns, are found at the same roosts day to day, and possibly even year to year. Antagonistic vocalizations have been heard near the apparent boundary between two roosting pairs, suggesting territorial defence of roosts. As many as five animals have been observed simultaneously in a single feeding tree, and although no hostile interactions were observed, each animal maintained a distance of at least 3 m from its nearest neighbour (authors' pers. obs.).
Population biology: The reproductive cycle is not known with certainty, but females have been observed carrying young in July and August (Engbring, 1989; J. Engbring, pers. comm.); a captive animal which was still pre-volant when acquired in July, was probably born in May or June. Since most Pteropus species have a well defined, synchronous breeding season, with one young per female per year (Baker and Baker, 1936; Falanruw, 1988a; Pierson and Rainey, 1992), it is probable that the majority of births occur between June and August.
Occurrence in protected areas: In 1988, the US Congress voted to establish a national park in American Samoa, which is planned to include about 3240 ha of forest habitat, containing both Samoan fruit bat species. In Western Samoa, during 1989 and 1990, two reserves (one of 12,150 ha and the other of 10,120 ha) were established on Savai'i as refugia for Samoan wildlife, particularly fruit bats. They incorporate the two largest tracts of unlogged lowland forest in Samoa. A long term research programme by the authors is monitoring fruit bat populations in both these areas. There is also O le Pupu-Pu'e National Park on the island of ‘Upolu, which contains suitable bat habitat.
Summary of threats:
Recommended action:
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Santa Cruz Islands).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: North-western, northern and eastern Australia (New South Wales, Northern Territory, Queensland, Victoria, Western Australia); southern New Guinea.
Status: Australia Apparently common throughout its geographic range (Strahan, 1983). Has the broadest range of any megachiropteran bat in Australia and overlaps with all other species. However, it is highly nomadic and most of the range covers underpopulated and remote regions of the continent where no information has been gathered and thus the sizes and movements of population have not been adequately assessed.
Not considered to be a serious pest to fruit-growers, although raids orchards and severely damages crops when native plants fail to flower. Since fruit bats appear to lead each other to ripening fruit, some fruit growers take advantage of this behaviour by poisoning fruit at the edge of an orchard, killing the scouts before they are able to return to the camp and communicate their discovery.
This species is unprotected in New South Wales, Queensland and Western Australia and listed as ‘noxious’ in Northern Territory.
New Guinea Rare (Flannery, 1990). Recorded from New Guinea on only one occasion, along the coastal strip between the Wassa Kussi and Morehead Rivers, in April-June 1973 (Waithman, 1979). Five skulls were collected by local hunters.
Ecology: Groups of individuals usually form camps in tall vegetation near water, although limestone cavities (Prociv, 1983) and sclerophyll forests (Hall and Richards, 1979) away from open water are also used. Camps more often in coastal mangroves, forest-lined banks of rivers and estuaries, and on islands in streams, Between 5000 and 1,000,000 may occupy one of these camps for periods ranging from a few days to 2 months.
Eucalyptus flowers (Myrtaceae: Eucalyptus spp.) form the greatest proportion of the diet. Fruits are of less importance. Leaves, growing shoots, bark and sap are also eaten. Feeding records are summarized below (data from Ratcliffe, 1931, 1932; Nelson, 1965a; Calaby, 1966; Armstrong, 1979; Friend and Braithwaite, 1986; McWilliam, 1985–86; author's data). Apocynaceae: Alstonia actinophylla (FL); Casuarinaceae: Casuarina cristata (C); Chrysobalanaceae: Maranthes corymbosa (FL); Cupressaceae: Callitris columellaris (S); Guttiferae: Calophyllum sil (FL); Lecythidaceae: Planchonia careya (FL); Leguminosae: Albizia umbellatum (FL), Bauhinia spp. (FL, S), Castanospermum australe (FL), Erythrophleum chlorostachys (FL); Moraceae: Ficus opposita (FR), F. racemosa (FR), F. scobina (FR), F. virens (FR); Myrtaceae: Angophora costata (syn: A. lanceolata costata) (FL), A. floribunda (syn: A. intermedia) (FL), A. woodsiana (FL), Eucalyptus bigalerita (FL), E. bleeseri (FL), E. camaldulensis (FL), E. citriodora (FL), E. clavigera (FL), E. confertiflora (FL), E. ferruginea (FL), E. fibrosa (syn: E. siderophloia) (FL), E. foelscheana (FL), E. grandifolia (FL), E. gummifera (FL), E. intermedia (FL), E. latifolia (FL), E. longifolia (FL), E. maculata (FL), E. microtheca (FL), E. miniata (FL), E. paniculata (FL), E. papuana (FL), E. pilularis (FL), E. polycarpa (FL), E. populnea (FL), E. porrecta (FL), E. ptychocarpa (FL, L), E. setosa (FL), E. tectfica (FL), E. tereticornis (FL), E. tessellaris (FL), E. tetrodonta (FL), E. trachyphloia (FL), Lophostemon grandiflorus (FL), L. lactifluus (FL), Melaleuca argentea (FL), M. cajuputi (FL), M. dealbata (FL), M. leucadendron (FL), M. nervosa (FL), M. quinquenervia (FL), M. symphyocarpa (FL, B), M. viridiflora (FL), Melaleuca sp. (FL), Syncarpia hillii (FL), Syzygium armstrongii (FR, FL), S. eucalyptoides bleeseri (FL), S. e. eucalyptoides (FR, FL), S. forte (FR, FL), S. suborbiculare (FL); Palmae: Carpentaria acuminata (FL); Proteaceae: Banksia integrifolia (FL), Grevillea pteridifolia (FL), G. robusta (FL); Rosaceae: Malus sylvestris (FR), Prunus persica, Pyrus communisRubiaceae: Aidia racemosa (syn: A. cochinchinensis) (FR, FL), Nauclea orientalis (FR, FL); Rutaceae: Citrus spp. (FR), Citrus reticulataSalicaceae: Populus spp. (FL, S, L); Tamaricaceae: Tamarix articulata (FL, S, L); Verbenaceae: Vitex glabrata (FR).
Fujita and Tuttle (1991) also listed the following food plants: Loranthaceae: Loranthus sp. (FL); Myrtaceae: Angophora subvelutina (FL), Eucalyptus acmenoides (FL), E. alba (FL), E. cloeiziana (FL), E. hemiploia (FL), E. planchoniana (FL), E. umbra (FL), Syncarpia glomulifera (FL), S. laurifolia (FL); Proteaceae: Banksia aemula (FL).
Important pollinators of native forests. Their diet of flowering species is extremely broad, including the dominant plant species of native forests. Therefore, the health of large tracts of native forests is maintained by the foraging behaviour of these animals (McCoy, 1990).
Nomadic and probably move into areas in response to the local availability of food, which fluctuates considerably from year to year. Most of the information about their movements comes from settled areas in the eastern part of their range during the summer months, between December and March, when camps form along the coastal strip. There is almost no information about their movements over most of their range for the rest of the year.
Population biology: The seasonal pattern of breeding and associated behaviour is similar to, but out of phase with, those of other Australian fruit bats. It has been suggested by McWilliam (1985–86) that the autumn parturition of this species compared to births in the spring for other Australian Pteropus species is linked to their need to be mobile in spring and summer when the majority of their food plants flower over vast areas. Camps are formed in November or December and territories are established within these roosts after the bats pair. After copulation, females leave to form their own groups and may establish new camps or even join the camps of other species of fruit bat. Young are born in April or May, when the population is dispersed, and are raised without the protection of huge numbers of cohorts. Young are occasionally seen in January and February, suggesting that there is a weak bimodal breeding cycle (Hall and Richards, 1979). Young are dependent on the adults for at least 3 months after birth until they learn to fly. Juveniles are frequently suckled and groomed by their mothers at the roost for a further 3–6 months after they fly. By the following November, when camps are again formed, the juveniles congregate in groups separate from the adults. They become sexually mature at 18 months of age.
Occurrence in protected areas: Makes use of suitable forests in national parks and reserves, although none of these has been established specifically for the protection of the species. In the northern part of their range, maternity camp sites in dense pockets of monsoon forest in national parks are used each year while the young are unable to fly.
Summary of threats:
Recommended action:
Principal author for this species: M. McCoy.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3. The subspecies P. s. aldabrensis and P. s. comorensis were formerly considered to be separate species. Recently, Bergmans (1990) has suggested that P. s. aldabrensis should again be considered as a separate species.
P. s. aldabrensis
Aldabra fruit bat
Priority Grade: 3 (Vulnerable: Limited Distribution).
Distribution: Aldabra Atoll.
Status: First collected by Abbot in 1893. There is a limited but healthy population (Cheke and Dahl, 1981). Carroll (1985) noted that although the population is only several hundred, it is apparently stable. Hutson (1991) also believed the population to be stable but limited by the size of the island (about 150 sq km) and food resources.
There are no known threats. However, the small size of the population and the limited land area available makes the bats vulnerable to the activities of man as well as to the effects of natural events such as cyclones.
Ecology: Feeds on the following plants (Fryer, 1911; Cheke and Dahl, 1981; Dobat and Peikert-Holle, 1985): Agavaceae: Agave sp. (FL); Celastraceae: Cassine sp. (FL); Combretaceae: Terminalia catappa (FR); Moraceae: Ficus avi-avi (FR); Palmae: Cocos nucifera (FL); Verbenaceae: Avicennia sp. (L).
Roberts and Seabrook (1989) reported bats feeding on coccids gleaned from the leaves of Ficus trees on Aldabra.
Population Biology: Females carrying young have been observed only in December and January, while copulation has been reported in October, November, March and June; this suggests a similar seasonality to that prevailing on the granitic islands (Cheke and Dahl, 1981).
Occurrence in protected areas: Aldabra Atoll was declared a Strict Nature Reserve on 17 February 1976, shortly before the return of Aldabra to the colony of the Seychelles on 23 June 1976. On 28 June 1976, the colony became independent as the New Republic of Seychelles and the status of Aldabra as a Strict Nature Reserve lapsed. But the government of Seychelles agreed to continue the policy of nature conservation on the atoll. On 9 September 1981 it was designated as a Special Reserve under the Seychelles National Parks and Nature Conservancy Act. In December 1982, The UNESCO World Heritage Committee agreed to its inclusion in the World Heritage List (Stoddart and Ferrari, 1983). The atoll is inhabited by only a small number of scientific personnel and their staff and so the usual pressures on the bat population do not exist (Carroll, 1985).
Summary of threats:
Recommended action:
P. s. comorensis
Comoros lesser fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: Comoros (Anjouan, Grand Comore, Mafia, Mayotte, Moheli); Tanzania (Mafia).
Status: The presence of ‘huge batts’ was first reported from Moheli in 1626 (Herbet, 1638). It was not until the 1860s that 2 species were known to be involved. There was little serious change in status over the first half of the 20th century.
Grand Comore Mountfort (1974) reported that in 1971, 1000 individuals were seen in the crater of the ‘Lac Salle’ in the north-eastern tip of Grand Comore. No bats were seen here in July 1975 or March 1977 (Cheke and Dahl, 1981). However, about 1500 individuals formed camps in the craters near Leoni and N'Bachile villages south of the capital (Moroni). Large numbers were seen around the crater lake of Dziani Boundouni in the east but no large camps were located. In 1975 and 1977, it was common in the west-central portion of Grand Comore up to 400 m. Bats were seen at a number of localities in the south and east but never in the northern third of the island. In 1988 The University of East Anglia Comoro Islands Expedition (Thorpe, 1988) reported bats as particularly common around Moroni with several large (50+ individuals) roosts in kapok trees (Bombacaceae: Ceiba pentandra) distributed around the town. Two roosts in tamarind trees (Leguminosae: Tamarindus indica) were seen at Lac Sal in late July, each containing approximately 15 individuals. A large roost of 150–200 individuals was present in a plantation of Casuarina equisetifolia (Casuarinaceae) at 610 m near the village of Maueni below the forest at La Grille. Another roost of 30 individuals was seen at 1000 m in the La Grille forest near Oussoudjou. No camps were seen close to Mitsamiouli on the north-east coast of the island, but bats were seen flying along the coast at dusk. A few individuals were seen flying at M'Vouni at 400 m on the east side of Mount Karthala, and some were seen along the road close to Foumboni on the south-east coast.
Anjouan Cheke and Dahl (1981) reported bats as numerous throughout the western slopes of the N'Tingui Massif. Several large (2000+) and smaller (<50) day camps were observed. Individuals were seen feeding on Kapok trees. In 1988 the UEA Expedition (Thorpe, 1988) noted that a large roost was probably present around the upper reaches of Mutsamudu. It was not located, although many bats could be seen flying at dusk. Three individuals were seen at dusk near Sima in the west of the island, and a few individuals were recorded at the lakes Dzialandze (900 m) and Dzialoutsounga (1000 m). Approximately 15 were seen feeding on the nectar of a clump of Erythrina fusca (Leguminosae) along the road between Moya and Sima and 25 individuals were seen in a roost close to the airport at Ouani.
Mayotte Cheke and Dahl (1981) reported one large feeding group near Coconi in the centre of the island. This island was not visited by the UEA Expedition.
Moheli Cheke and Dahl (1981) reported that sightings were less frequent than on the other islands but were made throughout the western half of the island between 0 and 300–400 m. Large numbers were seen around the crater lake of Dziani Boundouni in the east, but no large camps were found. In 1988 the UEA Expedition (Thorpe, 1988) reported that bats were seen feeding in the coconut palms (Palmae: Cocos nucifera) close to Fomboni, but no roosts were located. A few individuals were seen between 300–400 m but were not at all common. A roost of perhaps 20 individuals was seen in a ravine along the coastal road halfway between Fomboni and Miringoni.
Mafia N. Payne (pers. comm.) reported seeing two camps on Mafia in October 1990.
The UEA Expedition (Thorpe, 1988) noted that there are very specific requirements for roost sites; bats take advantage of topographical features to protect them from high winds, to maximize their thermoregulatory abilities, and to gain updrafts on the way to the feeding grounds. These requirements indicate that roost sites may be a limiting factor to the population size. Social interactions at the roost probably limit the density of individuals in a particular tree, and so if roosts are destroyed, disturbed, or have the local microclimate affected by deforestation, then populations may suffer. Deville (1977) noted that 44% of primary and secondary forest on the Comoros had been put under cultivation between 1968 and 1974. Current populations are patchily distributed compared to even 20 years ago (Cheke and Dahl, 1981).
There is little hunting on the islands because the majority of the population are Muslims who consider bats unclean. Away from the towns, there was evidence that bats were taken for food (Thorpe, 1988). The main methods of hunting were with catapults or sling-shots. It was unclear whether this practice was on the increase (Thorpe, 1988).
Cheke and Dahl (1981) considered that electricity wires were an important cause of mortality. The bats are electrocuted when both wires are touched simultaneously during take-off or landing. Verschuren (1985) considered it an important cause of mortality in the Seychelles. However, the UEA Expedition (Thorpe, 1988) observed few cases, and these were clustered around Moroni where both bats and electricity wires were common.
Cyclones affect Pteropus (Jones, 1980). Cheke and Dahl (1981) attributed the low numbers of bats seen on the west of Grand Comore in 1977 compared with 1975 to the cyclone, which caused high winds and heavy rain in early 1977. In 1950 a cyclone devastated Anjouan and Moheli and probably killed many bats (Cheke and Dahl, 1981).
Cheke and Dahl (1981) also noted that volcanic eruptions could be a threat to bat populations. A day after an eruption on Grand Comore, many bats were seen being sucked into the updraughts caused by the spout and burnt within it.
Ecology: The UEA Expedition observed bats feeding on the fruits of kapok (Ceiba pentandra), papaya (Caricaceae: Carica papaya) and the nectar from the flowers of Erythrina fusca. J. F. Dahl (pers. comm.) reports bats eating figs (Moraceae: Ficus sp.) and the fruits of badamier (Combretaceae: Terminalia catappa), and also the nectar from kapok and possibly also baobabs (Bombacaceae: Adansonia madagascariensis). This species is an important cross-pollinator for many fruit species (J. F. Dahl, pers. comm.).
Although occasional individuals could be seen flying during the day, most activity began just before sunset, when the large roosts would begin to disperse and start to forage. The bats usually return to the roosts before sunrise (Thorpe, 1988). Cheke and Dahl (1981) often recorded 5–6 km journeys, and suggested that much longer journeys were probably common.
An interesting behaviour is sea-dipping. At dusk, individuals were seen flying a short way out to sea, and following an incoming wave would dip their ventral surface or feet into the water before flying inland. This function appeared to be to collect sea-water only (Cheke and Dahl, 1981). Andersen (1912) suggested that it was to collect salts lacking in the fruit diet, which were ingested when the bat later preened itself, although an alternative suggestion was to rid the bats of ectoparasites (Bergmans, 1978b).
Cheke and Dahl (1981) suggested that strong flight (Dahl, 1979b) and the “time staggered” daily activity patterns (Dahl, 1979a) enabled this bat to exploit spatially and temporally separated food sources.
Population biology: Appears to be one annual breeding season extending over 2–3 months (Cheke and Dahl, 1981). This breeding season corresponds with the transition between the wet and dry seasons between April and June. The number of juvenile bats seen in June support this conclusion if a gestation of 150 days is assumed. Further confirmation is provided by Peters (1869) who shot pregnant females on Anjouan in October 1844.
Occurrence in protected areas: No protected areas have been established in the Comoros. The same seems to hold for Mafia (Stuart and Adams, 1989).
Summary of threats:
Recommended action:
P. s. seychellensis
Seychelles fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: Seychelles (Cousin, Curieuse, La Digue, Felicité, Mahé, Marianne, Praslin, Silhouette).
Status: Although first reported in 1742 by Picault, it was not described until 1877 (Milne-Edwards, 1877). It is considered to be common, with roosts reported on Mahé, Praslin, Silhouette and La Digue. Has also been seen flying over Felicité, Curieuse, Aride, Cousin, North and other islands to feed. There are sometimes temporary camps on these islands and there have been reports of flights from Praslin to Mahé (36 km).
Traditional roosts have been known for 30 years. The bats roost in tall trees, such as Casuarina (Casuarinaceae) and Albizia (Leguminosae). The roosts are mostly on north and west facing slopes during the south-east trade winds, which blow from May to October, the bats shifting to other sheltered sites during the north-west monsoon from November to April. This species has difficulty flying in moderate wind.
Racey (1979) reports a census of bats on the Praslin group of islands. Counts in July and August 1977 yielded figures of 1443 bats on Praslin (3 roosts) and 439 on La Digue (1 roost). The estimate of numbers on Praslin and surrounding islands was 2500. Nicoll and Racey (1981) report a further census on Felicité in 1978 when 170 bats (1 roost) were located. In September 1979, 2500 bats (10 roosts) were counted on Mahé. This figure may however be a substantial under-estimate and there may be as many as 10,000 on the island (Cheke and Dahl, 1981).
Man is the only predator (Nicoll and Racey, 1981). Bats are hunted for home consumption, but on Mahé and Praslin they are also sold to restaurants. Racey (in: Cheke and Dahl, 1981) reported that one restaurant on Mahé processed up to five bats a day, or 1500 or more per year. Nicoll and Racey (1981) judged that on Praslin there was little immediate cause for concern, but that the efforts to catch bats were much greater on Mahé because of the demands of tourist restaurants.
Nicoll and Racey (1981) found that, although the bats' diet included many cultivated fruits, the Seychellois generally did not consider it as a serious pest ‘and few kill it in order to protect fruit’. Until the government banned the carrying of firearms in 1977, the bats were shot while feeding. Since that time, netting and catapulting have been the principal methods of collecting bats. The banning of firearms has reduced the exploitation of the bats, but interest in increased development of commercial fruit growing poses another potential problem, in that bats might then be regarded as pests on fruit farms (Carroll, 1985).
Other human activities may adversely affect these bats (Nicoll and Racey, 1981). Accidental fires have occurred from time to time, killing food trees and destroying traditional roosts, forcing bats to seek alternative sites of poorer quality. Felling of roost trees is also a problem and was observed in one unoccupied roost on Praslin. A large Albizia tree was also felled within a large roost containing 2000–3000 bats inside the Morne Seychellois National Park on Mahé (Nicoll and Racey, 1981).
Verschuren (1985) noted that many bats in the Seychelles were killed by electicity wires (see under P. s. comorensis).
Ecology: Racey and Nicoll (1984) noted that there are several types of social group within roosts, although the first two comprise the majority of bats: 1) Family group always comprising one adult male and one adult female with a dependent juvenile; 2) Adult group comprising one adult male accompanied by one or two adult females with no juveniles present; 3) Juvenile aggregation; 4) Female groups; 5); Male groups; 6) Females and juveniles.
Following births in late November and December, most adults are organized into family groups. Each group occupies and defends a spherical territory about 2 m in diameter against other bats, most of the defence being carried out by the male. By April, when the bats move to the roosts they occupy during the south-east trade winds, the young form large aggregations in the centre of the roost and family groups are uncommon. The remaining bats form adult groups, which occupy similar territories to family groups, and are established and maintained by males. Adult groups account for the majority of bats throughout the south-east trade winds, but by the time females give birth, groups with more than one female lose one member to become family groups.
Individual bats occasionally hang on their own, often at the edge of the roost, at any time throughout the year. They can be of either sex, are occasionally joined by another bat, and are not territorial. They may have been unsuccessful in establishing a territory, or temporarily displaced from a territory following a disturbance in a roost. Juvenile aggregations are maintained throughout the south-east trade winds until the young bats are about 1 year old. There is no discernible territorial behaviour within these aggregations, where individuals frequently roost in association with neighbours. All male and all female groups are usually found on the edges of roosts, or as satellites of the main roosts.
Besides main seasonal roosts, groups of bats may be encountered occupying trees for from several hours to more than a month. These transient groups may comprise bats that have flown to alternative sites following disturbance within a roost. The numbers of bats involved are generally small but may occasionally reach several hundreds. Bats frequently fly between islands to feed and may be absent from their home roost for several days or weeks. These bats are normally well spaced, hang quietly among the foliage and are difficult to detect. Some roost sites are used only when the bats are in transit from one seasonal roost to another.
Racey and Nicoll (1984) listed the following food plants, although they considered this list to be incomplete: Anacardiaceae: Anacardium occidentale (FR), Mangifera indica (FR), Spondias dulcis (FR); Apocynaceae: Neisosperma oppositifolia (FR); Bombacaceae: Ceiba pentandra (FL); Caricaeae: Carica papaya (FR?, taken only in captivity); Combretaceae: Terminalia catappa (FR); Flacourtiaceae: Aphloia sp. (FR?, not confirmed); Guttiferae: Calophyllum inophyllum (FR), Pentadesma butyracea (FL); Meliaceae: Sandoricum indicum (FR); Moraceae: Artocarpus altilis (FR), A. heterophyllus (FR), Ficus avi-avi (FR), F. benghalensis (FR), F. natutarum (FR), F. reflexa (FR); Musaceae: Musa spp. (FR?, only taken in captivity); Myrtaceae: Eugenia jambos (FR?), E. javanica (FR), E. malaccensis (FR, FL), Psidium guajava (FR), P. littorale (FR); Palmae: Cocos nucifera (FL); Sapotaceae: Mimusops sechellarum (FR), Northea hornei (FR); Sonneratiaceae: Sonneratia caseolaris (FR?, not confirmed).
Some fruits may be carried away from the source tree and eaten elsewhere. For example, the fruits of the capucin, Northea hornei, were found under roost trees, although there were no capucin trees in the vicinity. It is also common to see flying bats carrying the fruits of the cashew (Anacardium occidentale). On Cousin the fruits of the bois chauve-souris, Neisosperma oppositifolia, could be found some distance from the trees after the bats had visited the island during the night and have even been found under bat roosts on Praslin, an island where Neisosperma has not been found. It is possible that these fruits are among several that are regularly dispersed by fruit bats. As well as a possible seed disperser, the bat may act as a pollinator of species such as Pentadesma butyracea, whose pale flowers with their copious nectar supply are borne on long, straight branches, and smell of rancid butter. Drops of nectar are present all over the petals, and to reach these, the fruit bat must brush against the anthers. Another very popular tree when in flower is kapok, Ceiba pentandra, which is known to be pollinated by fruit bats elsewhere in its range (Baker and Harris, 1959).
Population biology: Copulation occurs most commonly in June and July, although males may attempt to copulate at any time, and most young are born in November and December, suggesting a gestation similar to that recorded in other pteropodids (Racey, 1973). Births have also been recorded in September and October with a single occurrence in March. There is no evidence to suggest the litter size exceeds one.
Occurrence in protected areas: Nicoll and Racey (1981) mentioned a roost of 2000–3000 bats in the Morne Seychellois National Park on Mahé.
Summary of threats:
Recommended action:
Doubts exist as to the systematic status of this species: it may be synonymous with Pteropus griseus, which is relatively widely distributed (R. Wirth and W. L. R. Oliver, pers. comm.).
Priority Grade: 6 (Rare).
Distribution: Laut Kecil Islands and Masalembu Besar in the Java Sea; Philippines (Basilan, Malanipa, Mindanao, Sulu Archipelago). Records from Cebu and Negros in the Philippines are erroneous.
Status: Uncertain. It is rare in collections. Bräutigam (1989) noted that there is no good information on distribution or population numbers. It is dependent on forest trees for roosting and is assumed to be colonial (Bräutigam, 1989), making it particularly vulnerable to deforestation and hunting. It is subject to sport hunting and occurs in an area where forest is being cleared and hunting pressure is heavy (Bräutigam, 1989). Available evidence suggests that it is already experiencing population declines due to local hunting pressure and loss of suitable habitat (Bräutigam, 1989). Bräutigam (1989) believed local use to be limited because bats are forbidden food among the growing Muslim community within the species range.
Is known predominantly from ‘politically sensitive’ areas, which may preclude the initiation of any wide ranging surveys of status and management needs (R. Wirth and W. L. R. Oliver, pers. comm.).
Summary of threats:
Recommended action:
Principal authors for this species: P. D. Heideman, L. R. Heaney.
Priority Grade: 1 (Extinct).
Distribution: Mauritius; Réunion.
Status: Mauritius Cheke and Dahl (1981) concluded that it was extinct. First recorded on Mauritius by Desjardin in 1829/30, the last authentic record was in 1859. It is believed to have died out between 1864 and 1873. It roosted in trees and deforestation would have led to fewer roosts, and these would have been more accessible to hunters.
Réunion First described from Réunion by Brisson in 1756. La Nux (1772) noted that it had been common in 1722 but was by then rare. No new records appeared after 1862 when it was described as very rare. It seems probable, therefore, that it became extinct in the 1860s. The reasons for extinction may have been a combination of receding forest, a low birth rate and hunting. In the 1730s it had been common enough to consider using bat oil for trade. In 1772 both this species and P. niger were becoming rarer. It possible that it was able to exploit the higher altitude forest and thus survive longer than P. niger.
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Number of subspecies: 3.
P. t. capistratus
Priority Grade: 10 (No Data).
Distribution: Bismarck Archipelago (Duke of York, New Britain, New Ireland).
Summary of threats:
Recommended action:
P. t. liops
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Buru.
Summary of threats:
Recommended action:
P. t. temmincki
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Ambon; Seram.
Occurrence in protected areas:
Seram
Manusela National Park
Summary of threats:
Recommended action:
Priority Grade: 1 (Extinct?).
Distribution: Guam.
Status: Classified as endangered by the US Fish and Wildlife Service and the Government of Guam.
Guam Considered to be extinct on Guam (Wiles, 1987a). It had always been considered rare by hunters, elderly residents and scientific collectors (Baker, 1948; Perez, 1972). Only three specimens have ever been collected: two individuals were obtained in 1930 (Tate, 1934) and hunters shot a female in 1968 (Perez, 1972). The latter animal was the only one present among more than 100 P. mariannus killed and examined during legal hunting seasons in the 1960s. Aside from a possible sighting by M. Wheeler (pers. comm.) at Ritidian Point in June 1979, no observations have been made since 1968, despite intensive field work on the island's fruit bats since 1978 (Wiles, 1987a).
Other islands in the Marianas Several hunters have reported the presence of a small fruit bat, possibly P. tokudae, on Anatahan and several other islands in the northern Marianas. There are also conflicting reports about the historical presence of a small fruit bat on Rota, although recent interviews with elderly residents indicate that this was probably not true (J. Riechel, pers. comm.). Japanese collectors visited some of the islands in the 1920s and 1930s, but there have been few other biological investigations of the islands north of Saipan. The survey of Wiles et al. (1989) and a number of recent field trips to each of the islands by the staff of the Commonwealth of the Northern Mariana Island Division of Fish and Wildlife have not discovered any fruit bats other than P. mariannus.
There were a number of possible causes of the decline. Residents of Guam have long hunted Pteropus as a food source. The island's population of P. mariannus declined greatly as a result (Coultas, 1931; Baker, 1948; Perez, 1972; Wiles, 1987a), and hunting may have had a similar impact on P. tokudae. It may have been more vulnerable than P. mariannus to hunting with long-handled hoop nets, a technique widely used in the past.
The species was already rare by the time the brown tree snake (Boiga irregularis) was introduced to Guam after World War II. Snakes have become abundant and are a serious predator on young P. mariannus and other wildlife (Savidge, 1987; Wiles, 1987b). Snakes were found mainly in southern Guam for the first one or two decades after their arrival and did not reach the northern end of the island in large numbers until the early 1980s. Thus, snakes could have preyed on any P. tokudae remaining in the south, but it is doubtful that they ever did so in northern Guam.
Although some loss of forest occurred on Guam prior to World War II, large amounts of suitable habitat remain throughout the island. It seems unlikely that habitat destruction limited the small population of this species that remained in the 1930s and thereafter.
Population biology: The female killed in 1968 was carrying a juvenile that was large enough to fly away and escape capture (Perez, 1972, unpubl. data).
Summary of threats:
Recommended action:
Principal author for this species: G. Wiles.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
One of the most widely distributed of all Pteropus species, and in the eastern portions of its range (i.e., Niue, Cook Islands and Tonga) it is the only bat species. It is described by Koopman (1979) as a ‘supertramp’ species, referring to its absence from the largest, species-rich islands, and its occurrence on small, species-poor ones. There is a decrease in size across the range of P. t. geddiei and P. t. tonganus, with the Cook Islands population being the smallest (Wodzicki and Felten, 1980).
P. t. basiliscus
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Karkar Island off the north-east coast of New Guinea (Koopman, 1979), but also apparently occurs on Koil Island, Schouten Islands, about 200 km to the west (specimen in the British Museum [Natural History]).
Status: Current status is unknown.
The exact nature of threats are not known, but it is likely that they are similar to those facing other subpecies of P. tonganus. It is likely that it is a popular food item and as such may be threatened by commercial and non-commercial harvesting.
Deforestation poses a significant threat to fruit bat populations on many Pacific Islands and may similarly affect this bat.
Typhoons are also a recurring problem in the Pacific and can be devastating to local bat populations (e.g., P. samoensis in Samoa, hit by a storm in February 1990).
Epidemics and the effects of predators may also pose a threat (see P. t. tonganus).
Occurrence in protected areas: There are not thought to be any protected areas within the range of P. t. basiliscus.
Summary of threats:
Recommended action:
P. t. geddiei
Priority Grade: 11 (Not Threatened).
Distribution: New Caledonia (Loyalty Islands); Solomon Islands (Malaita, Rennell, Santa Cruz Islands); Vanuatu.
Status: New Caledonia Less common than the endemic P. ornatus in New Caledonia (Revilliod, 1914; Sanborn and Nicholson, 1950). It was described as ‘relatively abundant’ in New Caledonia in 1950, but limited, at least part of the year, to the central portion of the island (Sanborn and Nicholson, 1950). Recent reports on importation of P. tonganus for food from Vanuatu to New Caledonia (E. Banei, pers. comm.) raise serious questions about the current status of bat populations on New Caledonia.
The hunting of this species is regulated under local wildlife laws.
Solomon Islands Recent survey work by T. Flannery (pers. comm.) indicates that it is not common, but widespread, throughout the Solomon Islands.
Vanuatu Reportedly the most common bat on Vanuatu (Baker and Baker, 1936; Chambers and Esrom, 1988). In a recent questionnaire survey in Vanuatu, the majority (14) of respondents said bat densities had not changed; seven said they had increased, and five perceived a decrease. There was no evidence that local hunting pressure is causing population declines (Chambers and Esrom, 1988).
Fruit bats are a popular food item throughout most of the range. In Vanuatu, they occasionally appear in Port Vila market, and are served at local tourist hotels (Chambers and Esrom, 1988). In a survey questionnaire, 85% of the respondents said fruit bats were an important food item, and that people in the villages killed them using traditional methods (bow and arrow, sticks, stones, catapault, stick or string with hooks) as well as guns. Although taboos existed in some areas, there were generally few traditional controls on harvest (Chambers and Esrom, 1988). Bats are also widely eaten on New Caledonia (A. Brauer, pers. comm.).
The threat of non-commercial harvest should not be under-estimated, because overhunting is thought to have been the primary cause of extinction for at least two pteropodid species in recent years (P. tokudae on Guam [Wiles, 1987b];Dobsonia chapmani in the Philippines [Heaney and Heideman, 1988]), and currently places others at high risk, including the Niue and Cook Islands populations of P. tonganus, and P. voeltzkowi on Pemba off Tanzania (Seehausen, 1990).
Although commercial harvest for a luxury food market in Guam has caused serious depletions in some P. t. tonganus populations (see below), its impact on P. t. geddiei has probably been limited. A single bat was exported from the Solomon Islands to Guam in 1979. Also, 12 bats had been exported from Vanuatu to Guam up to the end of 1989 (Wiles, 1992). In addition, Vanuatu exported 365 P. tonganus to Noumea, New Caledonia in 1989 and early 1990 (E. Banei, pers. comm.).
Loss of native forest poses a significant threat to fruit bat populations on many Pacific and Indian Ocean islands, and has probably already affected P. t. geddiei on the Solomon Islands (Flannery, 1989).
Severe tropical storms are a recurrent threat. Evidence from the Marianas (P. O. Glass, pers. comm.), Vanuatu (Chambers and Esrom, 1988) and Samoa (N. Daschbach and B. Landin, pers. comm.) suggests, however, that the primary cause of storm-related mortality for fruit bats is intensive post-storm hunting. Following typhoons, food is in short supply for both bats and people. The animals are often forced to forage for fallen fruit on the ground, and roost without the concealment of canopy shelter, thus making themselves vulnerable to hunting.
On Fiji, pteropodids are an important food item for peregrine falcons (Falco peregrinus). In the range of P. t. geddiei, peregrine falcons are found in the Loyalty Islands, New Caledonia, the Solomon Islands and Vanuatu. Although no information is available, it is reasonable to expect that peregrines act as predators in these areas as well.
In the Solomon Islands, fruit bat teeth are used to make bracelets and necklaces, and on Vanuatu unspecified bat parts are used to make arrows (Chambers and Esrom, 1988).
Ecology: P. tonganus appears to be strongly colonial throughout most of its range (Baker and Baker, 1936; Degener, 1949; Medway and Marshall, 1975; Wodzicki and Felten, 1975; Pernetta and Watling, 1978; Wodzicki and Felten, 1980; Chambers and Esrom, 1988). Colonies are most often found in large, canopy trees, in mangrove or terrestrial forest, often near the edge of cliffs (Medway and Marshall, 1975; Wodzicki and Felten, 1980; authors' pers. obs.). Colony size and composition may vary seasonally. P. t. geddiei has been found roosting singly in the Cook Islands (Wodzicki and Felten, 1980) and singly, in pairs or small groups on Niue (Wodzicki and Felten, 1975), but such roosting behaviour may be attributable to depleted population levels.
In Vanuatu, Baker and Baker (1936) found that both sexes congregated together in camps from September to January, often in large Casuarina (Casuarinaceae) trees near the shore. When females became pregnant, in about February, they left the camps and became difficult to find, while the males stayed together. In June, in late pregnancy, females were found in inland camps, which at this time contained few males. At this time males lived separately, and were difficult to find. In New Caledonia as well, camp size fluctuated, and the animals moved seasonally (Sanborn and Nicholson, 1950). There P. t. geddiei was sometimes associated with the endemic P. ornatus, but the two species were almost always in separate trees (Sanborn and Nicholson, 1950).
P. tonganus feeds on the fruits and flowers of a wide range of native and introduced forest trees, plus a variety of agriculturally important plants, including banana (Musaceae: Musa spp.), coconut (Palmae: Cocos nucifera), papaya (Caricaceae: Carica papaya), mango (Anacardiaceae: Mangifera indica) and breadfruit (Moraceae: Artocarpus altilis). This species seems more willing to venture into agroforest and plantations than local endemics such as P. samoensis in Samoa, or P. anetianus in Vanuatu. Reported to feed on the fruits of Syzygium (Myrtaceae) and Ficus (Moraceae) and flowers of Eugenia malaccensis (Myrtaceae) in Vanuatu (Medway and Marshall, 1975). Chambers and Esrom (1988) noted that it was less frequently a blossom feeder than P. anetianus. Ridley (1930) reported that it fed on the following species: Leguminosae;Inocarpus fagifer (FR); Moraceae: Artocarpus incisor (FR); Sapotaceae: Sideroxylon sundaicum (FR).
Population biology: Baker and Baker (1936) noted that there was a general tendency for fruit bats to have a fixed, and synchronized breeding season, with young being born, after a six month gestation, in March-April north of latitude 4°N, and about September south of latitude 3°N. This pattern seems to hold for P. t. geddiei in Vanuatu (Baker and Baker, 1936) and New Caledonia (Sanborn and Nicholson, 1950).
Animals probably do not breed until they are 2 years old (Baker and Baker, 1936; Sanborn and Nicholson, 1950).
Occurrence in protected areas: No information is available on the status in reserves, but the following reserve areas may contain fruit bat populations:
New Caledonia
Parc Territorial de le Riviere Bleue
Solomon Islands
Queen Elizabeth Park (ca 6080 ha) (Singh, 1986)
Summary of threats:
Recommended action:
P. t. tonganus
Priority Grade: 9 (Indeterminate).
Distribution: American Samoa; Cook Islands (Mangaia, Rarotonga); Fiji; Niue; Tonga; Wallis and Futuna; Western Samoa.
The distribution includes the easternmost limit for the Pteropodidae.
Status: Protected by local hunting restrictions in American and Western Samoa, and Tonga (where the King's colony is protected), and excluded from commercial exploitation in Western and American Samoa.
American Samoa Amerson et al. (1982) estimated a population of 140,000 fruit bats in American Samoa in 1975– 76, but this is subject to question because the authors did not clearly specify their census techniques, and failed to distinguish between the two Samoan species (P. samoensis and P. tonganus). Knowles (1988) identified 11–14 roosts for P. tonganus on Tutuila, the largest island of American Samoa, with two large colonies (ca 3000 individuals in each), and other colonies in the low hundreds, yielding a total estimate of 10,000–13,000 animals on Tutuila. One of the large colonies contained no more than 1000 animals in July 1988. Populations appear to have been seriously depleted following Typhoon Ofa in February 1990.
Cook Islands Hunting for local consumption appears to be placing substantial pressure on populations in the Cook Islands. Wodzicki and Felten (1980) noted serious population declines in recent years, and D. Steadman (pers. comm.), who spent several months in the Cook Islands in 1989, reports that the population may number fewer than 1000 on Mangaia, and is considerably lower on Rarotonga. There are no local laws protecting fruit bats.
Fiji Widespread throughout the islands of Fiji (Pernetta and Watling, 1978), but current population trends are unknown. Degener (1949) described a roost of ‘thousands’ in the dense forest at Naruka. A brief survey by the US Fish and Wildlife Service and Bat Conservation International in 1989 located one colony near Suva with an estimated 5000 animals (D. Wilson, pers. comm.). Although Pernetta and Watling (1978) referred to it as abundant in 1978, Watling has recently expressed concerns about declines based on the disappearance of some colonies from traditional roosting sites (D. Bruning, pers. comm.).
International trade in this species is regulated under local wildlife laws.
Niue Apparently common at the turn of the century, when large flocks could be seen flying overhead, and still abundant in the 1920s. Reduction of native forest and the introduction of firearms, however, has led to drastic population reductions. On two extended visits in 1968 and 1969 Wodzicki and Felten (1975) did not observe any bats flying. A wildlife ordinance was passed in 1972 prohibiting the shooting of fruit bats, but was no longer in effect in 1980 (Wodzicki and Felten, 1980). Remaining animals were apparently restricted to the Tapu Forest Sanctuary and surrounding Huvalu Forest. Typhoon Ofa, which caused extensive damage to Niue in February 1990, probably provided additional threats to any remaining bat populations.
Tonga A brief faunal survey of Tonga in 1986 (Gill, 1987) noted large colonies at Tofoa and Kolovai on Tongatapu, and scattered individuals on ‘Eua. The Kolovai colony has been designated a fruit bat reserve, and is protected by the royal family. The status of populations on other islands, where the hunting ban does not apply, is unknown.
Wallis and Futuna A few specimens, now housed at the Paris Museum, were collected from Wallis and Futuna in the 1970s, but nothing is known of the current status of populations there.
Western Samoa Data are limited. Observations at flyways indicate that there is a small colony on the Falealupo Peninsula, and a colony near the Tafua Peninsula. There are also reports of a colony of about 100 near Fagafau on Savai'i.
Extensive hunting of fruit bats for local consumption has led to serious population declines on Niue (Wodzicki and Felten, 1975) and the Cook Islands (Wodzicki and Felten, 1980). Fruit bats are also a traditional food in Samoa, and eaten most commonly in the less westernized sections of Western Samoa. Here local chiefs report that bat populations began to decline with the introduction of firearms, but cultural recognition of the ecological importance of fruit bats provides a basis for instituting village controls against overharvest, as has been done now in two areas of Savai'i (Falealupo and Tafua).
A significant threat in recent years, particularly in Samoa, has been commercial exploitation for a luxury food market in Guam. As early as the late 1960s, following depletion of their own fruit bat resources, residents in Guam began importing bats from a number of islands in the Pacific (Wiles and Payne, 1986). American Samoa was a major supplier from 1980–86. and Western Samoa was the primary supplier from 1982–86. By the close of financial year 1989, at least 37,496 Samoan fruit bats (not distinguished by species) had been shipped to Guam. Over 5000 had come from Tonga and six from Fiji (Wiles, 1992). The amendment of the CITES regulations in 1989 made this international trade illegal.
Loss of native forest threatens P. t. tonganus on Niue (Wodzicki and Felten, 1975), the Cook Islands (Wodzicki and Felten, 1980) and Samoa (Cox, 1983, 1984b; Knowles, 1988). Substantial forest clearing in Wallis and Futuna has probably affected bat populations there. Even though P. tonganus is ecologically fairly adaptable, and is known to feed in agroforest, it shows a preference for native fruits in limited captive feeding trials, and roosts in emergent canopy trees at least part of the year. Deforestation and road construction also can pose an indirect threat by providing easier access to roost areas for hunters.
Severe tropical storms are a recurring problem. Evidence from Samoa (Daschbach, 1990; B. Landin, pers. comm.) suggests, however, that the primary source of storm-related mortality is intensive post-storm hunting (see under P. t. geddiei). Typhoon Ofa in February 1990 had a particularly severe impact on Samoa and Niue.
There are reports in the literature of epidemics decimating fruit bat populations on a number of islands, such as Pteropus neohibernicus hilli on Manus in the Admiralty Islands in 1985, P. rayneri grandis in Bougainville and Buka in the Solomon Islands in 1987 (Flannery, 1989) and P. mariannus ualanus in Kosrae in the 1930s (Coultas, 1931). Such an epidemic severely impacted P. t. tonganus populations near Savu Savu in Fiji sometime prior to 1949 (Degener, 1949). On Fiji, pteropodids are an important food item for peregrine falcons (Falco peregrinus). In rain-forest habitat, 56% of all prey remains collected in eyries contained bat parts (White et al., 1988).
Ecology: On Rarotonga, in the Cook Islands, this species roosts in hilly, deeply forested areas, in open trees, like Cananga odorata (Annonaceae), Cerbera manghas (Apocynaceae), Guettarda speciosa (Rubiaceae) and Homalium acuminatum (Flacourtiaceae). In Niue, it roosts primarily in Planchonella torricellensis (Sapotaceae), Ficus prolixa (Moraceae) and Syzygium richii (Myrtaceae) (Wodzicki and Felten, 1975). In Tonga, a large camp is located in Casuarina (Casuarinaceae) trees in the Kolovai fruit bat Sanctuary.
Combined data from the Cook Islands, Niue, Fiji and Samoa (Jaegar, 1954; Wodzicki and Felten, 1975, 1980; Cox, 1983; Ash, 1987; Cox et al., 1992) provided the following list of food plants: Anacardiaceae: Mangifera indica (FR), Rhus taitensis (FL); Annonaceae: Cananga odorata (FR, FL); Apocynaceae: Cerbera manghas (FR), Ochrosia oppositifolia (FR); Bombacaceae: Ceiba pentandra; Caricaceae: Carica papaya (FR); Combretaceae: Terminalia catappa (FR); Ebenaceae: Diospyros samoensis (FR); Elaeocarpaceae: Elaeocarpus rarotongensis (FL); Guttiferae: Calophyllum inophyllum (FR); Lauraceae: Persea americana (FR); Leguminosae: Inocarpus fagifer (FR); Loganiaceae: Fagraea beretiana (FR); Moraceae: Artocarpus altilis (FR), A. heterophyllus (FR), Ficus prolixa (FR); Musaceae: Musa paradisiaca (FR), M. nana (FR); Myrtaceae: Metrosideros villosa (FL), Psidium guajava (FL), Syzygium clusifolium (FR, FL), S. cumini (FR), S. inophylloides (FR, FL), S. jambos (FR, FL), S. malaccense (FR, FL), S. richii (FR, FL); Palmae: Cocos nucifera (FR, FL); Pandanaceae: Pandanus tectorius (FR); Rutaceae: Citrus sinensis (FR); Sapindaceae: Pometia pinnata (FR); Sapotaceae: Planchonella torricellensis (FR, FL); Solanaceae: Solanum lycopersicum (FR).
Observations of P. tonganus feeding at Ceiba pentandra trees indicate that animals stake out territories in the trees, and then fight with other animals to maintain their feeding post.
Population biology: The synchronized breeding season observed by Baker and Baker (1936) for P. t. geddiei (see above) seems to hold for P. t. tonganus populations in the Cook Islands (Wodzicki and Felten, 1980) but not for those in Niue or Samoa, where the limited data suggest an asynchronous breeding season as is found in P. mariannus on Yap and Guam (Falanruw, 1988a; Wiles, 1987a).
Wodzicki and Felten (1975) estimated that young were born on Niue from March to June, based on the weights of immature captured animals. A re-evaluation of their data, using the growth curve of a captive P. samoensis (a species of comparable size), however, suggested their specimens had been born in June, July and October. D. Wilson (pers. comm.) saw no young in Fiji in July but P. Cox and E. D. Pierson observed a roost in American Samoa in July 1987, which contained pregnant females and females carrying very small young. There are also observations of females carrying young in August, October and January (S. Gurr, pers. comm.). The estimated birth times for two immature animals that were netted in Samoa are October and April.
Occurrence in protected areas: Occurs within two recently established rain-forest reserves in Western Samoa (12,150 and 10,125 ha), and the 3250-ha area set aside in 1988 to become a United States National Park in American Samoa. There is a fruit bat sanctuary at Kolovai, on Tonga, where the animals are protected by the royal family. In 1975, occurred in the Tuvalu Forest sanctuary on Niue (Wodzicki and Felten, 1975), but no information is available on its current status there.
Summary of threats:
Recommended action:
Principal authors for this species: E. D. Pierson, W. E. Rainey, P. Cox, T. Elmqvist.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Solomon Islands (Santa Cruz Islands [Vanikoro]).
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Number of subspecies: 7.
Pteropus vampyrus (Photo by T. H. Kunz)
P. v. edulis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Savu; Timor.
Status: Timor Goodwin (1979) observed a spectacular colony of 2000 adults of both sexes near Metinar, Timor, in a dense mangrove forest which extended for about 8 km along the coast.
Ecology: The colony mentioned above on Timor fed on fruits of Ficus (Moraceae) in association with Pteropus griseus, Acerodon macklotii and Dobsonia peronii.
Summary of threats:
Recommended action:
P. v. intermedius
Priority Grade: 10 (No Data).
Distribution: Extreme south of Burma and adjacent areas of Thailand.
Summary of threats:
Recommended action:
P. v. lanensis
Priority Grade: 4 (Vulnerable).
Distribution: Philippines (Bohol, Canon, Catanduanes, Cebu, Dinagat, Leyte, Luzon, Mindanao, Mindoro, Negros, Palawan, Samar).
Status: Heavily hunted, both at its conspicuous roosts and in orchards. Individuals are sometimes captured by entanglement in weighted, fish hook-strung lines suspended from ropes stretched between tall trees in coconut groves (Palmae: Cocos nucifera). The declines in mixed Pteropus/Acerodon roosts from the reported 100,000 per camp in the 1920s to the 500–1000 reported currently indicate drastic falls in population numbers. It is possible that it could be extinct within the Philippines in the next 20 years, although it is more likely that small populations would persist in isolated areas. Although it may be able to persist in agricultural habitats, heavy hunting pressure is causing a serious decline on many Philippine islands. Most captures are for local consumption, but, in recent years, the large demand for fruit bats on Guam has resulted in heavy trade in large fruit bats, and a small number of these have been P. v. lanensis.
Ecology: Recorded from sea-level to at least 1300 m. Found in a variety of habitats, including primary forest, orchards and coconut groves. An important difference between this bat and Acerodon jubatus, which often uses the same roosts, is that P. v. lanensis can be found in orchards and coconut groves in agricultural areas.
Roosts in large colonies in canopy and emergent trees, often sharing roost sites with Acerodon jubatus. J. B. Steere (cited in Elliot, 1896) visited Negros in 1888, and wrote that ‘the fruit bats of the Philippines prefer small islands for their roosts, but will take up isolated localities. I found one roost on Negros occupying one immense hardwood tree standing by itself far from the forest on the plains of the western side of the island. Where they were not hunted for food by natives, they sometimes roost near native houses or villages for protection’. Taylor (1934) described a mixed colony of A. jubatus and P. v. lanensis on Mindanao made up of about 150,000 individuals, about one-third of which were the latter subspecies. Mudar and Allen (1986) reported specimens of both A. jubatus and P. v. lanensis from a roost of 800–1000 bats in north-eastern Luzon. There were similar mixed roosts in the Philippines that contained 500–1000 individuals.
Has been reported feeding on coconut pollen, nectar, and/or flowers and durian nectar and flowers (Bombacaceae: Durio zibethinus) (Gould, 1977) and ‘…flowering shoots and a wide variety of fruit, including rambutan (Sapindaceae: Nephelium lappaceum), langsat (Meliaceae: Lansium domesticum) and other cultivated fruit’ (Medway, 1978). Also feeds on fig fruits (Moraceae: Ficus spp.) (authors' pers. obs. and Goodwin, 1979), and probably eats a wide variety of other wild fruits.
Lawrence (1939) reported watching bats from a big colony ‘…in a tree in a large swamp near Zamboanga..’ (Mindanao) in April, when they flew about 27 km nightly to Basilan to feed. Rabor (1977) described commuting flights of these bats elsewhere in the Philippines, from camps to foraging grounds that must be at least 10–20 km distant, and this is consistent with reports for other large species in the genus (see review in Heideman and Heaney, 1989). Like A. jubatus, often feeds noisily in groups of a a few to 20 or more in large fruiting trees.
Population biology: In the Philippines, females appear to give birth once a year, in April or May. Timing of births is apparently similar throughout the Philippines (Rabor, 1977; Mudar and Allen, 1986; Heideman, 1987; authors' data), but more data are necessary.
Summary of threats:
Recommended action:
Principal authors for this subspecies: P. D. Heideman, L. R. Heaney.
P. v. malaccensis
Priority Grade: 11 (Not Threatened).
Distribution: Anamba Islands; Bangka; Lingga Islands; Mentawai Islands (Pagai Islands, Sipura); Peninsular Malaysia; Riau Archipelago; Singapore; Sumatra; Thailand; Vietnam.
Status: Peninsular Malaysia Widespread but declining in forest areas.
Sumatra Found quite commonly in the Padang Highlands up to 914 m.
Thailand Recorded from the coastal area of the peninsula and the south-east coast as far north as Korat, with records from the provinces of Chon Buri, Krabi and Hakhon Si Thammarat (Lekagul and McNeely, 1977; Yenbutra and Felten, 1986).
Vietnam Van Peenen (1969) noted that it had been recorded from only two localities in southern Vietnam. These were Phu Quoc Island and Hue in Thua Thien Province.
Threatened by the rapid loss and degradation of mangroves for coastal reclamation, aquaculture, and to logging and loss of lowland forest to commercial logging and land clearance for oil palm/rubber estates. It is also threatened by hunting (with or without licences). Is eaten throughout its range (in Malaysia more by Chinese, not Malays; elsewhere by almost everyone). It is also shot as a pest of fruit orchards despite its value as a pollinator.
Ecology: In Sumatra, roosts in kapok trees (Bombacaceae: Ceiba pentandra) in cultivation, and in remnant forest patches.
Feeds on cultivated durian (Bombacaceae: Durio zibethinus), rambutan (Sapindaceae: Nephelium lappaceum), mango (Anacardiaceae: Mangifera indica), banana (Musaceae: Musa spp.), langsat (Meliaceae: Lansium domesticum), and other orchard fruits. It also feeds on a wide variety of wild forest tree fruits as well as flowers and nectar.
Population biology: In Peninsular Malaysia, apparently pregnancies peak from November to January but can occur in other months. In Thailand, pregnancies occur in about the same period or later, with births in March–April. Females have one young each.
Occurrence in protected areas: In Peninsular Malaysia, occurs in the Taman Negara National Park, occasionally in the Krau Game Reserve, Pahang, and during 1987–88 a roost occurred in Templer's Park, Selangor. Widespread but declining in forest areas, most of which are forest reserves (i.e. forest under the jurisdiction of the Forest Department and used for logging). Also present in the area of the proposed Endau-Rompin Park, Johore/Pahang.
Colonies sometimes move, especially because of disturbance, and long-distance movements may carry them in and out of protected areas. Everywhere are more dependent on lowland forest over level ground than on hill forest, and become commoner towards the coast and mangrove areas.
Summary of threats:
Recommended action:
Principal author for this subspecies: G. W. H. Davison.
P. v. natunae
Priority Grade: 11 (Not Threatened).
Distribution: Borneo (Brunei, Central, East, South and West Kalimantan, Sabah [Balembangan, Banggi], Sarawak); Natuna Islands (Bunguran Besar).
Status: Payne et al. (1985) recorded that it was found throughout lowland coastal areas, occasionally invading the interior during the fruiting season. C. M. Francis (pers. comm.) reports that flock sizes in Sabah appear to have become smaller over the past 10 years, possibly indicating a decline.
Ecology: Eats both nectar and fruit including some orchard species such as rambutan (Sapindaceae: Nephelium lappaceum) and mangoes (Anacardiaceae: Mangifera indica). It pollinates the flowers of many forest trees such as durians (Bombacaceae: Durio zibethinus). It sometimes flies long distances to feed in flowering or fruiting trees (Payne et al., 1985).
Summary of threats:
Recommended action:
P. v. pluton
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Bali; Lombok; Sumbawa.
Summary of threats:
Recommended action:
P. v. vampyrus
Priority Grade: 11 (Not Threatened).
Distribution: Jawa; Krakatau Islands (Sebesi, Sebuku, Sertung).
Status: Tidemann et al. (in press) recorded it from islands in the Krakatau group off west Jawa. Bats were seen to move between the islands. A single specimen was seen roosting in a Casuarina (Casuarinaceae) tree and a colony of 250 roosted in Terminalia (Combretaceae) trees on Sertung in 1985 but was not seen in 1986. In the same region, Dammerman (1948) observed large numbers of Pteropus moving between Sebesi and Sebuku. At Bogor Gardens, west Java, it roosted in a variety of trees, including dead ones, in groups of hundreds of individuals (Kitchener et al., 1990).
Summary of threats:
Recommended action:
Priority Grade: 5 (Rare: Limited Distribution).
Distribution: New Caledonia (Loyalty Islands, New Caledonia).
Status: Sanborn and Nicholson (1950) regarded it as scarce on New Caledonia. There were indications of a wide distribution as regards both elevation and area. There is no information available regarding status in the Loyalty Islands.
The hunting of this species is regulated under local wildlife laws.
Summary of threats:
Recommended action:
Priority Grade: 1 (Endangered: Limited Distribution).
Distribution: Tanzania (Pemba).
Status: Andersen (1912) wrote that ‘only one colony of Pteropus voeltzkowi is said to exist in Pemba, at Fufuni, at the middle of the south coast’. Kingdon (1974) recorded it as occurring along the road between Mantangatuani and Mgogoni. Carroll (1985) quoted information supplied by R. H. W. Pakenham that there were several roosts in southern Pemba, and also on the islands off the south and north-west coasts. The bats on the islands off the south coast flew to forage on south Pemba. This is supported by specimens from various localities on Pemba in the British Museum (Natural History). From the available evidence, Carroll (1985) considered it to be ‘apparently abundant’. It is important to remember that Eidolon helvum also occurs on Pemba, and on at least one of the associated islands (Fundo). Thus, there could be confusion between species.
However, more recent evidence suggests it may have undergone a serious decline in numbers in the past few years. Seehausen (1990) reported on a visit made to Pemba in late October and early November 1989. Despite extensive searching, he could only locate one group of three bats on the central-west coast at Kisiwani; in addition on four occasions he saw a single bat in flight at dusk. At Mgogoni, it was reported that there was a small colony of about 50 bats in a section of natural forest on one of the steep, inaccessible hills. Seehausen (1990) could not confirm the presence of this colony. More recently, H. Bentjee (pers. comm.) reported seeing 150–200 Pteropus roosting in large trees in NW Ngezi Forest in December, 1989. He did not see any Pteropus in any other forest patches on Pemba (Msitu Mkuu and Ras Kiuyu).
Seehausen (1990) listed possible threats as predators, hunting and deforestation. Information on natural predators is limited, but there do not appear to be any that would constitute a serious threat. Hunting, on the other hand, may be a serious problem. In the past, traditional hunting methods, such as the use of long sticks, were used. But more recently, firearms have been a preferred method. The extent of hunting is unknown. Carroll (1985) thought that there was ‘some hunting’. It may be that, as in the Comoros, the Muslim population of Pemba consider bats unclean. Deforestation may also have played an important role in the decline. The western half of Pemba was formerly covered with tropical rain forest, whereas the eastern half was dominated by baobab (Bombacaceae: Adansonia digitata) and Borassus (Palmae) palm savannahs and coastal bush. Bats were endemic to the rain forest and formerly occurred throughout the west of the island but not in the east. Much of the original rain forest was cleared in the 18th Century, but bats continued to survive in remnant patches of forest on hill slopes. However, these remnants are now threatened and coupled with hunting this has reduced the population to a critical level (Seehausen, 1990).
Ecology: Is very social and it has been found in large colonies until very recently. It does not appear near human settlements unless large trees are fruiting. It is primarily a forest species and is now restricted to small remnants of forest in the north-west of the island.
Seehausen (in press) recorded the following food plants; Anacardiaceae: Mangifera indica (FR); Annonaceae: Cananga odorata (FL), Uvaria leptocladon (FR); Bombacaceae: Ceiba pentandra (FL); Caricaceae: Carica papaya (L); Moraceae: Artocarpus altilis (FR), ?A. heterophyllus (FR), Ficus capensis (FR), Ficus sp. (FR); Myrtaceae: Eugenia malaccensis (FR), Syzygium malaccense (FL), S. jambolanum (FR); Pandanaceae:?Pandanus spp. (FR).
Depending on the supply of fruit, colonies move from one site to another.
Population biology: Seehausen (1990) recorded births between June and August, between the rains. Assuming a gestation of 4–6 months, copulation probably takes place between January and April.
Occurrence in protected areas: Is said to occur in Ngezi Forest, which has the status of a forest reserve, but there is no legislation restricting hunting in that area. The forest is estimated at 5–12 sq km, probably nearer the former (Pakenham, 1984).
Summary of threats:
Recommended action:
Principal authors for this species: R. H. W. Pakenham, O. Seehausen.
Priority Grade: 10 (No Data).
Distribution: Solomon Islands (Fauro, Guadalcanal, Kolombangara, New Georgia, Nggela Sule, Russell Islands [Mbanika, Pavuvu]).
Summary of threats:
Recommended action:
Genus Rousettus (3 subgenera)
Subgenus Boneia (1 species)
Priority Grade: 8 (No Data: Limited Distribution).
The genus Boneia was recently synonymized with Rousettus by Bergmans and Rozendaal (1988). K. Koopman (pers. comm.) believes it should be included as a subgenus of Rousettus, while Hill and Corbet (1991) consider Boneia to be a separate genus.
Distribution: Gorontalo to Menado in north Sulawesi.
Status: Hunted for food in north Sulawesi (Bergmans and Rozendaal, 1988).
Ecology: Tambun, from where some specimens are said to have originated, is located in an area with a primary forest/farm bush mosaic. Boneia is a cave-dweller. Tambun area lies at an altitude of 200–500 m; the type specimen of Boneia menadensis was collected at an elevation of about 1000 m. From this information it may be expected to be a lowland forest bat, possibly restricted in its distribution by the presence of suitable caves (or substitutes), and foraging in partly cultivated areas as well (Bergmans and Rozendaal, 1988).
Population biology: Two females collected in June and July were pregnant (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Subgenus Lissonycteris (1 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
R. (L.) a. angolensis
Angolan soft-furred fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: North-west Angola; central and southern Cameroun; Central African Republic; central Congo; Equatorial Guinea (Bioko, Mbini); Gabon; eastern Nigeria; north-west and south-west Zaire.
Status: Unknown. Possibly quite common, but not known from many localities. In Nigeria, very localized in rocky habitats in the rain forest zone and relict forests in savannah (Happold, 1987). Happold (1987) thought it was probably uncommon. Threatened by deforestation. Locally, hunting may threaten populations (as in Congo; Bergmans, 1979).
Ecology: Occurs in forest habitats.
In Nigeria, habitually roosts in caves. Because it does not echolocate, it is found near cave entrances, which are not in total darkness and where it can use its eyes for orientation (Happold, 1987).
Eisentraut (1963) mentioned wild Ficus fruits (Moraceae) as a favourite food. Fedden and Macleod (1986) collected many specimens covered in sticky fig seeds and observed that much of the guano at roost sites also contained fig seeds.
Population biology: Data from Nigeria/Cameroun (Eisentraut, 1963; Fedden and Macleod, 1986) and from Congo (Bergmans, 1979) may indicate either continuous breeding with certain peaks connected with the rains, or two more or less distinct peaks separated by non-breeding periods. Eisentraut (1963) observed that breeding females form nurseries that tend to be avoided by adult males.
Summary of threats:
Recommended action:
R. (L.) a. ruwenzorii
Eastern soft-furred fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: Ethiopia; Kenya; Mozambique; Rwanda; southern Sudan; Tanzania; Uganda; north-east and south-east Zaire; Zambia; Zimbabwe. The assignment of populations beyond the Ruwenzori range is largely conjectural.
Status: Not known. Apparently quite common in eastern Zaire near the western Rift and in adjoining Uganda, but rather patchily distributed elsewhere. Threatened by deforestation. Locally, hunting may pose a threat.
Ecology: Gregarious, occurring in small colonies of up to about six together, and in the case of specimens from Umtali in Zimbabwe, associated with far larger numbers of Peter's epauletted fruit bat (Epomophorus gambianus crypturus), feeding on gauvas (Myrtaceae: Psidium guajava) in an orchard (Skinner and Smithers, 1990). In other parts of eastern Zimbabwe, eight were netted together in a forest clearing (Skinner and Smithers, 1990).
In Umtali in Zimbabwe, observed to eat figs (Moraceae: Ficus spp.), and orchard fruits such as guavas and mangoes (Anacardiaceae: Mangifera indica) (Skinner and Smithers, 1990).
Population biology: Ansell (1974) collected a pregnant female in Zambia in July. Data from museum specimens show that pregnant females were collected in Kenya in December and June, and in Uganda in March and December. Lactating females were collected in Uganda in March, April, June and December.
Occurrence in protected areas:
Kenya
?Mt Elgon National Park
Zaire
Parc National de l'Upemba
Summary of threats:
Recommended action:
R. (L.) a. smithii
Little soft-furred fruit bat
Priority Grade: 11 (Not Threatened).
Distribution: Burkina Faso; Ghana; Guinea; Guinea-Bissau; Ivory Coast; Liberia; western Nigeria; Senegal; Sierra Leone; Togo.
Status: See R. (L.) a. angolensis.
Ecology: Wolton et al. (1982) listed the fruits of Solanum torvum and S. erianthum (Solanaceae) as the favourite food at Mount Nimba between July and September. Dobat and Peikert-Holle (1985) listed Spathodea campanulata (Bignoniaceae), Ceiba pentandra (Bombacaceae) and Parkia roxburghii (Leguminosae) as trees whose flowers attract or may attract this species. In Liberia a specimen was netted in a banana (Musaceae: Musa spp.) plantation. Fujita and Tuttle (1991) also mentioned records of R. (L.) angolensis (subspecies unspecified but locality given as ‘West Africa’) feeding on the fruits of Adenia cissampeloides (Passifloraceae) and Solanum verbascifolium (Solanaceae). Thomas (1982) also mentions Chlorophora and Ficus (Moraceae) and Anthocleista (Loganiaceae) as food plants.
Population biology: Wolton et al. (1982), summarizing their own and other information, suggested an extended breeding season, beginning in July, peaking in September, and tailing off in the latter half of the dry season (March/April). They also found indications that adult males migrate away from the breeding area, resulting in all female nursery colonies (apart from juvenile males), as had also been found by Eisentraut (1963) in the Mount Cameroun region.
Occurrence in protected areas:
Ghana
Mole National Park
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Subgenus Rousettus (8 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 4.
R. (R.) a. aegyptiacus
Egyptian rousette
Priority Grade: 11 (Not Threatened). Distribution: Cyprus; Egypt; Israel; Lebanon; Syria; Turkey.
Status: Cyprus Boye et al. (1990) reported that specimens were recorded near Paphos, Episkopi and Kyrenia, north of Limassol, west of Nicosia and at Famagusta. Most observations concerned 1–3 animals and there were no colonies numbering more than 20 bats. In a cave in Limassol district, a colony of 300–400 was reported a few years ago by local people, but recently only one bat has been seen there (Boye et al., 1990). It seems likely that there has been a significant decline in population numbers, probably due to shooting. Spitzenberger (1979) found that the Ministry of Agriculture was responsible for a yearly decimation campaign, during which bats were disturbed at their roosts and subsequently shot. She observed that of a colony near Apsiou consisting of about 400–500 bats, about 200 were killed during one shooting session, but believed the colony would be able to recover from such tolls. These campaigns are no longer organized by local authorities but may still be practised privately by farmers. This is indicated by the presence of a number of abandoned roosts, which show marks of shot and soot (Boye et al., 1990).
On the whole probably not endangered, although regarded as a pest in Turkey (Spitzenberger, 1979), Israel (Spitzenberger, 1979; Makin and Mendelsson, 1986) and Egypt (K. Wassif, pers. comm.) and treated accordingly. Local abundance may vary with that of food (Lewis and Harrison, 1962).
Fruit farmers are the most important threat to populations. In Turkey and Israel, Rousettus caves have been fumigated or closed off by walls (Spitzenberger, 1979). In Israel, this species is listed as ‘noxious’ under wildlife laws and caves have been fumigated with Lindane (= gamma HCH) (Makin and Mendelsson, 1987) including some inside nature reserves, but recently attitudes towards bats have improved (Makin and Mendelssohn, 1987; Sowler, 1988a, 1988b). In Egypt, it is also destroyed by farmers (K. Wassif, pers. comm.), but no details are known.
A new threat to bats in Israel may arise through problems with bat droppings accumulating on the walls of buildings (D. Makin, pers. comm.).
Ecology: In Cyprus in the winter, roosts in caves, and in March some colonies move to open sites, such as tall trees or large buildings (Boye et al., 1990). In Israel has been found roosting in caves and deserted buildings (Makin, 1990).
Feeds on the following plants (Unger and Kotschy, 1865; Bate, 1903; Lewis and Harrison, 1962; Kaisila, 1966; Spitzenberger, 1979; Dobat and Peikert-Holle, 1985; Makin and Mendelsson, 1986): Bombacaceae: Bombax ceiba (FL); Leguminosae: Ceratonia siliqua (FR); Moraceae: Ficus religiosa (FR, L), F. sycamorus (FR), Morus alba (S); Musaceae: Musa sp. (FL); Palmae: Phoenix sp. (FR); Rutaceae: Citrus spp. (FR).
In Israel, bats show a high locality fidelity both in roosting and foraging. Migration from the north or within Israel has not been demonstrated (Makin, 1990).
In Israel, the causes of mortality are many. A large number of predators prey on Rousettus, cats being the most common (Makin, 1990). It appears that extreme winter conditions cause many casualties (Makin, 1990) and there is the added factor of persecution by fruit growers (see above).
Population biology: Spitzenberger (1979) found two breeding cycles per year in a Cyprus population. These cycles overlap: about half of adult females were about to give birth by the end of March or beginning of April although some had given birth already, while the other half and all subadult females each bore a small embryo (up to a fortnight old in a gestation of about 4 months). As a rule, a particular female would take part in only one cycle per year. Harrison (1964) observed suckling infants as well as half-grown young in Carmel Caves in Israel in October. Makin (1990) observed two main birth peaks in Israel, one in the first two weeks of April and the second in late August and early September. In June, most (67%) of the parous females were pregnant and lactating, indicating that there was a post-partum oestrus (Makin, 1990). Makin (1990) estimated that some females first conceived at the age of 7–8 months and others did so only at 15–16 months. Males born in the April season matured at the age of 18 months, those born in August–September matured earlier at the age of 14–18 months. Gaisler et al. (1972) collected 11 females, 8 of which carried one embryo each, and 8 males, 5 of which were in breeding condition, in April in Cairo, Egypt. According to Kingdon (1974), Rousettus would breed throughout the year in Egypt.
Occurrence in protected areas:
Israel
Occurs in all nature reserves (D. Makin, pers. comm.) including:
Beitan Aharon Nature Reserve
Carmel Nature Reserve
Summary of threats:
Recommended action:
R. (R.) a. arabicus
Arabian rousette
Priority Grade: 6 (Rare).
Distribution:?Eastern Ethiopia; Iran; Oman; Pakistan,; Saudi Arabia; Yemen.
Status: Not common, and apparently rare over a large part of its range.
Ethiopia Known only from a single record (Hayman and Hill, 1971).
Iran Could be abundant in certain caves on Qeshm Island (Blanford, 1876) but it is certainly not very numerous in the very few other localities where it has been found (DeBlase, 1980).
Oman Known from some localities in the north-east (e.g., Muscat; Harrison, 1964) and from several in Dhofar in the south-west, where it does not seem particularly rare (records in Harrison, 1980).
Pakistan Rare and locally distributed (Roberts, 1977).
Saudi Arabia Known from two localities (Yaman, 1966).
Yemen Known from three localities; Yerbury and Thomas (1895) wrote that it occurred ‘in great numbers in a cave on the banks of the Wady Jughur near Lahij’.
Threats would come from fruit growers. In Iran, DeBlase (1980) found that farmers were not aware of a fruit-eating bat. Roberts (1977) reported that in the fruit gardens of Malir, Pakistan, these bats caused ‘some damage’ but he did not mention whether they were being destroyed. From where it appears to be most numerous, in Oman, nothing has been reported on possible conflicts with man.
Ecology: Feeds on the following plants (Brosset, 1966a; Roberts, 1977; Nader, 1985a): Anacardiaceae: Mangifera indica (FL); Bombacaceae: Bombax malabarica (FL); Caricaceae: Carica papaya (FR); Meliaceae: Azadirachta indica (FR), Melia azerdarach (FR); Moraceae: Ficus religiosa (FR); Musaceae: Musa spp.; Myrtaceae: Psidium guajava (FR); Palmae: Phoenix dactylifera (FR); Rutaceae: Citrus spp. (FR).
Population biology Yerbury and Thomas (1895) found a female with a suckling infant near Lahij, Yemen, in March. In Pakistan the majority of young are born in the early spring with a few at the end of the monsoon season in September (Roberts, 1977).
Summary of threats:
Recommended action:
R. (R.) a. leachii
Cape rousette
Priority Grade: 11 (Not Threatened).
Distribution: Burundi; Ethiopia; Kenya; Malawi; Mozambique; Rwanda; South Africa; southern Sudan; Tanzania (mainland, Mafia, Pemba, Zanzibar); Uganda; north-east and south-east Zaire; Zambia; Zimbabwe.
Status: Locally abundant and not endangered.
Cave habitats are under pressure from tourists, particularly in Kenya (McWilliam, 1980a, 1980b). Large numbers are collected and used in research (e.g., Baranga, 1978, 1980). These bats are also killed in some areas because of orchard depredations.
Ecology: Highly gregarious, occurring in caves, buildings and tombs in very large numbers (up to several thousand). Highly adaptable and will even colonize newly built buildings in the middle of busy cities, feeding on the fruit of Ficus sp. (Moraceae). A primitive form of echolocatory click is used to navigate to roosting sites in dark caves/buildings.
Feeds on the following plants (Andersen, 1912; Cunningham van Someren, 1972a, 1972b; Start, 1972; Kingdon, 1974; Jacobsen and Du Plessis, 1976; Herzig-Straschil and Robinson, 1978; Thomas and Fenton, 1978; Dobat and Peikert-Holle, 1985; Herselman and Norton, 1985): Aloeaceae: Aloe dolmitica (FL); Anacardiaceae: Harpephyllum caffrum (FR), Mangifera indica (FR); Apocynaceae: Acokanthera oppositifolia (FR); Bignoniaceae: Kigelia pinnata (FL); Bombacaceae: Adansonia digitata (FL); Celastraceae: Cassine crocea (FR); Ebenaceae: Diospyros capensis (FR); Leguminosae: Erythrina abyssinica (L); Meliaceae: Ekebergia capensis (FR); Moraceae: Ficus capensis (FR), F. petersii (FR), F. sansibarica (FR); Myrtaceae: Syzygium cordatum (FR), S. gerrardii (FR), S. jamba (FR); Oleaceae:?Olea africana (FR), ?O. capensis (FR); Podocarpaceae: Podocarpus olinia (FR); Rosaceae: Eriobotrya japonica (FR), Malus sylvestris (FR), Prunus africana (FR), P. armeniaca (FR), P. persica (FR); Salvadoraceae: Salvadora persica (FL); Sapindaceae: Litchi chinensis (FR); Viscaceae: Viscum obscurum (FR).
Population biology: The breeding patterns change according to locality. From the northern part of its distribution, few data are known. In Ethiopia, a pregnant female was caught in February at Baro (60 km east of Gambela) and a lactating female in late April/May at Lake Abaya. In Sudan a pregnant female was caught on 1 July in Talanga Forest. In Uganda has been reported as bimodally polyoestrus with conceptions in December and June and births in March and September, occurring just before the peak of the two rainy seasons (Mutere, 1968). In south-east Zaire Anciaux de Faveaux (1978) concluded that reproduction was continuous. In the eastern Transvaal, South Africa, Jacobsen and Du Plessis (1976) reported copulation in June to mid-September and births from October to December, with a possible second minor birth peak in March/April. Herzig-Straschil and Robinson (1978), working in southern Cape Province, South Africa, observed births from October until June. The observation by Herselman and Norton (1985) of an infant in early January in this province confirms this.
Occurrence in protected areas:
Kenya
Amboseli Game Reserve
Marsabit National Reserve
Masai Mara Game Reserve
Meru Game Reserve
Mount Elgon National Park
Lake Nakuru National Park
Shimba Hills National Reserve
Tsavo National Park
South Africa
Transvaal
Kruger National Park
Wolkberg Wilderness Area (Rautenbach, 1982)
Natal
Harold Johnson Nature Reserve
Kosi Bay Nature Reserve
Ndumu Game Reserve
Oribi Gorge Nature Reserve
St Lucia Eastern Shores Nature Reserve
Umlalazi Nature Reserve
Cape
Keurbooms Reserve
Table Mountain Nature Reserve (Herselman and Norton, 1985)
Tsitsikama National Parks
Tanzania
Arusha National Park
Ruaha National Park (Kingdon, 1974)
Serengeti National Park
Uganda
Murchison National Park
Queen Elizabeth National Park
Zimbabwe
Chimanimani National Park
Great Zimbabwe Ruins (Smithers and Wilson, 1979)
Inyanga National Park
Summary of threats:
Recommended action:
R. (R.) a. unicolor
West African rousette
Priority Grade: 11 (Not Threatened).
Distribution: Angola; Cameroun; Congo; Equatorial Guinea (Bioko, Mbini); Gabon; Gambia; Ghana; Guinea; Ivory Coast; Liberia; Nigeria; São Tomé and Príncipe (São Tomé); Senegal; Sierra Leone; Togo; north-west and south-west Zaire.
Status: Locally abundant and not endangered. It could be threatened by deforestation and possibly locally by hunting.
Ecology: Recorded in both forests and savannahs. Hill and Carter (1941) quoted a field label of a specimen from Angola ‘Shot flying to banana flowers [ Musaceae: Musa spp.], not the fruit, which was still green’. Eisentraut (1963) collected a large series on Mount Cameroun chiefly in mist-nets set under or in wild fig trees [ Moraceae: Ficus spp.] in fruit. Rosevear (1965) reported that ‘cultivated fruit trees, from dates [ Palmae: Phoenix dactylifera] in the subdesert to guavas [ Myrtaceae: Psidium guajava] and mangoes [ Anacardiaceae: Mangifera indica] in the forest, have been recorded as attacked by this species… Wild figs are also a source of food.’ Brosset (1966a) wrote that it eats all kind of wild and cultivated fruits, including some that are inedible or even toxic to man. Wolton et al. (1982) observed this bat feeding extensively on the fruits of Solanum torvum (Solanaceae) and S. erianthum on Mount Nimba between July and September.
Population biology: Wolton et al. (1982) considered that on Mount Nimba the breeding season may be extended with a pregnancy peak between November and December at the end of the rains.
Eisentraut (1963) concluded that in the Mount Cameroun region the breeding period was not strictly seasonal but that it might be restricted to the dry season.
Occurrence in protected areas:
Ghana
Mole National Park
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
R. (R.) a. amplexicaudatus
Geoffroy's rousette, Common rousette
Priority Grade: 11 (Not Threatened).
Distribution: Alor; Ambon; Bagabag; Borneo (Brunei, West Kalimantan, Sabah, Sarawak); Burma; Cambodia; Enggano; Halmahera; Kisar; Lombok; Mentawai Islands; Ndao; New Guinea; Peleng; Peninsular Malaysia (including island of Langkawi); Philippines (Balabac, Caluya, Carabao, Cebu, Dinagat, Guimaras, Jolo, Lubang, Luzon, Mindanao, Mindoro, Negros, Palawan, Panay, Polillo, Samal, Samar, Siargao, Simunul, Sulu Archipelago [Bongao, Sanga Sanga, Sibutu, Tawitawi]; Rote; Savu; Seram; Sulawesi; Sumba; Ternate; Thailand; Timor; Yapen.
Rousettus amplexicaudatus amplexicaudatus (Photo by P. A. Morris)
Status: Borneo Payne et al. (1985) listed records: from Pulau Balembangan, Sukau, Madai Caves and Tawau in Sabah; Tasek Merimbun in Brunei; Sungai Baram and Niah Caves in Sarawak; Sungai Landak in West Kalimantan.
Lombok Kitchener et al. (1990) reported that it was moderately abundant in all habitats from sea-level to 400 m.
New Guinea Common (Flannery, 1990). Widespread on mainland New Guinea; Flannery (1990) did not record it above 300 m on the mainland, although Ziegler (1982) recorded it as possibly only below 600 m. Has been recorded as far east as Milne Bay Province. In Irian Jaya its western extent is not clear, although it is present in the Moluccas and could be expected to occur commonly as far west as the Vogelkop (Flannery, 1990).
Peninsular Malaysia Medway (1978) recorded it as known from scattered localities at all elevations, including roosts in Batu Caves, Selangor, and on Gunung Brinchang, Patang. It was also recorded on Langkawi. He considered it to be uncommon.
Philippines Rare in submontane forest on Negros, where it made up about 2% of captures (Heideman and Heaney, 1989). Captured three times more frequently in clearings and secondary growth than in forest. In contrast, 40–90% of captures in orchards and other agricultural habitats were of this species (Guerrero and Alcala, 1973; Heaney et al., 1989; unpubl. data). Apparently prefers disturbed habitats and areas with cultivated fruit trees (banana (Musaceae: Musa sp.), mango (Anacardiaceae: Mangifera indica), chico (Sapotaceae: Pouteria sapota), guava (Myrtaceae: Psidium guajava), and cultivated figs (Moraceae: Ficus spp.). Although successful in association with agriculture, populations on Negros and other areas of the Philippines are apparently declining. Caves that had held large numbers in 1981 contained very few batsin 1987; in one cave a large pile of dried wings cut from these and other species was found. Can be a pest in fruit orchards (Guerrero and Alcala, 1973), although mostly ripe or over-ripe fruit is taken and the fruits that are commercially most important are usually harvested while unripe. Where cave colonies have been destroyed because of supposed damage to commercial fruit crops, the small economic benefits of such measures are negated by the loss of associated Eonycteris spelaea as well as many species of insectivorous bats, all of which are beneficial.
Although populations in some areas of the Philippines appear to be declining, it remains abundant. Given the expansion of disturbed habitats and orchards, there are now probably as many or more individuals in the Philippines as there were several hundred years ago.
Sulawesi Hunted for food in north Sulawesi (Bergmans and Rozendaal, 1988).
Thailand Yenbutra and Felten (1986) listed records from the provinces of Chiang Mai, Chon Buri, Chanthaburi, Lop Buri, Saraburi, Phetchaburi and Ranong.
Timor Goodwin (1979) only found this species roosting in caves, concentrated in chambers close to entrances, with up to 800 individuals in a colony.
Ecology: Roosts in caves, and can echolocate in a rudimentary manner. One female netted at Yapsiei in 1986 was carrying a ripe fig (Moraceae: Ficus sp.) in its mouth. Often a lactating female and a subadult animal were caught close together in the same net, suggesting a close association between mother and quite advanced offspring (Flannery, 1990). Flannery (1990) netted bats most often in open areas with little tall or overhanging vegetation. In the Philippines it fed on a number of cultivated fruit trees. Presumably also feeds on the fruits of wild plants, including secondary growth species.
On Timor, Goodwin (1979) observed bats feeding on Muntingia fruit (Flacourtiaceae), with as many as 50 individuals feeding on a single tree at the one time.
May travel long distances to forage; Lekagul and McNeely (1977) collected evidence to suggest that it may commute 25 km a night to feed. At orchard sites in Dumaguete City in the Philippines, Rousettus did not arrive until one or more hours after dark, implying similarly long commuting distances (authors' pers. obs.).
In the Philippines, individuals fed almost silently in fruit trees. In orchards, they often remained in the fruit trees to consume fruit, perhaps because most of the cultivated fruits were too large to carry to a feeding roost. Heideman and Heaney (1989) estimated that at least 525, and possibly several thousand, individuals were feeding at least occasionally in an orchard of several hundred fruit trees on Negros, but there were many fewer there on any one night, and these individuals were probably foraging over a very wide area.
Population biology: In the Philippines, females probably produce two young per year, one in each of two birth periods separated by 4–5 months. There is some variation in timing of births among islands (Heideman, 1987). The duration of gestation is about 3.5–4.5 months; that of lactation approximately 2.5–3 months.
Young females produce their first young at an age of 8–12 months, females generally becoming pregnant at 6–9 months. Although males on Negros in the Philippines had moderately large and apparently functional testes at an age of 1 year, these animals weighed 30–50% less than older males and lacked their extensive orange-red throat ruff. On Negros, most or all of the females produced only one young in their first year (Heideman, 1987). Start (1974) reported that bats in Peninsular Malaysia were in reproductive condition all year, but he seems to have used a very broad definition of ‘breeding’ that implies either a synchronous or seasonally polyoestrus pattern or an asynchronous and aseasonal polyoestry. Lim (1973) reported pregnant females only from December through to June in a different Peninsular Malaysian population, suggesting a seasonal reproductive pattern. Kitchener et al. (1990) reported that in October 1987 on Lombok, it appeared that the birth season had recently finished. None of the nine females caught was pregnant, but seven were lactating. In May 1988 a single female caught on Lombok was lactating. A juvenile female was collected at the same time. On Timor Goodwin (1979) reported that most of the adult specimens collected in March, April and May were in breeding condition with many of the females pregnant, recently parturient, or lactating. On Sulawesi, two females caught in March were pregnant and an adult female caught in early December was apparently lactating at the time of capture (Bergmans and Rozendaal, 1988).
In New Guinea, out of seven females caught at Yapsiei, West Sepik Province in January 1984, four were in an advanced stage of pregnancy and the remainder were subadult (Flannery, 1990). On a return visit in late April 1986, a further 16 animals were taken. Only three of these were males, two definitely subadult. Of the 13 females, six were lactating, and four were subadult. These data suggest highly seasonal breeding in West Sepik Province, and a perplexing absence of adult males from the area after the birth of the young (Flannery, 1990). Gestation in South East Asia is said to last 15 weeks, and the young feed on solid food by the age of 3 months.
Occurrence in protected areas:
Seram
Manusela National Park
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Principal authors for this subspecies: T. Flannery, P. D. Heideman, L. R. Heaney.
R. (R.) a. brachyotis
Priority Grade: 11 (Not Threatened).
Distribution: Bismarck Archipelago (Duke of York, Emirau, New Britain, Tabar); Bougainville;?Halmahera; Solomon Islands (Choiseul, Fauro, Guadalcanal, Kolombangara, Malaita, Santa Isabel, Vella Lavella).
Population biology: On Bougainville, McKean (1972) reported pregnant females in July and September and a lactating female in September.
Summary of threats:
Recommended action:
R. (R.) a. infumatus
Priority Grade: 11 (Not Threatened).
Distribution: Bali; Flores; Jawa; Krakatau Islands (Lang); Penida; Sumatra.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: Sangir Islands (Sangir, Siau, Tahulandang); Sulawesi.
Status: Sulawesi Bergmans and Rozendaal (1988) commented that in the Tangkoko-Batuangus Nature Reserve, it was, along with Cynopterus brachyotis, the most numerous fruit bat mist-netted along the forest edge and in coastal bush. Within primary forest it was netted in smaller numbers. In this reserve R. (R.) celebensis was observed to roost in numbers in a shallow, coastal cave close to the tide line at Tanjung Mandera. Hunted for food in north Sulawesi.
Population biology: An adult female caught by Bergmans and Rozendaal (1988) in March 1985 in Sungei Tumpah (Dumoga-Bone National Park) gave birth to a single young while in captivity. In April 1983 a female nursing a young bat was observed in a roost in a shallow cave at Tanjung Mandera (Bergmans and Rozendaal, 1988). In lowland forest in the Tangkoko-Batuangus reserve two females carrying young were caught in April.
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Tangkok-Batuangus reserve
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened)
Number of subspecies: 2. The two currently recognized subspecies are separated by differences in dentition and fur colour. As many more localities have become known, and as these, because of the montane character of the species, tend to be isolated from one another, intraspecific variation is to be expected for other populations. Subspecific divisions can be assessed only by reviewing of all known material.
R. (R.) l. kempi
Priority Grade: 11 (Not Threatened).
Distribution: Eastern Ethiopia; Kenya; south-eastern Sudan; Tanzania.
Status: Unknown. Locally abundant but few observations.
Granvik (1924) related how on and around Mount Elgon in Kenya in 1920 a local wandering tribe, the Wanderoboos, ‘when sojourning for a longer or shorter time in the caves, live to a great extent on these fruit-bats.’ Allen and Loveridge (1927) reported that bats were also hunted and eaten by local people in the Uluguru Mountains in Tanzania. Locally, the easy hunting of cave-inhabiting fruit bats will certainly still occur and may pose a threat to populations.
Furthermore, the destruction of relict patches of montane forest will have an adverse effect. It is not known whether caves harbouring this bat are among those known to be affected by tourist pressures.
Ecology: Roosts in damp, dark caves in montane forest, where it assembles in hundreds and occasionally shares roosts with Rousettus (Rousettus) aegyptiacus and/or Rousettus (Lissonycteris) angolensis. Its long hair with fine woolly underfur is probably an adaptation to the cold regions in which it lives.
Occurrence in protected areas:
Kenya
Mount Elgon National Park
Summary of threats:
Recommended action:
R. (R.) l. lanosus
Priority Grade: 11 (Not Threatened).
Distribution: Burundi; Rwanda; Uganda; north-east and south-east Zaire.
Status: Unknown. Locally abundant, but few observations.
It presumably would be threatened by the same problems as those faced by R. (R.) l. kempi.
Ecology: In Rwanda seen feeding at the flowers of Lobelia (Campanulaceae) (J. R. Wilson, pers. comm.). At Fort Portal, in Uganda, a specimen was collected in a banana plantation (Musaceae: Musa sp.).
Occurrence in protected areas:
Rwanda
Parc National de Volcans
Uganda
Bwindi (Impenetrable) Forest Reserve
Queen Elizabeth National Park
Zaire
Parc National de Virunga
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
Number of subspecies: 3.
R. (R.) l. leschenaulti
Priority Grade: 11 (Not Threatened).
Distribution: Bangladesh; Burma; Cambodia; south China; Hong Kong; India; Laos; Nepal; Pakistan; Thailand; Vietnam.
Status: Bangladesh Uncommon, but found over a wide area. A large colony is known from Kudum Cave, near Whykeong (Khan, 1985).
Thailand Lekagul and McNeely (1977) commented that it was found throughout Thailand and was especially numerous in the fruit-growing areas of Nonburi and Chonburi. This is supported by Yenbutra and Felten (1986) who listed records from the provinces of Chaiyaphum, Chanthaburi, Chiang Mai, Chiang Rai, Chumphon, Kanchanaburi, Loei, Lop Buri, Nakhon Ratchasima, Nakhon Sawan, Nan, Phattalung, Phetchaburi, Prachin Buri, Prachuap Khiri Khan, Ranong, Saraburi, and Songkhla.
Vietnam Van Peenen (1969) listed records from the following localities in southern Vietnam: Phuochai, Mt Sontra and Fyan.
In Nepal this species is given partial protection under wildlife regulations.
Nothing is known of possible threats, although as a cave-dweller it would presumably face the same pressures as other cave species.
Ecology: In Bangladesh, distribution is limited to forested areas containing dilapidated buildings or ruins. Prefers to roost in dark places (Khan, 1985). In India, it roosts in colonies of 10–2000 in caves and man-made structures such as tunnels, rock-cut caves, wells and rooms in old ruins, not necessarily dark. There is movement between different roosting sites depending on the availability of food. There is usually no sexual segregation in the colonies; however, separate roosts of the sexes have been reported. Juveniles, once independent of their mothers, live in exclusive colonies (Prater, 1971).
Fujita and Tuttle (1991) listed the following food plants: Anacardiaceae: Anacardium occidentale (FR), Mangifera indica (FL); Annonaceae: Annonareticulata (FR), A. squamosa (FR); Bignoniaceae: Heterophragma roxberghii (FL), Oroxylum indicum (FL), Radermachera xylocarpum (FL); Bombacaceae: Adansonia digitata (FL), Ceiba pentandra (FL); Juglandaceae: Carya arborea (FL); Myrtaceae: Psidium guajava (FR,FL), Syzygium cumini (FR,FL); Sapotaceae: Madhuca indica (FL), Mimusops hexandra (FR,FL).
Population biology: Prater (1971) commented that in India the season of copulation is probably between November and March. Two discrete birth seasons, one in March and the second in August are reported. Gopalakrishna and Choudari (1977) found that pregnancies lasted from November to March and from March to July, with females having a post-partum oestrus. A single young is born to each female. The young are carried by their mothers for 2 months and adult size is reached after 1 year. Females are sexually mature at 5 months, while males do not reach sexual maturity until 15 months.
Summary of threats:
Recommended action:
R. (R.) l. seminudus
Priority Grade: 11 (Not Threatened).
Distribution: Sri Lanka.
Status: Colonies scattered sparsely over the greater part of the lowlands and lower hills, but never reported from Jaffna or northern districts (Phillips, 1980).
This species is fully protected under local wildlife laws.
Ecology: Roosts in the darkest recesses of large caves. Males sometimes live separately (Phillips, 1980).
Feeds on the fruit and flowers of guava (Myrtaceae: Psidium guajava), plantains (Musaceae: Musa sp.), loquats (Rosaceae: Eriobotrya japonica), soursops (Annonaceae: Annona muricata) and the kapok tree (Bombacaceae: Ceiba pentandra) (Phillips, 1980).
Population biology: Copulates in roosting caves. Gestation is 15 weeks (Phillips, 1980).
Summary of threats:
Recommended action:
R. (R.) l. shortridgei
Priority Grade: 10 (No Data).
Distribution:?Aru Islands; Bali; Jawa; Lombok; Simalue Islands; Sumatra.
Population biology: On Lombok, Kitchener et al. (1990) reported that two females caught in May 1988 were both pregnant and lactating.
Summary of threats:
Recommended action:
Priority Grade: 11 (Not Threatened).
Distribution: Madagascar.
Status: At least locally common (M. Nicoll, pers. comm.). Found in all lowland areas with (preferably moist) forest. It is very probably restricted by its need for cave habitats (Bergmans, 1977).
Threatened by deforestation, hunting and poor implementation of protection of reserves such as Ankarana (Wilson, 1987).
Ecology: A specimen was collected ‘near a banana plant’ (Musaceae: Musa sp.).
Occurrence in protected areas:
Madagascar
Ankarana Special Reserve
Summary of threats:
Recommended action:
Principal authors for this species : W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
Distribution: Comoros (Anjouan, Grande Comore, Moheli).
Status: Apparently common. The Belgian Zoological Missions in 1981 and 1983 collected 64 specimens ‘from the many that were caught’, and it was a regular catch in the mist-nets set up in the forests (Meirte, 1984b). Carroll (1985) quoted M. Louette, who reported that it was ‘quite common, at least on the three westernmost Comoros’. During a trip to the Comoros in late 1989, D. Meitre (pers. comm.) found large numbers on Grande Comore and Moheli.
Threatened by deforestation. The planting of bananas in the lower, deforested regions, where bats apparently forage in important numbers, results in enormous soil erosion. To compensate for this other areas are then deforested (D. Meitre, pers. comm.). Because the majority of the people of the Comoros are Muslims, bats are probably not threatened by hunting. Possible damage to fruit crops may lead to destruction of colonies in caves, but it is not known whether such damage occurs or whether bats are being destroyed. The Belgian expeditions of 1981 and 1983 (Meitre, 1984b) did not find where bats roost by day, which may indicate that they have retreated to higher altitudes.
Ecology: D. Meitre (pers. comm.) considered that bananas (Musaceae: Musa spp.) are probably the most important food.
Population biology: A female collected in October on Anjouan had one embryo.
Occurrence in protected areas: There are no protected areas in the Comoros.
Summary of threats:
Recommended action:
Priority Grade: 10 (No Data).
Distribution: Western Borneo (Sabah, Sarawak); northern Sumatra.
Status: A roost of about 300 individuals was found in a cave at Batu Timbang in central Sabah. Single individuals have been found in mixed roosts with R. (R.) amplexicaudatus at Niah Caves in Sarawak and along the Kinabatangan River in Sabah (Francis, 1989).
Occurrence in protected areas:
Borneo
Sarawak
Niah Caves
Summary of threats:
Recommended action:
Principal author for this species: C. M. Francis.
Genus Scotonycteris (2 species)
Priority Grade: 11 (Not Threatened).
Distribution: Cameroun; Congo; Ghana; Liberia.
Status: Known from a few scattered localities. As a forest dweller, it would be threatened by deforestation.
Ecology: A forest dweller. Eisentraut (1960) netted a specimen in a Ficus (Moraceae) tree, and found yellowish grains, which were probably fig seeds, in the stomach of another.
Population biology: Hayman (1946) described a female with a very young offspring collected at Oda, Ghana in December. Wolton et al. (1982) caught two pregnant females in August and one in September at Mount Nimba.
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Priority Grade: 11 (Not Threatened).
The subspecies S. z. occidentalis was described on the basis of material from eastern Zaire, but most authors consider this as a synonym of the typical form.
Distribution: Cameroun; Central African Republic; Congo; Equatorial Guinea (Bioko, Mbini); Gabon; Ghana; Ivory Coast; Liberia; Nigeria; north-east and south-east Zaire.
Status: Unknown. As a forest dweller, it would be threatened by deforestation.
Occurrence in protected areas:
Ivory Coast
Tai National Park (Roth and Merz, 1988)
Summary of threats:
Recommended action:
Principal authors for this species: W. Bergmans, S. Sowler.
Genus Sphaerias (1 species)
Priority Grade: 10 (No Data).
Distribution: Bhutan; Burma; south-west China; north India; north Thailand; south-west Tibet.
Summary of threats:
Recommended action:
Genus Styloctenium (1 species)
Priority Grade: 10 (No Data).
Distribution: Sulawesi.
Status: Hunted for food in north Sulawesi (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Dumoga-Bone National Park
Summary of threats:
Recommended action:
Genus Syconycteris (3 species)
Priority Grade: 11 (Not Threatened).
Number of subspecies: 7.
S. a. australis
Queensland blossom bat
Priority Grade: 11 (Not Threatened).
Distribution: Eastern coast of Australia (Queensland).
Status: Strahan (1983) considered it common in limited habitat.
Ecology: The protection offered by dense foliage in rain forest and adjacent wet sclerophyll forest is used for daytime roosting, but it has also been found in mango trees (Anacardiaceae: Mangifera indica) (Strahan, 1983).
Has been observed hovering in front of, or feeding on, the blossoms of paperbarks, bottlebrushes (Myrtaceae: Callistemon spp., Melaleuca spp.) banksias (Proteaceae: Banksia spp.), bloodwoods (Myrtaceae: Eucalyptus spp.) and cultivated bananas (Musaceae: Musa spp.) (Strahan, 1983). This small bat is highly specialized for a diet of nectar with a very pointed muzzle and a long, thin tongue. The tongue has minute, brush-like projections along the mid-line for gathering nectar and pollen and there is no doubt this species is an important pollinator.
Syconycteris australis (Photo by A. M. Hutson)
Summary of threats:
Recommended action:
S. a. crassa
Priority Grade: 10 (No Data).
Distribution: D'Entrecasteaux Islands (Fergusson Island); Trobriand Islands (Kiriwina).
Summary of threats:
Recommended action:
S. a. finschi
Priority Grade: 10 (No Data).
Distribution: Bismarck Archipelago (Admiralty Islands [Manus], New Britain, New Ireland).
Summary of threats:
Recommended action:
S. a. keyensis
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: Kai Islands.
Summary of threats:
Recommended action:
S. a. major
Priority Grade: 10 (No Data).
Distribution: Ambon; Seram. The range probably includes Buru and other small islands near Ambon.
Occurrence in protected areas:
Seram
Manusela National Park
Summary of threats:
Recommended action:
S. a. naias
Priority Grade: 8 (No Data).
Distribution: Muyua.
Summary of threats:
Recommended action:
S. a. papuana
Priority Grade: 11 (Not Threatened).
Distribution: Aru Islands; Bagabag; Crown; Long; Louisiade Archipelago (Misima Island, Rossel Island, Sudest Island); Misool; New Guinea; Sakar; Tolokiwa; Umboi.
Status: New Guinea Common (Flannery, 1990). When using mist-nets in New Guinea, it is the most commonly captured bat, being abundant at all altitudes, and above 1500 m is often the only species caught (Flannery, 1990).
Ecology: In New Guinea, found in most habitats (Flannery, 1990).
Feeds almost exclusively on nectar and pollen, and is probably an important pollinator (Richards, 1983). There is some evidence that it may become torpid at higher altitudes. In February 1984, Flannery (1990) collected some bats that had been caught in a mist-net at 2200 m on a very cold and rainy night. The animals were wet, cold and stiff, and appeared to be dead. They were placed by a fire and all recovered and became active within three-quarters of an hour.
Population biology: Reproduction appears to be aseasonal in New Guinea (McKean, 1972). A single embryo is usual (McKean, 1972; author's data).
Summary of threats:
Recommended action:
Principal author for this species: T. Flannery.
Priority Grade: 8 (No Data: Limited Distribution).
Distribution: North-west Halmahera.
Summary of threats:
Recommended action:
Priority Grade: 6 (Rare).
Distribution: New Guinea.
Status: Rare (Flannery, 1990). Reported only from high-altitude forest near the summit of Mount Kaindi, Morobe Province. In this area it is sympatric with S. australis papuana. In March 1986, a single specimen collected in a taro (Araceae: Colocasia esculenta) garden at 2300 m in the Telefomin area (West Sepik Province). Nyctimene cyclotis, Syconycteris australis and Pipistrellus collinus were taken in nets set in the same garden (Flannery, 1990).
Ecology: Presumed to feed on nectar and pollen, like S. australis papuana (Flannery, 1990).
Population biology: Of three females collected in June/July, 1967, one was possibly pregnant with an early stage embryo, while two were actively nursing or had recently ceased lactating (Ziegler, 1982).
Summary of threats:
Recommended action:
Genus Thoopterus (1 species)
Priority Grade: 10 (No Data).
Distribution: Morotai;?Philippines (Luzon); Sangir Islands; Sulawesi.
Status: Hunted for food in north Sulawesi (Bergmans and Rozendaal, 1988).
Ecology: In Sulawesi, has been collected in forest at altitudes of between about 50 m and 1800 m. At Bantimurong specimens were collected in a small patch of woodland on a narrow river plain wedged between limestone hills. It is thought that the bats roosted communally, possibly using the same roosts over long periods of time (Bergmans and Rozendaal, 1988). At Malino, specimens were collected while feeding in a fig tree (Moraceae: Ficus sp.) in a village area together with Cynopterus bbrachyotis brachyotis.
Population biology: In Sulawesi, subadult specimens were collected in October, December, January and March. Pregnant females were collected in January (Bergmans and Rozendaal, 1988).
Occurrence in protected areas:
Sulawesi
Bantimurong Protected Area
Dumoga-Bone National Park
Summary of threats:
Recommended action: