ORYX VOL 31 NO 3 JULY 1997 Flying foxes on Choiseul (Solomon Islands) the need for conservation action

E. Bowen-Jones, D. Abrutat, B. Markham and S. Bowe

The Choiseul 1995 project spent 6 months studying the Chiroptera of Choiseul, one of the least disturbed and most poorly known islands in the Solomons. Among the results were: the rediscovery of a member of the Megachiropteran genus Pteralopex, which had been thought possibly extinct; a new record of another endemic pteropodid with a limited distribution; and evidence of a decline in the 'commonest' member of the family in the archipelago. The threats, present and future, to these and the forests of the island are discussed. An overview of the status of Pteralopex spp. in the Solomons provides an insight into the need for conservation action in the country as a whole.

Introduction several areas, including Mt Maetambe, an area of cultural significance due to its height. Only Choiseul is a remote island in the Solomons several months of working in other areas and and, compared with other islands in the archi- talking with the inhabitants to build up trust pelago, is poorly known in terms of its bio- and thus be granted access to land made this geography. The fauna of the island is very possible. similar to that of neighbouring Bougainville Choiseul has been rated as being the fifth (Figure 1), which, although politically part of highest priority area in the world for fruit Papua New Guinea, is biogeographically part biodiversity and 10 taxa of Megachiroptera of the North Solomons (Flannery, 1995). have been recorded as present on the island Choiseul is approximately 120 km long and (Mickleburgh et al., 1992; Table 1). 40 km wide. The interior of the island is Of particular interest is the Bougainville mountainous, rugged and covered in rain for- monkey-faced flying fox Pteralopex anceps est, the highest point being Mt Maetambe at (Andersen, 1909), which is confined to 1060 m. Most of the island's 13,000 human in- Choiseul and Bougainville. This species of habitants live within 2 km of the coast, where Pteralopex is the largest member of a genus their coconut plantations are located and characterized by having reduced ears hidden where they were encouraged to relocate by the in the fur, and a specialist dentition composed missionaries who converted the various of multicuspate molars and bicuspate canines. groups of indigenous people. All land on the Pteralopex anceps is principally known from island is under customary ownership. specimens collected by the Reverend Poncelet To the authors' knowledge, only two sur- in the 1920s, and nothing was known of its veys with zoological content had been carried natural history. In 1992 Mickleburgh et al. out before the one described here. In 1927 the listed this species as Priority Grade 1: Whitney South Seas Expedition visited the Endangered, Limited Distribution, or possibly island collecting specimens and in 1991 an even extinct. Recent fieldwork had failed to lo- Australian Museum team working in the cate a single specimen and the possible reason Solomons collected limited data about the for the decline of the species was thought to island's fauna. However, according to the oral be due in part to the fact that the northern history of the tribes concerned, the Choiseul Solomons have the highest density of humans 1995 team were the first non-islanders to visit in the archipelago (Mickleburgh et al., 1992).

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135° 150° Papua New Guinea

Pteralopex anceps

New Georgia Pteralopex

P. atrata

P. pulchra

Figure 1. Map of the Solomon Islands, showing the main islands of the archipelago and the recorded distribution of Pteralopex species.

Villages on whose land netting was carried out (sometimes in several sites within the area): 1 Loimuni 2 Chivoko 3 Baukalo (old village site) 4 Olivetti 5 Tambarato 6 Sarelata Chivoko: 1/11/95 occupied 7 Sire 8 Loloko Choiseul Bay 9 Koa 10 Kolokopisi Kandova/Tambarato: 5/12/95, 11 Katurasele absent as of 1-5 years 12 Nanavule 13 Pisimoto Vanarave: 6/12/95, 14 Sassamunga absent as of 1-5 years 15 Voza Sassamunga: 11/10/95 occupied 16 Kaloni 17 Moli

Locality of observed Pteralopex anceps Kolokopisi: 10/1 /96, absent since 1994 Olokolata: 3/12/95, absent as of 2-5 years 10 km Rob Roy: not checked; possibly still occupied

Figure 2. Map of Choiseul, showing past and present myneri grandis roosts and date each was visited by the Choiseul 1995 team.

210 © 1997 FFI, Oryx, 31 (3), 209-217

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Another taxon of particular interest is the representative records for the year. The team largest flying fox on Choiseul, Pteropus rayneri consisted of the authors and two others. The grandis, the largest of the five subspecies of four authors carried out the bulk of the survey P. rayneri in the Solomons. Mickleburgh et al. work with the aid of local people from the 1992 classed this subspecies as Priority Grade various villages that were used as bases. 11: Not Threatened, but Flannery (1995) de- Throughout the survey permission was scribed it as Vulnerable because of local popu- sought from chiefs to work on their land- lation declines caused by disease. owning group's area. If permission was A third species of interest was Pteropus ma- granted, local people were questioned about haganus, which had not been recorded from bat sightings, caves and roosts in the area. Choiseul but had been recorded from neigh- These places were then checked to see if bouring Santa Isabel and Bougainville, and were present, what species they were and the was suspected as being present on Choiseul topography of the surrounding area to assess (Leary and Aujare, 1994). Mickleburgh et al. the best method of estimating populations. In (1992) listed it as Priority Grade 3: Vulnerable, addition, attempts were made to capture indi- Limited Distribution. viduals by setting mist nets around fruiting The aim of the fieldwork described here trees and in 'gardens' (areas used for shifting was to survey Choiseul to discover whether cultivation) as well as in primary forest. Pteralopex anceps was extinct, and the size of its Faeces and parasites were also collected for population if the species survived (an IUCN/ further analysis. SSC Chiroptera Specialist Group priority proj- Areas belonging to villages all round the ect; Mickleburgh et ah, 1992). At the same time coast of Choiseul were investigated (Figure 2) the other bat species on the island were to be but it was impossible to visit large tracts in- surveyed. land because the ownership of many areas is contentious. Additional information was collected at c. 25 Methods village lectures, which were attended by the majority of inhabitants, and during a lengthy Six months of fieldwork was undertaken be- discussion after a lecture at the Lauru (original tween July 1995 and February 1996, a period name for Choiseul) Land Conference, where that overlapped seasons to try and ensure most of the island's chiefs were assembled.

Table 1. Megachiropteran taxa from Choiseul

Taxon Priority grade Choiseul 1995 records

Dobsonia inermis minimus 8 No Data: Limited Distribution Recorded minimus microtus 10 No Data Recorded aurantius 8 No Data: Limited Distribution t Melonycteris woodfordi 11 Not Threatened Melonycteris woodfordi woodfordi Present major scitulus 8 No Data: Limited Distribution Not recorded Nyctimene vizcaccia bougainville 10 No Data Recorded Pteralopex anceps 1 Endangered: Limited Distribution Recorded Pteropus admiralitatum colonus 10 No Data Not recorded Pteropus rayneri grandis 8 No Data: Limited Distribution Recorded amplexicaudatus brachyotis 11 Not Threatened Recorded Pteropus mahaganus New Record

*Source: Mickleburgh et al. (1992). + Flannery (1993) considers M. aurantius to be a subspecies of M. woodfordi, with its distribution restricted to the Florida Islands in the Solomons.

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Pteropus rayneri grandis roosts in coconut and black fur on the head and dorsum with plantations, primary and secondary forest, the characteristic absence of fur between the either gregariously in large, obvious, noisy ag- wings (Andersen, 1909). The chest was con- gregations (McKean, 1972) or solitarily (Leary spicuously white in colour. All five individ- and Aujare, 1994). Estimation of numbers in uals took flight after 5 minutes because of the large roosts was carried out by walking disturbance and it was impossible to see how through the roost in a straight line, trying to far they flew in the dense vegetation. Another count the bats before they were disturbed. individual was seen on 26 December as it flew This often meant estimating the number of from a fig that a local helper was climbing in bats in a roost 'patch' and multiplying that by search of bats. It had been roosting in a small the number of patches. The technique is simi- hollow between branches. These were the only lar to the direct counts of Mutere (1980), observations of this species made during the 6 judged by Kunz (1988) to be fairly accurate. months of fieldwork. Additional dispersal counts were used where Pteralopex anceps might be confused with the possible to gauge and improve the accuracy of larger island form of Pteropus admiralitatum, the direct counts. Thus, a combination of the which is dark with relatively short ears, and most appropriate techniques, dependent on long silver hairs on the ventral side (T. the situation, was used to establish greater ac- Flannery, pers. comm.) and is known to roost curacy (after Mickleburgh et ah, 1992). Solitary in groups in large forest trees at higher elev- roosting individuals were observed in various ation (Flannery, 1995). However, the authors locations but transects through a variety of are convinced that the first sighting rep- habitats failed to reveal the presence of indi- resented P. anceps owing to the size - equal if vidual bats, which were thus taken to be at not larger than P. rayneri (the largest member non-significant numbers. of Pteropus in the region) and the absence of As a byproduct of the fact that mist nets fur between the wings. T. Flannery verified were a central part of the surveying tech- the second sighting of the single P. anceps niques used and that cave roosts were visited, from the photograph taken of this individual. a list of the Microchiroptera of Choiseul was also compiled. Pteropus rayneri grandis (Senga name, Kukuvai) Pteropus rayneri grandis roosts communally Results and roosts were only found close to the coast despite extensive work inland. Although the Pteralopex anceps (Senga name, Kunjulu) interior of Choiseul is rugged, the terrain The vast majority of people on Choiseul were around the swampy/mangrove areas that not aware of the existence of Pteralopex anceps, P. rayneri grandis favours is flat and covered including those familiar with the forest. This is with thick spiny vegetation in deep mud. The presumably because people use the coastal roosts may cover 2 sq km. areas where the species does not seem to Occupied and non-occupied roost sites are occur. Only two men from more than 60 shown in Figure 2 together with the dates elders interviewed knew the local name for when they were surveyed and last occupied. the . The majority of sites seemed to have been va- On the afternoon of 25 December 1995 the cated and only two areas had large colonies authors observed five individuals of P. anceps resident. This is surprising in view of reports hanging beneath the branches of a large fig, from other sources of the large numbers of Fkus sp., approximately 12 m above the colonies per island (Flannery, 1995), and nu- ground. The tree was on a ridge at 850 m in merous accounts from local people of seeing primary rain forest (Figure 2). Identification of the sky filled with bats in the past. the bats was enabled, with the use of binocu- Sassamunga and Chivoko roosts, respect- lars, by the absence of large protruding ears ively, held 7188 and 5200 bats. The only roost 212 © 1997 FFI, Oryx, 31 (3), 209-217

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site not checked was Rob Roy Island, which was active, according to the land owner. This roost was apparently approximately the same size as Sassamunga and Chivoko. Given a non-significant density of solitary animals, and the sizes of the active colonies, it seems reasonable to add a conservative 5000 animals for Rob Roy giving a maximum of 17,300, and a minimum of 12,300 (assuming Rob Roy vacant) for the current total Choiseul popu- lation of Pteropus rayneri grandis. Using the same cautious average size of 5000 animals per roost and seven past active roosts, there may have been a decline from perhaps at least 35,000 animals in the last 10 years. This con- firms that this subspecies is vulnerable.

Pteropus mahaganus (Babatana name, Gama) Our survey results filled in the previously dis- junct distribution of Sanborn's flying fox Pteropus mahaganus by recording this species from Choiseul for the first time. This species was caught on numerous occasions in many localities on Choiseul and observed at many Pteropus rayneri roosting (E. Bowen-Jones). others. Previously it had only been known from Santa Isabel and Bougainville, on either side of the study island. It is distinguishable in Pteropus admiralitatum life from the smaller inermis and We did not capture any Pteropus admiralitatum, larger P. rayneri grandis by its rapid flight with which had been collected from Reef Island its faster wing beat. When seen feeding its (Choiseul Bay, Figure 2) in the 1920s. As yellow/brown mantle makes identification Flannery (1995) noted, this small species is easy. The species appears to be fairly common more often found on islets than on large on the island. Despite this, no roost sites were islands. Perhaps this explains its absence from discovered, although local people said that it the survey, because the work concentrated on roosts in tree hollows in primary forest. mainland Choiseul. Observations showed that, as suspected by Flannery (1995), P. mahaganus feeds on the inflorescences of coconut rather than the fruit. Microchiroptera Additional data were gained on its role in the The results of our survey also included seven pollination of sago palm on the island, al- new records of Microchiroptera: Emballonura though the degree of exclusivity is unknown. dianae, Emballonura nigrescens, Hipposideros If, as seems likely, this species is an important dinops, Miniopterus propitristis, Miniopterus pollinator of sago palm and coconut, then it is schreibersii and Pipistrellus angulatus. important to local human economies and com- here has been kept to species level munities. In this subsistence-orientated region, because opinions differ about finer divisions; the copra harvest from coconuts is the pre- all distributions and nomenclature follow dominant source of income, while the sago Flannery (1995). The sixth-ever record of the palm provides the major building material Solomons' flower-nosed bat Anthops ornatus used for village housing. (Hipposideridae) was also made.

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Discussion the study and had been for as long as people could remember. Population status o/Pteralopex anceps The reduction of Choiseul P. rayneri grandis Fisher (1992) found that the most important populations could be due to several possible roost trees for the New Georgia Pteralopex (as factors. yet undescribed) were figs, and to a lesser ex- 1 Disease. Flannery (1990) described P. rayneri tent ngali nuts Canarium sp. The fact that the grandis populations being wiped out by epi- inland high ground of Choiseul, to which demic disease in Bougainville and Buka (an P. anceps now seems confined, is ultra-basic island off north-west Papua New Guinea) in rock, may partially determine a low popu- 1987. He also suggested that, because of its ex- lation. The small numbers of figs found on ceptionally high fatality, the disease may have this high ground is a consequence of the been introduced. In Kolokopisi, Choiseul, the edaphic features dictated by the bedrock. Such chief described to us how within a week or so areas are generally less productive forest, with in 1994 the large colony 'thousands' strong a low number of tree species, often in hom- 'fell down from the trees'. ogenous stands, and probably therefore offer a 2 Red ants. Many local people referred to P. r. lower availability of food (Whitmore, 1966). grandis roosting in large numbers in coconut Food availability in primary forest might be plantations in the past. However, this is not crucial to P. anceps, which was not netted or the case now and many people thought this observed during approximately 800 hours' was due to introduction in 1987 of red ants work in secondary forest and gardens. This from the Russell Island (central Solomons) fits with Fisher's (1992) reports that the New strain of coconuts. This occurred throughout Georgia Pteralopex is absent from these habi- the Solomons and would fit with Flannery's tats; she found that primary forest and old vil- description of population decimation in lage sites had a higher number of fig trees Bougainville in 1987. The ants are known to (roosts) and ngali nut trees (food). Pteralopex attack the eyes of cuscus Phalanger orientalis anceps probably has a similar diet to the New and the irritation and stress this induces may Georgia Pteralopex, and other species of the prevent bats roosting. In addition, it is poss- genus, which is predominantly based on nuts. ible that whatever disease is affecting This is based on the generic feature of special- P. rayneri, the vector is the introduced red ant. ist dentition. As a rule the genus is confined to Perhaps a more likely scenario is that another primary forest (Flannery, 1995). Fisher (1992) organism was introduced in the same way as described how the New Georgia bat seems to the ant and this is the source of the epidemics. associate with a roost site allowing recapture. 3 Cyclones. According to many people the The reverse may be true of P. anceps, where in- population of flying foxes on the island has dividuals disturbed from the roost had still not recovered from Cyclone Ida in 1972. This not returned 2 weeks later. destroyed large roost trees in several areas and apparently reduced the roost area at Chivoko to one-third the original. Absence o/Pteropus rayneri grandis from 4 Hunting and trapping. Hunting does not established roosts seem to be of primary concern on Choiseul The absence of P. r. grandis from past roost where the fruit bat is deemed to be an oc- sites may have been seasonal, but local people casional treat. Leg-hold snares are sometimes in all of these areas claimed that bats had not used to capture bats, as perceived pest control. been present for 2-5 years. It would seem un- However, there was no evidence of wide- likely that the absence of populations was due scale, intensive trapping. to inter-island seasonal migration, because of Perhaps a combination of these and other the reported length of absence and the fact events caused the decline in the island's bat that the roosts containing large existing popu- population while it was in a state of recovery lations were constantly occupied throughout from the 1972 cyclone. Population recovery 214 © 1997 FFI, Oryx, 31 (3), 209-217

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after such events is slow because of the low re- restriction that human disturbance places on productive rate of these . available habitat, P. anceps appears to be in low numbers. Thus, logging may now threaten the future of this species. A suitable Conservation action reserve of forest may be possible in the Mt Maetambe region, where there is local interest Recommendations have been made to rel- in setting up a community-owned conser- evant government and conservation organiz- vation area (Bowen-Jones et al., 1996). ations in the Solomons for protection of P. Mickleburgh et al. (1992) suggested that cap- rayneri grandis roosts on Choiseul. However, tive breeding should be considered as a poss- from the evidence of declining populations on ible management tool for P. anceps. Given the Malaita, Bougainville and Buka, north-west threats summarized above and the lack of in- Papua New Guinea (Flannery, 1990) it would formation about the populations, which are al- seem that such measures may be necessary most certainly in a similar condition on throughout the whole of the Solomons for all Bougainville, this is desirable. However, the subspecies. Community-based education cost of such a logistically difficult capture schemes on the ecological importance of bats process is likely to make such a scheme chal- and the establishment of protected areas lenging. Therefore, effort should be concen- would be the most appropriate measures. trated on habitat preservation and education Research also needs to be carried out to moni- schemes. tor populations of this species over a longer period of time and gauge the degree of threat it is under. Currently, rapid clearance of man- The Solomons: a conservation groves and other forests on many islands in imperative the archipelago threaten habitat and wild food resources. With the pressures discussed, plus Choiseul has particular relevance in being the an increased reliance on gardens as foraging least developed of the larger islands in the areas leading to more 'pest' control, P. rayneri Solomons. If its forests are clear-felled most of is under imminent threat. Perhaps education, the species found there will survive on leading to the realization of the true ecological Bougainville. Although this island is zoologi- value of fruit bats, would reduce the current cally almost identical to Choiseul (from what use of snares as 'pest' control. At the moment little is known of its fauna) it is politically part these are often placed indiscriminately, par- of Papua New Guinea. Pteralopex anceps, along ticularly on coconuts, affecting P. rayneri and with other species such as Poncelet's giant rat P. mahaganus. Solomys ponceleti, are confined to these two There are a great number of potential press- islands of the North Solomons. However, ures capable of reducing the extant population there is no reason to suggest that the more of P. anceps as socio-economic factors force vil- populous Bougainville will not be subject to lagers to sell customarily owned land for com- even more unsustainable development press- mercial clear felling. Some of Choiseul's forest ures once a solution has been found to the cur- cover has already been lost and many new li- rent conflict there, so there can be no cences are in the process of being granted. complacency. The other islands in the This destroys feeding and roosting habitat. Solomons also have limited-distribution, en- However, the more desirable selective logging demic species on them. may also reduce available roost trees for The general situation for Solomons' fauna Pteralopex and probably Pteropus mahaganus. If and flora (low diversity of canopy species but P. anceps is as sensitive to disturbance as it high endemicity; Whitmore, 1966) can be illus- seems to be, then the only hope for conser- trated with reference to the biogeography of vation is to preserve large areas of untouched the genus Pteralopex. All known species oc- forest. Whether due to natural densities or cupy one or two islands at the most (Figure 1).

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All are threatened by current logging prac- periods available to carry out this work, logis- tices. For example: the New Georgia Pteralopex tical, political and equipment factors (because is known to have occupied Kolombangara as of climate, plus distance and cost of resupply) well, but the clearance of 80 per cent of forest are likely to conspire against the collection of and replacement with plantations has caused data. its extinction there (Flannery, 1995). Now in- Environmental education is desperately creasing logging on New Georgia threatens to needed throughout the Solomons, and other extirpate the species. Pteralopex atrata on species of Megachiroptera and other mam- Guadalcanal faces a similar collapse due to the mals and birds await urgent investigation. In forecast of all forest below 800 m being lost many cases the time scale for this to be carried within the next decade. On this island it seems out is well inside a decade before all forest to be replaced by Pteralopex pulchra above this will have been lost, and with it undoubtedly height, the only record of altitudinal zonation many of the endemic species that inhabit these in Solomon mammals (Flannery, 1991). P. pul- small Pacific islands. chra is only known from one specimen. Although there is a complete absence of Acknowledgements ecological information on the role of fruit bats in the ecosystem of these islands, it is possible Thanks to Phil Richardson of The Bat Conservation that, as in other Pacific islands, a third of the Trust and Gilla Carstairs, who were project mem- canopy tree species rely on flying foxes for bers for the first few months. We would like to pollination or seed dispersal (Cox et ah, 1992); thank WWF Community Resource Development and they provide potential economic benefits Fund, Bristol University Expedition Fund, Alumni as pollinators of cash crops. There is urgent Foundation, Hosier Fund, British Ecological Society, need for action. Royal Geographic Society, Intervet Bursary, Institute of Biology, Gilchrist Trust, Bat Conservation Trust, BP Conservation Expedition Programme 1995 and others for making the necess- Further research ary funding available. In the Solomons we thank: Glynn Galo and Esau Tuza, at the University of the In order to ascertain more about the popu- South Pacific, Honiara Division; Tia Masolo and Moses Biliki, at the Ministry of Forests, lation of Pteralopex anceps further netting must Conservation and Education; Audrey Rusa, at the be carried out in Choiseul's interior with Ministry of Training and Education; Myknee large /owl-mesh mist nets. At the time that Sirikolo at the Botanical Gardens, Honiara and this work was carried out on Mt Maetambe everyone who allowed us to work on their land and only smaller mesh nets were available and capture animals in Choiseul. Finally we would like several large pteropodids 'bounced' off; one of to thank the following for their help: Professor Paul these was probably P. anceps. With the capture Racey, Professor Steven Harris, Simon Mickleburgh and FFI, Dr Gareth lones, and Dr Tim Flannery. of P. anceps there should be an effort made to radio-track them in the Maetambe region, which might enable the location of roost sites References and therefore estimation of numbers. Andersen, K. 1909. Two new bats from the Solomon Using the same techniques, an easier project Islands. Annals and Magazine of Natural History, would be to work on the more widely distrib- London, (8) 3,266-270. uted, little-known and potentially economi- Bowen-Iones, E., Abrutat D., Markham W. and cally important Pteropus mahaganus. With the Bowe, S. 1996. Report of the Bristol University, setting of mist nets at sufficient height around Choiseul 1995, Survey of Fruit Bats on Choiseul, suitable trees these were relatively easy to Solomon Islands. Unpublished. University of Bristol catch. If owl-mesh were used, there would be Cox, P.A., Elmquist T., Pierson E.D. and Rainey a higher rate of success. W.E. 1992. Flying Foxes as pollinators and seed dispersers in Pacific island ecosystems. In Pacific Much of this research will encounter prob- Island Flying Foxes: Proceeding of an International lems with land access. Without long time Conservation Conference (eds D. E. Wilson and G. 216 © 1997 FFI, Oryx, 31 (3), 209-217

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L. Graham), pp. 18-23. US Fish and Wildlife Leary, T. and Aujare, I. 1994. Review of the Status and Service Biological Report 90 (23). US Department Distribution of Pteropus Flying Foxes in Solomon of the Interior, Fish and Wildlife Service, Islands and a Preliminary Assessment of Fruit Crop Washington, DC. Damage in Temotu Province. The Nature Fisher, D. 1992. Investigation of the Ecology of a New Conservancy, Honiara Field Office, Solomon Species of Monkey-faced Bat from the Solomon Islands. Islands. Unpublished report, Section, Australian McKean, J.L. 1972. Notes on some Collections of Bats Museum from Papua New Guinea and Bougainville Island. Flannery, T.F. 1990. Flying foxes in Melanesia: CSIRO Division, of Wildlife Research Technical populations at risk. Bats, 7, 5-7. Paper No. 26. Flannery, T.F. 1991. A new species of Pteralopex Mickleburgh, S.P., Hutson, A.M. and Racey, P.A. (Chiroptera: Pteropididae) from montane 1992. Old World Fruit Bats: an Action Plan for their Guadalcanal, Solomon Islands. Records of the Conservation. IUCN, Gland, Switzerland. Australian Museum, 43,123-130. Mutere, F.A. 1980. helvum revisited. In Flannery, T.F. 1993. Revision of the fruit bats of the Proceedings 5th International Bat Research Conference genus Melonycteris (Pteropodidae: Mammalia) (eds. D. E. Wilson and A. L. Gardner), pp.145-150. Records of the Australian Museum, 45,59-80. Texas Tech University Press, Lubbock. Flannery, T.F. 1995. Mammals of the Southwest Pacific Whitmore, T.C. 1966. Guide to the Forests of the British and Moluccan Islands. Australian Museum/Reed Solomon Islands. Oxford University Press, Oxford. Books, Chatswood, NSW. Kunz, TH. 1988. Ecological and Behavioural Methods E. Bowen-Jones, D. Abrutat, B. Markham and for the Study of Bats. Smithsonian Institution Press, S. Bowe, 12 Aldersmead Road, Beckenham, Kent Washington, DC. BR3 1NA, UK.

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