The species is present from sea level to around 1,250 m asl and is found roosting in trees in primary and secondary forest, along the beach, in mangroves and inland, and on hill forests on small uninhabited islands, often near adjacent agricultural areas that it uses for feeding (Gumal et al. 1998; Mohd-Azlan et al. 2001, 2022; Mildenstein et al. 2005; Struebig et al. 2007 Harrison et al. 2011; Mildenstein 2012; Aziz et al. 2019a; Sheherazade unpublished data). In highly modified landscapes, colonies are often found in relatively undisturbed/protected sites, such as temple compounds/forests, parks, botanical gardens, cliff faces, or other socio-culturally significant grounds (S.A. Aziz and T. Mildenstein unpublished data). However, colonies have also been found in some urban areas (e.g., in the Philippines: in Bongao; >2 sites in Tawi-Tawi (Buan and Panglima Sugala); Pandan, Panay) (PBCFI unpublished data). In Malaysia, Sumatra and Java, the species is found in island, coastal and inland habitats, where it forages in both forests and orchards (S.A. Aziz unpublished data, Sheherazade unpublished data). In Peninsular Malaysia, it is possible that permanent or semi-permanent colonies may still roost in inaccessible forests (swamps, mangroves, rainforests) including sites further inland from the coast in Pahang, Johor and Perak, as foraging is still occasionally reported in cultivated fruit orchards (S.A. Aziz unpublished data). In Singapore, sightings of the species are primarily from temporary roosts in tall forest trees, and temporary roosts or observations of flight over remnant lowland dipterocarp forest (Yap 2016a,b; B. Lee unpublished data). In Sabah and Sarawak (Malaysian Borneo) the species is considered a visitor in interior parts of mixed dipterocarp and peat swamp forests and cultivated fruit orchards during flowering and fruiting seasons (Mohd-Azlan J. pers. comm. 2020, L.A. Bansa pers. comm. 2021, M. Gumal pers. comm. 2022). Forest areas in the Kinabatangan water basin of Sabah are important roosting sites. (L. Bansa pers. comm. 2021). In Kalimantan (Indonesian Borneo), colonies are patchily distributed in coastal peatlands, mangroves, and swamps (M. Struebig pers. comm. 2021). In the Philippines, the largest remaining colonies of the species are found roosting in mangroves but always within 50 km of lowland forest (Mildenstein 2012, Philippines Biodiversity Conservation Foundation, unpublished data).

Habitat selection studies demonstrate the species preferentially uses undisturbed native forest, including mangroves, peat swamps and freshwater swamps, over disturbed or agricultural areas for roosting, but it uses both forest and agricultural areas for foraging (Mildenstein et al. 2005). In both natural and disturbed landscapes, hemi-epiphytic figs are a staple food and substantial part of its diet (Shanahan 2001, Stier and Mildenstein 2005). The species is known to migrate long distances seasonally to track fruit and flower availability (Epstein et al. 2009, L.A. Bansa unpublished data). For example, in mid-May 2001 at least 100 P. vampyrus visited Singapore at Lower Peirce Reservoir in the Central Catchment Nature Reserve and fed on Syzygium grande which was fruiting profusely (B. Lee pers. obs. 2001). As such, it is likely to be vital for long-distance dispersal of both pollen and seeds (e.g. Ong et al. 2021), including between landmasses separated by sea (Aziz et al. 2021) – but data on these aspects are still lacking and urgently needed. Also, as Pteropus flying foxes are known to cease functioning as seed dispersers when population abundance is low (McConkey and Drake 2006), it is possible that functional extinction may have already occurred in places with drastic population declines.

Colonies of this species are often found roosting together with the Common Island Flying fox (Pteropus hypomelanus) (Pulliam et al. 2011, Philippines Biodiversity Conservation Foundation Inc. (PBCFI) pers. comm. 2021), and less frequently with the endemic Golden Crowned Flying Fox (Acerodon jubatus) (Mildenstein 2012, Philippines Biodiversity Conservation Foundation Inc. (PBCFI) unpublished data, M.R. Duya unpublished data). In eastern Thailand, it occasionally shares mainland roost sites with Pteropus lylei.

This species is generally found in forest from sea level to 1,250 m asl throughout much of continental and insular Southeast Asia, although its preferred lowland forest habitats are now fragmented across much of its historical range. On continental Southeast Asia, it has been reported from southern Viet Nam, Cambodia, Thailand, and through much of Peninsular Malaysia (Mohd-Azlan et al. 2001, Thong 2004, Epstein et al. 2009, Hul 2013, Ravon et al. 2014, Aziz et al. 2019b, T. Ngamprasertwong pers. comm. 2021, S. Bumrungsri pers. comm. 2021). Despite its historical ubiquity in Singapore, there are currently no roosts recorded from there, but visiting individuals have been spotted on rare occasions most recently (B. Lee pers. comm. 2021). Among the insular Southeast Asian countries, the species is found across the Philippines (except for the Batanes/Babuyan region) (Mildenstein 2012, Philippine Department of Environment and Natural Resources (DENR) unpublished reports, Anson Tagtag and Katherine Soriano pers. comm. 2021) and over much of western Indonesia and the Lesser Sunda Islands (Sheherazade pers. comm. 27 August 2021). On Borneo, the species is found throughout much of Kalimantan (Indonesia; Struebig et al. 2007, Harrison et al. 2011), Sabah (Malaysia; L. Bansa pers. comm. 2021), and some parts of Sarawak (Malaysia; Gumal 2001, Mohd-Azlan et al. 2022). Occasional sightings of small numbers of individuals are reported from across Brunei (M. Struebig pers. comm. 2021). Although this species is regional in nature, local/national distribution and range is much more limited, and generally restricted to localities that are geographically inaccessible to humans. Details on known populations by country/region are provided below.

Philippines:
Roosting colonies are found across 17 regions of the Philippines (except for the Batanes/Babuyan region) (DENR, unpublished data), including the newly documented presence in the following: National Capital Region, the Cordillera Administrative Region, the Bangsamoro Autonomous Region of Muslim Mindanao and Region IV into IVA (CALABARZON) and IVB (MiMAROPA – Mindoro, Masbate, Romblon, and Palawan) (DENR 2019). Specific island records include: Bohol, Bongao, Boracay, Cabo, Catanduanes, Culion, Leyte, Guimaras, Luzon (provinces: Abra, Bataan, Cagayan, Ilocos Norte, Isabela, La Union, Laguna, Pangasinan, Quezon, Tarlac, Zambales), Marinduque, Mindanao (Agusan del Norte, Bukidnon, Cotabato, Davao del Norte, Davao del Sur, Davao Oriental (Jojo Araujo pers. comm. 2021), Lanao del Norte, Lanao del Sur, Maguindanao, Misamis Occidental, Misamis Oriental, North Cotabato, South Cotabato (Abdulmukim Maruji pers. comm. 2021), Zamboanga del Norte, and Zamboanga del Sur provinces and Bongo Island (Dante Oporto, Michael dela Cruz and Georgina Fernandez, DENR Zamboanga, pers. comm. 2021), Mindoro (Katharine Cielo pers. comm. 2021), Negros, Palawan, Panay, Polillo (Alviola 2000), Tantangan, Dinagat (Gunther 1879), and Romblon (Timm and Birney 1980). Of the 173 colonies reported, only 86% (149 roosts) are found inside protected areas (DENR 2019).

Malaysia:
The species was historically widespread throughout the country, but reported sightings have decreased drastically by >50% (PERHILITAN 2017), and are now confined to only a few specific areas (Gumal 2001; Mohd-Azlan et al. 2001, 2022; Aziz et al. 2019a; L.A. Bansa unpublished data; S.A. Aziz unpublished data). In Peninsular Malaysia, the presence and movement of this species has been challenging to ascertain due its nomadic nature and inaccessible roost sites, with most roosts being only temporarily occupied, and most colonies comprising transboundary populations that are shared with southern Thailand, Indonesia (Sumatra), and Singapore (Epstein et al. 2009). Recent surveys (2018–2022) have only been able to confirm roosting colonies in unprotected areas in the states of Terengganu, Pahang, and off Langkawi Island in the state of Kedah, and flyout sightings (roost sites unknown) have been reported from Terengganu, Pahang, and Johor (S.A. Aziz unpublished data). One roost site location previously deemed to be permanent (Epstein et al. 2009) was not found to be occupied at the time of survey despite repeated visits (S.A. Aziz unpublished data). No recent roosts have been reported from protected areas in Peninsular Malaysia, although there are occasional sightings of flying or foraging individuals in the states of Perlis and Perak, and the protected areas of Kenyir State Park, Taman Negara National Park, and Royal Belum State Park (S.A. Aziz unpublished data). Local people at many previously recorded roost sites (Mohd-Azlan et al. 2001, Epstein et al. 2009) reported no recent sightings of the species for several years or even decades (Aziz et al. 2019a). However, in 2019 one new, previously unrecorded island roost site was discovered in Terengganu based on local knowledge (S.A. Aziz unpublished data). In Sabah, some colonies are only present on a seasonal basis, especially during fruiting season, as in Beaufort, Tenghilan, Kota Belud, Tamparuli, Tambunan and Ranau (L.A. Bansa unpublished data). Permanent colonies are only known from Deramakot Forest Reserve and Tangkulap Forest Reserve, but due to regular disturbance from selective logging, has not been seen roosting or flying in large numbers since 2020 (Sabah Forestry Department 2006–2017). However, individuals are still occasionally sighted along rivers in Kinabatangan, feeding in Maliau Basin Conservation Area (L.A. Bansa unpublished data), Silam Coast Conservation Area (SCCA), Lahad Datu (Pounsin et al. 2016). Nuluhon Trusmadi Forest Reserve and Sungai Kiluyu Forest Reserve, Tambunan, in 2015 (Wildlife Monitoring, 2018), and a colony has been reported in Imbak Canyon Conservation Area (L.A. Bansa unpublished data; Senawi et al. 2020; Bunya et al.2012). In Sarawak, several large roosts were reported in Bukit Sarang, Loagan Bunut, Pulau Bruit and Sedilu, (all now gazetted as National Parks) (Gumal 2004). Temporary, smaller roosts were reported at Bako, Limbang Mangroves, Lundu and Similajau. Appears sporadically around the western tip of Borneo, but also occasionally around the interior parts of western Sarawak (Simunjan, Lingga, Maludam, Roban and Lubok Antu), northern Sarawak (Limbang), national parks along the coast (Bako, Sedilu and Kubah), and further inland at Loagan Bunut and Batang Ai.

Indonesia:
The species is found over much of western Indonesia and the Lesser Sunda Islands, being recorded from parts of Sumatra and its surrounding islands, including Bangka; the southern Mentawai Islands of Sipura, North Pagai and South Pagai; Riau Islands (including Natuna Islands), Krakatau Islands (including Sebesi), Java, Bali, West Nusa Tenggara (Lombok and Sumbawa islands), and East Nusa Tenggara (Flores, Sumba, Timor, and Komodo islands) (G. Fredriksson pers. comm. 2021; Sheherazade pers. comm. 2021). In Kalimantan (Indonesian Borneo), roosting colonies are reported around Sebangau, Sampit, Buntok, Ampah, Kuala Pembuang and Kasongan in Central Kalimantan, Ketapang in West Kalimantan, and Balikpapan Bay in East Kalimantan (G. Fredriksson pers. comm. 2020, M. Struebig pers. comm. 2021). Individuals have also been infrequently sighted in other areas as far inland as the Upper Barito (Puruk Cahu, Tumbang Topus) in Central Kalimantan, Landak in West Kalimantan, upper and middle Mahakam (Muara Kaman) in East Kalimantan, and Kayan Mentarang in North Kalimantan, though no colonies have been reported in these areas.

Singapore:
This species is considered native to Singapore, but all roosting colonies are reportedly extirpated from the main island and offshore islands; it is unknown when this occurred. Currently, it is only a visitor, and from historical records, is sighted in small numbers almost every month of the year except January and February. On the main island, the species is occasionally sighted flying over the Central Catchment Nature Reserve (Yap 2016a, 2016b) and a remnant forest next to the Swiss Club area (B. Lee unpublished data). It is also occasionally sighted on Ubin Island in the northeast, St John’s Island (Ollington and Subaraj 1988) and Sisters’ Islands (Shivaram Rasu pers. comm. 2021), with the last visitation in early June 2020 when 100 bats appeared for a week and roosted in the Central Catchment Nature Reserve (Tan 2020); the bats subsequently left Singapore.

Viet Nam:
This species was previously found in different areas from the Mekong River Delta up to the central region of the country (Huynh et al. 1994). Currently, it is only found in southern Viet Nam where there is a single remnant roosting colony of ca 300 individuals in a mangrove area of Can Gio district, Ho Chi Minh city (Thong 2018). Between 1995 and 2019, a few colonies of this species (estimated less than 100 individuals per colony) were occasionally sighted at several localities within the Mekong Delta Region of southern Viet Nam. However, apart from the roosting colony in Ho Chi Minh City, this species has not been reported from any other locality in the country since 2019 (Vu Dinh Thong pers. comm. 2022).

Thailand:
This species has been found in south and southeast Thailand, below the Isthmus of Kra, and in eastern Thailand, near the Thailand-Cambodia border (S. Bumrungsri pers. comm. 2021, T. Ngamprasertwong pers. comm. 2021). Several new roost sites were recently reported in southern Thailand, reflecting the movement of some adjacent colonies, and it has been recorded in Chonburi, Trat, Krabi, Chumporn, Narathiwat, Suratthani and Satun. Several populations are on islands, and at least one population is in a protected area in Satun (S. Bumrungsri pers. comm. 2021).

Cambodia:
A rapid survey identified 12 roosting colonies of this species along the Mekong River and Tonle Sap lake and drainage basin, which likely represent most of the flying fox populations in the country (Ravon et al. 2014). One of the roosting sites where this species was identified is the Wat Pi Chey Sa Kor temple in Kandal Province (Hul 2013).

Brunei:
Occasional sightings of small numbers of individuals have been reported from across the country, including Ulu Temburong, Muara, Menunggul and Berambang island (M. Struebig pers comm. 2021, Mohd-Azlan J. pers. obs. 2021).

Timor-Leste:
No information is available, but as the species is known to occur in Timor, Indonesia (Sheherazade pers. comm. 2021), it is likely to be extant here as well.

While some colonies in some countries are known to roost in protected areas (e.g. in Sarawak where there was a direct conservation initiative to create protected areas around these roosts), many others roost in legally unprotected habitat (e.g., most roost sites in the Philippines, Peninsular Malaysia and Cambodia are not found in protected areas), and many range countries either have insufficient or a complete lack of conservation activities to address population declines and threats. Conservation actions to protect and manage roosts/habitats, address hunting and persecution, and mitigate climate change impacts, are urgently needed. This species is listed on Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), which restricts sale and transport internationally, and permits are needed – but most trade appears to be local or confined within national borders, rather than international (K. Krishnasamy pers. comm. 2018). Protection on a national scale is dependent on each country in which the species occurs and is described below.

Philippines:
The Wildlife Conservation and Protection Act (Republic Act 9147) lists the species as Endangered through Department Administrative Order (Gonzalez et al. 2018; DENR 2019), and hunting is prohibited for this, and all fruit bat, species in the Philippines. Most of the Pteropus vampyrus populations in the Philippines are found on private lands (PBCFI unpublished data). Some populations of the species are found within the many protected areas throughout the country, but the amount of active protection these populations receive varies greatly with the resources and management capacity of each protected area. In a few cases, roosting sites are actively protected (e.g. Dagat, Cebu; Mambukal, Negros OR, Subic Bay), however, the majority of roosting populations, even in protected areas, are not shielded from disturbance and hunting. Some populations on Mindanao Island are relatively protected because of cultural beliefs and practices. In 2017, the Philippines’ Department of Environment and Natural Resources (DENR) launched a country-wide conservation management action for its native large fruit bats, including Pteropus vampyrus (Anson Tagtag and Katherine Soriano, Biodiversity Management Bureau pers. comm. 2021). The DENR’s fruit bat conservation action plan involves managers from regional and municipal offices across the country, who are directed to survey for, and record the location and size of, roosting populations of large fruit bats within their jurisdictions. The results of this enormous effort have provided a 5-fold increase in the number of known roosting populations of large fruit bats, baseline information on the species present and their population sizes, as well as time series data that will be essential to detecting population trends. With this demonstrated commitment to conservation of large fruit bats at the national government level, there is potential for management for the long-term persistence of Pteropus vampyrus in the country with the largest population of the species.

Malaysia:
In Peninsular Malaysia the species, along with P. hypomelanus, is now listed as Endangered in Version 2.0 of the Red List of Mammals for Peninsular Malaysia based on Criteria B2 and E (PERHILITAN 2017), but based on the results of recent field surveys (Aziz et al. 2019a), the status of P. vampyrus may have to be revised to Critically Endangered. Both species are on the Protected list under Schedule II of the Wildlife Conservation Act 2010, whereby hunting without a license is an offence subject to a fine of up to fifty thousand ringgit (~USD 12,500), imprisonment of up to two years, or both. However, enforcement of the law has been challenging, as is detection and monitoring of hunting and trade activities. In 2012 the Terengganu state government officially banned hunting of Pteropus spp. in the state (Heng 2012a), and from 2015 onwards the Department of Wildlife and National Parks Peninsular Malaysia (PERHILITAN) initiated a moratorium on all flying fox hunting. They also announced that they would move Pteropus spp. from the ‘Protected’ to the ‘Totally Protected’ (Schedule I) list, meaning that flying fox hunting would be officially prohibited under the law in all states across the peninsula (Chu 2017). However, this latter action is still pending as of April 2022. In 2019 the local NGO Rimba collaborated with PERHILITAN to produce the ‘Conservation Roadmap for Flying Foxes Pteropus spp. in Peninsular Malaysia’ (Aziz et al. 2019b) and is also currently working with the Terengganu State Government and PERHILITAN to legally protect and manage roost sites and foraging habitat (e.g. Kenyir State Park); currently no other organisations or initiatives exist for flying fox conservation on the peninsula (S.A. Aziz pers. comm. 2022). Conservation of this species on the peninsula requires a regional effort involving collaboration, cooperation and communication with Indonesia (Sumatra), Singapore, and southern Thailand (Epstein et al. 2009). In Sabah (Malaysian Borneo) this species is protected under Schedule 3 of the Wildlife Conservation Enactment 1997, for which a hunting license is required (Section 2, 25 (2)). Hunting is subject to the quota listed on the license, and is not allowed on alienated land, forest reserves (Forest Enactment 1968), parks (Parks Enactment 1984), reserved land, or state land that requires the owner’s or authorities’ approval for entry (Wildlife Conservation Enactment 1997). Possession and selling of this species also require permits or certificates of legal ownership. In Sabah there is still a lack of ecological information on this species such as population size (e.g. number and sizes of colonies), spatio-temporal patterns (e.g. roosting patterns, roosting sites, and movement patterns) (L.A. Bansa pers. comm. 2021). Statewide surveys on this species are needed to provide a baseline database for conservation action. In Sarawak (Malaysian Borneo), this species is currently protected under the First Schedule (Part II) of the Sarawak Wild Life Protection Ordinance (1998) where all bats are given similar protection status. No hunting or trapping are allowed except under and in accordance with the terms and conditions of a license issued under the Ordinance, carrying the offence: Penalty, imprisonment for one year and a fine of ten thousand ringgit. All known roosts (Limbang Mangroves, Loagan Bunut, Pulau Bruit, Sarang and Sedilu), are also under protection as National Parks. In the next five years, the Wild Life Protection Ordinance (1998) is anticipated to be amended to mandate higher fines for killing Totally Protected and Protected Animals. Sarawak intends to increase the protection for P. vampyrus by listing it as Totally Protected (from merely Protected, which permits regulated hunting). Follow-up roost surveys are anticipated to take place by 2025 (M. Gumal pers. comm. 2021).

Indonesia:
In Indonesia, P. vampyrus did not get included in the revised list of protected species (2018), and as it took close to 20 years for this revision to come about, it is not likely to see the species listed as protected by national law in the near future (G. Fredriksson pers. comm. 2021). Although harvesting and trade are allowed in central Java through an annual quota system, this has been difficult to enforce, and without proper management these quotas are always exceeded (Sheherazade pers. comm. 2021).

SINGAPORE: P. vampyrus was placed under the category ‘Endangered’ in the first edition of the Singapore Red Data Book (SRDB) (Ho et al. 1994) but was not listed in the second edition. For the third edition, currently scheduled to be published in 2022, the species has been reviewed and up-listed to ‘Critically Endangered’ due to the loss of permanent roosts and the rare occurrences in recent years, presumably due to population declines in neighbouring countries (B. Lee pers. comm 2021). The Wildlife Act (2020) has increased punishment for wildlife offences related to protected species (Wildlife (Protected Wildlife Species) Rules 2020), which includes all Pteropus species. The Endangered Species (Import and Export) Act gives effect to CITES by controlling the importation and exportation of certain animals, plants, and parts thereof. Apart from the continued protection of key habitats such as the rainforest nature reserves and Ubin Island, there is also habitat restoration and intensive tree-planting by the National Parks Board, which would hopefully support P. vampyrus when using Singapore as a transient habitat in the wider region of the Riau Archipelago, Sumatra, and Peninsular Malaysia (B. Lee pers. comm. 2021).

Viet Nam:
All three flying fox species in Vietnam (Pteropus hypomelanus, P. lylei and P. vampyrus) are listed as “protected species” under Vietnamese law and regulation systems including the Decree 84/2021/NĐCP, dated 22 September 2021, for strict management of wildlife and implementation of the current CITES regulations. It is also listed as a Critically Endangered species in the Vietnam Red Data Book which is still in press (V.D. Thong pers. comm. 2022, Thong and Son in press).

Thailand:
This species was listed as a “protected species” in the Thailand Wildlife Protection Act (1960, 1992) including the most recently revised version (2019). However, it was categorized as Least Concern in 2004, and that status was maintained in 2017 (S. Bumrungsri pers. comm. 2021).

Cambodia:
Under the Cambodian Law on Forestry in 2002 (Forestry Administration 2003), all wildlife species are regarded as state property and for legal purposes must be categorized as either endangered species, rare species or common species (Article 48). Pteropus vampyrus is regarded as a ‘common species’ under the related proclamation (Ministry of Agriculture, Forestry and Fisheries 2007) which consequently makes it illegal to hunt, harm or harass the taxon in ‘protected zones and special public areas’ (Article 49, 2002 Law on Forestry). However, because only one of the 12 extant flying fox roosts identified by Ravon et al. (2014) are located inside a protected area (Ang Trapeang Thmor, Banteay Meanchey province) and ‘special public areas’ are apparently undefined in practice, this means domestic law enforcement agencies have no legal mandate to address hunting of P. vampyrus (and other flying foxes also categorized as ‘common species’ such as P. lylei) at most of the other colonies currently known. As such, it would appear that religious and sociocultural prohibitions against animal hunting associated with many of the roost sites (10 colonies located on the grounds of Buddhist temples and official government buildings) and remoteness, to a lesser extent (two colonies on isolated islands), presently constitute the main deterrence to hunting of flying foxes in the country, though these are evidently less than absolute and do not apply to individuals outside of roost sites (Ravon et al. 2014).

RESEARCH NEEDS

Roost site identification and protection
There is a need to identify what constitutes the best roosting site and protect important roosting sites for this species. In view of the species’ wide range, it seems probable that it has roosts in many protected areas. However, the presence of roosts appears to be determined less by whether the area is legally protected and more by whether the area is geographically inaccessible to humans (Mildenstein 2012, S.A. Aziz per. obs. in Peninsular Malaysia 2019).

Habitat, ecology and ecosystem roles
Comprehensive dietary information is still lacking for this species across most range countries apart from the Philippines and Sarawak (Malaysian Borneo) and is necessary in order to identify important wild food sources and foraging grounds for protection. The vital ecological roles of this species in maintaining healthy ecosystems (e.g. Ong et al. 2021) can currently be inferred from studies on other Pteropus species in other countries (Aziz et al. 2021). However, these roles are still poorly understood for this species across its range, as there is insufficient research on these aspects. Such studies are urgently needed for this species in order to strengthen arguments and support for conservation action, but also to avoid functional extinction due to population declines (McConkey and Drake 2006).

Population monitoring
Regular, coordinated field surveys and long-term monitoring of known roost sites and populations should be undertaken for all countries across the species’ range, ideally, simultaneously for transboundary populations. Without such long-term population data, assessments of hunting impacts are impossible to conduct (Harrison et al. 2011).

Hunting
The effects of hunting pressure on population growth rates of fruit bats needs to be studied, so managers can identify populations in need of protection. As the seed-dispersing function of Pteropus flying foxes is known to be negatively impacted by low population abundance (McConkey and Drake 2006), any hunting assessments must also include such aspects of ecological roles in order to avoid functional extinction. Methods for protecting bats at the roost need to be developed, so bat population can rebound without disturbance. Reasons for hunting need to be better understood, so awareness campaigns can be developed aimed at changing attitudes and behaviors. If livelihoods are dependent on fruit bat harvesting, sustainable hunting programs must be pursued through research into population growth rates and methods for controlling mortality rates due to hunting. However, alternative livelihood sources should be explored first wherever possible. Hunting has not been successfully regulated by any government so far, and yet market demand for fruit bats remains high. Having legal protection for fruit bats has not proved effective in curbing hunting and persecution of this species, so more research is needed to better understand how to balance public needs with fruit bat population needs. Unless local community livelihoods are genuinely dependent on harvesting and/or consumption of flying foxes, with little-to-no alternatives available, the safest practice is banning hunting of the rapidly declining populations of this species.

Values/Perceptions of bats
There is a need to understand the specific contextual drivers of flying fox conflict, including perceptions, awareness and taboos of local communities related to their dependency on flying foxes, in order to prescribe relevant conservation actions. Social and communication psychology research is needed for interventions to encourage behavioural change, as is ecological research on the beneficial ecosystem services this species provides to humans, in parallel with research on conflict, fruit raiding, and viable mitigation methods.

Management and Legal Protection
There is also a need to revisit its legal protection status and develop conservation action plans for this species in all countries as well as coordinated across all countries in its range. Similarly, management should be coordinated at the local, national, and regional levels. Efforts should be made to have a regional-level understandings or agreements on the welfare of the species. A review should be conducted on the policies of each country at the regional and international levels e.g. CITES. Perhaps this can be included (when enough data are available) as part of the discussion on the Convention of Migratory Species/Convention on Biological Diversity/at the ASEAN level. Finally, there needs to be mechanisms for regional-level communication, collaboration and management when transboundary issues are involved.

Taxonomy
Further studies are needed to clarify the taxonomic status of populations currently allocated to this species, as P. vampyrus might represent a species complex. Collaboration needs to be initiated among countries for taxonomic studies to resolve taxonomic issues. Discussions along these lines can be initiated through the ASEAN Centre for Biodiversity which has a strong research component.