Cambodian Journal of Natural History New orchid records Ethnobotanical knowledge Carbon stocks and dynamics A homage to Pauline Dy Phon National Biodiversity Action Plan Movement of Siamese crocodiles Payments for Ecosystem Services Camera trapping of large mammals June 2017 Vol. 2017 No. 1 Cambodian Journal of Natural History Editors Email: Editor.CJNH@gmail.com • Dr Neil M. Furey, Chief Editor, Fauna & Flora International, Cambodia. • Dr Jenny C. Daltry, Senior Conservation Biologist, Fauna & Flora International, UK. • Dr Nicholas J. Souter, Mekong Case Study Manager, Conservation International, Cambodia. • Dr Ith Saveng, Project Manager, University Capacity Building Project, Fauna & Flora International, Cambodia. International Editorial Board • Dr Stephen J. Browne, Fauna & Flora International, U.K. • Dr Sovanmoly Hul, Muséum National d’Histoire Naturelle, France. • Dr Martin Fisher, Editor of Oryx – The International Journal of Conservation, U.K. • Dr Andy L. Maxwell, World Wide Fund for Nature, Cambodia. • Dr L. Lee Grismer, La Sierra University, California, USA. • Dr Brad Pettitt, Murdoch University, Australia. • Dr Knud E. Heller, Nykøbing Falster Zoo, Denmark. • Dr Campbell O. Webb, Harvard University Herbaria, USA. Other peer reviewers • Prof. Henrik Balslev, Aarhus University, Denmark. • Dr Chou Ly, Virginia Tech, USA. • Dr J.W. Duckworth, IUCN SSC Asian Species Action Partnership, UK. • Jonathan Eames, BirdLife International Cambodia Programme. • Dr Le Phat Quoi, Institute for Environment and Natural Resources, Ho Chi Minh National University, Vietnam. • Dr Benjamin Rawson, World Wide Fund For Nature, Vietnam. • Dr Sasaki Nophea, Asian Institute of Technology, Thailand. • Dr Tracy Farrell, Conservation International, Cambodia. • Dr André Schuiteman, Royal Botanic Gardens, Kew, UK. • Paul Herbertson, Fauna & Flora International, UK. • Boyd Simpson, Copenhagen Zoo, Malaysia Programme. • Dr Toyama Hironori, Kyushu University, Japan. • Ben Smitheram, Scientific Capacity Development Initiative, Cambodia. • Jeremy Holden, Independent, UK. • Josh Kempinski, Fauna & Flora International, Vietnam. • Dr Somran Suddee, Department of National Parks, Wildlife and Plant Conservation, Thailand. • Dr Kikuzawa Kihachiro, Ishikawa Prefectural University, Japan. • Dr Tagane Shuichiro, Kyushu University, Japan. • Dr Hubert Kurzweil, Singapore Botanic Gardens, Singapore. • Prof. Phan Ke Loc, Hanoi University of Science, Vietnam. • Dr Rajindra Puri, University of Kent at Canterbury, UK. • Dr Toriyama Jumpei, Forestry and Forest Products Research Institute, Japan. • Dr Tran Triet, International Crane Foundation, Southeast Asia Program. • Dr Jaap Vermeulen, J.K. Art and Science, Netherlands. • Dr Santi Watthana, Suranaree University of Tecnology, Thailand. The Cambodian Journal of Natural History (ISSN 2226–969X) is an open access journal published by the Centre for Biodiversity Conservation, Royal University of Phnom Penh. The Centre for Biodiversity Conservation is a non-profit making unit dedicated to training Cambodian biologists and to the study and conservation of Cambodian biodiversity. Cover image: Mangrove in Peam Krasaop Wildlife Sanctuary, Koh Kong Province (© Jeremy Holden/Global Wildlife Conservation). The carbon stock of peat soils in mangrove forests at the sanctuary is explored by Taing et al. in this issue (pages 55–62). Editorial Edit oria l — T he fut ure of Pa ym e nt s for Ec osyst e m Se r vic e s in Ca m bodia Virginia SIMPSON & Nicholas J. SOUTER* Conservation International, Greater Mekong Program, 4th Floor, Building E1, Phnom Penh Center, Sothearos Boulevard, Tonle Bassac, Phnom Penh, 12000, Cambodia. * Corresponding author. Email nicholas.souter@alumni.adelaide.edu.au In 2016 Cambodia’s protected area system increased significantly in size. The establishment of five new protected areas (Souter et al., 2016) and the declaration of an additional 1,427,940 ha of ‘biodiversity conservation corridors’ (RGC, 2017) increased Cambodia’s coverage of terrestrial protected areas to 42% of its land surface, up from 26% in 2014 (World Bank, 2017). This places Cambodia in the top 4% of nations worldwide in terms of the percentage of land under protection (World Bank, 2017). However, providing adequate oversight is proving difficult, with Cambodia experiencing high rates of both forest and biodiversity loss (Souter et al., 2016). Financing protected area management is a formidable task for the already under-resourced Ministry of Environment (MoE). Current levels of financial support from government and development partners are significantly below that needed (Souter et al., 2016). There are very few policies or regulations that enable collection of revenues from forests, protected or otherwise, and the revenues that are collected are remitted to the national treasury, rather than directed back into natural resource management. Private sector engagement in sustainable forest management is also very low. Consequently, Cambodia relies heavily on support from development partners, especially bilateral and multilateral donors and large NGOs, to fund protected area management. However, as continued investment by donors and NGOs is not sustainable (Souter et al., 2016), there is increasing pressure for Cambodia to devise independent, long term strategies for funding the management of its natural resources. Payments for ecosystem services (PES) offer a promising source of revenue which could be directly tied to conservation and management of Cambodia’s protected areas. PES is a financial model through which people who benefit from an ecosystem service (like water), provide financial recompense to people whose lands or activities provide that service (such as forest-dwelling communities). Cambodia’s urgent need for sustainable Cambodian Journal of Natural History 2017 (1) 1–3 finance is not the only priority that PES could help the country to address: poverty reduction, species conservation and boosting the agricultural sector are amongst the others. The people contributing to the maintenance of Cambodia’s forest ecosystems or threatened species are often among the nation’s poorest and have limited income sources. A PES programme could offer a new, continuous source of revenue and provide an alternative to non-renewable income sources such as unsustainable logging or mining. Forest ecosystems provide four major ecosystem services (Pagiola, 2008) to which PES could be applied in Cambodia: greenhouse gas mitigation, hydrological services, biodiversity conservation, and scenic vistas for recreation and tourism. PES has previously been used or assessed for some of these purposes in Cambodia. Reducing Emissions from Deforestation and Forest Degradation (REDD+) demonstration sites have been assessed in Mondulkiri, Oddar Meanchey, and Preah Vihear (Cambodia REDD+, 2017) with the aim of generating revenue through reducing greenhouse gas emissions and sequestering carbon. Carbon credits have been sold to protect the Keo Seima Wildlife Sanctuary (Wildlife Conservation Society, 2017). The Wildlife Conservation Society’s Ibis Rice programme, bird nest protection, and ecotourism programme have all improved biodiversity conservation (Clements & Milner-Gulland, 2015). Conservation International’s conservation agreement programme in the Central Cardamom Mountains has reduced deforestation (Chervier & Costedoat, 2017). Also, the hydrological services provided by the forest catchment of the Stung Atay hydro-power dam have been assessed (Fauna & Flora International, 2014). There is considerable scope to build upon these efforts and expand the scope and coverage of PES in Cambodia. Providing incentives to improve agricultural productivity and add value to Cambodian farms, including maintaining and increasing forest cover could result in medium- and long-term gains, including increasing the © Centre for Biodiversity Conservation, Phnom Penh 1 2 Editorial production of traditional (timber) and non-traditional (carbon, firewood, water and biodiversity) goods and services. The growth of Cambodia’s agricultural sector (the largest contributor to the national economy) has lagged behind that of the industrial and service sectors. This indicates a real potential for PES to improve agriculture’s contribution to GDP, which the government hopes to maintain at 7–8% per annum. PES could also help the government reach water security goals. The Rectangular Strategy for Growth, Employment, Equity and Efficiency Phase III recognizes the critical role of freshwater ecosystems for ensuring food security as well as sustaining economic activities such as hydroelectricity production and servicing a burgeoning tourism sector. Ongoing water provision through incentivized forest conservation and restoration will be critical for social security, for traditional and emerging economic activities, and for human health. A well-designed national PES scheme could also facilitate the government’s efforts to meeting its international commitments under the Convention on Biological Diversity, the United Nations Framework Convention on Climate Change, and the Sustainable Development Goals. Under the direction of Minister Say Samal, the MoE is drafting a national policy on PES, and a workplan to undertake the studies required to develop a national PES scheme. So far, two PES pilot sites have been identified for further investigation—Kbal Chhay in Sihanoukville, and Phnom Kulen in Siem Reap—both critical watersheds for important tourist destinations, and both supplying water to large beverage companies. In developing its policy, the Royal Government of Cambodia has been examining the highly successful use of PES in the small Central American nation of Costa Rica. Costa Rica’s PES programme is funded by a gas tax, a water tax, protected area entry fees and payments from hydropower operators (Pagiola, 2008). It is credited with contributing to the country’s economic success. In only 25 years Costa Rica has tripled its GDP, doubled its forest cover, and won acclaim as an ecotourism destination (Guerry et al., 2015). From 1986 to 2012, national forest cover increased from 21% to 52% (JICA, 2016), and Costa Rica has pledged to become the first carbon neutral country by 2021. In September 2016, a Cambodian government delegation, sponsored by Conservation International, led by Minister Say Samal and comprising senior officials from MoE, the Ministry of Economics and Finance and the Ministry of Agriculture, Forestry, and Fisheries, visited Costa Rica to examine its PES approach. The visit was © Centre for Biodiversity Conservation, Phnom Penh hosted by the former Costa Rican Environment Minister, Carlos Manuel Rodriguez, who shared the Costa Rican experience with the Cambodian delegation. On receiving a report of the trip Prime Minister Hun Sen responded with an official order to set Cambodia’s PES development in motion. Cambodia’s adoption of PES also needs to be informed by the experience of neighbouring Vietnam. The Government of Vietnam implemented a pilot policy framework on Payments for Forest Ecosystem Services in 2008 which aims to strengthen forest conservation, improve local livelihoods and generate external revenue for nature conservation. The policy focuses on water supply and regulation, soil conservation and landscape conservation for tourism (To et al., 2012). Buyers of ecosystem services in Vietnam include the government, and hydropower, water supply, and tourism companies (To et al., 2012; Surhardiman et al., 2013). However, PES schemes in Vietnam have been compromised by insecure land tenure, high transaction and opportunity costs, benefit-capture by local elites and lack of a market structure and other regional PES schemes (To et al., 2012; Surhardiman et al., 2013). Lack of monitoring has also made it difficult to determine whether these PES schemes have succeeded in protecting forests. Indeed, it is believed that two of the main drivers of forest degradation in Vietnam—uneven land tenure and lack of community participation in forest protection—cannot be solved by PES as it is currently practiced (McElwee, 2012). These are all problems which, unless carefully managed, could undermine PES in Cambodia. Conservation International is continuing to support the Royal Government of Cambodia’s PES programme, and the Costa Rican government has extended an invitation to develop a bilateral memorandum of understanding with Cambodia to formalize ongoing technical assistance. The road to a national scale PES scheme will be long and difficult—Costa Rica’s success took 25 years to realize—and there are lessons from neighbouring Vietnam that need to be learned. PES will only be one tool in the range of approaches needed to secure Cambodia’s natural capital. But if, in the long term, we can implement market-based incentives that result in forest and waterway conservation, and a healthy, prospering rural population, the journey will be well worth it. Re fe re nc e s Cambodia REDD+ (2017) REDD Implementation: Demonstration Projects. Http://www.cambodia-redd.org/category/reddimplementation/demonstration-projects [accessed 30 May Cambodian Journal of Natural History 2017 (1) 1–3 Editorial 2017]. Chervier, C. & Costedoat, S. (2017) Heterogeneous impact of a collective payment for environmental services scheme on reducing deforestation in Cambodia. World Development. doi: 10.1016/j.worlddev.2017.04.014. Clements, T. & Milner-Gulland, E.J. (2015) Impact of payments for environmental services and protected areas on local livelihoods and forest conservation in northern Cambodia. Conservation Biology, 29, 78–87. Fauna & Flora International (2014) A Feasibility Study Summary for an Incentives for Ecosystem Services Approach in the Stung Atay Catchment in the Cardamom Mountains, Cambodia. Http://www. ffi-spes.org/feasibility-study.html [accessed 30 May 2017]. Guerry, A.D, Polasky, S., Lubchenco, J., Chaplin-Kramer, R., Daily, G.C., Griffin, R., Ruckelshaus, M., Bateman, I.J., Duraiappah, A., Elmqvist, T., Feldman, M.W., Folke, C., Hoekstra, J., Kareiva, P.M., Keeler, B.L., Li, S., McKenzie, E., Ouyang, Z., Reyers, B., Ricketts, T.H., Rockström, J., Tallis, H. & Vira, B. (2015) Natural capital and ecosystem services informing decisions: from promise to practice. Proceedings of the National Academy of Sciences of the United States of America, 112, 7348–7355. Pagiola, S. (2008) Payments for environmental services in Costa Rica. Ecological Economics, 65, 712–714. RGC—Royal Government of Cambodia (2017) Establishing Protected Areas’ Biological Diversity Conservation Corridor System, Subdecree No. 7 Or Nor Kro Bor Kor (26 January 2017). Royal Government of Cambodia, Phnom Penh, Cambodia. Souter, N.J., Simpson, V., Mould, A., Eames, J.C., Gray, T.N.E., Sinclair, R., Farrell, T., Jurgens, J.A., & Billingsley, A. (2016) Editorial — Will the recent changes in protected area management and the creation of five new protected areas improve biodiversity conservation in Cambodia? Cambodian Journal of Natural History, 2016, 1–5. Suhardiman, D., Wichelns, D., Lestrelin, G. & Hoanh C.T. (2013) Payments for ecosystem services in Vietnam: market-based incentives or state control of resources? Ecosystem Services, 6, 64–71. To P.X., Dressler, W.H., Mahanty, S., Pham T.T. & Zingerli, C. (2012) The prospects for Payment for Ecosystem Services (PES) in Vietnam: a look at three payment schemes. Human Ecology, 40, 237–249. JICA—Japan International Cooperation Agency (2016) With Citizen Participation, Costa Rica Documents Biodiversity Nationwide. Https://www.jica.go.jp/english/news/ field/2015/160205_01.html [accessed 5 June 2017]. Wildlife Conservation Society (2017) Cambodia’s Keo Seima Wildlife Sanctuary Sells First Carbon Credits. Https://newsroom.wcs.org/News-Releases/articleType/ArticleView/ articleId/9125/Cambodias-Keo-Seima-Wildlife-SanctuarySells-First-Carbon-Credits.aspx [accessed 30 May 2017]. McElwee, P.D. (2012) Payments for environmental services as neoliberal market-based forest conservation in Vietnam: panacea or problem? Geoforum, 43, 412–426. World Bank (2017) Terrestrial Protected Areas (% of Total Land Area). Http://data.worldbank.org/indicator/ER.LND.PTLD. ZS [accessed 30 May 2017]. Cambodian Journal of Natural History 2017 (1) 1–3 © Centre for Biodiversity Conservation, Phnom Penh 3 4 A. Schuiteman et al. Shor t Com m unic at ion N e w re c ords of Orchida c e a e from Ca m bodia I V André SCHUITEMAN1,*, Rudolf JENNY2, KHOU Eang Hourt3, NAY Sikhoeun4 & ATT Sreynak4 1 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom. 2 Moosweg 9, 3112 Allmendingen, Switzerland. 3 Department of Environment, Forest and Water, National Authority for the Protection and the Development for the Cultural and Natural Site of the Preah Vihear Temple, Eco-village, Sra-Aem Commune, Choam Ksant District, Preah Vihear Province, Cambodia. 4 Department of Wildlife & Biodiversity, Forestry Administration, Ministry of Agriculture Forestry and Fisheries, 40 Preah Norodom Boulevard, Phnom Penh, Cambodia. * Corresponding author. Email a.schuiteman@kew.org Paper submitted 3 April 2017, revised manuscript accepted 27 April 2017. The previous article in this series on new orchid records from Cambodia is Schuiteman et al. (2016). We here report on eight new records, including the following three new generic records: Ania, Cylindrolobus, and Hetaeria. All were found by the authors, together with Mr Neang Thy, during field trips to western and southern Cambodia in November 2016. Two species (Cylindrolobus biflorus and Hetaeria oblongifolia) were not seen in flower in the field but flowered in cultivation at the Royal Botanic Gardens, Kew, from collected living specimens, while a third, Tropidia angulosa, was observed only as sterile specimens. In the interests of conservation we do not provide exact localities. Global distribution data follow Govaerts et al. (2017), unless indicated otherwise. Vouchers of all specimens mentioned are kept in the Kew Spirit Collection. Species recorded Ania penangiana (Hook.f.) Summerh. (voucher specimen: Schuiteman et al. 16-111; Figs 1 & 2) This terrestrial orchid was found in flower on Mt Bokor on 27 November 2016, growing in shallow humus in rocky places under trees in open forest near a waterfall at 910 m above sea level (asl). Only few specimens were seen. This is a widespread, but not common, species that has been recorded from all neighbouring countries of Cambodia, and is distributed from Northeast India to New Guinea. Cylindrolobus biflorus (Griff.) Rauschert (voucher specimen: Schuiteman et al. 16-90; Figs 3 & 4) Formerly known as Eria biflora Griff., this appears to be a relatively common species on Mt Bokor, growing on branches of small, almost shrub-like trees in open, heathlike vegetation at 1,060 m asl. The short-lived flowers were photographed in cultivation at Kew. The species is, like the previous one, widespread, occurring from NE India, through Myanmar and Indochina to Sumatra, Java and Borneo. Dendrobium metrium Kraenzl. (voucher specimen: Schuiteman et al. 16-124; Figs 5 & 6) Whereas almost all species of Dendrobium are epiphytes, D. metrium is one of very few terrestrial species. On Mt Bokor, where we found it in flower on 27 November 2016, only a few specimens were seen, growing on the ground in open, scrub-like forest on rocky outcrops with e.g., Rhododendron moulmainense Hook.f., at 980 m asl. The plants have an unusual growth habit in that new growths are often formed from the distal part of the stems, producing a somewhat scrambling habit. Seidenfaden (1992) included D. metrium (under the synonym D. sociale J.J.Sm.) in the section Pedilonum, but the papillose- CITATION: Schuiteman, A., Jenny, R., Khou E.H., Nay S. & Att S. (2017) New records of Orchidaceae from Cambodia IV. Cambodian Journal of Natural History, 2017, 4–9. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 4–9 Eight new orchid records Fig. 2 Ania penangiana (Hook.f.) Summerh., flower. Voucher specimen: Schuiteman et al. 16-111. Fig. 1 Ania penangiana (Hook.f.) Summerh., in situ. Voucher specimen: Schuiteman et al. 16-111. Fig. 3 Cylindrolobus biflorus (Griff.) Rauschert, in situ. Voucher specimen: Schuiteman et al. 16-90. Fig. 5 Dendrobium metrium Kraenzl., in situ. Voucher specimen: Schuiteman et al. 16-124. Cambodian Journal of Natural History 2017 (1) 4–9 Fig. 4 Cylindrolobus biflorus (Griff.) Rauschert, flowering stem. Voucher specimen: Schuiteman et al. 16-90. © Centre for Biodiversity Conservation, Phnom Penh 5 6 A. Schuiteman et al. Fig. 6 Dendrobium metrium Kraenzl., flower. Voucher specimen: Schuiteman et al. 16-124. Fig. 8 Eulophia graminea Lindl., flower. Voucher specimen: Schuiteman et al. 16-57. Fig. 7 Eulophia graminea Lindl., in situ. Voucher specimen: Schuiteman et al. 16-57. Fig. 9 Habenaria hosseusii Schltr., in situ. Voucher specimen: Schuiteman et al. 16-0. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 4–9 Eight new orchid records Fig. 10 Habenaria hosseusii Schltr., flower. Voucher specimen: Schuiteman et al. 16-0. Fig. 11 Hetaeria oblongifolia Blume, flowers. Voucher specimen: Kew cult. 2016-2586. Fig. 12 Hetaeria oblongifolia Blume, in situ. Voucher specimen: Kew cult. 2016-2586. Fig. 13 Phaius indochinensis Seidenf. & Ormerod, flowers. Voucher specimen: Schuiteman et al. 16-51. Cambodian Journal of Natural History 2017 (1) 4–9 © Centre for Biodiversity Conservation, Phnom Penh 7 8 A. Schuiteman et al. Eulophia graminea Lindl. (voucher Schuiteman et al. 16-57; Figs 7 & 8) specimen: In general, this is a common, even somewhat weedy species of open, disturbed places, although it also occurs in closed forest (Pedersen et al., 2014). Outside its natural range it is naturalized in Australia, South Africa and USA (Pemberton et al., 2008). We found this species in humid, old-growth, evergreen forest in Pursat Province near Pramaoy, at 350 m asl. Only a single specimen was seen in flower, on 24 November 2016. It occurs over much of tropical Asia, from Pakistan to China, as well as Sumatra, Java, Borneo and the Philippines. Eulophia graminea was already listed as occurring in Cambodia by Govaerts et al. (2017), but it was not recorded from Cambodia by Seidenfaden (1992). As we have not seen a reference to an exact locality for Cambodia, the present record may be the first from a known locality in this country. Habenaria hosseusii Schltr. (voucher Schuiteman et al. 16-0; Figs 9 & 10) Fig. 14 Phaius indochinensis Seidenf. & Ormerod, in situ. Voucher specimen: Schuiteman et al. 16-51. specimen: Dozens of flowering specimens of this long-spurred orchid were found on 20 November 2016 on an eroded limestone hill near Takream Village in Battambang Province, at 150 m asl. Two other species of Habenaria were also observed in flower there: H. lindleyana Steud. and H. dentata (Sw.) Schltr. The vegetation was an open, secondary forest of young trees without epiphytes, and there were signs of burning. The present species differs in minute details from H. dentirostrata Tang & F.T.Wang, which has already been recorded from Cambodia (Leti et al., 2013), and the two species may in fact be conspecific (Kurzweil, 2009). Habenaria hosseusii has been recorded from Thailand and Laos. Hetaeria oblongifolia Blume (voucher specimen: Kew cult. 2016-2586; Figs 11 & 12) Fig. 15 Tropidia angulosa (Lindl.) Blume, in situ. pubescent adaxial surface of the lip is more typical of the section Dendrobium, to which it may well belong. So far, this species has not been included in any phylogenetic analyses. It was previously recorded from Thailand, Vietnam, Peninsular Malaysia, and Sumatra, but in spite of its relatively wide distribution it appears to be a rare species (Averyanov, 2012). © Centre for Biodiversity Conservation, Phnom Penh The flowers of this inconspicuous terrestrial orchid are only about 3.5 mm long. As in all species of the genus Hetaeria, they are not resupinated, which means that the lip is held uppermost; in most other orchids, the lip is held lowermost. Like Eulophia graminea mentioned above, this species was found in humid, old-growth, evergreen forest in Pursat Province near Pramaoy, at 350 m asl. It is widespread throughout tropical and subtropical Asia, from Sri Lanka to Japan, and south-eastwards to Australia and New Caledonia. Phaius indochinensis Seidenf. & Ormerod (voucher specimen: Schuiteman et al. 16-51; Figs 13 & 14) This is undoubtedly the showiest orchid reported in the present paper, with flowers 6–8 cm across. It proved to be common in humid evergreen montane forest in Pursat Province, about 24 km SW of Pramoy, at 870 m Cambodian Journal of Natural History 2017 (1) 4–9 Eight new orchid records asl, growing in leaf litter in dense shade. Only two or three specimens out of hundreds were seen in flower on 23 November 2016. It often grew together with Plocoglottis bokorensis Seidenf., which is superficially similar in appearance when not in flower. However, the latter has distinct, ovoid pseudobulbs at the base of the slender stems, whereas the stems of P. indochinensis are uniformly terete. Phaius indochinensis is also known from Laos (Schuiteman et al., 2008), Thailand and Vietnam, and has been misidentified as P. indigofer Hassk. (Seidenfaden, 1992, as “indigoferus”), a species that probably does not occur in Thailand and Indochina. Tropidia angulosa (Lindl.) Blume (not vouchered; Fig. 15) Several specimens, long past flowering, were discovered not far from a population of Hetaeria oblongifolia, mentioned above. A few living specimens that we collected for cultivation did not survive for more than a few weeks, with the stems rapidly turning black. It has been our experience that, unlike most orchids, Tropidia does not respond well to being bare-rooted. Consequently, it is difficult to bring the species into cultivation from wild-collected material. The present species, with its two broad, almost opposite leaves and short terminal raceme, has been reported from all of Cambodia’s neighbouring countries, and there can be little doubt that the plant we photographed is indeed T. angulosa. A good photograph of a flowering specimen can be found in Seidenfaden (1992) and Averyanov (2008). The species is widely distributed from Northeast India to South Japan, and south to Java, Bali and the Philippines. It is common in Vietnam in “all kinds of lowland and submontane forests on any soils. 0–1,600 m” (Averyanov, 2008). Conclusions Fieldwork in almost any area in Cambodia where there is still natural vegetation left reveals new orchid records. At the same time, suitable habitats are still being destroyed, and there can be little doubt that species have already been lost before they could be recorded. Most of the limestone hills that we visited in Battambang Province showed signs of heavy disturbance, with little if any of the original forest cover left. We can only suspect that sensitive and endemic species may have disappeared from these hills, which are entirely surrounded by cultivated land. We will probably never know. Cambodian Journal of Natural History 2017 (1) 4–9 Ack now le dge m e nt s We thank the Director of the Department of Terrestrial Protected Areas, Ministry of Environment, Cambodia, for granting us access to protected areas, and Dr Keo Omaliss of the Cambodia Forestry Administration, for his invaluable help before and during our visit. We are grateful to Mr Neang Thy (MoE) for his assistance in the field. André Schuiteman received a travel grant from the Bentham-Moxon Trust. We are grateful to the CITES authorities in Cambodia and UK for providing the necessary permits. The living specimens were imported into the UK under Defra Plant Health Licence Number 2149/194627/4. All photos were taken by André Schuiteman. Re fe re nc e s Averyanov, L.V. (2008) The orchids of Vietnam illustrated survey, part 1: subfamilies Apostasioideae, Cypripedioideae and Spiranthoideae. Turczaninowia, 11, 5–168. Averyanov, L.V. (2012) New orchid taxa and records in the flora of Vietnam. Taiwania, 57, 127–152. Govaerts, R., Bernet, P., Kratochvil, K., Gerlach, G., Carr, G., Alrich, P., Pridgeon, A.M., Pfahl, J., Campacci, M.A., Holland Baptista, D., Tigges, H., Shaw, J., Cribb, P., George, A., Kreuz, K. & Wood, J.J. (2017) World Checklist of Orchidaceae. Facilitated by the Royal Botanic Gardens, Kew. Http://apps.kew. org/wcsp/ [Last accessed 3 April 2017]. Kurzweil, H. (2009) The genus Habenaria (Orchidaceae) in Thailand. Thai Forest Bulletin (Botany), special issue, 7–105. Leti, M., Hul S., Fouché, J.-G., Chéng S.K. & David, B. (2013) Flore Photographique du Cambodge. Editions Privat, France. Pedersen, H.Æ., Kurzweil, H., Suddee, S., de Vogel, E.F., Cribb, P.J., Chantanaorrapint, S., Watthana, S., Gale, S.W., Seelanan, T. & Suwanphakdee, C. (2014) Flora of Thailand, Volume 12, Part 2: Orchidaceae 2 (Epidendroideae P.P.: Neottieae, Tropideae, Nervilieae, Gastrodieae, Thaieae, Calypsoeae, Arethuseae, Collabieae, Cymbidieae). The Forest Herbarium, Royal Forest Department, Bangkok, Thailand. Pemberton, R.W., Collins, T.M. & Koptur, S. (2008) An Asian orchid, Eulophia graminea (Orchidaceae: Cymbidieae), naturalizes in Florida. Lankesteriana, 8, 5–14. Schuiteman, A., Bonnet, P., Svengsuksa, B. & Barthélémy, D. (2008) An annotated checklist of the Orchidaceae of Laos. Nordic Journal of Botany, 26, 257–314. Schuiteman, A., Ryan, C., Nut M., Nay S., & Att, S. (2016) New records of Orchidaceae from Cambodia III. Cambodian Journal of Natural History, 2016, 84–89. Seidenfaden, G. (1992) The orchids of Indochina. Opera Botanica, 114, 1–502. © Centre for Biodiversity Conservation, Phnom Penh 9 10 Suetsugu K. et al. Shor t Com m unic at ion Ga st rodia ex ilis (Orchida c e a e ), a ne w ly re c orde d m yc ohe t e rot rophic ge nus a nd spe c ie s in Ca m bodia SUETSUGU Kenji1,*, HSU Tian-Chuan2, TAGANE Shuichiro3, CHHANG Phourin4 & YAHARA Tetsukazu3 1 Department of Biology, Graduate School of Science, Kobe University, 1-1 Rokkodai, Nada, Kobe, 657-8501, Japan. 2 Herbarium of Taiwan Forestry Research Institute, No. 53, Nanhai Road, Taipei 100, Taiwan. 3 Center for Asian Conservation Ecology, Kyushu University, 744 Motooka, Fukuoka, 819-0395, Japan. 4 Institute of Forest and Wildlife Research and Development, Forestry Administration, 40 Preah Norodom Boulevard, Phnom Penh, Cambodia. * Corresponding author. Email kenji.suetsugu@gmail.com Paper submitted 26 December 2016, revised manuscript accepted 18 February 2017. The genus Gastrodia (Orchidaceae) comprises approximately 90 species of mycoheterotrophic orchids distributed throughout the temperate and tropical regions of Asia, Oceania, Madagascar and Africa (Govaerts et al., 2016; Hsu et al., 2016). Within Indochina, Gastrodia has been recorded from Thailand and Vietnam but not yet from Cambodia and Laos (Schuiteman & de Vogel, 2000; Suddee & Harwood 2009; Averyanov, 2011; Suddee, 2014; Schuiteman et al., 2015). The genus is characterised by fleshy tubers, the absence of leaves, as well as the fusion of its sepals and petals, and the presence of two mealy pollinia that lack caudicles. As is the case in most other mycoheterotrophic species, most Gastrodia species occur in small populations and appear above the soil surface only during their brief reproductive period (Suetsugu, 2016). Therefore, the diversity and distribution patterns of Gastrodia species are easily underestimated. Our recent botanical surveys have resulted in the discovery of many new species and distribution records for this genus (e.g., Suetsugu, 2012, 2013, 2014, 2015, 2016; Hsu & Kuo, 2010, 2011; Hsu et al., 2012, 2016). We document the occurrence of Gastrodia exilis here as a newly recorded genus and species for Cambodia. The following description is based on our Cambodian material. Gastrodia exilis Hook.f., The Flora of British India, 6, 123 (1890) Gastrodia siamensis Rolfe ex Downie, Bulletin of Miscellaneous Information, 1925, 416 (1925); Gastrodia hayatae Tuyama, Journal of Japanese Botany, 17, 580 (1941). Terrestrial, mycoheterotrophic herb. Rhizome tuberous, fusiform or cylindrical, 7–12 mm long, 3–5 mm in diameter. Inflorescence erect, white to pale brown, glabrous, 14–17 cm long, 1.5 mm in diameter, with 4–6 nodes, with tubular, membranous sheaths. Bracts up to 3 mm long, 1.5 mm wide. Pedicel and ovary up to 10 mm long. Flowers 2–4, 7–9 mm long, white, tubular, slightly turned upwards, resupinate. Sepals and petals connate, forming a five-lobed perianth tube. Sepals ca. 7 mm long, connate basally 3/4–4/5 their length with petals, lateral ones connate basally 2/3–3/4 with each other; free tips of sepals ovate-triangular, ca. 2 mm long, 2 mm wide, apex obtuse, margins slightly undulate; free tip of petals ovate-triangular, ca. 1.5 mm long, 1.5 mm wide, apex obtuse, margins slightly undulate. Lip 4–4.5 mm long, 1.5–2.0 mm wide, hypochile with two globose calli; epichile ovate-elliptic, base contracted, disc 5-ridged with two parallel erect longitudinal keels extending toward apex in the distal half, margin slightly undulate. Column straight, semi-cylindrical, 4–4.5 mm long, white; CITATION: Suetsugu K., Hsu T.C., Tagane S., Chhang P. & Yahara T. (2017) Gastrodia exilis (Orchidaceae), a newly recorded mycoheterotrophic genus and species in Cambodia. Cambodian Journal of Natural History, 2017, 10–13. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 10–13 A new orchid record A B C Fig. 1 Flowering and fruiting plant of Gastrodia exilis in Bokor National Park. A) Flowering plant, B) Flowers, C) Immature fruit. lateral wings narrow, the edges parallel to column, acute; rostellum absent. Anther hemispherical, pollinia 2. Specimen examined: Cambodia, Kampot Province, Bokor National Park, evergreen forest around sphagnum bog, 10°39’9.06”N, 104°03’38.68”E, alt. 935 m, 17 October 2012, Tagane S., Fuse K., Choeung HN 4275 (deposited in the Forestry Administration herbarium in Cambodia). Distribution: Cambodia (new record), India, Thailand and Sumatra. Habitat and ecology: Fewer than 10 individuals were found in a small population in the wet understorey of tropical lower montane forest on the plateau of Bokor National Park. The site is close to the collection locality of Aphyllorchis pallida Blume and dominant trees were reported by Tagane et al. (2015). The flowering time is in early to mid-October. Conservation status: While Gastrodia exilis has not been assessed by the IUCN Red List of Threatened Species (IUCN, 2016), this species has been reported from only a few scattered localities in India, Thailand, and Sumatra (Hooker, 1890; Downie, 1925; Tuyama, 1941; Seidenfaden, 1978; Joseph et al., 1980; Kumar & Kumar, 2001; Suddee, 2004). In addition, in Cambodia the species is only known from Mt Bokor (see also Note 2 below), where the aforemenCambodian Journal of Natural History 2017 (1) 10–13 tioned specimen was collected. Given that mycoheterotrophic plants are highly dependent on the activities of both the fungi and the trees that sustain them (e.g., Suetsugu et al., 2014), they are particularly sensitive to environmental disturbance. Because deforestation for resort development is rapidly expanding very near to the locality of this species on Mt Bokor, urgent attention is needed to conserve the Cambodian population. This is also in the interest of many other rare plants growing nearby, such as Nepenthes bokorensis Mey and Dipodium paludosum (Griff.) Rchb.f. Notes: 1) Compared with published descriptions and illustrations (Hooker, 1890; Downie, 1925; Tuyama, 1941; Seidenfaden, 1978, Joseph et al., 1980; Kumar & Kumar, 2001; Pedersen et al., 2004), the Cambodian plants we observed were smaller than those from previously reported localities (e.g., India and Thailand), producing shorter rhizomes (0.7–1.2 cm vs. 1.0–5.5 cm), shorter inflorescences (14–17 cm vs. 15.5–112 cm) and smaller number of flowers per inflorescence (2–4 vs. 3–25). However, there are no significant differences in the morphology of the lip and column, which are the most important characters used to classify Gastrodia species. Furthermore, given that plant size can be dramatically affected by the availability of nutrients, it would not be surprising to observe variability in plant size among different popula© Centre for Biodiversity Conservation, Phnom Penh 11 12 Suetsugu K. et al. tions reflecting their different nutritional resources, such as the activity of their mycorrhizal fungi. Therefore, we consider that the differences most likely represent an example of intraspecific variation. The overlap in the size range of plants between the Cambodian and previously reported populations supports this assumption. 2) On 27 November 2016, Dr André Schuiteman and co-workers found another population of G. exilis nearby (in evergreen forest near Popokvil Waterfall): this consisted of at least a dozen plants, most growing close together, but some about 100 m away from the others. Most were in fruit, but one specimen still had flowers (voucher specimen: Schuiteman et al. 16-120A, spirit mat. K; Schuiteman, pers. comm.). Ack now le dge m e nt s The authors would like to thank the Cambodian Ministry of the Environment and Ministry of Agriculture, Forestry and Fisheries for permitting our botanical inventories in Bokor National Park and staff from these organizations who assisted our field surveys. This study was supported by the Environment Research and Technology Development Fund (S9) of the Japanese Ministry of the Environment and also by a JSPS grant from the Global Center of Excellence Program ‘Asian Conservation Ecology as a basis of human-nature mutualism’. We also wish to thank Dr André Schuiteman and two anonymous reviewers for their constructive comments on an earlier version of the manuscript. Re fe re nc e s Averyanov, L.V. (2011) The orchids of Vietnam illustrated survey. Part 3. Subfamily Epidendroideae (primitive tribes— Neottieae, Vanilleae, Gastrodieae, Nervilieae). Turczaninowia, 14, 15–100. Downie, D.G. (1925) Contributions to the flora of Siam. Bulletin of Miscellaneous Information, 1925, 404–423. Govaerts, R., Bernet, P., Kratochvil, K., Gerlach, G., Carr, G., Alrich, P., Pridgeon, A.M., Pfahl, J., Campacci, M.A., Holland Baptista, D., Tigges, H., Shaw, J., Cribb, P.J., George, A., Kreuz, K. & Wood, J.J. (2016) World Checklist of Orchidaceae. Royal Botanic Gardens, Kew. Http://apps.kew.org/wcsp/. [accessed on 14 December 2016]. Hooker, J.D. (ed) (1890) The Flora of British India 6: 220. L. Reeve & Co, London, U.K. Hsu T.C. & Kuo C.M. (2010) Supplements to the orchid flora of Taiwan (IV): four additions to the genus Gastrodia. Taiwania, 55, 243–248. Hsu T.C. & Kuo C.M. (2011) Gastrodia albida (Orchidaceae), a new species from Taiwan. Annales Botanici Fennici, 48, 272–275. © Centre for Biodiversity Conservation, Phnom Penh Hsu T.C., Chung S.W. & Kuo C.M. (2012) Supplements to the orchid flora of Taiwan (VI). Taiwania, 57, 271–277. Hsu T.C., Fanerii, M., Yang T.Y.A., Pitisopa, F. & Li C.W. (2016) Gastrodia isabelensis and G. solomonensis (Gastrodieae, Epidendroideae, Orchidaceae): two new species representing a new generic record in the Solomon Islands. Phytotaxa, 270, 137–145. IUCN (2016) The IUCN Red List of Threatened Species. Version 2014.2. Http://www.iucnredlist.org [accessed 10 November 2016]. Joseph, J., Abbareddy, N.R. & Haridasan, K. (1980) Gastrodia exilis Hook.f.—a rare and interesting orchid from Khasi and Jaintia Hills, Meghalaya, India. Nelumbo, 22, 203–205. Pedersen, H.A., Watthana, S., Suddee, S. & Sasurat, S. (2004) Breeding system, post-pollination growth, and seed dispersal in Gastrodia exilis (Orchidaceae). Natural History Bulletin of the Siam Society, 52, 9–26. Sathish Kumar, C. & Suresh Kumar, P.C. (2001) Gastrodia exilis Hook.f. (Orchidaceae), a new genus and species record for South India. Rheedea, 11, 49–52. Schuiteman, A. & de Vogel, E.F. (2000) Orchid Genera of Thailand, Laos, Cambodia and Vietnam. Nationaal Herbarium Nederland, Leiden, The Netherlands. Schuiteman, A., Ryan, C. & Nut, M. (2015) New records of Orchidaceae from Cambodia I. Cambodian Journal of Natural History, 2015, 131–138. Seidenfaden, G. (1978) Orchid genera in Thailand VI. Neottioideae Lindl. Dansk Botanisk Arkiv, 32, 1–195. Suddee, S. (2014) Gastrodia. In Flora of Thailand 12(2) (eds H. Pendersen, H. Kurzweil, S. Suddee, E.F. de Vogel, P.I. Cribb, S. Chantanaorrapint, G. Watthana, S.W. Gale, T. Seelanan & C. Suwanphakdee), pp. 525–531. The Forest Herbarium, Department of National Parks, Wildlife and Plant Conservation, Bangkok, Thailand. Suddee, S. & Harwood, B. (2009) Gastrodia verrucosa (Orchidaceae), a new, but not unexpected, record for Thailand. Thai Forest Bulletin (Botany), 37, 144–146. Suetsugu K. (2012) A new form of Gastrodia confusa (Orchidaceae). Journal of Phytogeography and Taxonomy, 59, 125–126. Suetsugu K. (2013) Gastrodia takesimensis (Orchidaceae), A new mycoheterotrophic species from Japan. Annales Botanici Fennici, 50, 375–378. Suetsugu K. (2014) Gastrodia flexistyloides (Orchidaceae), a new mycoheterotrophic plant with complete cleistogamy from Japan. Phytotaxa, 175, 270–274. Suetsugu K. (2015) First record of the mycoheterotrophic orchid Gastrodia uraiensis (Orchidaceae) from Yakushima Island, Japan. Acta Phytotaxonomica et Geobotanica, 66, 193–196. Suetsugu K. (2016) Gastrodia kuroshimensis (Orchidaceae), a new mycoheterotrophic and complete cleistogamous plant from Japan. Phytotaxa, 278, 265–272. Suetsugu K., Kawakita A. & Kato M. (2014) Evidence for specificity to Glomus group Ab in two Asian mycoheterotrophic Burmannia species. Plant Species Biology, 29, 57–64. Cambodian Journal of Natural History 2017 (1) 10–13 A new orchid record Tagane S., Yukawa T., Chhang P., Ogura-Tsujita Y., Toyama H. & Yahara T. (2015) A new record of Aphyllorchis pallida (Orchidaceae) from Cambodia. Cambodian Journal of Natural History, Cambodian Journal of Natural History 2017 (1) 10–13 2015, 128–130. Tuyama T. (1941) Notes on the genus Gastrodia of Southeastern Asia. Journal of Japanese Botany, 17, 579–586. © Centre for Biodiversity Conservation, Phnom Penh 13 14 M. Maltby et al. Shor t Com m unic at ion Revision of Ca m bodia ’s N at iona l Biodive rsit y St rat e gy & Ac t ion Pla n t o int e grat e c onse r vat ion a nd sust a ina ble use of biologic a l re sourc e s int o nat iona l de c ision m a k ing Matthew P. MALTBY1,*, CHAN Somaly2 & Jady SMITH3 1 Forestry & Natural Resource Management Unit, Winrock International, 2121 Crystal Drive, Suite 500, Arlington, Virginia 22202, USA. 2 National Council for Sustainable Development, Ministry of Environment, Morodok Techo Building (Lot 503), Tonle Bassac, Chamkarmorn, Phnom Penh, Cambodia. 3 Live & Learn Environmental Education, Ross House, 4th Floor, 247–251 Flinders Lane, Melbourne Victoria 3000, Australia. * Corresponding author. Email mmaltby82@gmail.com Paper submitted 7 April 2017, revised manuscript accepted 13 June 2017. Cambodia became a signatory and acceded to the United Nations Convention on Biological Diversity (CBD) in 1995 (CBD, 2017a), under which the country is obligated to develop and adopt a National Biodiversity Strategy & Action Plan (NBSAP). An NBSAP is the principal means of implementation of the CBD at a national level (CBD, 2017b). It reflects how Cambodia intends to fulfil the objectives of the CBD in light of its specific national circumstances, and sets out a sequence of steps to meet these goals. The present NBSAP for Cambodia includes the following vision for biodiversity: “Equitable economic prosperity and improved quality of life through sustainable use, protection and management of biological resources”; and a mission statement, namely “To use, protect and manage biodiversity for sustainable development in Cambodia” (MoE, 2016a). The first NBSAP for Cambodia appeared in 2002 (MoE, 2002). Since then, knowledge of the country’s natural history has expanded significantly. A combination of applied research, new species discoveries and far-ranging biodiversity conservation programmes have yielded a wealth of new information on the country’s flora and fauna, and great progress has been made in developing national capacity and human resources for biodiversity conservation. On 5th February 2016, The Royal Government of Cambodia approved the country’s updated NBSAP (Fig. 1). To meet the complex and often changing challenges facing biodiversity and ecosystem conservation, the essence of a NBSAP is to create a policy requirement that integrates consideration of conservation and sustainable use of biological resources into national decision-making across all sectors of the national economy and policymaking framework. In parallel to the NBSAP, the National Biodiversity Status Report (MoE, 2016b) was also updated for the first time since 2001. The status report provides a snapshot of the current status of biodiversity in Cambodia, and inventory lists of all species known in major groups (including 162 mammals, 601 birds, 173 reptiles, 72 amphibians, 1,357 fish, 3,113 plants and 671 invertebrates), plant and animal genetic resources, and ecosystem diversity. Verified species lists are now held by the Department of Biodiversity of the National Council for Sustainable Development (Ministry of Environment), and details are available through the National Clearing House Mechanism website (MoE, 2017b). The clearing house mechanism provides information services to facilitate implementation of the NBSAP, which in turn supports CITATION: Maltby, M.P., Chan S. & Smith, J. (2017) Revision of Cambodia’s National Biodiversity Strategy & Action Plan to integrate conservation and sustainable use of biological resources into national decision making. Cambodian Journal of Natural History, 2017, 14–16. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 14–16 National Biodiversity Strategy & Action Plan implementation of the global Strategic Plan for Biodiversity 2011–2020 (CBD, 2017b). The newly updated NBSAP reflects how Cambodia intends to fulfil the three objectives of the CBD in light of its specific national circumstances and governmental reform, and identifies steps to meet these goals. The purpose of this article is to inform a wider audience of the significance and importance of Cambodia’s NBSAP and to briefly outline the key thematic objectives which Cambodia aims to meet over the coming years. Purpose of the NBSAP The CBD is a binding agreement that obligates countries to ensure they are conserving biodiversity, which includes having effective national biodiversity planning. Cambodia’s new NBSAP also constitutes a direct contribution to Aichi Biodiversity Target 17 under the CBD, which states that by 2015 each Party should have developed, adopted as a policy instrument, and commenced implementing an effective, participatory and updated NBSAP. Indeed, Cambodia was one of the few 150+ signatories to the convention to update its NBSAP prior to the 21st Conference of Parties in Paris, 2015. Following extensive inter-ministerial meetings, national and sub-national consultations and technical review by national and international experts, the updated NBSAP now includes plans for the conservation and sustainable use of Cambodia’s biological diversity. These plans are integrated with relevant sectoral and cross-sectoral programmes, related national policies and also provide significant contributions to national sustainable development goals. Relevance to biodiversity stakeholders The NBSAP provides a framework for action at all levels to ensure the productivity, diversity and integrity of natural systems and, as a result, Cambodia’s ability as a nation to develop sustainably. It promotes the conservation of biodiversity and ecosystems, the sustainable use of biological resources and describes national contributions to international efforts to implement the CBD, including 20 Aichi Biodiversity Targets. The NBSAP lists priority actions to be undertaken by the various ministries, departments and agencies during the implementation phase. A series of strategic objectives and priority actions are presented under 24 themes, including: Protection of Natural Resources (Protected areas, Threatened species, Ex situ Conservation); Animal Cambodian Journal of Natural History 2017 (1) 14–16 Fig. 1 Cambodia’s updated National Biodiversity Strategy and Action Plan. Wildlife Resources; Freshwater Fisheries and Aquaculture; Coastal and Marine Resources; Forest and Wild Plant Resources; Agriculture and Animal Production; Energy Resources; Mineral Resources; Industry, Technology and Services (Manufacturing, Biotechnology and Biosafety, Tourism); Environmental Security; Land Use Planning; Water Resources; Climate Change and Biodiversity; Community Participation; Awareness, Education, Research Coordination and Development; Legislation and Institutional Structures; and Quality of Life and Poverty Reduction (MoE, 2016a). Priority actions adopted by the government are grouped into three broad categories: i) Actions promoting awareness and building the capacity of government staff and local communities; ii) Actions promoting community-based natural resource management; and, iii) Actions aimed at clarifying ministerial jurisdictions, reducing responsibility overlap and promoting interministerial coordination and collaboration in a sustainable development perspective (MoE, 2016a). © Centre for Biodiversity Conservation, Phnom Penh 15 16 M. Maltby et al. Monitoring and evaluation Implementation of the NBSAP is an ongoing, continuous and cyclical process. Proposed mechanisms for implementation are identified in the NBSAP, such as coordination of national and international elements of the strategy through a permanent Inter-ministerial Biodiversity Steering Committee (IBSC) and National Secretariat for Biodiversity. However, as with any policy initiative, rigorous monitoring and evaluation must accompany implementation to measure progress towards overall goals. Long-term success will be determined by the degree to which all parts of society adopt the NBSAP vision and principles and contribute to achieving its goals. The IBSC is charged with monitoring, evaluation and reporting of progress towards NBSAP targets. Reporting can take place annually and also as a part of national reports to the CBD, the next of which is due in 2018. While 20 biodiversity targets have been adopted, baseline information still needs to be collected to allow accurate reporting and with this in mind, the IBSC will focus its efforts on: i) Developing a strategy for obtaining information from various NGOs and government departments; ii) Strengthening technical skills of nationals on management, collection and processing of data; and, iii) Ensuring complementarity of NBSAP monitoring and evaluation efforts with other national monitoring activities to avoid duplication of effort. Ultimately, conservation of biodiversity and sustainable use of biological resources will require the support and participation of individual citizens, local communities, urban and regional governments, conservation groups, business and industry, and educational and research institutions. In addition to regular reporting requirements and biennial revision of the NBSAP, an actions and indicators matrix has been developed to support evaluation of effectiveness and can be found in Appendix I of the full document. Looking forward The strategic objectives (indicators) and associated priority actions under each NBSAP theme serve as a valuable planning tool and guide for conservation practitioners, policymakers and stakeholders alike. Indeed, the NBSAP provides a national mandate for the establishment of numerous new priority initiatives, as well as continuation and enhancement of many existing longterm conservation activities. © Centre for Biodiversity Conservation, Phnom Penh It should remain a priority of the Royal Government of Cambodia, development partners and NGOs to revise the NBSAP periodically, ideally every two years, to ensure that biodiversity data are current and that action plans remain appropriate as threats and challenges to biodiversity protection evolve over time. In particular, the recent environmental reforms in 2016, such as the proclamation of numerous new protected areas, should be a priority to include in future NBSAPs. The complete text of Cambodia’s NBSAP can be found online at https:// www.cbd.int/doc/world/kh/kh-nbsap-v2-en.pdf. Ack now le dge m e nt s The revision of the NBSAP was conducted by the Department of Biodiversity of the National Council for Sustainable Development (Ministry of Environment) and supported by development partners United States Agency for International Development and Global Environment Facility through the United Nations Environment Programme. Special thanks must go to the Minister of Environment H.E. Say Samal for his political and technical support; to the team of consultants who supported NBSAP development, including Mr Jo Mulongoy, Ms Ken Bopreang and Mr Ou Ratanak; and all individuals and organizations who shared biodiversity data and technical reports. Re fe re nc e s MoE—Ministry of Environment (2002) National Biodiversity Strategy and Action Plan. Http://www.chm.gdancp-moe.org/ convention-and-policy/cambodia-nbsap.html [accessed 15 March 2017]. MoE—Ministry of Environment (2016a) National Biodiversity Strategy and Action Plan, February 2016. Https://www.cbd.int/ doc/world/kh/kh-nbsap-v2-en.pdf [accessed 4 April 2017]. MoE—Ministry of Environment (2016b) National Biodiversity Status Report, February 2016. Http://www.chm.gdancp-moe. org/publications/national-reports.html [accessed 10 March 2017]. MoE—Ministry of Environment (2017b) Cambodia Clearing House Mechanism. Http://www.chm.gdancp-moe.org/ [accessed 31 March 2017]. CBD—Convention on Biological Diversity (2017a) Treaty State Description. Https://www.cbd.int/world/ratification.shtml [accessed 15 March 2017]. CBD—Convention on Biological Diversity (2017b) ClearingHouse Mechanism. Https://www.cbd.int/chm/ [accessed 14 May 2017]. Cambodian Journal of Natural History 2017 (1) 14–16 Flora of Bokor Plateau T he fl ora of t he Bokor Plat e a u, sout he a st e r n Ca m bodia : a hom a ge t o Pa uline Dy Phon Philip W. RUNDEL1,* & David J. MIDDLETON2 1 Department of Ecology and Evolutionary Biology, University of California, Los Angeles, California 90095, USA. 2 Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore 259569, Singapore. * Corresponding author. Email rundel@biology.ucla.edu Paper submitted 7 April 2017, revised manuscript accepted 6 June 2017. ɊɮɍɅʂɋɑɳȶſɆ ȳƕȶɽǍɆɆɮȲɳƵɑƏɩɁɳǷNJȴǕɳȴƒɋɿɵɅƙɆɳɃɑȲɊƕɭƺ ƺɃɪȹƙɊȲɌɆɑɽƙɆɳɉɃɌɭȲƺ ſ ɁɩȲƙɊɅɩȶɑɒȴɊɅɿɌȲ ɭ ƺ ſ ɁɩɑɸƴɅɽʉɵɅɵƙɈƙȲɩɅ ɅɩȶɵƙɈɍɮɁɌɒʂɑ ƙɈɊDŽɸȶNjɅɌɭȲſƺɁɩɑɊƓɮɌɴɆɆʆ ɈƙȶɪȲɆɴɅƏɊɳɍˊƳɌƷɌɌɆɑɽ PaulineȱDyȱPhon ɴȼɍNjɅǕɋɭƳɍƺȶ ljȲɽȲǁ Ǝ ɍɑɁɎɁƞȲɅƚȶɊȲɳɒˊɋ ɳɋˊȶLJɅɳɄƛˊɆȷƃɭɆƓɅƒNJɈɳɍˊɌȲ ɭ ſƺɁɩNjɅɑɌɵɑdžɸɳǷɁɸɆɅɽȳȶ ƕ ɽǍɆɳɅɹʆ ɆȥƅɪȲɸɀɁɽƙǂɌɭȲſ ƺɁɩɌɆɑɽɳɋˊȶNjɅʓʕʙƙɆɳɉɃ ɴȼɍʒʒƙɆɳɉɃƺƙɆɳɉɃɴȼɍNjɅɴɁȲƒɭȶɁɸɆɅɽɆɮȲɳƵɆɻɭɳǁƍɹʆ Abst ra c t The Bokor Plateau in southeastern Cambodia is home to rare and significant plant communities of stunted forest and heathland, as well as a rich flora. Expanding on the pioneering work of Pauline Dy Phon more than half a century ago, we update the current knowledge of the vascular plant flora of the plateau. Our checklist includes 359 species, with 22 of these endemic to Bokor. Ke yw ords Bokor Plateau, Cambodia, stunted forest, heathland, Preah Monivong National Park, endemic species. I nt roduc t ion Bokor National Park (officially known as Preah Monivong National Park) in southeastern Cambodia represents a biodiversity hotspot with a rich plant diversity and a high level of endemism. Within this park the sandstone massif of the Elephant Mountains rises very steeply from a narrow coastal plain along the Gulf of Thailand to an elevation of 1,080 m (Fig. 1). The combination of the steep south-facing slopes of the range and close proximity of the ocean produces unusually wet conditions on the southwestern slopes and upper plateau. Tall species-rich wet forests are present on the lower and middle elevation slopes at Bokor. However, as with tropical montane forests in many other areas of the world, the shallow soils and water-logged depressions at higher elevations on the gently sloping Bokor Plateau exhibit dwarf forests with relatively low sclerophyllous evergreen trees (Dy Phon, 1970; Rundel et al., 2016). A complex interaction of high winds, saturated soils, impeded root respiration, physiological drought, high soil leaching with low nutrient availability, limited rooting volume of shallow soils, reduced solar insolation, and high humidity combine to produce these low forests (Grubb, 1971; 1977; Weaver et al., 1973). Much of what we know about the plant ecology and flora of Bokor National Park broadly, and the Bokor Plateau more specifically, comes from the remarkable work conducted by Pauline Dy Phon which was carried CITATION: Rundel, P.W. & Middleton, D.J. (2017) The flora of the Bokor Plateau, southeastern Cambodia: a homage to Pauline Dy Phon. Cambodian Journal of Natural History, 2017, 17–37. Cambodian Journal of Natural History 2017 (1) 17–37 © Centre for Biodiversity Conservation, Phnom Penh 17 18 P. Rundel & D. Middleton out in the context of her PhD studies at the University of Toulouse in the late 1950s. This detailed study (Dy Phon, 1970) has long formed the basis for understanding the rich plant diversity of global significance in Bokor National Park. Her work has been the stimulus for our studies and for others as well, and we are pleased to dedicate this article to Pauline Dy Phon (see box). T he Bokor Plat e a u Despite its seasonal pattern, rainfall on the Bokor Plateau reaches very high levels. Records at 950 m elevation at the southern end of the plateau had a mean annual rainfall of 5,309 mm (Tixier, 1979), while the Val d’Emeraude on the southeast margin of the plateau was reported to receive a mean of 5,384 mm (Dy Phon, 1970). The distribution of this rain peaks sharply in July and August, dropping to a mean of 50 mm or less in January and February at both stations. The Val d’Emeraude experiences rain virtually every day from May through October, but on only 12 days on average in March (Dy Phon, 1970). During the dry season mornings are semi-sunny with scattered clouds moving overhead, while heavier overcast and brief periods of intense showers can occur in the afternoon. Mean monthly temperatures are relatively constant throughout the year, varying only from a low of 19.2°C in July and August to a high of 21.5°C in April (Dy Phon, 1970). The sandstone substrate of the plateau of the Elephant Mountains weathers into an acidic coarse white sand with a pH of 4.6. Soil profiles of the sphagnum bog as described by Dy Phon (1970) consist of upper sandy A horizons 90 cm in thickness with declining organic matter and increasing saturation with depth. Even in forested areas of the plateau there is often a B horizon at 90–105 cm consisting of an indurated layer of white sand, with yellowish sandstone parent material below this level. As a result there are mosaics present of seasonally water-logged soils. The wet forests encountered at middle elevations below the plateau on Bokor were termed forest submontagnardee a fagacées et cibotium by Dy Phon (1970). These replace the lower wet evergreen forests at elevations of 500–800 or higher. In many ways these are comparable to the hill evergreen forests described in Thailand with a dominant role of Fagaceae and Podocarpaceae in the canopy and the relative absence of Dipterocarpaceae (Rundel et al., 1999). This community changes at about 920 m with the transition from montane wet forest to the Bokor Plateau with its associated edaphic and climatic conditions. © Centre for Biodiversity Conservation, Phnom Penh Our study has focused on the gently sloping and weathered Bokor Plateau itself. Here a distinct community of stunted forest (Fig. 2) appears, termed forêt sempervirente basse de montagne by Dy Phon (1970). This transition between forest types can be seen near Popokvil Waterfall at about 920 m with mosaics of taller wet forest and lower stunted forest (Fig. 3). The dwarfing of what are commonly tall trees at lower elevations results from a complex interaction of soil depth, elevation, wind exposure, and distance from the coast. This community, which we have called stunted forest, commonly has a matrix that reaches no more than 4 m, while the canopy dominant Dacrydium elatum (Roxb.) Wall. ex Hook. with Dacrycarpus imbricatus (Blume) de Laub., Tristaniopsis merguensis (Griff.) Peter G.Wilson & J.T.Waterh., and Vaccinium viscifolium King & Gamble can reach greater heights. The gradient of height for D. elatum across the plateau illustrates the effect of environment on stature. Trees near Popkvil Waterfall are 14–16 m in height, but mean height drops to 8–10 m moving across the plateau, and finally only 4–6 m to the south near the developed area (Rundel et al., 2016). Despite changes in commuity structure, tree diversity and density remain relatively unchanged across this gradient (Zhang et al., 2016). Lianas are common in taller forest stands, including the notable presence of spiny rattans. In addition, there is a moderate diversity of epiphytic and lithophyllic orchids and ferns present. However as soil and wind conditions produce a lower forest canopy, the lianas largely disappear and Pandanus cupribasalis H.St.John and Pinanga sylvestris (Lour.) Hodel appear in the semi-open understorey. As soils become shallower and winds increase moving from Popokvil south across the plateau toward the coastal escarpment, the stunted forest is replaced by an irregular cover of sclerophyllous shrubland with a typical height of 1–2 m. Dy Phon (1970) termed this la lande de myrtacées et vacciniacées, which we have translated as sclerophyllous heathland. As Dy Phon’s name suggests, this community is dominated by species of Myrtaceae and Ericaceae. The former include Rhodamnia dumetorum (DC.) Merr. & L.M.Perry, Rhodomyrtus tomentosa (Aiton) Hassk., and Syzygium antisepticum (Blume) Merr. & L.M.Perry together with Vaccinium bracteatum Thunb., V. viscifolium King & Gamble, and Rhododendron moulmainense Hook. Epiphytes are rare. Open rocky areas that are waterlogged for major portions of the year have a scattered cover of herbaceous perennial such as Hedyotis rosmarinifolia (Pit.) Craib and Polygonum chinense in a matrix of the graminoids Carex indica L., Fimbrystylis eragrostis (Nees) Hance, and Dapsilanthus disjunctus (Mast.) B.G.Briggs & L.A.S.Johnson. Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Localized bog communities (Fig. 4) on the plateau are dominated by a diverse community of low-growing herbaceous perennials 20–30 cm in height. Scattered through this matrix are small islands of shrub establishment where soils have built up to allow better drainage. Four graminoid species provide the major part of the matrix cover (Rundel et al., 2003). These are Eremochloa eriopoda C.E.Hubb. (Poaceae), Eriocaulon ubonense Lecomte (Eriocaulaceae), and Dapsilanthus disjunctus and Centrolepis cambodiana Hance (Restionaceae). Small shrub islands scattered across the bog are dominated by single or multiple species reaching to no more than 30–50 cm in height and low mounds 0.5–2.0 m across. Rings of Sphagnum spp. are commonly present around the edges of these shrub islands at the edge of the canopy. Tixier (1979) identified these as S. beccarii Hampe. and S. cuspi- datum Ehr. ex Hoffm. Low shrubs forming these islands include Hedyotis rosmarinifolia (Rubiaceae), Ploiarium alternifolium (Vahl) Melch. (Bonnetiaceae), Calophyllum calaba L. var. cuneatum (Symington ex M.R.Henderson & Wyatt-Smith) P.F.Stevens (Calophyllaceae), Ilex wallichii Hook.f. (Aquifoliaceae), Syzygium antisepticum (Myrtaceae), and Hygrophila ringens (L.) R.Br. ex Spreng. (Acanthaceae). Species of Cladonia and other macrolichens and microbial crusts of cyanobacteria cementing areas of open soil are also present. History of Bokor Development The decision by the French colonial government to establish a tourist resort in the uplands of the Elephant Mountains in southeastern Cambodia was made in 1917. The Pa uline Dy Phon, 1 9 3 3 –2 0 1 0 Pauline Dy Phon made major contributions to our understanding of Cambodian plant ecology, systematics and economic botany, but the significance of her work has not been broadly or appropriately recognized. Although a plant taxonomist and biogeographer by training, her career demonstrated broad interests beyond these fields to include pioneering work on the economic and medicinal uses of Cambodian plants and their cultural significance. Pauline was born in Cambodia in 1933 to a successful Catholic family. The nation gained independence from France as a constitutional monarchy in 1953, at a time of uncertain political future for the former French colony. With strong family support, Pauline was encouraged to follow her interests and left the country to study in France, obtaining her bachelor’s degree in 1959 at the Faculty of Natural Sciences in Paris. She returned to Cambodia to accept a high school teaching position at the Lycée Sisowath in Phnom Penh. Ambitious though to expand her background in botany, she traveled again to France for graduate work. She obtained her doctorate at the University of Toulouse in 1969, working with Jules Émile Vidal, returning to teach at the Faculty of Sciences in Phnom Penh. It was under her graduate programme that she completed her remarkable work on the vegetation of southeastern Cambodia, which was published in 1970. It was in this era that she also wrote Guide botanique de la ville de Phnom Penh (1972) with M.A. Martin, a work republished in 2009. She was recognized by her peers at this time with an appointment by the Cambodian government as chair of the National Commission on Science and Culture. Pauline’s teaching and research career in Phnom Penh was sharply interrupted by the takeover of the country by the Khmer Rouge in April 1975. There followed four difficult years of genocidal government policies that virtually emptied the city of Phnom Penh. Her only written account of the Khmer Rouge period, published in 1982, was a study of “plants in the Khmer diet in normal times and in times of famine”. The Vietnamese invasion in 1979 forced out the Khmer Rouge government but there followed a challenging period of Vietnamese occupation. It was at this time that Pauline sought refuge in France, accepting a position at the Laboratory of Botany at the Natural History Museum in Paris in 1980. This was the beginning of two decades of work in Paris where her research contributed significantly to identifying and classifying the poorly known flora of Cambodia and other countries in Indochina. It was at this time that she completed and published her classic trilingual Dictionary of Plants Used in Cambodia (Édition Olympic, Phnom-Penh, 915 pp.). After the Khmer Rouge period, Pauline returned for the first time to Cambodia in 1994. She died at her home in Paris on 21 May, 2010. Cambodian Journal of Natural History 2017 (1) 17–37 © Centre for Biodiversity Conservation, Phnom Penh 19 20 P. Rundel & D. Middleton Fig. 1 South-facing escarpment of the Bokor Plateau looking out to the Gulf of Thailand with Phu Quoc Island (Vietnam) in the distance, March 2001 (© Rasoul Sharifi). Fig. 3 Transition area near Popokvil Waterfall from montane wet evergreen forest to stunted forest on the Bokor Plateau with Dr Kansri Boonpragob of Ramkhamhaeng University, Bangkok, March 2001 (© Rasoul Sharifi). Fig. 2 Stunted forest of the Bokor Plateau with emergent trees of Dacrydium elatum, March 2001 (© Philip Rundel). Fig. 4 Sphagnum bog and wetland in areas of poor drainage on the Bokor Plateau, March 2001 (© Rasoul Sharifi). © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Bokor Plateau was selected, largely for the French elite, for a variety of reasons including its proximity to coastal cities, the cooler upland climate above 1,000 m elevation, the spectacular panorama of the Gulf of Thailand and Phu Quoc Island from the plateau, and the presence of the picturesque Popokvil Waterfall. A steep, winding road to the plateau was completed in 1921, and the construction of buildings on the plateau began the following year. These included a Catholic church (1922), a Buddhist temple (1924), and finally the luxurious Bokor Hotel Palace at the edge of the escarpment in 1925. The opening of the hotel was associated with the development of associated infrastructure to supply water and electricity and a school and hospital, altogether covering an area of about 5 km2 (Kowlalcyzk, 2009). The tourist resort was short-lived, however, as the casino was closed in 1940 with the Japanese occupation of Indochina. The end of World War II brought little change to Bokor as an extended war of liberation began throughout the region. Vandalism and gang activity had taken over during the war when some of the buildings were utilized as a military hospital and sanitarium for wounded soldiers. Two decades passed before an independent Cambodia renovated the hotel and casino in 1959, with Bokor quickly becoming the leading tourist attraction in the country. This golden era for the resort lasted only until 1972 when regional wars made its continued operation untenable. With the coming to power of the Khmer Rouge in 1975, all tourism ceased and Bokor was essentially abandoned. The resort facilities were heavily damaged in 1979 during fighting between Khmer Rouge soldiers and the advancing Vietnamese army who captured the area in 1982. With the establishment of Bokor National Park in 1993, Cambodian forest rangers established a presence in the park and a small trickle of tourism returned and grew slowly. As political stability returned to Cambodia, there was renewed interest in developing the Bokor Plateau again. In January 2008, the Sokimex Group, a large corporate entity, announced that they had obtained a 99-year lease on 5 km2 of the plateau for an international tourist development. A new access road up the mountain was soon completed and the large Thansur Bokor Highland Resort and casino constructed with 418 rooms. These developments greatly increased the ease of access and scientific work at Bokor, with increased visits by botanical collectors. In these early stages of resort development, planning is ongoing for extensive expanded facilities including golf courses, individual villas, and agricultural operations, with a proposed total expendiCambodian Journal of Natural History 2017 (1) 17–37 ture of US$1 billion (see http://www.sokimex.com/ourbusiness/casino/thansur-bokor-highland-resort). Conservation Despite a diverse and ecologically significant flora existing on the Bokor Plateau, conservation needs have not been given serious attention under strong pressures for development. Despite the massive scope of the project, forest and animal protection groups were quiet about the development’s potential impact before its opening (Phnom Penh Post, 2012). The scale of ongoing construction can be readily seen in comparing Google Earth images of the plateau before and under current development. Much of the area between the new Thansur Bokor Highland Resort and Popokvil Waterfall has been graded into a network of access roads for housing and infrastructure development. These projects have already impacted the natural communities on the plateau, leading to a strong need to incorporate conservation planning and education into the development process. The Bokor Plateau includes scattered small sphagnum bogs among its ecological communities, a rare habitat in mainland Southeast Asia (Rundel et al., 2003). The plateau is also home to at least 22 endemic plant species known only from Bokor National Park: Schefflera cambodiana Yahara & Tagane, Argostemma fasciculata Sridith & Larsen, Impatiens bokorensis S.H.Cho & B.Y.Kim, Garcinia bokorensis H.Toyama & Yahara, Diospyros elephasii Lecomte, Elaeocarpus bokorensis Tagane, Croton phourinii H.Toyama & Tagane, Lithocarpus eriobotryifolius Yahara, Gentiana ting-nung-hoae Halda, Cinnamomum bokorense Tagane & Yahara, Lindera bokorensis Yahara & Tagane, Machilus bokorensis Yahara & Tagane, Neolitsea bokorensis Yahara & Tagane, Memecylon bokorense Tagane, Sonerila bokorense S.H.Cho and Y.D.Kim, Syzygium bokorense W.K.Soh & J.Parn., Nepenthes bokorensis Mey, Cleyera bokorensis Nagam. & Tagane, Phyllanthus bokorensis Tagane, Helicia elephanti Sleumer, Wikstroemia bokorensis E.Oguri & Tagane, and Globba bokorensis Nob.Tanaka & Tagane. The Bokor Plateau and its associated development offers significant educational opportunities for Cambodian students at all levels to better appreciate the conservation and sustainability of biodiversity of the country. We hope that there will be broad interest in expanding existing programmes and developing new opportunities for environmental education at Bokor. T he Che ck list Our checklist includes taxa mentioned by Dy Phon (1970) as present on the plateau, our own collections, Sridith & © Centre for Biodiversity Conservation, Phnom Penh 21 22 P. Rundel & D. Middleton Larsen (2004), Averyanov et al. (2013, 2016), Nuraliev et al. (2015), Cho et al. (2015, 2017), Schuitemen et al. (2016), and reports by Tagane et al. (2017) for taxa occurring at or above 920 m. This is the elevation of Popokvil Waterfall. Excluded are non-native plant taxa previously or currently used for landscaping around the developed areas on the plateau. Also excluded are several collections present in the Muséum National d’Histoire Naturelle in Paris for species that are not otherwise accounted for in the collections cited below or in the works of Dy Phon (1970) and Tagane (2017) and for which no altitude data are given. Although some are clearly collections from the plateau area and already accounted for in the list below, many are just as equally clearly taxa of lower altitudes. Although it is possible that the collections include taxa of higher elevations, better data are needed to justify their inclusion in a list accounting only for plants growing above 920 m. Our checklist is comprised of 359 species with 29 ferns and lycophytes, 4 gymnosperms, and 326 angiosperms. The largest family in this checklist is the Rubiaceae with 30 species, followed by the Orchidaceae (28 species), Lauraceae (20 species), and Myrtaceae (13 species). For this flora, 22 species are believed to be endemic to Bokor, as indicated in the text below, highlighting the biodiversity significance of the Bokor Plateau. We hope that our article will serve to stimulate new research that can add to or correct this checklist. extensive surveys of the flora of the southern slope of Bokor National Park in a series of collecting trips from December 2011 to December 2013. This work, which covered the gradient from the coast to the plateau area, recorded 747 species in 105 families including 24 new species (Tagane et al., 2017). The first set of their collections is deposited in the Kyushu University herbarium in Fukuoka (FU) with a second set in the herbarium of the Forest Administration of Cambodia. Partial sets of collections were distributed to the Forest Herbarium Bangkok (BKF), the Kyoto University Museum (KYO), Royal Botanic Gardens, Kew (K), Naturalis Biodiversity Center (L) and Muséum National d’Histoire Naturelle (P). More recently there have been large collections made by Nguyen Van Du and by Su Kung Wu, with records in TROPICOS database. The inclusion of taxa within families follows the Pteridophyte Phylogeny Group (PPG I, 2016) for ferns and lycophytes and the Angiosperm Phylogeny Group (APG IV, 2016) for angiosperms. For ferns and lycophytes, the generic delimitations also follow PPG I (2016), except in Cyathea pending clarification of the current placement of the Southeast Asian taxa. For angiosperms, the generic delimitation follows the recent literature for each group of plants most of which is summarised by Stevens (2017). If the spelling or authorship of a taxon name was given incorrectly by an earlier author, we correct it here without comment. What we term stunted forest in the list below is equivalent to what Dy Phon termed forêt sempervirente basse de montagne. We have translated her lande a myrtacées et á vacciniacées as heathland. Phlegmariurus squarrosus (G.Forst.) Á.Löve & D.Löve. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970) as Lycopodium squarrosum G.Forst. Our work is the outcome of field excursions in 1999 and 2001. The cited Middleton and Monyrak specimens are deposited in the herbarium of the Arnold Arboretum at Harvard University Herbaria (A) and in the Herbarium of the Ministry of the Environment in Phnom Penh. Duplicates of many collections can also be found in the herbarium of the Muséum National d’Histoire Naturelle in Paris (P), which also houses some of the original collections by Dy Phon. Shuichiro Tagane from Kyushu University in Japan and his research group conducted © Centre for Biodiversity Conservation, Phnom Penh LYCOPHYTES Lycopodiaceae Huperzia serrata (Thunb. ex Murray) Trevis. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970) as Lycopodium serratum Thunb. ex Murray. Palhinhaea cernua (L.) Vasc. & Franco — Herbaceous perennial in stunted forest and open wetlands. Included by Dy Phon (1970) and Rundel et al. (2003) as Lycopodium cernuum L. Selaginellaceae Selaginella sp. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). FERNS Aspleniaceae Asplenium nidus L. — Uncommon epiphyte in stunted forest. Noted in our work but not collected. Cyatheaceae Cyathea gigantea (Wall. ex Hook.) Holttum — Tree fern in understorey of stunted forest. Reported as rare by Dy Phon (1970) under the misapplied name Cyathea glabra (Blume) Copel. Cyathea podophylla (Hook.) Copel. — Tree fern in understorey of stunted forest. Reported by Dy Phon (1970). Davalliaceae Davallia repens (L.f.) Kuhn — Lithophytic fern on rocks in exposed area beside track on way to Popokvil Waterfall, Middleton and Monyrak 652. Also reported by Dy Phon (1970) as Humata repens (L.f.) Diels. Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Davallia solida (G.Forst.) Sw. — Epiphytic fern in stunted forest. Reported by Dy Phon (1970). Dennstaedtiaceae Pteridium aquilinum (L.) Kuhn — Terrestrial fern widespread in open grassy areas across the plateau. Noted as common in our work and also noted by Dy Phon (1970) Pteridium esculentum (G.Forst.) Cockayne — Terrestrial fern at margin of stunted forest. Reported by Dy Phon (1970). Dicksoniaceae Cibotium barometz (L.) J.Sm. — Tree fern in understorey of stunted forest. Reported by Dy Phon (1970). Gleicheniaceae Dicranopteris linearis (Burm.f.) Underw. — Trailing or scrambling fern common along stunted forest margin. Observed but not collected by us and reported by Dy Phon (1970). Aglaomorpha rigidula (Sw.) Hovenkamp & S.Linds. — Terrestrial or epiphytic fern in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 608. Microsorum scolopendria (Burm.f.) Copel. — Epiphytic or lithophytic fern in areas of stunted forest. Reported by Dy Phon (1970) as Phymatodes scolopendria (Burm.f.) Ching. Oreogrammitis dorsipila (Christ) Parris — Lithophytic fern on shaded rocks beside Popokvil Waterfall at 920 m, Middleton and Monyrak 618. Pyrrosia lingua (Thunb.) Farw. var. heteractis (Mett. ex Kuhn) Hovenkamp — Terrestrial fern in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 582. Reported by Dy Phon (1970) as Pyrrosia eberhardtii (Christ) Ching. Although terrestrial on the Bokor plateau this species is usually epiphytic. Diplopterygium norrisii (Mett. ex Kuhn) Nakai — Climbing fern at margin of stunted forest. Reported by Dy Phon (1970) as Gleichenia norrisii Mett. Selliguea triloba (Houtt.) M.G.Price — Epiphytic or lithophytic fern in areas of stunted forest. Reported by Dy Phon (1970) as Phymatodes triphylla (Jacq.) C.Chr. & Tardieu. Lindsaeaceae Pteridaceae Lindsaea ensifolia Sw. — Terrestrial fern in stunted forest. Reported by Dy Phon (1970) as Schizoloma ensifolium (Sw.) J.Sm. Pityrogramma calomelanos (L.) Link — Delicate lithophytic fern. Sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 584. Lygodiaceae Lygodium flexuosum (L.) Sw. — Herbaceous climbing fern. Reported by Dy Phon (1970) from heathland areas. Nephrolepidaceae Nephrolepis brownii (Desv.) Hovenkamp & Miyam. — Terrestrial fern on sandy soil in scrubland near top of plateau, Middleton and Monyrak 600. Taenitis blechnoides (Willd.) Sw. — Terrestrial fern in stunted forest. Reported by Dy Phon (1970). Thelypteridaceae Cyclosorus interruptus (Willd.) H.Ito — Short erect fern on mossy rock in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 586. Nephrolepis hirsutula (G.Forst.) C.Presl — Epiphytic fern in stunted forest. Reported by Dy Phon (1970). GYMNOSPERMS Oleandraceae Gnetaceae Oleandra musifolia (Blume) C.Presl — Large terrestrial fern in understorey of stunted forest. Reported by Dy Phon (1970). Gnetum latifolium Blume — Woody climber, occasional at higher elevations around 1,014 m (Tagane et al., 2017). Oleandra neriiformis Cav. — Terrestrial fern in stunted forest on sandy soil beside stream on way from road to Popokvil Waterfall at 934 m, Middleton and Monyrak 617. Also reported by Dy Phon (1970). Dacrycarpus imbricatus (Blume) de Laub. — Tree, common in moist evergreen forest on the plateau. Noted in our work and by Dy Phon (1970) and Tagane et al. (2017). Polypodiaceae Aglaomorpha coronans (Wall. ex Mett.) Copel. — Epiphytic or terrestrial fern in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 583. Cambodian Journal of Natural History 2017 (1) 17–37 Podocarpaceae Dacrydium elatum Wall. ex Hook. — Dominant canopy tree over much of the stunted forest on the plateau. Noted by Dy Phon (1970) and Tagane et al. (2017). Discussed in Rundel et al. (2016) and Tagane et al. (2017) Podocarpus pilgeri Foxw. — Sclerophyllous shrub to small tree in stunted forest among rocks on sandy soil near field © Centre for Biodiversity Conservation, Phnom Penh 23 24 P. Rundel & D. Middleton station, at the top of the plateau at 1,042 m, Middleton and Monyrak 613. ANGIOSPERMS Acanthaceae Hygrophila ringens (L.) R.Br. ex Spreng. — Small shrub to tiny herb in seasonally inundated vegetation on sandy soil beside track near Popokvil Waterfall and near field station at 936—1,059 m, Middleton and Monyrak 638, 674. Also listed by Rundel et al. (2003) as Hygrophila angustifolia R.Br. Justicia ventricosa Wall. ex Hook.f. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Phlogacanthus geoffrayi Benoist — Shrub at margin of moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Strobilanthes aff. lilacina C.B.Clarke — Woody subshrub in heathland and stunted forest areas. Reported by Dy Phon (1970). Thunbergia grandiflora Roxb. — Open ground beside path to Popokvil Waterfall, scrambling over shrub at 1,034 m, Middleton and Monyrak 626. Adoxaceae Viburnum sambucinum Reinw. ex Blume — Small tree in scrubby vegetation and stunted forest on sandy soil near summit on roadside towards research centre at 936 m, Middleton and Monyrak 634. Also reported as common by Tagane et al. (2017). Altingiaceae Liquidambar siamensis (Craib) Ickert-Bond & J.Wen — Tree in stunted forest. Reported by Tagane et al. (2017). Reported as Altingia siamensis Craib by Dy Phon (1970) and treated as a synonym of Altingia excelsa Noronha (= Liquidambar excelsa (Noronha) Oken) in the Flora of Thailand. Anacardiaceae Toxicodendron succedaneum (L.) Kuntze — Small tree, common in moist evergreen forest on the plateau (Tagane et al., 2017). Annonaceae Uvaria hamiltonii Hook.f. & Thomson — Woody climber, occasional on the plateau (Tagane et al., 2017). Apiaceae Centella asiatica (L.) Urb. — Herbaceous creeper. Reported by Dy Phon (1970) from heathland areas. © Centre for Biodiversity Conservation, Phnom Penh Apocynaceae Alyxia reinwardtii Blume — Woody climber, occasional in moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Chilocarpus denudatus Blume — Woody climber, common in moist evergreen forest at higher elevations around 941 m (Tagane et al., 2017). Hoya multiflora Blume — Epiphytic or lithophytic herb scattered at higher elevations around 925 m (Tagane et al., 2017). Tabernaemontana bufalina Lour. — Small shrub, occasional in evergreen forest at higher elevations around 1,014 m (Tagane et al., 2017). Tabernaemontana pauciflora Blume — Small shrub, occasional in evergreen stunted forest at higher elevations around 1,014 m (Tagane et al., 2017). Tylophora ovata (Lindl.) Hook. ex Steud. — Climber in moist evergreen forest on the upper elevation of the plateau at 1,043 m (Tagane et al., 2017). Urceola micrantha (Wall. ex G.Don) Mabb. — Woody climber, common in evergreen forests, around 1,014– 1,043 m (Tagane et al., 2017). Aquifoliaceae Ilex annamensis Tardieu — Shrub in understorey of stunted forest. Reported by Dy Phon (1970). Ilex cymosa Blume — Tree, common in evergreen forest at higher elevations at 970 m (Tagane et al., 2017). Ilex excavata Pierre — Tree, fairly common in moist evergreen forest on the top of the plateau at 970–1,043 m (Tagane et al., 2017). Ilex triflora Blume — Small tree, occasional in moist evergreen forest, common along streamside at 960–1,014 m (Tagane et al., 2017) Ilex viridis Champ. ex Benth. — Subshrub in heathland areas. Reported by Dy Phon (1970) as Ilex triflora Blume var. viridis (Champ. ex Benth.) Loes. Ilex wallichii Hook.f. — Shrub in dry sandy soil on open roadside beside inundated area on track towards Popokvil Waterfall at 936 m, Middleton and Monyrak 627. Also reported by Tagane et al. (2017) as a shrub to a tree and locally common in moist evergreen forest and open bog on the plateau at 1,005–1,043 m. N.B. Ilex is a genus that deserves more study at Bokor. In addition to the species above, Tagane et al. (2017) report two additional unnamed taxa. Araceae Pothos chinensis (Raf.) Merr. — Herbaceous climber in stunted forest. Reported by Dy Phon (1970) as Pothos cathcartii Schott. Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Scindapsus hederaceus Miq. — Herbaceous climber in stunted forest. Reported by Dy Phon (1970) as Scindapsus polanei Gagnep. Araliaceae Dendropanax lancifolius (Ridl.) Ridl. — Small tree, occasional in moist evergreen forests on the plateau at 935–1,043 m (Tagane et al., 2017). Dendropanax maingayi King — Shrub to small tree, common in dense evergreen forest at higher elevations at 962–1,014 m (Tagane et al., 2017). Polyscias diversifolia (Blume) Lowry & G.M.Plunkett — Small tree, common in moist evergreen forest on the plateau at 970–1,014 m (Tagane et al., 2017). Schefflera cambodiana Yahara & Tagane — Tree, occasional in moist evergreen forest at higher elevations, especially common by the stream below Popokvil Waterfall at 970 m (Tagane et al., 2017). Endemic to Bokor. Schefflera pueckleri (K.Koch) Frodin — Tree in stunted forest. Reported as Tupidanthus calyptratus Hook. & Thomson by Dy Phon (1970). Schefflera schizophylla (Hance) Frodin — Tree in stunted forest on sandy soil beside track on way to Popokvil Waterfall at 1,000 m, Middleton and Monyrak 651. Reported by Dy Phon (1970) as Schefflera incisa R.Vig. Schefflera subintegra (Craib) C.B.Shang — Tree in thin soil on rocky ground beside Popokvil Waterfall at 920 m, Middleton and Monyrak 620. Also reported by Tagane et al. (2017) from moist evergreen forest at 1,014 m. Arecaceae Areca triandra Roxb. ex Buch.-Ham. — Erect palm, occasional in higher elevation (Tagane et al., 2017). Calamus bousigonii Becc. — Climbing palm in stunted forest. Reported by Dy Phon (1970). Also reported by Tagane et al. (2017) as locally common on the plateau at 1,014 m, as well as lower elevations. in moist evergreen forest at higher elevations, around 935–1,043 m. Plectocomia elongata Mart. ex Blume — Stout, climbing palm, somewhat common at higher elevations, around 1,014 m (Tagane et al., 2017). Plectocomia pierreana Becc. Climbing palm in stunted forest. Reported by Dy Phon (1970) as Plectocomia cambodiana Gagnep. ex Humbert. Also reported by Tagane et al. (2017) as somewhat common at higher elevations, around 930–1,014 m. Asphodelaceae Dianella ensifolia (L.) DC. — Herbaceous perennial in stunted forest understorey. Reported by Dy Phon (1970). Asparagaceae Dracaena elliptica Thunb. & Dalm. — Shrub, common in moist evergreen forest on the plateau, and also found in middle elevation. Reported by Dy Phon (1970) and Tagane et al. (2017) as Dracaena gracilis (Baker) Hook.f., an illegitimate name. Dracaena reflexa Lam. var. angustifolia Baker — Trailing shrub, reported by Dy Phon (1970) from the understorey of stunted evergreen forest. Chlorophytum orchidastrum Lindl. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Cordyline fruticosa (L.) A.Chev. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Asteraceae Ageratina adenophora (Spreng.) R.M.King & H.Rob. — Herbaceous perennial on roadsides in heathland area. Reported by Dy Phon (1970) as Eupatorium adenophora. Naturalized non-native. Camchaya kampotensis Gagnep. — Herbaceous perennial in wetland and stunted forest areas. Reported by Dy Phon (1970). Calamus palustris Griff. — Climbing palm common at middle to high elevations (Tagane et al., 2017). Gynura divaricata (L.) DC. — Herbaceous perennial in heathland area. Reported by Dy Phon (1970) as Gynura auriculata. Calamus rudentum Lour. — Spiny climbing palm in stunted forest. Reported by Dy Phon (1970). Elephantopus scaber L. — Herbaceous perennial in heathland area. Reported by Dy Phon (1970)). Daemonorops jenkinsiana (Griff.) Mart. — Climbing palm in stunted forest. Reported by Dy Phon (1970) as Daemonorops pierreana Becc. Also reported by Tagane et al. (2017) as occasional in evergreen forest and damp sites in middle and higher elevations, to 928 m. Spilanthes iabadicensis A.H.Moore — Herbaceous perennial in heathland area. Reported by Dy Phon (1970) as equal to S. acmella (L.) L. Spilanthes iabadicensis is placed in synonymy of Acmella uliginosa (Sw.) Cass. in the Flora of Thailand. Pinanga sylvestris (Lour.) Hodel — Erect palm, growing in open areas of the understorey in stunted evergreen forest, often abundant with Pandanus cupribasalis H.St. John. Reported by Dy Phon (1970) as Pinanga cochinchensis Blume. Also reported by Tagane et al. (2017) as common Balanophoraceae Cambodian Journal of Natural History 2017 (1) 17–37 Balanophora fungosa J.R.Forst. & G.Forst. subsp. indica (Arn.) B.Hansen. — Herbaceous root parasite in stunted forest. Reported by Dy Phon (1970) as Balanophora gracilis Tiegh. and B. sphaerica (Tiegh.) Lecomte. © Centre for Biodiversity Conservation, Phnom Penh 25 26 P. Rundel & D. Middleton Balsaminaceae Celastraceae Impatiens angustisepala Tardieu — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Euonymus indicus B.Heyne ex Wall. — Shrub on sandy soil in scrubland near the top of the plateau at 1,056 m, Middleton and Monyrak 598. Reported by Tagane et al. (2017) as Euonymus javanicus Blume var. talungensis Pierre and said to be common in moist evergreen forest on the plateau at 1,014 m. Impatiens bokorensis S.H.Cho & B.Y.Kim — Herbaceous perennial known only from the type locality on the Bokor plateau at 1,050 m (Cho et al., 2017). Endemic to Bokor. Impatiens muelleri Tardieu — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). The type material of this species from Bokor has been variously labelled with other names in the genus but we continue to recognize it pending a definitive revision of the species. Impatiens velaxata Hook.f. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). N.B. The genus Impatiens is in need of revision for the region. Bonnetiaceae Ploiarium alternifolium (Vahl) Melch. — Shrub in seasonally inundated area on sandy soil near top of plateau at 944 m, Middleton and Monyrak 591. Also reported by Tagane et al. (2017) as common in open sunny bogs at 926 m. Listed by Rundel et al. (2003). Burmanniaceae Burmannia disticha L. — Herb in seasonally inundated area on sandy soil near top of plateau at 944 m, Middleton and Monyrak 590. Also listed by Rundel et al. (2003). Calophyllaceae Calophyllum calaba L. var. cuneatum (Symington ex M.R.Hend. & Wyatt-Sm.) P.F.Stevens — Shrub in sclerophyllous stunted forest on rocky sandy soil near field station, near top of plateau at 1,042 m, Middleton and Monyrak 656. Also reported by Tagane et al. (2017) as a shrub or tree, common in open sunny bog on the plateau and included by Rundel et al. (2003). Dy Phon (1970) reports this as Calophyllum saigonense Pierre var. nanum Gagnep. Calophyllum dryobalanoides Pierre — Shrub or tree, rare in moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Calophyllum tetrapterum Miq. — Shrub or tree, common in evergreen forest at higher elevations at 933 m (Tagane et al., 2017). Cannabaceae Microtropis discolor (Wall.) Wall. ex Meisn. — Small tree, occasional in evergreen forest, often found in humid sites near streams at 1,014 m (Tagane et al., 2017). Clusiaceae Garcinia bokorensis H.Toyama & Yahara — Tree, common in moist evergreen forest on and near the top of the plateau at 935–1,041 m (Tagane et al., 2017). Endemic to Bokor. Garcinia celebica L. — Tree, occasionally at middle elevations but reaching 928 m (Tagane et al., 2017) Garcinia hanburyi Hook.f. — Tree in forest from the foot to the top of Mt Bokor. Reported by Dy Phon (1970) and Tagane et al. (2017). Garcinia merguensis Wight — Tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 644, 646. Tagane et al. (2017) report that it is widely found from the foot to the top in Mt Bokor. N.B. The genus Garcinia would be appropriate for more detailed study as Tagane et al. (2017) list two unidentified species, one from the plateau. Convolvulaceae Argyreia longipes (Gagnep.) Traiperm & Staples — Climber, common in moist evergreen forest at higher elevations around 1,014 m (Tagane et al., 2017). Argyreia scortechinii (Prain) Prain ex Hoogland — Climber in dwarf forest on sandy soil beside track on way to Popokvil Waterfall at 1,000 m, Middleton and Monyrak 647. Cyperaceae Carex indica L. — Clump forming sedge in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 581. Reported by Dy Phon (1970). Caprifoliaceae Fimbristylis dichotoma (L.) Vahl — Clump forming sedge in boggy vegetation indicating seasonal waterlogging on sandy soil beside track towards Popokvil Waterfall at 936 m, Middleton and Monyrak 654. Also listed by Rundel et al. (2003). Lonicera cambodiana Pierre ex P.Danguy — Climber, locally common in moist evergreen forest and its margins on the plateau at 1,011–1,043 m (Tagane et al., 2017). Fimbristylis eragrostis (Nees & Meyen) Hance — Reported by Dy Phon (1970) as Fimbristylis lepidota E.G.Camus from bogs as well as dry places. Gironniera subaequalis Planch. — Small tree in thin soil on rocky ground beside Popokvil Waterfall at 920 m, Middleton and Monyrak 621. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Scleria ciliaris Nees — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Scleria harlandii Hance — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Scleria terrestris (L.) Fassett — Clump forming sedge in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 603. Daphniphyllaceae Daphniphyllum beddomei Craib — Tree, occasional in moist evergreen forests at higher elevations at 970–1,014 m (Tagane et al., 2017). This species is included as a synonym of Daphniphyllum paxianum Rosenthal in the Flora of China but is recognized as distinct in the Flora of Thailand. Daphniphyllum sp. — Dy Phon (1970) reported a species she called Daphniphyllum roxburghii H. Br. for which we can find no record, either of a specimen or publication. There is an illegitimate name Daphniphyllum roxburghii Baill. ex Rosenthal, now treated as D. oldhamii (Hemsl.) Rosenthal from Japan, Korea and China, but there is no evidence this species occurs in Cambodia. We are unsure of the identity of this species. N.B. The genus Daphniphyllum deserves more study at Bokor. Tagane et al. (2017) list two additional species in middle elevation forests. Droseraceae Drosera burmannii Vahl — Small insectivorous herb in seasonally inundated area on sandy soil at 944 m, Middleton and Monyrak 588. Also listed by Rundel et al. (2003). Drosera peltata Thunb. — Small insectivorous herb. Not seen by us but reported by Dy Phon (1970) from areas of seasonal wetlands. Ebenaceae Diospyros elephasii Lecomte — Small tree, common in moist evergreen forest on the top plateau. 962–1,043 m (Tagane et al., 2017). Endemic to Bokor. Diospyros venosa Wall. ex A.DC. — Tree in stunted forest. Reported by Dy Phon (1970). Tagane et al. (2017) used this name for a taxon at middle elevations below the plateau. Elaeocarpaceae Elaeocarpus bokorensis Tagane — Tree, common on the plateau from 800–1,000 m. (Tagane et al., 2015). Endemic to Bokor. Elaeocarpus dubius Aug.DC. — Tree, common in evergreen forest and its margin and roadside at 450–900(<1,000) m (rare on the plateau) (Tagane et al., 2017). Cambodian Journal of Natural History 2017 (1) 17–37 Elaeocarpus griffithii (Wight) A.Gray — Tree, rare in moist evergreen forest on the plateau around 928 m (Tagane et al., 2017). Elaeocarpus lanceifolius Roxb. — Tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau around 1,055 m, Middleton and Monyrak 670. Elaeocarpus thorelii Pierre — Small tree, rare, at higher elevation, around 970 m (Tagane et al., 2017). N.B. The genus Elaeocarpus deserves more study at Bokor to evaluate records. Tagane et al. (2017) list two unidentified taxa collected on the plateau. There are other species at middle elevations. Ericaceae Lyonia ovalifolia (Wall.) Drude — Tree in stunted forest. Reported by Dy Phon (1970) as Pieris ovalifolia. Also reported as rare in open bog on the plateau at 926 m by Tagane et al. (2017). Rhododendron moulmainense Hook. — Shrub in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau around 1,047 m, Middleton and Monyrak 662. Tagane et al. (2017) identify this as Rhododendron klossii Ridl. Vaccinium bracteatum Thunb. — Shrub or treelet in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau around 1,044 m, Middleton and Monyrak 659. Also reported by Tagane et al. (2017). Reported by Dy Phon (1970) as Vaccinium cambodianum Dop. Vaccinium viscifolium King & Gamble — Small tree in thin soil on rocky ground by Popokvil Waterfall and near field station at the top of the plateau, 920–1,046 m, Middleton and Monyrak 622, 661. Also reported by Tagane et al. (2017) as occasional in moist evergreen forest on the plateau. Eriocaulaceae Eriocaulon ubonense Lecomte f. kradungense (Satake) A.Prajaksood & J.Parn. — Small herb in boggy vegetation indicating seasonal waterlogging on sandy soil beside track towards Popokvil Waterfall at 936 m, Middleton and Monyrak 628. Included as Eriocaulon cf. henryanum Ruhland in Rundel et al. (2003). Escalloniaceae Polyosma integrifolia Blume — Tree, occasional in evergreen forest at higher elevations around 944 m (Tagane et al., 2017). Euphorbiaceae Croton phourinii H.Toyama & Tagane — Shrub, locally common in moist evergreen forest on the plateau at 930 m (Tagane et al., 2017). Dy Phon (1970) reported a Croton sp. in the understorey of stunted forest. Endemic © Centre for Biodiversity Conservation, Phnom Penh 27 28 P. Rundel & D. Middleton to Bokor although its distinction from Croton phaenodon Airy Shaw was questioned by an anonymous reviewer. (1970) and Tagane et al. (2017) as Quercus cambodiensis Hickel & A.Camus. Gymnanthes remota (Steenis) Esser — Small tree, occasional in moist evergreen forest at higher elevations around 960–1,014 m (Tagane et al., 2017). Quercus sp. — Tree, rare in evergreen forest at 970 m. Reported by Tagane et al.(2017). Macaranga andamanica Kurz — Small tree, common in moist evergreen forest on the plateau, especially frequent along streams at higher elevations, around 975–1,014 m (Tagane et al., 2017). Fabaceae Codariocalyx gyroides (Roxb. ex Link) Hassk. — Shrub, occasional at semi-shaded forest margin on the plateau at 960 m (Tagane et al., 2017). Ormosia fordiana Oliv. — Tree in thin soil on rocky ground beside Popokvil Waterfall at 920 m, Middleton and Monyrak 624. Tagane et al. (2017) report an unidentified Ormosia on the plateau at 933 m. Fagaceae Castanopsis acuminatissima (Blume) A.DC — Tree, common at higher elevations around 970 m (Tagane et al., 2017). Castanopsis cambodiana A.Chev. ex Hickel & A.Camus — Tree, occasional in moist evergreen forest on the plateau at 935 m (Tagane et al., 2017). Lithocarpus elegans (Blume) Hatus. ex Soepadmo — Tree, rare in moist evergreen forest on the plateau at 1,000 m (Tagane et al., 2017). Lithocarpus elephantum (Hance) A.Camus — Tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 585. Also reported by Tagane et al. (2017) from moist evergreen forest at higher elevations, around 1,014–1,043 m. Lithocarpus eriobotryifolius Yahara — Tree, occasional in moist evergreen forest at higher elevations around 930 m (Tagane et al., 2017). Endemic to Bokor. Lithocarpus farinulentus (Hance) A.Camus — Tree in stunted forest. Reported by Dy Phon (1970). N.B. The Fagaceae would appear to be in need of more detailed study at Bokor and throughout Indochina. Flagellariaceae Flagellaria indica L. — Woody climber in stunted forest. Reported by Dy Phon (1970). Gentianaceae Fagraea auriculata Jack — Tree, common in humid evergreen forest on the top plateau around 1,014 m. Reported y Dy Phon (1970) and Tagane et al. (2017). Fagraea ceilanica Thunb. — Scandent tree, occasionally epiphyte, common in middle to high elevations around 1,014 m (Tagane et al., 2017). Gentiana greenwayae Merr. — Herbaceous perennial in heathland and stunted forest areas. Reported by Dy Phon (1970). Sometimes treated as Gentiana praticola subsp. greenwayae (Merr.) Halda. Gentiana ting-nung-hoae Halda — Herbaceous perennial in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 607. Also listed by Rundel et al. (2003). Gentiana bokorensis Hul is a synonym of this species. It has also been treated as a subspecies of Gentiana nudicaulis Kurz. Endemic to Bokor. Hanguanaceae Hanguana cf. malayana (Jack) Merr. — Herb on sandy soil in stunted forest beside track on way to Popokvil Waterfall at 1,000 m, Middleton and Monyrak 604. N.B. The genus Hanguana was, until recently, considered to be monotypic. Recent research in Malaysia and Singapore has led to the description of many new species and the conclusion that Hanguana malayana itself is likely quite narrowly distributed. It is quite possible that the Bokor plant is an undescribed species. Juglandaceae Lithocarpus leiophyllus A.Camus — Tree in sclerophyllous stunted forest on rocky sandy soil beside track towards Popokvil and near field station, at the top of the plateau at 1,043 m, Middleton and Monyrak 658. Also reported by Tagane et al. (2017) as fairly common and one of the dominant species in moist evergreen forest on the plateau. Engelhardia roxburghiana Lindl. ex Wall. — Tall tree found from low to high elevations (Tagane et al., 2017). Quercus augustinii Skan — Tree, occasional at higher elevations around 970 m (Tagane et al., 2017). Lamiaceae Quercus langbianensis Hickel & A.Camus — Tree in stunted forest on the top plateau. Reported by Dy Phon © Centre for Biodiversity Conservation, Phnom Penh Juncaceae Juncus prismatocarpus R.Br. — Herbaceous perennial. Reported by Dy Phon (1970) from wetland areas with scattered subshrubs. Anisochilus cambodianus Murata — Woody herb, rarely found on open sunny rocks on the plateau at 1,014 m (Tagane et al., 2017). Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Clerodendrum smitinandii Moldenke — Small tree, fairly common along forest edge and roadside around 970 m (Tagane et al., 2017). Premna interrupta Wall. ex Schauer — Scandent shrub, often epiphytic on tree trunks, occasional at higher elevations around 935 m (Tagane et al., 2017). Lauraceae Beilschmiedia gammieana King ex Hook.f. — Tree, rare, at 1,014 m (Tagane et al., 2017). Beilschmiedia penangiana Gamble — Small tree, common in moist evergreen forest at higher elevations around 970–1,043 m (Tagane et al., 2017). Cassytha filiformis L. — Herbaceous scrambling parasite in stunted forest. Reported by Dy Phon (1970). Cinnamomum bokorense Tagane & Yahara — Tree, occasional at middle elevations and reaching 935 m (Tagane et al., 2017). Endemic to Bokor. Cinnamomum curvifolium (Lour.) Nees — Tree, somewhat common in evergreen forest in middle and higher elevations around 970 m (Tagane et al., 2017). Cinnamomum dimorphandrum Yahara & Tagane — Small tree, somewhat common in moist evergreen forest at higher elevations, 941–1,043 m (Tagane et al., 2017). Cinnamomum iners Reinw. ex Blume — Treelet in heathland areas. Reported by Dy Phon (1970). Cinnamomum litseifolium Thwaites — Tree in stunted forest. Reported by Dy Phon (1970). Lindera bokorensis Yahara & Tagane — Small tree, rare, at 970 m (Tagane et al., 2017). Endemic to Bokor. Litsea cambodiana Lecomte — Tree in stunted forest. Reported by Dy Phon (1970). Tagane et al. (2017), however, describes this as a middle elevation tree not present on the plateau. Litsea martabanica (Kurz) Hook.f. — Small tree, common in middle elevation evergreen forests and a single collection from 970 m (Tagane et al., 2017). Litsea monopetala (Roxb.) Pers. — Tree locally common in moist evergreen forest on the plateau around 928 m (Tagane et al., 2017). Litsea verticillata Hance — Small tree, somewhat common in evergreen forest at higher elevations around 1,014 m (Tagane et al., 2017). (2017). Reported by Dy Phon (1970) as Machilus odoratissima Nees. Endemic to Bokor. Neolitsea aff. alongensis Lecomte — Tree in stunted forest. Reported by Dy Phon (1970). Neolitsea bokorensis Yahara & Tagane, ined. — Small tree, common in moist evergreen forest on the plateau around 1,011–1,043 m (Tagane et al., 2017). Presumably endemic to Bokor. Neolitsea cambodiana Lecomte var. cambodiana — Small tree, locally common at higher elevations around 1,014– 1,043 m (Tagane et al., 2017). Neolitsea cambodiana Lecomte var. glabra C.K.Allen — Tree in stunted forest. Reported by Dy Phon (1970). Neolitsea zeylanica (Nees & T.Nees) Merr. — Sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 611. Tagane et al. (2017) restricts this name to a low elevation tree species. N.B. The Lauraceae is a diverse family on both the plateau and lower elevations at Bokor. The Lauraceae of Cambodia, like the rest of Southeast Asia, is poorly known and in need of greater study. Lentibulariaceae Utricularia bifida L. — Tiny herb in boggy vegetation indicating seasonal waterlogging on sandy soil at 936 m, Middleton and Monyrak 641. Also listed by Rundel et al. (2003). Utricularia caerulea L. — Small herb in boggy vegetation indicating seasonal waterlogging on sandy soil at 936 m, Middleton and Monyrak 640. Utricularia delphinioides Thorel ex Pellegr. — Wet grassland. Reported by Dy Phon (1970). N.B. In addition to the above species, blogs by F.S. Mey in 2011 report four additional species — Utricularia minutissima Vahl, U. odorata Pellegr., U. striatula Sm., and U. uliginosa Wight. Loganiaceae Mitrasacme pygmaea R.Br. — Small herb growing in cracks in disintegrating road beside sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,058 m, Middleton and Monyrak 673. Loranthaceae Litsea vang Lecomte — Tree in stunted forest. Reported by Dy Phon (1970). Barathranthus axanthus (Korth.) Miq. — Small parasitic shrub, scattered in evergreen forest at stunted and high elevation, around 940 m (Tagane et al., 2017). Machilus bokorensis Yahara & Tagane — Small tree, common in moist evergreen and stunted forest on the top plateau on sandy soil at 936–1,056 m. Middleton and Monyrak 597, 629, 643. Also reported by Tagane et al. Macrosolen cochinchinensis (Lour.) Tiegh. — Semi-woody epiphytic parasite in stunted forest. Reported by Dy Phon (1970). Reported by Tagane et al. (2017) as occasional in hill evergreen forest at 800–1,000 m. Cambodian Journal of Natural History 2017 (1) 17–37 © Centre for Biodiversity Conservation, Phnom Penh 29 30 P. Rundel & D. Middleton Lythraceae Ammannia baccifera L. — Herbaceous perennial in wetland areas. Reported by Dy Phon (1970). Magnoliaceae Magnolia duperreana Pierre — Tree in stunted forest. Reported by Dy Phon (1970) as Kmeria duperreana. Also reported by Tagane et al. (2017) as common in moist evergreen forest on the plateau around 939–1,014 m. Sonerila bokorense S.H.Cho and Y.D.Kim — Herbaceous perennial in stunted forest at 950–1,050 m. Endemic to Bokor and described by Cho et al. (2015). N.B. Tagane et al. (2017) describes one unidentified species of Memecylon from 970 m on the plateau. More work on the Melastomataceae would help resolve species limits, with many taxa present at middle elevations. Meliaceae Magnolia liliifera (L.) Baill. — Small tree, common in moist evergreen forest on the plateau, around 1,014–1,043 m (Tagane et al., 2017). Aglaia spectabilis (Miq.) S.S.Jain & Bennet — Tree, occasional at higher elevations, around 970 m (Tagane et al., 2017). Magnolia sp. — A treelet in stunted forest on sandy soil beside a stream on way from road to Popokvil Waterfall at 934 m, Middleton and Monyrak 616. Dysoxylum cauliflorum Hiern — Tree, common in evergreen forest at all elevations, in particular along a stream at 970 m (Tagane et al., 2017). Malvaceae Dysoxylum sp. — Tall tree, occasional in primary forest at all elevations (Tagane et al., 2017). Pavonia rigida (Wall. ex Mast.) Hochr. — Herbaceous creeper. Reported by Dy Phon (1970) from heathland area as synonymous with Urena rigida Wall. Both genera may now be synonyms of Hibiscus. Sterculia parviflora Roxb. ex G.Don — Tree, occasional in evergreen forest from middle to high elevations, around 1,014 m (Tagane et al., 2017). Melastomataceae Medinilla rubicunda (Jack) Blume — Small tree, often epiphytic on tree trunks and rocks, somewhat common in moist evergreen forest on the plateau, around 1,014 m (Tagane et al., 2017). Melastoma malabathricum L. subsp. normale (D.Don) Karst. Mey. — Shrub in sclerophyllous stunted forest on rocky sandy soil near field station, near top of plateau at 1,049 m, Middleton and Monyrak 664. Reported by Dy Phon (1970) as Melastoma normale. Melastoma pellegrinianum (H.Boissieu) Karst.Mey. — Shrub to small tree, common in disturbed semi-evergreen forest in lowland and open sunny bogs on the plateau, around 1,014 m (Tagane et al., 2017). Melastoma saigonense (Kuntze) Merr. — Wet grassland. Reported by Dy Phon (1970) as Melastoma villosum Sims which is a later homonym of M. villosum Aublet. Melastoma sanguineum Sims — Woody shrub in heathland areas. Reported by Dy Phon (1970). Tagane et al. (2017) report this species only from disturbed forest margins at low and middle elevation. Memecylon bokorense Tagane — Shrub to small tree, occasional in understorey of moist evergreen forest at higher elevations, especially common around Popokvil Waterfall (Tagane et al., 2017). Endemic to Bokor. Memecylon lilacinum Zoll. & Moritzi — Shrub in montane forests collected at 928 m (Tagane et al., 2017). © Centre for Biodiversity Conservation, Phnom Penh Toona ciliata M.Roem. — Tree, rare at higher elevations, around 970 m (Tagane et al., 2017). Menispermaceae Hypserpa nitida Miers — Climber, occasional in moist evergreen forest on the plateau at 1,014–1,043 m (Tagane et al., 2017). Moraceae Ficus consociata Blume — Small tree, often found near streams at higher elevations, around 970 m (Tagane et al., 2017). Ficus heteropleura Blume — Hemi-epiphytic shrub or tree, common in moist evergreen forest on the plateau, around 962–1,014 m (Tagane et al., 2017). Ficus ischnopoda Miq. — Subshrub in stunted forest. Reported by Dy Phon (1970). Tagane et al. (2017) records this species from open rapidly flowing streams at middle elevations but not the plateau. Ficus sumatrana (Miq) Miq. — Tree in stunted forest. Reported by Dy Phon (1970). Ficus sundaica Blume — Small tree, common in moist evergreen forest on the plateau, and also found in the lowlands (Tagane et al., 2017). Streblus indicus (Bureau) Corner — Small tree in stunted forest. Reported by Dy Phon (1970). Tagane et al. (2017) report it as a scandent or erect tree, common in moist evergreen forest, especially abundant along lower streamside of Popokvil Waterfall below 930 m. Myristicaceae Horsfieldia amygdalina (Wall.) Warb. — Tree, common in lowland, occasional in middle elevation, and rare on the plateau, around 970–1,014 m (Tagane et al., 2017). Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Myrtaceae Decaspermum montanum Ridl. — Small tree, somewhat common in dense evergreen forest on the plateau, around 970–1,043 m (Tagane et al., 2017). Melaleuca leucadendra (L.) L. — Tree in boggy areas with saturated soils. Reported by Dy Phon (1970). Rhodamnia dumetorum (DC.) Merr. & L.M.Perry — Shrub in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 657. Also reported by Tagane et al. (2017) as locally common in somewhat disturbed areas in both lower and higher elevations. Rhodomyrtus tomentosa (Aiton) Hassk. — Shrub to small tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,048 m, Middleton and Monyrak 663. Also reported by Tagane et al. (2017) as occasional in disturbed areas in both lower and higher elevations. Syzygium antisepticum (Blume) Merr. & L.M.Perry — Prostrate shrub to small tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,054 m, Middleton and Monyrak 669. Also reported by Tagane et al. (2017) as fairly common in evergreen forest at higher elevations and open sunny bog on the plateau. In the latter environment, this species grows as a dwarf shrub, 40 cm tall, having smaller and thicker leaves (Tagane et al., 2017). Included as Syzygium zeylanicum (L.) DC. by Rundel et al. (2003). Syzygium attenuatum (Miq.) Merr. & L.M.Perry — Small tree, occasional in dense evergreen forest at higher elevations, around 1,014–1,043 m (Tagane et al., 2017). Syzygium bokorense W.K.Soh & J.Parn. — Shrub on sandy soil in high rainfall area on way to Popokvil Waterfall at 936 m, Middleton and Monyrak 610, 630. Also reported by Tagane et al. (2017) as common in moist evergreen forest and its margins on the plateau, around 1,014 m. Endemic to Bokor. Syzygium claviflorum (Roxb.) Wall ex Steud. — Shrub to small tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,045 m, Middleton and Monyrak 660. Also reported by Tagane et al. (2017) as common in open bogs on the plateau, around 938–1,043 m. Syzygium formosum (Wall.) Masam. — Small tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,056 m, Middleton and Monyrak 671. Also reported by Tagane et al. (2017) as fairly common along rapid streams and in swamp forests on the plateau. Syzygium hancei Merr. & L.M.Perry — Small tree in thin soil on rocky ground beside Popokvil Waterfall at 920 m, Middleton and Monyrak 623. Tagane et al. (2017) treated Cambodian Journal of Natural History 2017 (1) 17–37 this species as Syzygium mekongense (Gagnep.) Merr. & L.M.Perry. Syzygium jambos (L.) Alston var. sylvaticum (Gagnep.) Merr. & L.M.Perry — Tree, somewhat common in middle and higher elevations, around 970 m (Tagane et al., 2017). Syzygium lineatum (DC.) Merr. & L.M.Perry — Tree in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 605. Also reported by Tagane et al. (2017) as somewhat commonly found in evergreen forest from low to high elevations on the plateau, around 1,014 m. Tristaniopsis merguensis (Griff.) Peter G.Wilson & J.T.Waterh. — Common tree in stunted forest. Reported by Dy Phon (1970). Also reported by Tagane et al. (2017). N.B. The genus Syzygium is a diverse group at Bokor, and questions remain about the species present on the plateau. Soh and Parnell (2015) recently revised the species for Indochina. Nepenthaceae Nepenthes bokorensis Mey — Herb in seasonally inundated area on sandy soil at around 944 m, Middleton and Monyrak 587, 589, 592, 602. Reported by Dy Phon (1970) as Nepenthes thorelii Lecomte. Listed as Nepenthes kampotiana Lecomte in Rundel et al. (2003). Endemic to Bokor. Nyssaceae Nyssa javanica (Blume) Wangerin — Tall tree in moist evergreen montane forests and reaching 928 m (Tagane et al., 2017). Ochnaceae Campylospermum serratum (Gaertn.) Bittrich & M.C.E. Amaral — Shrub or treelet in scrubland or stunted forest in sandy soil near top of plateau at 936–1,056 m, Middleton and Monyrak 596, 612, 636. Also reported by Tagane et al. (2017) under the name Gomphia serrata (Gaertn.) Kanis as common in evergreen forest on the plateau. Oleaceae Jasminum lanceolaria Roxb. — Climber, somewhat common in both lower and higher elevations (Tagane et al., 2017). Jasminum nobile C.B.Clarke — Woody climber in stunted evergreen sclerophyllous forest along edge of track on plateau at 989–1,042 mm, Middleton and Monyrak 593, 625. Also reported by Tagane et al. (2017) as common in edge of evergreen forest on the plateau, around 1,014– 1,043 m. Olea brachiata (Lour.) Merr. — Tree, occasional in evergreen forest and its margins on the plateau around 970–1,043 m (Tagane et al., 2017). Reported by Dy Phon (1970) as Olea maritima Wall. ex G.Don. © Centre for Biodiversity Conservation, Phnom Penh 31 32 P. Rundel & D. Middleton Olea salicifolia Wall. ex G.Don — Small tree, common in moist evergreen forest on the plateau, around 1,014–1,043 m (Tagane et al., 2017). Orchidaceae Appendicula hexandra (J.Koenig) J.J.Sm. — Epiphytic orchid in stunted forest. Listed by Averyanov et al. (2013). Reported by Dy Phon (1970) as Appendicula koenigii Hook.f. Bulbophyllum lobbii Lindl. — Lithophytic epiphyte in areas of stunted forest. Reported by Dy Phon (1970). Bulbophyllum physocoryphum Seidenf. — Epiphytic orchid in evergreen forest on the plateau at 1,000 m. Reported by Averyanov et al. (2013). Bulbophyllum retusiusculum Rchb.f. — Epiphytic orchid in evergreen forest on the plateau. Reported by Averyanov et al. (2013). Bulbophyllum tenuifolium (Blume) Lindl. — Epiphytic orchid in evergreen forest on the plateau and common along streams. Reported by Averyanov et al. (2013). Calanthe cardioglossa Schltr. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970) and collected by Averyanov et al. (2012). Calanthe lyroglossa Rchb.f. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970) as Calanthe nephroidea Gagnep. Calanthe spathoidea — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). This name does not appear to have ever been published and it is not known to what it refers. Ceratostylis subulata Blume — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970) as Ceratostylis teres (Griff.) Rchb.f. Cleisostoma birmanicum (Schltr) Gerey — Epiphytic orchid. Stunted forest at 1,000 m (Schuiteman et al., 2016). Cleisostoma fuerstenbergianum Kraenzl. — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970) as Sarcanthus geoffrayi Guillaumin. Coelogyne parishii Hook.f. — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970). Conchidium muscicola (Lindl.) Rauschert — Epiphyte in areas of stunted forest. Reported by Dy Phon (1970) as Eria muscicola (Lindl.) Lindl. Dendrobium revolutum Lindl. — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970). Dendrobium scabrilingue Lindl. — Epiphytic orchid in evergreen forest. Reported by Averyanov et al. (2016). Dendrobium tenellum (Blume) Lindl. — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970). © Centre for Biodiversity Conservation, Phnom Penh Eria biflora Griff. — Lithophytic or epiphytic orchid in stunted forest. Reported by Dy Phon (1970) Eria lasiopetala (Willd.) Ormerod — Lithophytic or epiphytic orchid in stunted forest. Reported by Dy Phon (1970) as Eria albidotomentosa (Blume) Lindl. Eria tenuiflora Ridl. — Epiphytic orchid in cloud forest on the plateau (Averyanov et al., 2013). Liparis acuminata Hook.f. — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970). Mycaranthes floribunda (D.Don) S.C.Chen & J.J.Wood. — Lithophytic or epiphytic orchid in stunted forest. Reported by Dy Phon (1970) as Eria paniculata Lindl. Oberonia falcata King & Pantl. — Epiphytic orchid in evergreen montane forest at 940 m (Schuiteman et al., 2016. Papilionanthe pedunculata (Kerr) Garay. — Terrestrial orchid climber in stunted forest. Reported by Dy Phon (1970) as Aerides pedunculata Kerr. Pholidota articulata Lindl. — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970). Plocoglottis bokorensis (Gagnep.) Seidenf. — Terrestrial herbaceous perennial. Submontane forests of Bokor. Reported by Nuraliev (2014) without elevation, but collected at 1,300 m at Khao Yai, Thailand. Spathoglottis pubescens Lindl. — Terrestrial orchid in wetland areas and stunted forest. Reported by Dy Phon (1970). Stichorkis gibbosa (Finet) J.J.Wood — Epiphytic orchid in evergreen montane forest at 940 m (Schuiteman et al. 2016). Trichotosia velutina (Lodd. ex Lindl.) Kraenzl. — Epiphytic orchid in stunted forest. Reported by Dy Phon (1970) as Eria velutina Lodd. ex Lindl. N.B. Averanov et al. (2013) collected a number of orchid species from Bokor without elevation data. Pandanaceae Pandanus capusii Martelli — Subshrub up to 80 cm. Stunted forest and heathland areas. Reported by Dy Phon (1970). Pandanus cupribasalis H.St.John — Understorey of open forest stands, especially near Popokvil Waterfall. Typically 2–3 m in the lower forest stands but as tall as 8 m in wet forest at lower elevations (Stone, 1971). Pentaphylacaceae Anneslea fragrans Wall. — Tree, occasional on the top plateau, and rare in the lowland, around 1,011–1,043 m (Tagane et al., 2017). Reported by Dy Phon (1970) as Anneslea sp. Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Cleyera bokorensis Nagam. & Tagane, ined. — Rheophytic shrub to small tree, common along streams at higher elevations, around 991–1,043 m (Tagane et al., 2017). Endemic to Bokor. Eurya nitida Korth. var. nitida — Shrub in sclerophyllous stunted forest near field station, near top of plateau at 1,050 m, Middleton and Monyrak 665. This appears to be what Dy Phon (1970) reports as Eurya japonica Thunb. Tagane et al. (2017) report only Eurya trichocarpa Korth. from upper middle elevations and not the plateau. Ternstroemia gymnanthera (Wight & Arn) Bedd. — Small tree, occasional in moist evergreen forest on the plateau, around 928–1,043 m (Tagane et al., 2017). Phyllanthaceae Antidesma montanum Blume var. montanum — Small tree in dwarf forest on sandy soil beside track on way to Popokvil Waterfall at 1,000 m, Middleton and Monyrak 650. Also reported as Antidesma montanum by Tagane et al. (2017) as fairly common in moist evergreen forest on the plateau, around 930–1,014 m. Aporosa yunnanensis (Pax & K.Hofm.) F.P.Metcalf — Tree, rare in evergreen forest at higher elevations, around 970 m (Tagane et al., 2017). Glochidion lanceolarium (Roxb.) Voigt — Tree 3–4 m tall. Stunted forest. Reported by Dy Phon (1970). Tagane et al. (2017) record this species only at low elevation. Glochidion hypoleucum (MIq.) Boerl. — Tree in stunted forest. Reported by Dy Phon (1970) as Glochidion glaucifolium Müll.Arg. Glochidion rubrum Blume — Small tree, occasional in edge of evergreen forest, around 930–1,014 m (Tagane et al., 2017). Reported by Dy Phon (1970). Phyllanthus bokorensis Tagane — Small tree, common at streamside, especially along lower stream of Popokvil Waterfall, and in open areas on the plateau, around 1,014 m (Tagane et al., 2017). Endemic to Bokor. Phyllanthus kampotensis Beille — Shrub in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 645. Phyllanthus roseus (Craib & Hutch.) Beille — Small tree, occasional in understorey of moist evergreen forest and its margin on the plateau at 1,014 m (Tagane et al., 2017). N.B. Both Glochidion and Phyllanthus require more careful study. Pittosporaceae Pittosporum balansae A.DC. — Shrub in understorey of stunted forest. Reported by Dy Phon (1970) as Pittosporacus balansae, possibly a typological error. Cambodian Journal of Natural History 2017 (1) 17–37 Pittosporum pauciflorum Hook. & Arn — Small tree, common in moist evergreen forest on the plateau, around 975–1,043 m (Tagane et al., 2017). N.B. More study may determine that these two taxa are the same. Poaceae Bambusa sp. — Edge of shrub stands in heathland area. Reported by Dy Phon (1970). Eremochloa eriopoda C.E.Hubb. — Common grass in bog areas and seasonally waterlogged soils. Listed by Rundel et al. (2003). Imperata cylindrica (L.) P.Beauv. — Forest edge and roadside. Reported by Dy Phon (1970). Panicum sp. — Roadside. Reported by Dy Phon (1970). Also listed by Rundel et al. (2003). Phragmites aff. karka (Retz.) Trin. ex Steud. — Common in disturbed areas at forest edge and roadside. Reported by Dy Phon (1970). Polygalaceae Polygala arillata Buch.-Ham. ex D.Don — Shrub, locally common around moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Polygala tonkinensis Chodat — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Salomonia longiciliata Kurz — Boggy vegetation indicating seasonal waterlogging on sandy soil beside track towards Popokvil Waterfall at 936 m, Middleton and Monyrak 639. Xanthophyllum ellipticum Korth. ex Miq. — Small tree, scattered in moist evergreen forest and its vicinity on the plateau at 930 m (Tagane et al., 2017). Polygonaceae Polygonum chinense L. — Herbaceous perennial. Reported by Dy Phon (1970) as common in heathland areas. Primulaceae Ardisia crenata Sims subsp. crassinervosa (E.Walker) C.M.Hu & Vidal — Treelet in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 642. Also reported by Tagane et al. (2017) as common in moist evergreen forest and its vicinity on the plateau around 1,014 m. Ardisia quinquegona Blume var. quinquegona — Small tree, occasional in evergreen forest from middle to high elevations, around 970–1,043 m (Tagane et al., 2017). Ardisia sanguinolenta Blume — Small tree, widely and commonly found from middle to high elevations (Tagane © Centre for Biodiversity Conservation, Phnom Penh 33 34 P. Rundel & D. Middleton et al., 2017). Reported by Dy Phon (1970) as Ardisia colorata Roxb. Ardisia smaragdina Pit. — Shrub on sandy soil in scrubland near top of plateau at 1,056 m, Middleton and Monyrak 595. Also reported by Tagane et al. (2017) as common in moist evergreen forest on the plateau, around 1,014–1,043 m. Labisia pumila (Blume) Fern.-Vill. — Shrub, occasional in moist evergreen forest, often occurs near streams, around 941 m (Tagane et al., 2017). Maesa ramentacea (Roxb.) A.DC. — Shrub in understorey of stunted forest. Reported by Dy Phon (1970). Tagane et al. (2017) report this species from middle elevations. Rapanea neriifolia (Siebold & Zucc.) Mez var. macrocarpa (Pit.) C.M.Hu — Tree, occasional in moist evergreen forest on the plateau, around 1,005–1,043 m (Tagane et al., 2017). Proteaceae Helicia elephanti Sleumer — Shrub in stunted forest on the plateau. Reported by Dy Phon (1970). Also reported by Tagane et al. (2017) as a small tree, occasional along streams at higher elevations and particularly abundant along the lower stream of Popokvil Waterfall. Endemic to Bokor. Helicia vestita W.W.Sm. — Tree, somewhat common in hill evergreen forest at 800–940 m (Tagane et al., 2017). Restionaceae Centrolepis cambodiana Hance — Tufted herb on sandy soil in stunted forest beside inundated area on way to Popokvil Waterfall at 936 m, Middleton and Monyrak 632. Also listed by Rundel et al. (2003). Dapsilanthus disjunctus (Mast.) B.G.Briggs & L.A.S.Johnson — Clump forming herb in boggy vegetation in seasonally inundated area beside track towards Popokvil Waterfall at 936 m, Middleton and Monyrak 655. Included as Leptocarpus disjunctus Mast. in Rundel et al. (2003). Rhamnaceae Frangula crenata (Siebold & Zucc.) Miq. — Shrub to small tree, common in open bog and its surroundings on the plateau, at 926 m (Tagane et al., 2017). Rosaceae Prunus grisea (Blume ex Müll.Berol.) Kalkman var. tomentosa (Koord. & Valeton) Kalkman — Small tree, fairly common in moist evergreen forest on the plateau, and along the stream at higher elevations, 930–1,043 m (Tagane et al., 2017). Rhaphiolepis indica (L.) Lindl. — Small tree in thin soil on rocky ground beside Popokvil Waterfall at 920 m, Middleton and Monyrak 619. Also reported by Tagane © Centre for Biodiversity Conservation, Phnom Penh et al. (2017) as fairly common in moist evergreen forest on the plateau, and along the stream at higher elevations, around 933–1,043 m. Rhaphiolepis mekongensis (Cardot) Tagane & H.Toyama — Tree, common in moist evergreen forest on the plateau. Rubus rugosus Sm. — Woody creeper. Reported by Dy Phon (1970) from heathland areas. Rubus rosaefolius S.Vidal — Woody subshrub. Heathland areas. Reported by Dy Phon (1970). Sorbus corymbifera (Miq.) T.H.Nguyên & Yakovlev — Tree in stunted forest. Reported by Dy Phon (1970) as Sorbus granulosa (Bertol.) Rehder. Also reported by Tagane et al. (2017) as occasional in moist evergreen forest on the plateau, around 988–1,014 m. Rubiaceae Argostemma fasciculata Sridith & Larsen — Perennial herb, in mixed shrubby sclerophyllous montane forest rich in epiphytes (Sridith & Larsen, 2004). Endemic to Bokor. Canthium cambodianum Pit. — Small tree, rare in moist evergreen forest on the plateau, around 970–1,014 m (Tagane et al., 2017). Chassalia curviflora (Wall.) Thwaites — Treelet in heathland areas. Reported by Dy Phon (1970). Also reported by Tagane et al. (2017) as a shrub, commonly and widely found in the understorey of evergreen forest from middle to high elevations. Coelospermum truncatum (Wall.) Baill. ex K.Schum. — Woody climber, somewhat common at higher elevations, at forest edge along roadside, around 1,014 m (Tagane et al., 2017). Gaertnera sralensis (Pierre ex Pit.) Kerr — Shrub in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,053 m, Middleton and Monyrak 668. Also reported by Tagane et al. (2017) as fairly common in understorey of moist evergreen forest at higher elevations, around 1,014–1,043 m. Gynochthodes sublanceolata Miq. — Climber, occasional in moist evergreen forest at higher elevations, around 970 m (Tagane et al., 2017). Hedyotis rosmarinifolia (Pit.) Craib — Herb with woody base in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,065 m, Middleton and Monyrak 580. Also listed by Rundel et al. (2003). Hedyotis scandens Roxb. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Ixora brunonis Wall. ex G.Don subsp. kratensis (Craib) Chamch. — Shrub, somewhat common in understorey of moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Ixora villosa Roxb. var. chevalieri Pit. — Shrub in stunted evergreen sclerophyllous forest along edge of track on plateau at 989 m, Middleton and Monyrak 594. Psychotria asiatica L. — Shrub, occasional in evergreen forest and its vicinity at higher elevations, around 960 m (Tagane et al., 2017). Lasianthus cambodianus Pit. — Shrub, occasional in moist evergreen forest at higher elevations, around 935 m (Tagane et al., 2017). Psychotria sarmentosa Blume var. membranacea (Pit.) P.H.Hô — Climber, scattered in moist evergreen forest at higher elevations, around 1,014 m (Tagane et al., 2017). Lasianthus chinensis (Champ.) Benth. — Shrub, occasional in moist evergreen forest and its vicinity, around 960–1,014 m (Tagane et al., 2017). Psychotria serpens L. — Climber, occasional in moist evergreen forest at higher elevations, around 974–1,043 m (Tagane et al., 2017). Reported by Dy Phon (1970). Lasianthus chrysoneurus (Korth.) Miq. — Shrub in understorey of stunted forest. Reported by Dy Phon (1970) as Lasianthus hoaensis Pierre ex Pit. Tagane et al. (2017) record this only at upper middle elevations. Psychotria sp. — Treelet in stunted forest on sandy soil beside track on way to Popokvil Waterfall at 1,000 m, Middleton and Monyrak 653. Lasianthus curtisii King & Gamble. — Shrub occasional at higher elevations around 928 m (Tagane et al., 2017). Lasianthus fordii Hance. — Shrub, common in understorey of evergreen forest at higher elevations, around 962 m (Tagane et al., 2017). Reported by Dy Phon (1970) as Lasianthus kamputensis Pierre ex Pit. Lasianthus giganteus Naiki — Treelet in moist evergreen forest, locally abundant near Popokvil sphagnum bog at 960 m (Tagane et al., 2017). Lasianthus hirsutus (Roxb.) Merr. — Shrub, commonly and widely found from stunted to high elevations, around 970–1,014 m (Tagane et al., 2017). Lasianthus inodorus Blume — Shrub, occasional at higher elevations, around 935 m (Tagane et al., 2017). Reported by Dy Phon (1970) as Lasianthus poilanei Pit. Lasianthus sp. — Shrub, occasional at higher elevations, around 941–970 m. Reported as “Lasianthus sp. 3” by Tagane et al. (2017). Mussaenda cambodiana Pierre ex Pit. — Climber, common at the margin of evergreen forest in middle to high elevations, around 930–1,043 m (Tagane et al., 2017). Probably the same as Middleton and Monyrak 672 from sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,057 m. Oldenlandia tenelliflora (Blume) Kuntze — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970) as Borreria stricta (L.f.) K.Schum. Ophiorrhiza sanguinea Blume — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970) Pavetta graciliflora Wall. ex Ridl. — Shrub to small tree, common in moist evergreen forest and its margin at higher elevations, around 940–1,014 m (Tagane et al., 2017). Prismatomeris tetrandra (Roxb.) K.Schum. subsp. tetrandra — Small tree, somewhat common and widely found in evergreen forest, around 970 m (Tagane et al., 2017). Reported by Dy Phon (1970) as Prismatomeris albidiflora Thwaites. Cambodian Journal of Natural History 2017 (1) 17–37 Psydrax sp. — Climber, occasional in moist evergreen forest at higher elevations, around 1,014 m (Tagane et al., 2017). Tarenna quocensis Pit. — Shrub to small tree, occasionally found in middle to high elevations (Tagane et al., 2017). N.B. The Rubiaceae is a particularly large and difficult family. Further study is necessary to ensure the same species concepts are used by each author. Rutaceae Acronychia pedunculata (L.) Miq. — Shrub or treelet in sclerophyllous stunted forest on rocky sandy soil near field station, at the top of the plateau at 1,042 m, Middleton and Monyrak 609, 667. Also reported by Tagane et al. (2017) as common in moist evergreen forest at higher elevations, around 1,014–1,043 m . Melicope lunu-ankenda (Gaertn.) T.G.Hartley — Tree, locally common in moist evergreen forest on the plateau at 928 m (Tagane et al., 2017). This appears to be what was reported by Dy Phon (1970) as Euodia triphylla (Lam.) DC., although Tagane et al. (2017) also report another species, Melicope pteleifolia (Champ. ex Benth.) T.G.Hartley, in the upper montane forest. Salicaceae Casearia grewiifolia Vent. var. grewiifolia — Small tree, rare in moist evergreen forest and its margin on the plateau at 970 m (Tagane et al., 2017). Homalium cochinchinensis (Lour.) Druce — Small tree, locally common in moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Santalaceae Dendrotrophe varians (Blume) Miq. — Woody climbing root parasite in moist evergreen forest at higher elevations, around 1,014 m (Tagane et al., 2017). Reported by Dy Phon (1970) from heathland areas. Sapindaceae Guioa diplopetala (Hassk.) Radlk. — Treelet on sandy soil in high rainfall area in stunted forest beside inundated © Centre for Biodiversity Conservation, Phnom Penh 35 36 P. Rundel & D. Middleton area on way to Popokvil Waterfall at 936 m, Middleton and Monyrak 631. Also reported by Tagane et al. (2017) as common in moist evergreen forest on the plateau at 1,014 m. N.B. The genus Smilax is diverse at Bokor and deserving of more detailed study. Tagane et al. (2017) report four undescribed species, including two that occur on the plateau. Mischocarpus pentapetalus (Roxb.) Radlk. — Tree, commonly found in middle to high elevations, around 970–1,043 m (Tagane et al., 2017). Stemonuraceae Mischocarpus sundaicus Blume — Tree, occasional at higher elevations, around 935 m (Tagane et al., 2017). Nephelium hypoleucum Kurz — Tree in stunted forest on sandy soil beside track on way to Popokvil Waterfall at 1,000 m, Middleton and Monyrak 649. Also reported by Tagane et al. (2017) as common in evergreen forest at 970 m. Schisandraceae Illicium cambodianum Hance — Small tree, common in moist evergreen forest at higher elevations, around 935–1,043 m (Tagane et al., 2017). Illicium griffithii Hook.f. & Thomson — Treelet to 1.5 m in heathland areas. Reported by Dy Phon (1970). This may be the same as Illicium tenuifolium (Ridl.) A.C.Sm., a locally common shrub in moist evergreen forest in upper middle elevations reported by Tagane et al. (2017). Schoepfiaceae Schoepfia fragrans Wall. — Small tree, scattered in moist evergreen forest on the plateau, around 1,014–1,043 m (Tagane et al., 2017). Smilacaceae Heterosmilax paniculata Gagnep. — Climber, common in evergreen forest from middle to high elevations (Tagane et al., 2017). Smilax cambodiana Gagnep. — Semi-woody climber. Reported by Dy Phon (1970) from stunted forest areas. Smilax corbularia Kunth subsp. corbularia — Climber, common in moist evergreen forest on the plateau, around 941–1,043 m (Tagane et al., 2017). Smilax davidiana A.DC. — Climber on sandy soil in scrubland near the top of the plateau at 1,056 m, Middleton and Monyrak 601. Smilax glabra Roxb. — Climber, common in moist evergreen forest at higher elevations, around 991–1,014 m (Tagane et al., 2017). Reported by Dy Phon (1970) from heathland areas. Gomphandra cambodiana Pierre ex Gagnep. — Tree, occasional in evergreen forest at higher elevations, around 935 m (Tagane et al., 2017). Symplocaceae Symplocos caudata Wall. ex G.Don — Small tree, occasional in moist evergreen forest at middle and high elevations, 975–1,014 m (Tagane et al., 2017). Symplocos theifolia D.Don — Small tree, occasional in evergreen forest from middle to high elevations (Tagane et al., 2017). This species was reported as Symplocos lucida (Thunb.) Siebold & Zucc. by Nooteboom & Vidal (1977) but that is an illegitimate name. Theaceae Schima wallichii (DC.) Korth. — Shrub to small tree in stunted forest on sandy soil near summit on roadside towards research centre, 936–1,060 m, Middleton and Monyrak 633, 648. 675. Tagane et al. (2017) and Dy Phon (1970) treat this as Schima crenata Korth., a name considered to be a synonym in the Flora of Thailand. Thymeleaceae Eriosolena composita (L.f.) Tiegh. — Shrub or tree, occasional in edge of moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Reported by Dy Phon (1970) as Daphne composita (L.f.) Gilg. Wikstroemia bokorensis E.Oguri & Tagane, ined. — Shrub, rare in edge of moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Presumably endemic to Bokor. Wikstroemia longifolia Lecomte — Shrub, occasional in edge of moist evergreen forest on the plateau, 963–1,014 m (Tagane et al., 2017). Verbenaceae Lantana camara L. — Stunted woody subshrub along roadsides in heathland area. Reported by Dy Phon (1970). Naturalized non-native. Vitaceae Smilax inversa T.Koyama — Climber, rare in moist evergreen forest on the plateau at 1,014 m (Tagane et al., 2017). Cayratia japonica (Thunb.) Gagnep. var. mollis (Wall. ex M.A.Lawson) Momiy. — Herbaceous climber, common along edge of evergreen forest in middle elevation at 970 m (Tagane et al., 2017). Smilax lanceifolia Roxb. — Climber, common in moist evergreen forest and its vicinity on the plateau, around 975–1,014 m (Tagane et al., 2017). Reported by Dy Phon (1970) from heathland areas. Tetrastigma ramentaceum Planch. — Semi-woody climber, somewhat common in moist evergreen forest on the plateau, 970–1,063 m (Tagane et al., 2017). Reported by Dy Phon from heathland and stunted forest areas. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 17–37 Flora of Bokor Plateau Xyridaceae Xyris complanata R.Br. — Herbaceous perennial in open wetlands and bogs. Reported by Dy Phon (1970) and Rundel et al. (2003). Zingiberaceae Alpinia oxyphylla Miq. — Rare herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Amomum repoeense Pierre ex Gagnep. — Herbaceous perennial in stunted forest. Reported by Dy Phon (1970). Globba bokorensis Nob.Tanaka & Tagane — Herbaceous perennial, occasional in open Sphagnum bog and semishaded moist evergreen forests on the plateau, often epiphytic on trunk and rocks (Tanaka et al., 2015). Endemic to Bokor. Ack now le dge m e nt s We thank Meng Monyrak, Sok Sothea, and Hong Lork for field assistance, and the Department of Nature Conservation and Protection in the Cambodian Ministry of the Environment for arranging permission to work in Bokor National Park. Our field work was greatly assisted by M. Rasoul Sharifi, Kansri Boonpragob, Judy King-Rundel, and the late Mark Patterson. We gratefully acknowledge the logistic support of the national park staff in providing housing. We thank Hans-Joachim Esser for help on the correct name for Schefflera incisa. We also thank two anonymous reviewers for their valuable comments. This project was funded in part by the UCLA Asian Studies Center and Arnold Arboretum of Harvard University. Re fe re nc e s APG IV—Angiosperm Phylogeny Group (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society, 181, 1–20. Averyanov, L.V., Nguyen K.S., Maisak, T.V., Konstantinov, E.L., Nguyen T.H. & Bounphanmy, S. (2013) New and rare orchids (Orchidaceae) in the flora of Cambodia and Laos. Turczaninowia, 16, 26–46. Averyanov, L.V., Ponert, J., Nguyen P.T., Duy N.V., Khang N.S. & Nguyen V.C. (2016) A survey of Dendrobium Sw. sect. Formosae (Benth. & Hook. f.) Hook. f. in Cambodia, Laos and Vietnam. Adansonia, 38, 199–217. Cho S.-H., Kim B.-Y., Park H.-S., Chhang P. & Kim Y.-D. (2017) Impatiens bokorensis (Balsaminaceae), a new species from Cambodia. PhytoKeys, 77, 33–39. Cho S.H., Lee J.H., Won H., Chhang P., C. & Kim Y.D. (2015) Sonerila bokorense (Melastomataceae), a new species from Cambodia. Phytotaxa, 222, 295–299. Dy Phon P. (1970) La vegetation du sud-ouest du Cambodge. Annales de la Faculté Sciences de Phnom Penh, 3, 1–136; 4, 1–59. Grubb, P.J. (1971) Interpretation of the “Massenerhebung” effect on tropical mountains. Nature, 229, 44–45. Grubb, P.J. (1977) Control of forest growth and distribution on wet tropical mountains with special reference to mineral Cambodian Journal of Natural History 2017 (1) 17–37 nutrition. Annual Review of Ecology and Systematics, 8, 83–107. Kowalczyk, A. (2009) ‘Mountain resorts’: origins and evolution. Tourism, 19, 33–41. Nooteboom, H.P. & Vidal, J.E. (1977) Flore du Cambodge, du Laos et du Viet-nam (revision de la flore generale de l’Indochine): 16. Symplocacees. Museum National d’Histoire Naturelle, Paris, France. Nuraliev, M.S., Averyanov, L.V., Kuznetsov, A.N. & Kuznetsova, S.P. (2015) Review of the genus Plocoglottis (Orchidaceae) in Cambodia, Laos and Vietnam. Wulfenia, 22, 189–199. Phnom Penh Post (2012) City on a hill sparks little talk. March 12, 2012. Http://www.phnompenhpost.com/index. php/2012031655065/National-news/city-on-a-hill-sparkslittle-talk.html [accessed 1 March 2017]. PPG I—Pteridophyte Phylogeny Group (2016) A communityderived classification for extant lycophytes and ferns. Journal of Systematic Evolution, 54, 563–603. Rundel, P.W. (1999) Forest habitats and flora in Lao PDR, Cambodia, and Vietnam. WWF Indochina Programme, Hanoi, Vietnam. Rundel, P.W., Middleton, D.J., Patterson, M. & Monyrak M. (2003) Structure and ecological function in a tropical montane sphagnum bog of the Elephant Mountains, Bokor National Park, Cambodia. Natural History Bulletin of the Siam Society, 51, 185–195. Rundel, P.W., Sharifi, M.R., King-Rundel, J. & Middleton, D.J. (2016) Dacrydium elatum (Podocarpaceae) in the montane cloud forest of Bokor Mountain, Cambodia. Cambodian Journal of Natural History, 2016, 90–97. Schuiteman, A., Ryan, C., Nut M., Nay S. & Att S. (2016) New records of Orchidaceae from Cambodia II. Cambodian Journal of Natural History, 2016, 7–14. Soh W.K. & Parnell, J. (2015) A revision of Syzygium Gaertn. (Myrtaceae) in Indochina (Cambodia, Laos and Vietnam). Adansonia, 37, 179–275. Sridith, K. & Larsen, K. (2004) Argostemma fasciculata (Rubiaceae), a new species from Cambodia. Nordic Journal of Botany, 23, 169–171. Stevens, P.F. (2017) Angiosperm Phylogeny Website. Version 13. Http://www.mobot.org/MOBOT/research/APweb/ [acccessed 1 March 2017]. Stone, B.C. (1971) A preliminary survey of the Pandanaceae of Thailand and Cambodia. Natural History Bulletin of the Siam Society, 24, 1–32. Tagane S., Toyama H., Fuse K., Chhang P., Naiki A., Nagamasu H. & Yahara T. (2017) A Picture Guide of Forest Trees in Cambodia IV ~ Bokor National Park. Center for Asian Conservation Ecology, Kyushu University, Fukuoka, Japan. Https:// sites.google.com/site/pictureguides/home/cambodia/bokornational-park [accessed 1 May 2017]. Tanaka N., Tagane S., Chhang P. & Yahara T. (2015) A purple flowered new Globba (Zingiberaceae), G. bokorensis, from southern Cambodia. Bulletin of the National Museum of Natural Sciences, Series B, 41, 155–159. Tixier, P. (1979) Bryogéographie du Mont Bokor (Cambodge). J. Cramer, Vaduz, Liechtenstein. Weaver, P.L., Medina, E., Pool, D., Dugger, K., GonzalesLiboy, J. & Cuevas, E. (1986) Ecological observations in the dwarf cloud forest of the Luquillo Mountains in Puerto Rico. Biotropica, 18, 79–85. Zhang M., Tagane S., Toyama H., Kajisa T., Chhang P. & Yahara T. (2016) Constant tree species richness along an elevational gradient of Mt. Bokor, a table-shaped mountain in southwestern Cambodia. Ecological Research, 31, 495–504. © Centre for Biodiversity Conservation, Phnom Penh 37 38 T. Gray et al. St at us a nd c onse r vat ion signifi c a nc e of ground-dw e lling m a m m a ls in t he Ca rda m om Ra infore st La ndsc a pe , sout hw e st e r n Ca m bodia Thomas N.E. GRAY1,*, Andrew BILLINGSLEY2, Brian CRUDGE3, Jackson L. FRECHETTE4, Romica GROSU1, Vanessa HERRANZ-MUÑOZ5, Jeremy HOLDEN4, KEO Omaliss7, KONG Kimsreng6, David MACDONALD8, NEANG Thy6, OU Ratanak6, PHAN Channa6,8 & SIM Sovannarun3,9 1 Wildlife Alliance, No. 86, Street 123, Toul Tompong, Chamkarmorn, Phnom Penh, Cambodia. 2 Conservation International – Greater Mekong Program, 4th floor, Building B1, Phnom Penh Center, Sihanouk Boulevard, Tonle Bassac, Chamkarmorn, Phnom Penh, Cambodia. 3 Free the Bears Fund Inc., PO Box 723, Phnom Penh, Cambodia. 4 Fauna & Flora International, No. 19, Street 360, Boeung Keng Kang 1, Phnom Penh, Cambodia. 5 Bastet Conservation, No. 143, Street 105, Toul Tompong, Chamkarmorn, Phnom Penh, Cambodia. 6 Ministry of Environment, Morodok Techo Building, Chaktomuk, Daun Penh, Phnom Penh, Cambodia. 7 Forestry Administration, Ministry of Agriculture Forestry and Fisheries, No. 40, Preah Norodom Boulevard, Phnom Penh, Cambodia. 8 Wildlife Conservation Research Unit, Department of Zoology, University of Oxford, The Recanati-Kaplan Centre, Tubney House, Tubney, Oxon OX13 5QL, England. 9 Royal University of Phnom Penh, Confederation of Russia Boulevard, Toul Kork, Phnom Penh, Cambodia. * Corresponding author. Email gray@wildlifealliance.org Paper submitted 30 March 2017, revised manuscript accepted 20 June 2017. ɊɮɍɅʂɋɑɳȶſɆ ɁɸɆɅɽɳɃɑNJɈɵƙɈɌȶɃɫȲɳɉƚȣȶȹɯɌɉƒɸƙȲǏȻNjɅɃɸɒɸʑʗ.ʐʐʐȴɊʒ ɴȼɍƺɁɸɆɅɽƳɌljɌɳɃɑNJɈɑƏɩɁɳǷNJȴɅɩɌɁɪɵɅƙɆɳɃɑ ȲɊƕɭƺ ɴȼɍNjɅɌɋɺȲɊƕɑɽƸɆɽɈɪȲɊƕɑɽɅɪɎɑ ɮ ɻ ɊɭƙɃ ɌɒɮɁȼɍɽƺȶ ʑ.ʗʐʐɊʆ ɳDŽɹɆɪƺ ɁɸɆɅɽɳɃɑNJɈɳɅɹNjɅɁɵɊƚɴɇƒȲɔɉɩɌȲƞȲʁ ɳƽɋ ȲʁɂɪƗʉɳɅɹNjɅɈʂɁɾNjɅɁɩȷɁɯȷɳǷɳɓˊɋƙɁȪɎLJɅɳLJɹɈɭɊƖɑɪɈ Ǝ ɪǒƏɅNJɈɅɩȶǒɌɺɑɸƴɅɽɵɅƳɌɔɉɩɌȲƞɆɻɮɈɭɋǔɑƘɭȶɂɅɩȲ ɑɁƛɳƵȲʆ ɳɋˊȶǍɋƳɌɀɿɍɃƑɇɍƳɌɑɩȲǜɳƽɋNjɻ ɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩȷɸɅɯɅƙLJɸɈɪɌɳǷȲƒɭȶƙLJɸɁɸɆɅɽƳɌljɌɳɇƞȶʉƵƒɳǷȲƒɭȶɁɸ ɆɅɽɳɃɑNJɈDŽɸȶɊɮɍ ɌǏȶƹƒɸʒʐʑʒɅɩȶʒʐʑʖ ɴȼɍNjɅ ʒʕʕ ɃɪǂɸȶNjɻ ɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩɅɩȶƙɁȪɎƺƺȶʓʐ.ʐʐʐ ɔdžƐȲɽ ɋɆɽʆ njɻ ȶɁɩȷǁɑɽɂɅɩȲɑɁƛɳƵȲ ʓʐƙɆɳɉɃ ɈɪNjȾɊɄƘɊɳǵɄɸƙɁȪɎLJɅȲɁɽƙǂ ȲƒɭȶɳdžɹNjɅɊɯɋƙɆɳɉɃƙɁȪɎLJɅȷɭɹȲƒɭȶɆȥƅɪ ƙȲɒɊɌɆɑɽɔȶƀƳɌ IUCNƺƙɆɳɉɃȹɩɁɇɭɁɈɮȹɄƂɅɽɄƂɌ ƺɑȲɍʆ ƴƚȵƗɸɭɁɮȷ ɈɪɌƙɆɳɉɃȹɩɁɇɭɁɈɮȹ ƙLJɸɆɪƙɆɳɉɃƷɋɌȶɳƙƵɹɅɩȶɆɪƙɆɳɉɃȹɩɁɌȶɳƙƵɹ Helarctosȱ malayanus ƴƚɈɈȲ Neofelisȱ nebulosa ɅɩȶɴȸžɵƙɈ Cuonȱ alpinus ƙɁȪɎLJɅȲɁɽƙǂƙLJɸɊɯɋ ȼȶɞɳƙȷˊɅƺȶɳɅɹ ɳǷȲƒɭȶɔɸɓɭȶɳɈɍɑɩȲǜȷɸɅɯɅƙLJɸɈɪɌȼȶʆ ɳDŽɹɆɪƺɆɻɮɈɭɋǔɑƘɭȶɵɅƙɆɳɉɃDŽɸȶɆɪɳɅɹɳǷDŽɆƺȶɑɊɁƏNJɈ CITATION: Gray, T.N.E., Billingsley, A., Crudge, B., Frechette, J.L., Grosu, R., Herranz-Muñoz, V., Holden, J., Keo O., Kong K., Macdonald, D., Neang T., Ou R., Phan C. & Sim S. (2017) Status and conservation significance of ground-dwelling mammals in the Cardamom Rainforest Landscape, southwestern Cambodia. Cambodian Journal of Natural History, 2017, 38–48. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 38–48 Status of ground-dwelling mammals ƙɃƙɃȶɽɵɅɳɔȲɮɓɮɑɭɪȲɭƒȶɁɸɆɅɽȲʁɳƽɋ ȲʁɈɯȲǏNjɅǒɌɺɑɸƴɅɽǁɑɽɳǷȲƒɭȶɁɸɆɅɽDŽɸȶɊɮɍʆ ɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩɃɪǂɸȶǁɊɯɋLJɅȲɁɽƙǂɅɮɎɈɈɯȲƴƚ ɳDŽɹƺnjɻ ȶǁȲʁɳƽɋ ƵƗɅNjɻ Panthera ɳdžɹɳɃ ɳɅɹɆƷƟȻǃƙɆɳɉɃƴƚɄɸɅɩȶƴƚɌȳɩɅɃɸɅȶƺɇɭɁɈɮȹ ɳǷȲƒɭȶɁɸɆɅɽɳɅɹɳǵɳɒˊɋʆ ɳɍˊȲɴɍȶɴɁƴƚɈɪɌƙɆɳɉɃɳɅɹɳȷȻ Ʌɩȶ ƙɆɳɉɃɑɁƛljɒɅɺNjɅƙȲȷȲ (ungulates) ɴȼɍƺɔƒȲɡȲ ɳɃɑɳǷȲƒɭȶɵƙɈɌɳLJɹ ƙɆɳɉɃɂɅɩȲɑɁƛɳƵȲɌȶƳɌȴɸǍɊȲɸɴɒȶȲƒɭȶɆȥƅɪƙȲɒɊɌɆɑɽɔȶƀƳɌ IUCN ɅɩȶɂɅɩȲɑɁƛɃɫȲǒɆ ɃɸɅȶƺNjɅɎɁƎNjɅȲƒɭȶɁɸɆɅɽɳɃɑNJɈɵƙɈɌȶɃɫȲɳɉƚȣȶȹɯɌɉƒɸƙȲǏȻ ɳɈɍɂƗɪʉɳɅɹʆ ɳƽɋɈɯȲǏƙɁȪɎLJɅɳȵˊȻǂɊɌɋɺNjɻ ɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩdž ȼɮɳȷƒɹȹɯɌɉɸƒƙȲǏȻNjɅǒɌɺɑɸƴɅɽǁɑɽɑƙNjɆɽƳɌɔɉɩɌȲƞƺɑȲɍʆ ɇƐɭɋɊȲɎ ɩȻ ƳɌƙɆNjȻɽƺɈɩɳɑɑ ƳɌƽȲɽɔdžƐȲɽƙɈɊƺɊɯɋɅɫȶƳɌɳƙɆˊɑɁƛɴȸžȯɑȩȲȹɯɋƙɆNjȻɽȲɭƒȶɁɸɆɅɽɳɃɑNJɈɃɸɅȶƺǕȷɳɄƛˊɤƘNjɅƳɌɆɻɹljɍɽnjɻȶƴƚɸȶɳɍˊ ɁɵɊƚɴɇƒȲɔɉɩɌȲƞDŽɸȶɳɈɍɆȷƃɭɆƓɅƒɅɩȶɔdžȴɁʆ ɳȼˊɊƓɪɌȲǜɤƘȴȶɽɎȶƞɅɮɎNJɈȷƙɊȩɹɵɅɂɅɩȲɑɁƛɑɸƴɅɽʉLJɅȴɬ ɁƙɊȪɎɤƘNjɅƳɌ LjƚɑɽɆɮƎɌɤƘƙɆɳɑˊɌɳɓˊȶɅɮɎƳɌɃɃɯɍȳɭɑƙɁȪɎDŽɸȶɌƽƊɉɩLJɍɅɩȶɑȶƀɊɑɭɪɎ ɩɍɳȼˊɊƓɪɳȸƚˊɋɁɆɳǵɅɫȶƙɆɉɈɵɅƳɌɆɌLJȻɽɳɅɹʆ Abst ra c t The Cardamom Rainforest Landscape (CRL) is a 17,000 km2 protected landscape in southwestern Cambodia spanning an elevation range from sea-level to above 1,700 m. Despite the conservation value of the landscape there is little recent published information on the status and conservation significance of the ground-dwelling mammal populations. We report on seven camera trap studies conducted in five protected areas across the landscape between 2012 and 2016 with 255 trap-stations and >30,000 trap-nights. At least 30 species of medium to large ground-dwelling mammals were detected including one species included on the IUCN Red List as Critically Endangered, two as Endangered, eight as Vulnerable, and three as Near Threatened. Sun bears Helarctos malayanus, mainland clouded leopards Neofelis nebulosa, and dholes Cuon alpinus were detected from six or more of the seven studies. Populations of these three species in the landscape, though below ecological carrying capacity, are regionally significant. However we did not detect any Panthera cats, confirming that tigers P. tigris and leopards P. pardus are likely to have been extirpated. With the exception of these two species, and deciduous dipterocarp forest specialist ungulates, all globally threatened ground-dwelling and freshwater mammals likely to occur in the CRL have been detected in recent camera trapping surveys. The Cardamoms are thus of global conservation significance. However, poaching, particularly snaring, combined with the presence of domestic dogs across the landscape is likely to be impacting current and future conservation value strongly. The persistence of significant mammalian biodiversity requires a paradigm shift in both governmental and civil society responses to the drivers of poaching. Ke yw ords Asian elephant, by-catch, camera trap, protected area, snare, small carnivore. I nt roduc t ion The Cardamom Rainforest Landscape (CRL) is a conservation landscape covering >17,000 km2 of protected areas in the southwestern Cambodian provinces of Koh Kong, Pursat, Kompong Speu, Preah Sihanouk, Battambang, and Kompong Chhnang (Table 1; Fig. 1). The landscape spans a large elevation range from sea level to the peak of Phnom Aural—at >1,700 m Cambodia’s highest mountain—and consequently a diversity of habitat types from mangroves and lowland rainforest to limited areas of montane cloud forest. The CRL forms part of a larger conservation landscape in southern and western Cambodia with 12 largely contiguous protected areas, from Bokor National Park to Samlaut Multiple Use Cambodian Journal of Natural History 2017 (1) 38–48 Area, covering 20,680 km2. Since April 2016, the management of all protected areas in the landscapes has been under the General Department of Administration for Nature Conservation and Protection of the Ministry of Environment (MoE) (Souter et al., 2016). A number of international conservation NGOs, including Wildlife Alliance, Fauna & Flora International, and Conservation International, are active in some of the protected areas in the CRL, supporting the MoE with protected area management, law enforcement, biodiversity monitoring, and conservation outreach and community development activities. Nevertheless, despite the presence of conservation activities in the landscape and presumed significance for biodiversity, little has been published on the conservation status of the landscape’s mammals © Centre for Biodiversity Conservation, Phnom Penh 39 40 T. Gray et al. since pioneering surveys conducted at the turn of the century (e.g., Boonratana, 1999; Daltry & Momberg, 2000; Daltry & Traeholt, 2003; but see Holden & Neang, 2009; Royan, 2010; Coudrat et al., 2011). The aim of this paper is to provide a compilation of recent (post 2012) camera trapping data from the landscape in order to provide an update on the status, and conservation significance, of the CRL’s ground-dwelling mammals. M e t hods We collated data from seven discrete systematic camera trap studies conducted between 2012 and 2016 within five of the protected areas in the CRL (Table 2). Whilst camera trapping occurred in the landscape prior to this, 2012 was chosen as a start date for our analysis because data during the study period (2012–2016) were available to the authors and did not require significant additional analysis. All of the studies deployed at least 10 camera trap stations within clearly defined survey areas of between 10 and 200 km2. Camera trapping on Phnom Dalai (site A; Fig. 1) was part of a monitoring programme for Asian elephant Elephas maximus conducted between 2010 and 2013; however we only use data from this site from between February 2012 and March 2013. Results from a camera trapping study in Peam Krasaop Wildlife Sanctuary between January and May 2015, which detected a number of threatened species, are being published separately (Thaung et al., unpublished data). All of the studies had different objectives (Table 2), used Fig. 1 Protected areas and locations of camera-trap studies within the Cardamom Rainforest Landscape, southwestern Cambodia. Abbreviations are given in Table 1 and individual letters refer to studies detailed in Table 2. Table 1 Protected areas of the Cardamom Rainforest Landscape, southwestern Cambodia. Protected Area Southern Cardamom National Park (SCNP) Size (km2) Elevation range (m a.s.l.) % deforestation 2000–20151 % Economic Land Concession2 4,104 10–980 2.7 0.3 Central Cardamom National Park (CCNP) 4,013 20–1,540 1.2 0 Phnom Samkos Wildlife Sanctuary (SWS) 3,338 10–1,717 8.2 2.3 Phnom Aural Wildlife Sanctuary (AWS) 2,538 60–1,740 8.6 20.2 Botum Sakor National Park (BSNP) 1,472 0–420 15.2 36.9 Tatai Wildlife Sanctuary (TWS) 1,443 10–520 3.1 0.5 238 0–240 5.7 0 Peam Krasop Wildlife Sanctuary (PKWS) 1 Estimated following Hansen et al. (2013). 2 From datasets held by the Ministry of Agriculture, Forestry and Fisheries and the Ministry of Environment. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 38–48 Status of ground-dwelling mammals Table 2 Camera trap studies in the Cardamom Rainforest Landscape (2012–2016) included in this paper. NP = National Park; WS = Wildlife Sanctuary. Figures for survey area and elevation are approximate. Locations Survey & traparea nights (km2) Elevation range (m) Methodology & target species Study Protected area Dates A1 Phnom Dalai, Phnom Samkos WS February 2012–March 2013 13 & 3,923 10 540–1,040 Asian elephant-targeted camera trapping with locations chosen to maximise detection of elephants B2 Central Cardamom NP December 2012–March 2013 81 pairs & 8,152 95 460–1,220 Camera trap grid for clouded leopard capture-mark-recapture with cameras set as pairs in locations to maximise clouded leopard detections. C3 Central Cardamom NP June 2015– June 2016 31 & 4,599 185 100–820 Approximate grid formation with cameras set at locations to maximise detections of large mammals. D4 Southern Cardamom NP December 2015–June 2016 67 & 8,236 200 105–620 Random grid with cameras set within 50 m of predetermined random points. E5 Tatai WS March–May 2016 14 & 969 20 140–440 Approximate grid formation with cameras targeting bears. F5 Central Cardamom NP March–May 2016 14 & 865 30 540–660 Approximate grid formation with cameras targeting bears. G3 JW Concession, Botum Sakor NP August– December 2016 35 & 3,425 180 10–380 Approximate grid formation with cameras set along trails (50%) and at random locations (50%). Organisations leading data collection: 1 Fauna & Flora International; 2 Wildlife Conservation Research Unit; 3 Conservation International; Wildlife Alliance; 5 Free the Bears. All work was done with the support of the Royal Government of Cambodia. different methodologies, equipment, and survey teams. As such, their results are not directly comparable. Nevertheless they provide a useful summary of the current status of the ground-dwelling large mammal community across the landscape. For every camera trap study we extracted records of all mammals detected excluding Scandentia (treeshrews) and Rodentia apart from the two species of Hysticidae (Malayan porcupine Hystrix brachyura and Asiatic brush-tailed porcupine Atherurus macrourus). All records of primates were retained. Throughout the paper the taxonomy and nomenclature of IUCN (2016) is used. All photographs were verified for identification by three authors (TNEG, VHM, PC) with experience of camera trapping in the region. However, as many regional studies have shown (e.g., Hla Naing et al., 2015), this approach is not foolproof and misidentifications may be present in the dataset. Images in which identifications were not possible (~6% of all encounters with ‘mammals’) Cambodian Journal of Natural History 2017 (1) 38–48 4 were excluded. The percentage of functioning (>20 nights of usable photographs) camera trap stations in each study from which each species was recorded was then calculated. We subsequently refer to this metric as ‘trapprevalence’. For each camera trap site we also calculated a camera trap encounter rate (i.e. the number of photographic events per 100 trap-nights) for each species. Re sult s Between February 2012 and December 2016, the seven studies generated data from 255 camera trap stations deployed for 30,169 trap-nights across approximately 720 km2 of the CRL (Table 2). At least 30 species of mammal were detected including one listed as Critically Endangered (Sunda pangolin Manis javanica), two as Endangered (Asian elephant Elephas maximus, dhole Cuon alpinus), eight as Vulnerable, and three as Near Threatened (Table 3). Seven species were detected from © Centre for Biodiversity Conservation, Phnom Penh 41 42 T. Gray et al. Table 3 IUCN status (CR = Critically Endangered; EN = Endangered; VU = Vulnerable; NT = Near Threatened, LC = Least Concern), trap-prevalence (% of functioning camera traps recorded from) and camera trap encounter rate (number of photographic events per 100 trap-nights) of ground-dwelling mammals across seven camera trap studies in the Cardamom Rainforest Landscape, southwestern Cambodia (2012–2016). Study sites are identified in Table 2. Study sites IUCN status A B C D E F G Northern pig-tailed macaque Macaca leonina VU 100 / 6.4 27 / 0.4 35 / 0.5 38 / 0.7 36 / 1.3 64 / 3.4 51 / 0.7 Nicobar crab-eating macaque Macaca fascicularis LC 7 / 0.9 3 / <0.1 Sunda pangolin Manis javanica CR Species 1 / <0.1 17 / 0.1 Malayan porcupine Hystrix brachyura LC Asiatic brush-tailed porcupine Atherurus macrourus LC 77 / 2.1 Dhole Cuon alpinus EN 77 / 0.4 Sun bear Helarctos malayanus VU Asiatic black bear Ursus thibetanus VU Yellow-bellied weasel Mustela kathiah LC 1 / <0.1 16 / 0.2 15 / 0.4 21 / 0.7 7 / 0.3 14 / 0.9 37 / 1.1 35 / 0.6 50 / 1.9 21 / 0.1 7 / 0.1 20 / 1.8 32 / 0.5 26 / 0.2 21 / 0.4 14 / 0.5 23 / 0.4 38 / 0.2 26 / 0.4 19 / 0.2 14 / 0.3 11 / 0.3 54 / 0.2 1 / <0.1 29 / 0.3 4 / <0.1 LC 62 / 1.5 17 / 0.2 Yellow-throated marten Martes flavigula LC 8 / <0.1 36 / 0.8 6 / 0.1 2 /<0.1 Ferret-badger Melogale sp. LC 8 / <0.1 77 / 0.6 4 / <0.1 3 / <0.1 2 / <0.1 71 / 0.8 83 / 4.0 13 / 0.1 6 / 0.1 Greater hog badger Arctonyx collaris VU VU Common palm civet Paradoxurus hermaphroditus LC Masked palm civet Paguma larvata LC Spotted linsang Prionodon pardicolor LC Large Indian Civet Viverra zibetha LC Small Indian civet Viverricula indica LC Spotted linsang Prionodon pardicolor LC Clouded leopard Neofelis nebulosa VU Asiatic golden cat Catopuma temminckii NT 21 / 0.7 2 / <0.1 Crab-eating mongoose Herpestes urva Binturong Arctictis binturong 3 / <0.1 6 / <0.1 3 / <0.1 21 / 0.4 34 / 0.9 2 / <0.1 54 / 1.0 47 / 1.3 50 / 1.2 71 / 4.7 74 / 5.8 4 / <0.1 4 / <0.1 69 / 2.1 58 / 2.0 7 / 0.3 7 / 0.1 3 / <0.1 7 / 0.1 9 /<0.1 4 / <0.1 31 / 0.3 14 / 0.1 3 / <0.1 14 / 0.2 9 / 0.1 6 /<0.1 Marbled cat Pardofelis marmorata NT Leopard cat Prionailurus bengalensis LC 38 / 0.3 Asian elephant Elephas maximus EN 85 / 1.9 Wild pig Sus scrofa LC 100 / 27.6 57 / 1.5 58 / 0.7 39 / 2.7 29 / 1.0 57 / 6.4 37 / 1.4 Lesser Oriental chevrotain Tragulus kanchil LC 23 /<0.1 41 /2.0 65 / 0.6 68 / 5.6 7 / 0.4 43 / 0.8 63 / 3.9 Sambar Rusa unicolor VU 10 / 0.2 10 / 0.1 11 / 0.4 14 / 0.5 7 / 0.2 Northern red muntjac Muntiacus muntjak LC 100 / 26.9 78 / 3.7 61 / 0.9 58 / 1.8 36 / 1.3 50 / 2.1 Chinese serow Capricornis milneedwardsii NT 8 / 0/6 10 / 0.1 3 / <0.1 15 / 0.7 Gaur Bos gaurus VU 85 / 11.0 © Centre for Biodiversity Conservation, Phnom Penh 23 / 0.3 3 /<0.1 11 / 0.1 36 / 0.7 39 / 0.3 35 / 0.9 7 / 0.1 19 / 0.2 6 / <0.1 14 / 0.2 43 / 0.7 29 / 0.7 49 / 1.6 7 / 0.1 Cambodian Journal of Natural History 2017 (1) 38–48 Status of ground-dwelling mammals all studies including northern pig-tailed macaque Macaca leonina, sun bear Helarctos malayanus (both listed as Vulnerable), lesser Oriental chevrotain Tragulus kanchil, and northern red muntjac Muntaicus vaginalis. Mean trap prevalence varied from 64% for common palm civet Paradoxurus hermaphroditus to <0.5% (i.e. detected from a single camera trap location) for ferret badger Melogale sp. Disc ussion Few places in tropical Asia support a near-intact mammal species complement (Wilcove et al., 2013) and our camera trap records confirm that the Cardamom Rainforest Landscape (CRL) is one such region. Rhinoceroses (Rhinocerotidae) have been extinct in Cambodia since at least the 1980s and the last record of tiger Panthera tigris from the country was in 2007 (Gray et al., 2012). Nevertheless we present records of 11 globally threatened species and the CRL remains nationally and regionally significant for large mammal conservation. Although significant portions of the landscape were unprotected prior to the creation of the 4,100 km2 Southern Cardamom National Park in May 2016, the CRL appears to have avoided the ecological extirpation of many medium to large mammals that has occurred due to hunting in many other protected landscapes in Indochina (Wilcox et al., 2014; Harrison et al., 2016). Almost all of the studies detected mainland clouded leopards Neofelis nebulosa (6 out of 7 studies; Fig. 2), dholes Cuon alpinus (6 out of 7; Fig. 2), and sun bears Helarctos malayanus (7 out of 7), which demonstrates that habitat quality and prey base remain reasonable within the CRL and that the pervasive snaring, which is impacting much of Southeast Asia (Gray et al., 2017), has yet to drive massive declines in the populations of these moderately hunting-sensitive species. We present camera trapping data in two forms: trap prevalence and encounter rate. However these metrics are unlikely to be directly correlated with species abundance or density. The term ‘Relative Abundance Index’ (sensu O’Brien et al., 2003) for camera trap encounter rates is highly misleading and is increasingly regarded as a meaningless measures of species abundance or status (Sollmann et al., 2013; Burton et al., 2015). We therefore recommend trap-prevalence and encounter rate be used as the terms to report by-catch information from camera trap studies when more robust methodologies to account for non-detection (e.g., Capture Mark Recapture: Gray & Prum, 2012; Occupancy: Gray, 2012; Random Encounter Model: Rowcliffe et al., 2008) are not employed. However, both trap-prevalence and encounter rate are likely to be biased. The former is likely a function of the size of the study area and, particularly, the duration of camera trap Cambodian Journal of Natural History 2017 (1) 38–48 deployment and the latter by camera trap placement in relation to a species’ daily movements amongst a myriad of other factors (Sollmann et al., 2013). Status of selected species The majority of the CRL comprises hilly evergreen forest and thus would have supported historically lower densities of ungulates and carnivores than the open deciduous dipterocarp forests of the northern and eastern plains (Gray et al., 2013). The landscape’s largest mammalian predators, tiger and leopard Panthera pardus, are likely to have been extirpated. Neither species has ever been recorded by camera trap from the CRL (though tiger was camera trapped from the adjoining Bokor National Park between 2000 and 2004) whilst there are no confirmed 21st century records of leopard from the Cardamom Mountains. Reliable surveys in the early 2000s recorded tiger pug-marks in a small number of locations (Daltry & Momberg, 2000; J. Holden pers. obs.) but there have been no records since 2005. With the exception of bears, the dhole therefore remains the largest carnivore present in the landscape, as indeed Boonratana (1999) speculated was the case as long ago as the late 1990s. Dholes still appear to be relatively widespread: detected from six of the seven studies but pack size appears low (<5; many photographs show single individuals). However Kawanishi & Sunquist (2008) suggested dholes persist in smaller packs in the evergreen forests of Southeast Asia than in the Indian subcontinent probably due to the low prey biomass and small size of ungulate prey. It is also unclear how effective camera trapping is for estimating group size of species in dense evergreen forest. Nevertheless, videos (e.g., from sites D and G; Fig. 1) often show an individual dhole limping, presumably as a result of snare injuries. It is possible that dhole numbers in the CRL are depressed due to a combination of accidental mortality from snaring, interactions with domestic dogs, and reduced prey densities: threats which impact the species across its Asian range (Kamler et al., 2015). Although clouded leopards (Fig. 2) were detected in six of our studies, trap prevalence (10%) was lower than reported elsewhere in the species’ range. For example, Tan et al. (2016) detected the species from 233 of 894 camera trap stations (trap-prevalence 26%) across nine sites in Peninsular Malaysia, with approximately 200 trap-nights required per clouded leopard photograph (compared to >750 trap-nights across our studies). This suggests that clouded leopard densities in the Cardamoms are likely to be below estimates from elsewhere in the species’ range (e.g., between 2 and 5 individuals per © Centre for Biodiversity Conservation, Phnom Penh 43 44 T. Gray et al. Fig. 2 Threatened and Near Threatened mammals in the Cardamom Rainforest Landscape. Clockwise from top-left: Asiatic golden cat Catopuma temminckii (site B, © WildCRU); Greater hog badger Arctonyx collaris (site G, © Wildlife Alliance); Marbled cat Pardofelis marmorata (site C, © Conservation International); Dhole Cuon alpinus (site G, © Wildlife Alliance); Chinese serow Capricornis milneedwardsii (site D, © Wildlife Alliance); Clouded leopard Neofelis nebulosa (site D, © Wildlife Alliance). 100 km2: Borah et al., 2014; Mohamad et al., 2015). Nevertheless the species is absent or very rare across many areas in Indochina (Wilcox et al., 2014) including much of Cambodia (e.g., Gray et al., 2014). Thus, given the size of the landscape, and detections of clouded leopard relatively close to villages and National Road 48 (e.g., <6 km at site G; Fig. 1), the CRL still seems likely to support a regionally significant population of the species. Asian elephants remain in the landscape with the species detected from three of the seven studies, plus additional ad-hoc camera trapping in the core of Botum Sakor National Park (58 ‘encounters’ from five camera trap stations between December 2013–January 2014: Fauna & Flora International, unpublished data) and Tatai Wildlife Sanctuary (two camera trap locations between December 2014–March 2015: Wildlife Alliance, unpublished data). While Daltry & Traeholt (2003) reported strong local community support for Asian elephant conservation in the CRL, an estimated 38 indi© Centre for Biodiversity Conservation, Phnom Penh viduals were poached between 2000 and 2004 (Gray et al., 2016). However we believe poaching of elephant has been extremely limited in the landscape since 2006 and therefore elephant populations may be recovering with evidence of breeding (Gray et al., 2016). Whilst population estimates and demographic data on the landscape’s elephant population have yet to be collated or analysed, field data collection for a faecal DNA capture-markrecapture study was conducted across the core of the landscape during the 2015–2016 dry season by Fauna & Flora International. A population estimate is expected during 2017. The second largest herbivore extant in the landscape, the gaur, was only recorded from three of the seven camera trap studies in the landscape and its population seems likely to be small and potentially fragmented. Sambar Rusa unicolor and Chinese serow Capricornis milneedwardsii were recorded slightly more widely, particularly in more remote areas, but detection levels were low. Nevertheless the CRL is likely to support the most nationally important populations of these two Cambodian Journal of Natural History 2017 (1) 38–48 Status of ground-dwelling mammals species, which are rarely detected in camera trap studies elsewhere in Cambodia (e.g., Phan et al., 2010; Gray & Phan, 2012). Sambar are likely to have declined significantly throughout the country as much of Cambodia is suitable for the species (Timmins et al., 2015). In contrast, mountainous habitat for Chinese serow (Fig. 2) is limited in Cambodia and the species is unlikely to have declined as precipitously. Nevertheless, recent camera trap records of serow appear restricted to the Cardamoms and Virachey National Park, where they were detected from 15 of 26 camera traps in 2014–2015 (G. McCann / HabitatID, pers. comm. 2017). The status of small carnivores in the CRL warrants further research and more detailed analysis. The high trap-occupancy of the globally Vulnerable greater hog badger from Phnom Dalai (site A; Figs 1 & 2) and the JW Concession in Botum Sakor National Park (site G), at opposite ends of the range of elevations we camera trapped, is noteworthy. Only camera trapping in Virachey National Park, northeastern Cambodia (from 500–1,400 m. a.s.l., trap-occupancy 35% from 26 stations: McCann & Pawlowski, 2017) has detected the species as frequently in Cambodia in recent years. Believed to be highly susceptible to snaring (Duckworth et al., 2016), the low number of detections of this species from the other sites is likely to reflect genuine declines driven by hunting. The JW Concession has the highest densities of patrol staff in the CRL (>8 per 100 km2). Combined with a unique management status (as an Economic Land Concession for ecotourism with Wildlife Alliance providing technical support for law enforcement), and the surrounding forest areas in Botum Sakor National Park receiving only nominal protection, such patrol levels may mean illegal activity, particularly snaring, may be low by regional and even landscape levels. It is possible that these levels of enforcement, instituted in 2014, may have allowed the recovery of hog badger and not slower breeding species (e.g., sambar). Alternatively, the JW Concession may represent prime habitat for the species differing in some respect from the majority of the landscape. Our higher elevation sites (A and B; Fig. 1) produced a wider variety of small carnivores including the first national record of yellow-bellied weasels Mustela kathiah (for more details see Phan et al., 2014) and supporting the finding of Holden & Neang (2009) that masked palm civets Paguma larvata, spotted linsangs Prionodon pardicolor, and ferret-badgers Melogale sp. are present at higher elevations within the Cardamom Mountains. Large-spotted civets Viverra megaspila was not detected in any of our studies despite records from other studies in the landscape with considerably less effort. Royan (2010) Cambodian Journal of Natural History 2017 (1) 38–48 reported a single camera trap photograph from Botum Sakor National Park in 2005, Timmins & Sechrest (2010) camera trapped two ‘in the Andoung Teuk area’ in 2008– 2009, while the species has also been camera trapped around the Wildlife Release Station, Tatai Wildlife Sanctuary (N. Marx, pers. comm. 2017) and in areas of Central Cardamom National Park (Holden & Neang, 2009; Conservation International, unpublished data). Finally Thaung et al. (unpublished data) obtained 22 records from three of their six camera trap stations in and around Peam Krasaop Wildlife Sanctuary in 2015. Many of these sites are closer to villages and thus may experience higher hunting pressure than our study sites, making it appear unlikely that the absence of this species is due to hunting. The lack of large-spotted civet detections from the relatively well protected lowland and largely flat forests of the JW Concession (site G; Fig. 1) is perplexing. Large Indian civets Viverra zibetha were detected at more than 50% of camera trap stations in studies A and B (the two highest altitude sites) but only rarely elsewhere with no detections in studies E and G (Fig. 1). In sum, most of the survey areas did not record any Viverra at all despite recording a large complement of species conventionally considered to be more hunting sensitive than this genus. The reasons for the observed patterns of Viverra civet detections are unclear and may represent complex interactions involving the detectability of these species from large mammal focused camera trapping studies, hunting pressure, and habitat preferences. The Critically Endangered Sunda pangolin Manis javanica was detected from two of the seven camera trap studies, both of which randomly deployed the cameras (sites D and G; Fig. 1). It is possible that detectability of pangolins from conventional large mammal focused camera trapping (in which cameras are often placed on trails, paths, water features etc.) may be very low and this would explain the paucity of records from camera trapping throughout tropical Asia. Sun bears Helarctos malayanus were detected from all camera trap studies and thus appears relatively widespread throughout the landscape. In contrast, the Asiatic black bear Ursus thibetanus, with only two records (at ~500 m a.s.l. at site D and ~800 m a.s.l. at site B; Fig. 1), was rarely detected away from Phnom Dalai (site A; Fig. 1) where the species was recorded by more than half of camera traps. This relatively high altitude site was the only study where Asiatic black bears were recorded more widely than sun bears. The northern pig-tailed macaque was the most widely recorded globally threatened species (detected from all studies and on average from 50% of camera trap stations) and the CRL likely supports a large population. In Laos, Vietnam, and Myanmar, the species is predominantly © Centre for Biodiversity Conservation, Phnom Penh 45 46 T. Gray et al. associated with lowland forests below 500 m (Boonratana et al., 2008). However, our study includes multiple records above this elevation in sites A and B (Fig. 1) and the species was found across the full range of elevations camera trapped. Our analysis also supports the assertion of Coudrat et al. (2011) that stump-tailed macaque Macaca arctoides may not occur in the CRL. All confirmed Cambodian records are from east of the Mekong and it seems possible that previous claims (e.g., Kong & Tan, 2002) and assertions of occurrence in southwestern Cambodia (e.g., Walston, 2001) may have been in error. Our studies did not detect any otter species but targeted camera trapping in the landscape recorded both hairy-nosed otter Lutra sumatrana and smooth-coated otter Lutrogale perspicillata between 2006 and 2012 (Heng et al., 2016). As far as we can ascertain there are no records of Eurasian otter Lutra lutra from Cambodia and also no reliable (c.f. Daltry & Momberg, 2000) records of Asian small-clawed otter Aonyx cinereus west of the Mekong in the country. There has been little camera trapping in remnant areas of deciduous dipterocarp forest or grassland in the CRL. However, Thaung et al. (unpublished data) recorded large-spotted civets (see above), hog deers Hyelaphus (porcinus) annamiticus, and Sunda pangolins Manis javanica from grassland–Melaleuca–mangrove mosaics around Peam Krasaop Wildlife Sanctuary in 2015 (six camera trap stations; 511 trap-nights). More effort in remnant areas of deciduous dipterocarp forest is required, but it seems unlikely that significant, if any, populations of dry forest specialist species (e.g., banteng Bos javanicus, Eld’s deer Rucervus eldii, jungle cat Felis chaus, golden jackal Canis aureus, small Asian mongoose Herpestes javanicus and Burmese hare Lepus peguensis) remain in the landscape. The former two species and leopards are thus the only globally threatened terrestrial or freshwater mammals known to have occurred historically in Cambodia west of the Mekong, and still extant in the country, without recent (post 2012) camera trap records from the CRL. Threats to Cardamom rainforest mammals As is the case throughout Southeast Asia (Hughes, 2016) the mammal populations of the CRL are threatened by deforestation and hunting. Cambodia experienced the most rapid growth in deforestation rates globally between 2001 and 2014 (Petersen et al., 2015) and approximately 1,150 km2 of the CRL’s protected areas have been lost to Economic Land Concessions for industrial agriculture (Table 1; Davis et al., 2015). This deforestation has disproportionately impacted lowland deciduous dipterocarp forest, particularly in protected areas with © Centre for Biodiversity Conservation, Phnom Penh limited NGO support for enforcement, and may have also isolated the core of Botum Sakor National Park from the rest of the landscape. The widespread presence of domestic dogs in accessible areas (e.g., dogs were detected from 15 of 35 camera trap stations in site G; Fig. 1) is also an issue, particularly given the landscape’s dhole population. Domestic dogs are a significant threat to wildlife through disease transmission, predation, and non-lethal effects (SilvaRodríguez & Sieving, 2012; Hughes & Macdonald, 2013). Free-ranging domestic dogs in the landscape require lethal management and protected area management authorities should be given the authority to implement this. Illegal commercial hunting, particularly snaring, remains the major threat to the CRL’s ground-dwelling mammals and is likely to be impacting populations of most such species in the landscape (Gray et al., 2017). For example, more than 109,000 snares were removed from the Southern Cardamom National Park and Tatai Wildlife Sanctuary between 2010 and 2015 (Wildlife Alliance, unpublished data) and law enforcement elsewhere in the landscape, with the notable exception of the JW Concession (see above), is minimal. This needs urgent attention through legislative reform criminalising the possession of materials used to construct snares and greater numbers of, and more efficient, protected area staff and rangers. It is hoped that the Natural Resource and Environmental Code of the MoE will sufficiently strengthen the Protected Area Law to ensure that snaring can be severely punished. Long-term social behaviour change communication, targeting the emotional and functional drivers of wild meat consumption in largely urban centres across Southeast Asia, is also critical. Any move to normalise wild meat consumption through wildlife farming needs to be strongly resisted given the potential for extremely negative impacts on biodiversity (Brooks et al., 2010; Livingstone & Shepherd, 2016). Despite the extirpation of some of Asia’s largest and most charismatic species of large mammal (e.g., two species, presumably, of rhinoceros, leopard, and tiger), our camera trap records show that the CRL remains regionally significant for the conservation of medium to large ground-dwelling mammals. However without urgent strengthening of legislation and law enforcement to reduce levels of snaring, and concurrent allocation of conservation resources to effective results-based protected area management and enforcement (Gray et al., 2016), many of these species may soon disappear from the landscape and the spectre of “empty forests” will be realised. Cambodian Journal of Natural History 2017 (1) 38–48 Status of ground-dwelling mammals Ack now le dge m e nt s All work across the Cardamom Rainforest Landscape is with the support and cooperation of the Royal Government of Cambodia, particularly the Ministry of Environment, and Ministry of Agriculture, Forestry and Fisheries. We thank HE Say Samal and HE Veng Sakhon. Chheng Tim, Chum Sokkhheng, Jan Kamler, Lim Thona, Eduard Lefter, La Peng Ly, Peng Sovannarin, Jo Ross, Tun Setha, and Ung Vises assisted with fieldwork and data management. J.W. Duckworth and an anonymous reviewer provided comments which improved the quality of the manuscript. Greg McCann, Nick Marx, and Thaung Ret provided details of recent unpublished camera trap results from the landscape and Virachey National Park. Funding came from various sources including (in alphabetical order): Australia Zoo, Barbara Delano Foundation, Daikin Corp, Golden Triangle Asian Elephant Foundation, Los Angeles Zoo, Minor Hotel Group, Perth Zoo Wildlife Conservation Action, Rainforest Trust, Tamaki Foundation, USAID (Fintrac), and U.S. Fish and Wildlife Service. Re fe re nc e s Boonratana, R. (1999) A Preliminary Wildlife Survey in the Kravanh Range of Southwestern Cambodia. Fauna & Flora International, Hanoi, Vietnam. Boonratana, R., Das, J., Yongcheng L., Htun, S. & Timmins, R.J. (2008) Macaca leonina. In The IUCN Red List of Threatened Species 2008: e.T39792A10257933. Http://dx.doi.org/10.2305/ [accessed 20 IUCN.UK.2008.RLTS.T39792A10257933.en February 2017]. 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Cat News, Special Issue 8, 53–61. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 38–48 Water quality at Anlung Pring T he im pa c t of shrim p fa r m ing on w at e r qua lit y in Anlung Pring, a prot e c t e d la ndsc a pe in Ca m bodia YAV Net1, SENG Kimhout2,4, NHIM Sophea3, CHEA Vannara3, BOU Vorsak1 & Tomos AVENT5 1 BirdlIfe International–Cambodia Programme, No.2, Street 476, Sangkat Toul Tompong 1, Khan Chamkarmon, PO Box 2686, Phnom Penh, Cambodia. 2 Forestry Administration, No. 40, Preah Norodom Boulevard, Phsar Kandal 2, Khann Daun Penh, Phnom Penh, Cambodia. 3 Department of Hydrology and River Works, Ministry of Water Resources and Meteorology, No. 364, Monivong Boulevard, Phsa Demthkov, Chamkamon, Phnom Penh, Cambodia. 4 Wildfowl & Wetlands Trust, No. 2, Street 476, Toul Tompung 1, Chamkarmon, Phnom Penh, Cambodia. 5 Wildfowl & Wetlands Trust, Slimbridge, Gloucestershire, GL2 7BT, United Kingdom. * Corresponding author. Email yavnet2011@gmail.com Paper submitted 27 December 2016, revised manuscript accepted 21 March 2017. ɊɮɍɅʂɋɑɳȶſɆ ɁɸɆɅɽȼɪɳɑˊɊɃɫȲNJƚɎɇƎɍɽƺɃɪȹƙɊȲnjɻ ȶɑɸƴɅɽɑƙNjɆɽɑɁƛǒƚɆɃɫȲ ɅɩȶƙɆɳɉɃȹɍƺɁɩɳɇƞȶʉɳɃȢɁ ɆɻɭɴɅƎɳǷNJȴƴȶɁƓɮȶ ɵɅƙɆɳɃɑ ȲɊƕɭƺ ɁɸɆɅɽɳɅɹȲɸɈɭȶɃɃɯɍɌȶɅɮɎƳɌȴɸǍɊȲɸɴɒȶƳɅɽɴɁƴƚɸȶɳɓˊȶʉ ɳƽɋǒɌƳɌɔɉɩɎȾƌɅɿǏɌ ɪɎɆƓȲɊƗʆ ɁɸɆɅɽɃɫȲNJƚɎNJȴɳƙȷˊɅɳǷ ȹɭɸɎ ɩȻɁɸɆɅɽƳɌljɌɳɃɑNJɈɔɅƚȶɽƙɈɪȶ ɴȼɍNjɅɃɪǂɸȶɉɮɊɩǒȝɑƎɑɩɁ Ə ɳǷɳȳɁƎȲɸɈɁƙɁȪɎLJɅɴȲɴƙɆɳǵƺȲɑɩƽƊɅȷɩȥƃɫɊɆƷƀɴȼɍ ɇƎɍɽɅɮɎȲƎɪȲȶƛɍɽnjɻȶƴƚɸȶǃ ǏɅɫȶǕȷɇƎɍɽɇɍɆɻɹljɍɽȼɍɽȴɭɀNJɈɃɫȲɳǷȲƒɭȶɁɸɆɅɽɔɉɩɌȲƞʆ ɳȼˊɊƓɪɳɄƛˊƳɌɳɑɭˊɆɔɳȶžɁǂɊƽɅ ɈɪɆȦ Ɵ ɳɅɹ ɳɋˊȶLJɅƙɆɊɮɍɑɸǁȲɃɫȲȷɸɅɯɅ ʓ ɴȳ (ɴȳɊȲǍ Ɋɪdž ɅɩȶɴȳəɑNJ ƹƒɸʒʐʑʖ) ɈɪƙLJɸȲɴɅƚȶɳɇƞȶʉƵƒDŽɸȶȲƒɭȶ ɅɩȶɳƙǤ ɁɸɆɅɽƳɌljɌ ɳɒˊɋɳɄƛˊƳɌɎ ɩNJȴɳȼˊɊƓɪǏɋɁɵɊƚȴɭɀNJɈɃɫȲ ɅɩȶɇɍɆɻɹljɍɽɵɅɃɫȲɑɸɀɍɽ ɴȼɍƙɁȪɎLJɅɆɳȥƃȻɈɪȲɑɩƽƊɅȷɩȥƃɫɊ ɆƷƀɳǷɴȲƓɌɳdžɹʆ ȲƙɊɩɁɵɅȷɌɅƎ (conductivity) ȲƙɊɩɁɍơȲɽ (turbidity) ɑɸɀɍɽɌȶ ɫ ɌǎɋɑɌɭɆ (TDS) ɑɼɭɍLjɁ (SO4) ǕɊɻɮȻɮ ɻɊ (NH4-N) ȴɪɊɪǒȝɑƎNjɅɁƙɊȪɎƳɌɔɭȲɑɭɪɴɑɅ (COD) ɅɩȶȹɪɎǒȝɑƎNjɅɁƙɊȪɎƳɌɔɭȲɑɭɪɴɑɅ (BOD) NjɅȲƙɊɩɁȳƕɑɽƺȶȲƙɊɩɁ ɑƎȶɽƽɌɳǷǂɊȲɴɅƚȶƙɆɊɮɍɑɸǁȲƺɳƙȷˊɅ ƺɈɩɳɑɑȲɴɅƚȶDŽɸȶɳdžɹɑƏɩɁɳǷȲƒɭȶ ɅɩȶɴȲƓɌȲɑɩƽƊɅȷɩȥƃɫɊɆƷƀʆ LJɻ Ǎɻ ɴɊɻƙɁɆɪ (ɑɸɀɍɽɌ ɫȶɌǎɋɑɌɭɆ ȴɪɊɪǒȝɑƎ ɅɩȶȹɪɎǒȝɑƎɴȼɍNjɅɁƙɊȪɎƳɌɔɭȲɑɭɪɴɑɅ) ȴɬNjɅȲƙɊɩɁȳƕɑɽƴƚɸȶɳǷȲƒɭȶȲɑɩƽƊɅȷɩȥƃɫɊɆƷƀ ɴȼɍƙɁȪɎLJɅȴɩɁǃƺɊɮɍɳɒɁɭɵɅƳɌɆɸɈɭɍʆ ȲɴɅƚȶɑɸǁȲɴȼɍƙɆɊɮɍɳǷȲƒɭȶɁɸɆɅɽɔɉɩɌȲƞ ɆɻɭɴɅƎNjɅɃɫȲƽȷɽɳƽɋɴɓȲɈɪ ȲɑɩƽƊɅȷɩȥƃɫɊɆƷƀɆƷƟȻǃ ȲƙɊɩɁɵɅLJɻ Ǎɻ ɴɊɻƙɁɅɪɊɯɋʉNJȴɳƙȷˊɅɴȲƓɌ ɅɩȶȲƒɭȶȲƙɊɩɁɑƎȶɽƽɌȷɸɳljɹLJɻǍɻ ɴɊɻƙɁȼɮȷƵƒʆ ɳɋˊȶɑɸɀɮɊɈɌ ɌȲȼɸɳǁɹȯǒɋ ɳȼˊɊƓɪƳɁɽɆɅƏɋɇɍɆɻɹljɍɽɵɅȲɑɩƽƊɅȷɩȥƃɫɊɆƷƀɳǵɳɍˊȴɭɀNJɈɃɫȲ ɅɩȶDŽɊDŽɌɣƘɳɄƛˊƳɌɑɩȲǜȯǒɎƙƺɎ ɆɴɅƏɊɳɃȢɁɔɸɈɪȲƙɊɩɁɵɅɇɍɆɻɹljɍɽɳɍˊƙɆɈʂɅƑɳɔȲɮɓɮɑɭɪʆ Abst ra c t Estuaries provide a critically important habitat for waterbirds and other aquatic species, but in southern Cambodia they are increasingly threatened by aquaculture development. Much of the area around Anlung Pring Protected Landscape in Kampot Province has been converted to shrimp farming and there are concerns that these farms may negatively impact water quality inside the protected area. To investigate this, we collected water samples (in January, March and CITATION: Yav N., Seng K., Nhim S., Chea V., Bou V. & Avent, T. (2017) The impact of shrimp farming on water quality in Anlung Pring, a protected landscape in Cambodia. Cambodian Journal of Natural History, 2017, 49–54. Cambodian Journal of Natural History 2017 (1) 49–54 © Centre for Biodiversity Conservation, Phnom Penh 49 50 Yav N. et al. May, 2016) from five discrete locations inside and outside of the reserve and analyzed these to determine the effects of waste water from nearby shrimp farms on local water quality. Levels of conductivity, turbidity, total dissolved solids, sulphate, ammonium, chemical and biological oxygen demand were higher than recommended levels at many sampling points, especially those within and adjacent to the shrimp farms. Three parameters (coliforms, chemical and biological oxygen demand) were substantially higher within the shrimp farms, suggesting that they were the cause of this pollution. Sampling points that were in the reserve, but hydrologically isolated from shrimp farms, showed levels much closer to, and often within, recommended guidelines for the same parameters. We suggest that solutions to minimize the impact of shrimp farms on water quality, and further research on their ecological impact, are required. Ke yw ords Water quality, shrimp farming, environmental impact. I nt roduc t ion Estuaries are important ecosystems and provide critical habitat for waterbirds and other aquatic species. They are also excellent areas for aquaculture development (Hossain, 2001; Wouter & Patrick, 2003; Primavera, 2006) and land conversion for shrimp farming is a cause for environmental concern, especially in relation to water quality (Flaherty & Karnjanakesorn, 1995; Graslund & Bengtsson, 2001; Hossain, 2001; Jones et al., 2001; Paezosuna et al., 2003; Islam & Tanaka, 2004; Primavera, 2006; Crab et al., 2007) and the subsequent potential for longterm damage to aquatic food webs (Kennish, 1997). Many studies have reported significant environment degradation due to shrimp farming (Paezosuna et al., 2003), including increased organic pollution (Bui et al., 2012; Wilbers et al., 2014), creation of favorable habitat for pathogenic microorganisms (Anh et al., 2010), high nutrient waste (Bui et al., 2012), increased phytoplankton productivity (Islam & Tanaka, 2004), removal of nutrients from culture water (Crab et al., 2007), and decline of aquatic ecosystem health (Naylor et al., 2000; Bui et al., 2012). As a consequence, the Thai government has identified areas where shrimp farming is permitted and areas where it is banned (Flaherty et al., 2000). Many shrimp farms have recently been developed along canals in coastal and inland areas in Cambodia and Vietnam (Anh et al., 2010) and it is likely that these have caused dramatic declines in numbers of sarus cranes (Grus antigone) visiting Hon Chong in southern Vietnam (van Zalinge et al., 2011). Water quality in Cambodia is protected by a sub-degree on pollution which was issued in 1999 (Royal Government of Cambodia, Sub-decree 27, 1999). Anlung Pring Protected Landscape (APPL) is one of three protected non-breeding areas of the globally Endangered eastern sarus crane G. a. sharpii in Cambodia (Yav et al., 2015). Most wetlands to the south of APPL have been converted to shrimp farming and the landscape © Centre for Biodiversity Conservation, Phnom Penh has directly abutted one shrimp farm since 2013 (Yav, 2014). Concerns regarding the farms include the impact of water pollution on the wetland ecosystem of the area, especially as changes to water quality may affect growth of Eleocharis spp., which are the main food for the cranes (Yav, 2014; Yav et al., 2015). Rigorous understanding of the influence of water waste discharged from shrimp farming is consequently required, especially regarding chemical substances that may affect the ecology of the wetland. This study consequently assessed water quality inside and outside of APPL and investigated the effects of shrimp farms on levels of pollutants entering the site. M e t hods Study site APPL covers 217 ha in Kampong Trach District of Kampot Province (10°28’40’’N, 104°31’32’’E) in southern Cambodia, approximately one kilometre from the border with Vietnam within the lower Mekong floodplain (Fig. 1). The site is divided into two parts by a road embankment, and the northern part covers 33 ha and southern section 184 ha. As many as 342 sarus cranes foraged in the site in 2013 (Yav, 2014). The cranes mainly feed upon Eleocharis tubers and preferentially select areas where these occur (Yav et al., 2015). The area also supports the livelihoods of surrounding communities through the provision of fire wood, wild food, tourism and livestock fodder (van Zalinge et al., 2013) The area of APPL is low-lying with an elevation range of 0.0–3.5 m above sea level and is next to a small river that experiences tidal influences, even though the site is approximately 20 km from the Gulf of Thailand (Yav et al., 2015). The area around the southern section of APPL is dominated by shrimp farms, some of which are contiguous with the conservation area (Fig. 1). Waste water is regularly discharged from these shrimp farms into a Cambodian Journal of Natural History 2017 (1) 49–54 Water quality at Anlung Pring canal which is influenced by tidal movements, allowing waste water to flow into the conservation area (Yav et al., 2015). The eastern, northern and western sections of APPL are mainly surrounded by rice fields and settlements. Sample collection Water samples were collected from five discrete areas within APPL during the sarus crane feeding season on 11 January, 11 March and 16 May 2016 (Fig. 1). With the exception of one sampling area in the northern section of the reserve, all other sampling areas were hydrologically linked to the shrimp farms. The five areas comprised: 1) the northern part of the reserve (NPR), which is hydrologically separated from the rest of the reserve by an embankment and receives fresh water from northern uplands during the wet season; 2) canals inside the central section of the reserve (CIR); 3) the southern part of the reserve, which is adjacent to but separated by an embankment from the shrimp farms (SPR); 4) inside the shrimp farms (SF); and finally, 5) canals outside the reserve and adjacent to shrimp farms, which receive water from the farms and tidal flows (COR). Two points at least 200 m apart were sampled in each of the five areas every month, resulting in a total of 30 samples. Samples were collected by filling containers with water taken from a depth of 30–50 cm (approximately the middle of the water column). Different containers were used to collect water for different analyses. Water for coliform analysis was collected in a 250 ml glass bottle, sterilized at 120°C. Water for biochemical oxygen demand tests was collected in a 1,000 ml glass bottle, filled with no air bubbles remaining. Water for total phosphorus, iron and chemical oxygen demand tests were collected in 500 ml polyethylene bottles and preserved with H2SO4. Water for other tests including pH, EC, turbidity, sulphate, ammonium, nitrate as nitrogen and nitrite-nitrogen were collected in 1,000 ml polyethylene bottles with no preservation. All samples were kept in ice boxes at temperatures of 4–6 °C. Sample tests and data analysis Samples were tested by the Department of Hydrology and River Works of the Ministry of Water Resources and Meteorology for pH, conductivity, turbidity, salinity, total dissolved solids (TDS), sulphate (SO4), ammonium (NH4-N), nitrate as nitrogen (NO3-N), nitrite-nitrogen (NO2-N), total phosphorus (TP), iron (Fe), coliforms, chemical oxygen demand (COD), biochemical oxygen demand (BOD) and aluminum (Al). Cambodian Journal of Natural History 2017 (1) 49–54 Fig. 1 Locations of water sampling at Anlung Pring Protected Landscape, Kampot Province. SF = shrimp farm, COR = canal outside reserve, SPR = southern part of reserve, CIR = canals inside reserve, NPR = northern part of reserve. The mean values from the three sampling events at each sample location were calculated. Data normality was tested using one-sample Kolmogorov-Smirnov tests and Shapiro-Wilk tests (Dytham, 2011; Stowell, 2014). A one-way analysis of variance (ANOVA) was used for group comparisons and Tukey HSD tests for pairwise comparisons (Stowell, 2014) if data were normally distributed (p>0.05), whereas Kruskal-Wallis tests and MannWhitney U tests were used if data were not normally distributed (p<0.05). The statistical packages SPSS vers. 20 and R vers. 3.3.2 were employed for this purpose. Re sult s Significant variation was found among the five areas sampled in terms of pH, conductivity, salinity, TDS, NO3-N, coliforms, COD and BOD (all values of p<0.01) (Table 1). No significant differences were found between © Centre for Biodiversity Conservation, Phnom Penh 51 52 Yav N. et al. Table 1 Water quality parameters (Mean ±SE) sampled in and around Anlung Pring Protected Landscape in 2016. Bold font indicates significantly different values, and dashes indicate a parameter was not recorded. SF = shrimp farm, COR = canals outside reserve, SPR = southern part of reserve, CIR = canals inside reserve, NPR = northern part of reserve. Parameter Mean ±SE Advised levels Associated references SF COR SPR CIR NPR pH 8.00 ±0.14 7.02 ±0.09 7.10 ±0.14 6.62 ±0.03 2.30 ±0.77 6–9 Conductivity (mS/cm) 49.85 ±6.78 40.38 ±6.61 40.75 ±3.07 42.97 ±5.08 10.15 ±6.64 <0.01 Turbidity (NTU) 29.63 ±11.15 133.00 ±40.20 60.97 ±21.10 26.40 ±2.86 19.33 ±7.03 <5 Salinity (g/l) 32.98 ±4.95 25.48 ±1.93 26.65 ±2.71 27.80 ±3.72 6.28 ±4.21 No limit TDS (g/l) 23.23 ±5.24 21.16 ±1.56 21.18 ±1.89 22.47 ±2.63 5.18 ±3.38 <0.6 WHO (1996, 2011) SO4 (mg/l) 1,869.28 ±451.45 1,674.00 ±392.69 1,846.20 ±482.44 1,948.87 ±622.77 757.65 ±456.83 100 Chapman (1996) NH4-N (mg/l) 0.19 ±0.07 0.16 ±0.02 0.27 ±0.07 0.46 ±0.12 1.95 ±0.68 <0.1 Bui et al. (2012) NO3-N (mg/l) 0.05 ±0.01 0.07 ±0.02 0.08 ±0.02 0.04 ±0.01 0.01 ±0.00 <0.06 Pulatsu et al. (2004); Bui et al. (2012) NO2-N (mg/l) 0.05 ±0.01 0.06 ±0.01 0.06 ±0.01 0.04 ±0.01 0.02 ±0.01 0.1–0.75 Boyd & Green (2002) TP (mg/l) 0.09 ±0.04 0.15 ±0.04 0.09 ±0.02 0.05 ±0.01 A 0.05 ±0.02 <0.3 Boyd (2003); Bui et al. (2012) Fe (mg/l) 0.63 ±0.21 0.72 ±0.05 0.88 ±0.13 1.04 ±0.12 1.87 ±0.96 0.03 Ramakrishnaiah et al. (2009) Coliforms (MPN/litre) 280.00 ±149.55 146.67 ±23.90 78.33 ±26.26 76.67 ±28.60 31.67 ±15.79 <500 Anh et al. (2010) COD (mg/l) 25.48 ±2.11 14.65 ±1.37 15.27 ±1.28 17.03 ±1.41 7.07 ±4.16 <3 Bui et al. (2012); Ly & Larsen (2012) BOD (mg/l) 20.72 ±4.88 7.18 ±1.16 6.27 ±0.66 7.38 ±0.55 2.33 ±0.86 <6 Boyd & Green (2002); Bui et al. (2012) Aluminum (mg/l) 0.03 ±0.01 0.03 ±0.00 0.06 ±0.03 0.04 ±0.01 - 0.2 Chapman (1996) Anh et al. (2010); Bui et al. (2012) Bartram & Ballance (1996) Chapman (1996); WHO (2011) Boyd & Green (2002) B sampling areas in the remaining parameters (turbidity, SO4, NH4-N, NO2-N, TP, Fe, and Al) (all values of p>0.05). Compared to the northern section of the reserve, levels of conductivity, salinity, TDS, and NO3-N were significantly higher within the shrimp farm and all hydrologically connected areas (all values of p<0.01). Levels of coliforms were also significantly higher within shrimp farms and adjacent canals outside the reserve compared to other hydrologically linked areas (southern section of the reserve and central canals) and the northern section of the reserve (p<0.01). Compared to all other areas, pH © Centre for Biodiversity Conservation, Phnom Penh levels were significantly higher within the shrimp farms (p<0.05) and significantly lower within the northern section of the reserve (p<0.01) (Table 1). The same was true of BOD and COD (all values of p<0.05). Disc ussion Our results show significant variation in water quality between the shrimp farms, hydrologically linked areas and a hydrologically unconnected area in APPL. Because the northern section of Anlung Pring is hydrologiCambodian Journal of Natural History 2017 (1) 49–54 Water quality at Anlung Pring cally isolated from all other areas of the site by a road embankment, water quality in this section is unaffected by discharge from shrimp farms adjacent to the southern section of the reserve. Values for pH were very low in the northern section and fall outside recommended ranges for protecting farming and aquatic ecosystems (Anh et al., 2010; Bui et al., 2012). This may be due to the embankment separating the reserve and allowing chemical fertilizers from rice fields in northerly catchment areas to accumulate in the northern section, creating increased acidification. This is supported by the fact that levels of ammonia nitrogen (NH4-N) were also much higher in the northern section of the site. Ammonia can acidify the environment through the release of H+ ions during the biochemical conversion to nitrate (Schuurkes & Mosello, 1988). Conductivity, TDS, SO4, NH4-N, NO3-N, Fe, COD and BOD were higher than recommended levels for a variety of water uses at most sampling points outside of the hydrologically isolated northern section of Anlung Pring (Lloyd, 1987; Bartram & Ballance, 1996; Chapman, 1996; World Health Organization, 1996, 2011; Boyd & Green, 2002; Ramakrishnaiah et al., 2009; Bui et al., 2012; Ly & Larsen, 2012). In addition, the highest levels of conductivity, TDS, BOD and COD were found in shrimp farms, suggesting that effluent from these may be affecting water quality within the protected landscape. TDS is normally positively correlated with conductivity (Ansari et al., 2015) and indicates the degree of dissolved substances such as metal ions in water (Efe et al., 2005; Ubwa et al., 2013). BOD measures of the amount of oxygen required by bacteria and other microorganisms that stabilize decomposable organic matter (Ubwa et al., 2013). High levels of COD and BOD indicate reductions in water quality (Ubwa et al., 2013; Ansari et al., 2015) by organic compounds (Bui et al., 2012; Ubwa et al., 2013) and reflect high organic and inorganic matter (Boyd & Green, 2002; Bui et al., 2012; Ansari et al., 2015). This has negative impacts on ecosystem health (Anh et al., 2010; Bui et al., 2012) and reduces habitat quality for fish (Nguyen et al., 2006). High levels of both COD and BOD can result from organic waste from abattoirs (Bartram & Ballance, 1996), pellet feed use and discharge from shrimp farming (Anh et al., 2010). Our analyses indicates that many water quality parameters fall outside of a variety of recommended guidelines at APPL and may therefore be negatively affecting the quality of wetland habitats at the site. In Cambodia, Yang & Guo (2003) suggest that water pollution mostly occurs in transnational waters in the Lower Mekong Basin. Poor water quality affects habitat integrity (Simeonov et al., 2003), fish and aquatic producCambodian Journal of Natural History 2017 (1) 49–54 tion (Lloyd, 1987), causes acid sulphate soils and algae blooms (Boyd & Green, 2002; Bui et al., 2012), and creates favourable conditions for pathogenic microorganisms (Anh et al., 2010). Though some aspects of Cambodia’s environment have remained cleaner compared to some neighbouring countries (Monirith et al., 2000), wetland pollution along the coastline threatens habitat integrity and could have a highly adverse impact on biodiversity (Naylor, 1998). We consequently suggest monitoring of water quality and engagement with shrimp farmers to minimize the impact of their practices on protected areas. The development of structures to prevent polluted waters from entering APPL may also prove necessary, and further research on the ecological impacts of water pollution is required to assess the effectiveness of related conservation interventions. 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Cambodian Journal of Natural History 2017 (1) 49–54 Carbon stock of peat soils Ca rbon st ock of pe at soils in m a ngrove fore st in Pe a m K ra sa op Wildlife Sa nc t ua r y, Koh Kong Provinc e , sout hw e st e r n Ca m bodia TAING Porchhay, EANG Phallis, TANN Sotha & CHAKRABORTY Irina* Faculty of Mathematics, Sciences and Engineering, Paññāsāstra University of Cambodia, No. 184, Preah Norodom Blvd (41), Phnom Penh, Cambodia. * Corresponding author. Email cambodiapeatland@gmail.com Paper submitted 10 October 2016, revised manuscript accepted 29 March 2017. ɊɮɍɅʂɋɑɳȶſɆ NjɅƳɌɋɍɽȼɫȶɁɩȷɁɯȷǁɑɽɈɪɁɸɆɅɽȼɪɊɳNjȲ (Peatland) ɳǷȲɊƕɭƺʆ ɁɸɆɅɽȼɪɊɳNjȲɴȼɅȹƙɊȲɑɁƛɵƙɈljɊƙȲɳǒɆ ȲƒɭȶɳȳɁƎɳƳɹȲɭȶƙɁȪɎLJɅɌȲɳȵˊȻȲƒɭȶƹƒɸʒʐʑʔ ɴȼɍƙȴɆȼɀƎɆɽȼɪʔʙʗʖɒɩȷǂ (ȴɩɁDŽɸȶʓʘɒɩȷǂɳǷɳƙǤɴȼɅȹƙɊȲ) ɵɅɵƙɈɳƳȶƳȶɁɸɆɅɽɳȸƒɌʆ ɳƙǤɈɪɊɭȳƷɌƺɃɪȹƙɊȲ ɅɩȶɌȲǜȴɭɀNJɈɃɫȲ ȼɪɊɳNjȲȲʁƺǕȶɑƎɭȲƳɆɮɅȼʁɑɸƴɅɽ ɳɒˊɋ ɳȼˊɌɁɯnjɻȶɑɸƴɅɽȲɭƒȶƳɌɴƙɆƙɆȫɍǕƳɑDžɁɭʆ ƳɌȲɸɀɁɽɈɪɆɌ ɩNjɀƳɆɮɅɑƎɭȲȲƒɭȶȼɪɊɳNjȲɳǷɴȼɅȹƙɊȲɑɁƛɵƙɈljɊ ƙȲɳǒɆLJɅɇƎɍɽƺȷɸɳɀɹȼɫȶȼʁNjɅɁɵɊƚɳȼˊɊƓɪƻɅȼɍɽƳɌɋɍɽȼɫȶɈɪɍɃƑNJɈɇƐɭȲƳɆɮɅȲƒɭȶɁɸɆɅɽȼɪɊɳNjȲȲɊƕɭƺʆ ɳɋˊȶLJɅLJɻ ɅɽƙɆNjɀɈɪɆɌ ɩNjɀƳɆɮɅɑƎɭȲȲƒɭȶɁɸɆɅɽȼɪɊɳNjȲȲƒɭȶɵƙɈɳƳȶƳȶɵɅɴȼɅȹƙɊȲʆ ɑɸǁȲȼɪɊɳNjȲƙɁȪɎLJɅ ƙɆɊɮɍ ɅɩȶɎ ɩNJȴʆ ȷɸɀɭɹƳɆɮɅȲƒɭȶȼɪɊɳNjȲȴɬȷɳdžƚɹʑʙ,ʖ ɳǵ ʒʒ,ʙ% ɅɩȶNjɅȼȶɽɑɭɪɳɁNjȾ ʐ,ʓʔʗƙȲ/ɑɊʓʆ ɴɇơȲɳɍˊ ƳɌɑɩȲǜɳɅɹ ɅɩȶƳɌɑɩȲǜɈɪɊɭɅʉ ȹɳƙǼƺɊɄƘɊɵɅȯɑDŽɆɽȼɪɊɳNjȲȴɬ ʑʑʐɑɊ ɅɩȶNjɅNjȾɑɌɭɆȴɬƙɆɴɒɍ ʕ,ʘʓ x ʑʐʗɊʓʆ ɳɋˊȶLJɻ ɅɽƙɆNjɀǃNjɅƳɆɮɅƙɆɴɒɍʔ,ʔʗ x ʑʐʖɊƙȲ ɴȼɍɑƎɭȲȲƒɭȶȼɪɊɳNjȲɵɅɴȼɅȹƙɊȲɑɁƛɵƙɈljɊƙȲ ɳǒɆʆ Abst ra c t Very little is known about peatlands in Cambodia. The peatland in Peam Krasaop Wildlife Sanctuary (PKWS), Koh Kong Province, was discovered in 2014 and covers 4,976 ha (including 38 ha outside the sanctuary) in a coastal mangrove forest. In addition to their functions as habitats and maintaining water quality, peatlands are significant carbon sinks and therefore play important roles in mitigating climate change. Determining the size of the carbon stock in peat in PKWS is consequently valuable for understanding the sequestration capacity of Cambodian peatlands. We estimated the amount of carbon stock of peat soils in the mangrove forest of the sanctuary. Peat cores were collected and analysed. The carbon content of the peat was between 19.6 and 22.9%, and its bulk density was 0.347 g/cm3. Based on our work and previous studies, the average depth of the peat layer is 110 cm and the total peat volume is about 5.83 × 107 m3. We consequently estimate that approximately 4.47 × 106 Mg of carbon is stored in the peatlands of PKWS. Ke yw ords Bulk density, carbon stock, mangrove, organic carbon content, peat soil, wetlands. CITATION: Taing P., Eang P., Tann S. & Chakraborty, I. (2017) Carbon stock of peat soils in mangrove forest in Peam Krasaop Wildlife Sanctuary, Koh Kong Province, southwestern Cambodia. Cambodian Journal of Natural History, 2017, 55–62. Cambodian Journal of Natural History 2017 (1) 55–62 © Centre for Biodiversity Conservation, Phnom Penh 55 56 Taing P. et al. I nt roduc t ion Wetland environments are abundant in Cambodia, covering 30% of the country (Kol, 2003; Mak, 2015). Peatlands, a type of wetland ecosystem characterized by accumulated organic matter (Parish et al., 2008), have not been well studied in Cambodia and mangrove peatlands in particular have been neglected (Donato et al., 2011). Peat is a soil type dominated by decomposing plant materials, and contains more than 18% and 30% of organic carbon and organic matter respectively (Agus et al., 2011). Peat is formed from decomposing plant materials under saturated conditions (Parish et al., 2008), as dead vegetation layers on top of the soil. Benefits provided by peatlands include climate regulation—in the form of carbon sequestration—and other ecosystem services (Parish et al., 2008). The latter include the roles peatlands play in the hydrological cycle by removing nutrients and sequestering large volumes of water and provision of habitat to diverse animal and plant species. Peatlands cover around 3% (400 x 106 ha) of the global land area and occur in most areas of the world (Strack, 2008). Most are found in the boreal and temperate zones (3.57 x 108 ha: Page et al., 2010), but tropical and subtropical zones are also important peatland regions, because of the high rates of plant production and high rainfall which reduces rates of decay (Parish et al., 2008). Peatlands cover around 2.5 x 107 ha in Southeast Asia, almost 60% of peatlands within the Tropics. More than 70% of these occur in Indonesia. Malaysia, Brunei, and Thailand also have significant peatland areas, while smaller areas are found in Vietnam, the Philippines, Cambodia, Laos, Myanmar, and Singapore (ASEAN Secretariat, 2014). The world’s peatlands contain a carbon pool of about 550 Gt, which is twice that of above-ground forest biomass (Parish et al., 2008). As a consequence, these play a major role in regulating climate through carbon dioxide storage, but also as a source of methane, another greenhouse gas (GHG). Loss of carbon storage caused by peatland fires or inappropriate management practices can lead to GHG emissions which contribute to climate change. When peat is exposed to oxygen, it oxidizes and releases carbon dioxide into the atmosphere. Climate change also affects the GHG cycle of peatlands by transforming their carbon sinks into sources of carbon emissions due to changes in temperature and rainfall, whereas their carbon content remains constant if they are protected and water levels remain unchanged. For instance, climate change is currently predicted to severely degrade 60% of Canadian peatlands and further contribute to global warming by releasing carbon dioxide and methane into the atmosphere (Tarnocai, 2006). © Centre for Biodiversity Conservation, Phnom Penh Degradation of peatlands is commonplace despite their many documented benefits, with human activities having a significant impact. A major cause of peatland degradation is their conversion to agricultural land by draining or burning, with over 12% (3 x 106 ha) of peatlands having being converted in Southeast Asia (ASEAN Secretariat & GEC, 2011). In the absence of disturbance, peatlands continuously accumulate carbon by storing slowly-decaying plant materials in the anaerobic peat layer. Because carbon sequestration in peatlands plays an important role in climate regulation, restoration of peatland is among the most cost-effective ways to mitigate climate change (Bain et al., 2011). The objective of our study was to estimate the amount of carbon stored in the coastal mangrove peatland in Peam Krasaop Wildlife Sanctuary, Koh Kong Province, southwestern Cambodia. M e t hods Study site Peam Krasaop Wildlife Sanctuary (PKWS) is a protected area established by Royal Decree in 1993. It includes 23,750 ha of coastal mangrove in Koh Kong Province (Fig. 1), although the sanctuary area is larger (25,897 ha) according to an official map approved in 2003 (An et al., 2009). Part of PKWS lies within the boundary of the Koh Kapik and Associated Islets Ramsar Site, which was designated as a result of supporting a significant mangrove ecosystem (criteria 1), endangered and rare species (criteria 2), and providing a site for feeding, breeding, and nursery grounds for fish and shellfish species (criteria 8) (Srey, 2012). The main tree species at the site are Lumnitzera racemosa, Excoecaria agallocha, Rhizophora apiculata, R. mucronata, Brugueira gymnorrhiza, Melaleuca cajuputi, Heritiera littoralis, Xylocarpus granatum, L. littorea, Ceriops tagal, Avicennia alba, Scyphiphora hydrophyllacea, Glochidion littorale, Phoenix paludosa, Nypa fruticans, Acrostichum speciosum, and Pandanus sp. (Lo et al., 2014). In 2014, as part of the ASEAN SEApeat project, activities were undertaken to assess peatlands in Cambodia. Based on satellite images, it was determined that PKWS had a high likelihood of containing peat (Lo et al., 2014). Following interpretation of satellite imagery, on-site assessments were conducted to verify the presence of peat. Our study was conducted in 2016 within the mangrove peatland in PKWS. Prior to fieldwork, peat depth measurements from 22 peat cores and a map of the peatland were obtained from the SEApeat project. The mangrove peat layers ranged from 44–200 cm, with an Cambodian Journal of Natural History 2017 (1) 55–62 Carbon stock of peat soils Fig. 1 Location of Peam Krasaop Wildlife Sanctuary and study samples. average depth of 115 cm, and a total of 4,976 ha of the area were estimated to be peatland (Lo et al., 2014). Sampling methods To select sampling locations, maps were created by transferring polygon outlines of areas of peatland identified in PKWS by Lo et al. (2014) into ArcGIS software and overlaying these with images from GoogleEarth. Because the characteristics of peat across PKWS were expected to be similar to Lo et al. (2014), it was assumed 14 peat cores would be sufficient to estimate its carbon stock. These were collected in PKWS on 23–24 January 2016. A soil auger (made by Eijkelkamp, the Netherlands) was used to collect the 14 core samples. The volume of the soil auger (half-cylinder) was 1,410 cm3 (height = 100 cm, radius = 3 cm). The depth of the peat layer was measured in each core. Each core was divided into 25 cm vertical sections. From each vertical core section, 5 cm samples were cut to produce 46 samples, these representing the length of each core at 25 cm intervals. Each of the 46 samples was analysed for bulk density and organic carbon. An additional 14 samples of the top layer of peat (1–25 cm) were collected using a soil ring (height = 4 cm, radius = 2.1 cm, volume = 55.4 cm3) constructed by the Cambodian Journal of Natural History 2017 (1) 55–62 authors from a stainless steel pipe, with a thickness of 1 mm. A GPS was used to record the locations of samples. Samples were wrapped in aluminium foil, placed in individual plastic bags labelled with their respective locations, and packed into an ice box for transport. These were stored at 4°C for three days prior to analyses for bulk density and organic carbon content. Bulk density The volume of the soil sample was calculated from the diameter of the auger and soil ring. Peat sample bulk density (BD) was determined using the gravimetric method (Agus et al., 2011). To determine dry mass, samples were dried at 105°C for six hours and weighed, and this process was repeated up to four times until constant mass was achieved. BD was defined as the dry weight of soil per unit volume. This was calculated as BD = Ms/Vt, where BD was bulk density (g/cm3), Ms was the mass of the dry peat soil (g), and Vt was the volume of the soil sample (cm3). Peat depth and area The depth of the peat layer was defined as the length of peat cores obtained in the field. The area of peatland in PKWS was digitized from the SEApeat project map into © Centre for Biodiversity Conservation, Phnom Penh 57 58 Taing P. et al. 71 polygons using ArcGIS, which was used to calculate the area of each polygon. The locations for our peat depth data were also included in ArcGIS, and the depth of each polygon was estimated as the product of its area and respective peat depth. Because of the limited number of samples, peat depths were assigned to each polygon as follows: 1) If a single core sample had been taken in the polygon, the value for that core was used; 2) If more than one core sample was taken, the mean of these was used; and 3) If no core sample was available for the area, the value of the nearest similar area with a known depth was used. Similarity was determined subjectively, based on vegetation cover and contiguousness. Organic carbon content Total organic carbon (TOC) was measured by loss on ignition at 550°C (Agus et al., 2011). Two grams of each sample were dried at 105°C for 15 minutes, then weighed. To obtain a dry mass value (Mdry), this process was repeated until mass did not change between drying cycles. The dry samples were then transferred into a combustion oven at 550°C for four hours (Santisteban et al., 2004; the method was modified according to Agus et al., 2011, reducing the combustion time from six hours). The samples were cooled in a desiccator and their mass recorded as ash mass (Mash). TOC was calculated as TOC = (Mdry – Mash/ Mdry) / 1.724, where TOC was total organic carbon (g/g), Mash was ash mass (g), Mdry was dry mass (g) and 1.724 was the conversion factor for organic matter to organic carbon (Agus et al., 2011). Carbon stock The total carbon stock of the PKWS peatland was calculated as Cstock = A x D x BD x TOC, where Cstock was carbon stock (Mg), D was average peat depth (m), BD was average bulk density (Mg/m3), TOC was organic carbon content (Mg/Mg), and A was the peatland area (m2) (Weissert et al., 2013). auger (see Discussion). There was no significant difference between bulk densities and depth (p=0.309) and there was no consistent trend in BD variation with depth, some cores having increased BD at greater depth, some decreased BD, and others similar BD (Fig. 2). Peat depth and volume Combining data from Lo et al. (2014) with our study, the average depth of the peat layer was estimated as 1.10 m (n=35, SD=0.41) (Fig. 3). The total area of the peatland in PKWS was estimated at 4,938 ha (exluding 38 ha of peatland outside the sanctuary). Using these data, the total volume of peatland within PKWS is estimated to be approximately 5.83 × 107 m3 (Table 2). Organic carbon content Organic matter values for the peat soils in PKWS ranged from 33.8–40.2%. The organic carbon content of peat by depth is shown in Table 2 and averaged 22.2% overall. There were no significant difference in organic carbon content with depth (p>0.05). Carbon stock The carbon stock of peatland in PKWS was estimated to be 4.47 x 106 Mg (Table 1). As BD and carbon content did not vary significantly with depth, we conclude that there is no significant difference in carbon stock with depth. Disc ussion Bulk density The bulk density (BD) of peat in PKWS appears to be typical of mangrove systems. Previous reviews indicate that the BD of mangrove peat in Indo-Pacific oceanic and Table 1 Bulk density values for core sections from soil Re sult s ring samples (surface samples only) and auger samples (25 cm subsections of each core). Bulk density Bulk density values for core sections obtained from soil ring samples were on average 26% lower than auger samples taken at the same depth, at 0.347 (n=14, SD=0.11) and 0.436 (n=14, SD=0.14) g/cm3 respectively (Table 1). However, this difference was only significant at p=0.037 due to high variance within the data. The value obtained from the soil ring is used in subsequent calculations because this sampling method is less disruptive than the © Centre for Biodiversity Conservation, Phnom Penh Sampling depth n Mean BD (g/cm3) Soil ring 14 0.347 0.118 1 to 25 cm 14 0.436 0.149 25 to 50 cm 14 0.398 0.086 50 to 75 cm 9 0.410 0.121 SD 75 to 100 cm 6 0.446 0.085 >100 cm 3 0.338 0.045 Cambodian Journal of Natural History 2017 (1) 55–62 Carbon stock of peat soils Fig. 2 Normalized bulk density of core sections from Peam Krasaop Wildlife Sanctuary. Each line represents one core. Fig. 3 Frequency of peat layer depths (n=36) at 15 cm intervals at Peam Krasaop Wildlife Sanctuary. Table 2 Carbon stock in different peat layers at Peam Krasaop Wildlife Sanctuary. Depth (cm) n Area (m2) Volume (m3) Bulk density (Mg/m3) Organic matter (Mg/Mg) Carbon content (Mg/Mg) Carbon stock (Mg) 1 to 25 14 4.94 x 107 1.23 x 107 0.347 0.387 0.224 9.62 x 105 25 to 50 14 7 4.94 x 10 1.22 x 10 7 0.347 0.402 0.233 9.89 x 105 50 to 75 9 4.46 x 107 1.11 x 107 0.347 0.372 0.216 8.32 x 105 75 to 100 6 7 4.42 x 10 1.04 x 10 7 0.347 0.338 0.196 7.07 x 105 >100 cm 3 4.03 x 107 1.23 x 107 0.347 0.394 0.229 9.76 x 105 Total 5.84 x 107 Cambodian Journal of Natural History 2017 (1) 55–62 4.47 x 106 © Centre for Biodiversity Conservation, Phnom Penh 59 60 Taing P. et al. estuarine systems ranges from 0.35 to 0.55 g/cm3 (Donato et al., 2011). The BD of peat in U Minh Ha National Park (Vietnam) ranges from 0.19 to 0.26 g/cm3 with an average of 0.23 g/cm3 (Quoi, 2010), compared to 0.347 g/cm3 in PKWS. According to Andriesse (1988), the BD of peat soil ranges from 0.05 g/cm3 in very fibric (i.e. containing undecomposed plant fibres) soils to around 0.5 g/cm3 in well-decomposed materials. The high BD of mangrove peat in PKWS thus suggests it mainly comprises welldecomposed material, with only some areas including fibric peat. Based on our observations, the presence of plant roots within the well-decomposed peat of some of our samples came from live mangrove trees growing in the area (Fig. 4). The BD of samples taken with our soil ring were more accurate than samples taken using the auger, which tends to distort soil cores. Agus et al. (2011) recommend using a soil ring to sample peat soil for BD analysis. It was not possible to use a soil ring for deeper layers of peat in PKWS due to the presence of mangrove root structures and overlaying water. Peat cores taken to estimate the depth of the peat layer using the auger were consequently less than optimal for the purposes of calculating BD. The mean BD of auger samples in the 1–25 cm peat layer was 0.436 g/cm3, 26% higher than the value of samples obtained with the soil ring. We suspect that distortion occurred because the auger compressed the soil cores when rotated to obtain samples. Tides can affect accumulation rates of soil organic carbon due to regular water movement disturbing and washing away decomposing material, and depositing mineral sediments. Water movement and mixing may also increase oxygenation of organic material and reduce accumulation rates by increasing aerobic mineralization (Alongi, 2009). Carbon stock The carbon stock of peatlands in PKWS is 904 Mg/ha, which is typical of estuarine (1,074 Mg/ha) and marine (990 Mg/ha) mangroves (Donato et al., 2011). In contrast, because peat thickness varies at U Minh Ha, this affects the amount of carbon stored per area. For instance, where peat layers reach a depth of 70 cm, carbon content is about 814 Mg/ha and where these reach 120 cm, carbon content is about 1,480 Mg/ha (Quoi, 2010). Carbon storage in peat Peat depth Peatlands sequester more carbon per area than terrestrial ecosystems (Parish et al., 2008). According to Toriyama et al. (2011), soil carbon stocks range from 56.9–108 Mg/ha in evergreen forest soils and 34.9–53.2 Mg/ha in deciduous forest soils in Cambodia. In addition, the carbon stock of forest soils in the Mondulkiri and Kompong Thom provinces was at most 114 Mg/ha (Toriyama et al., 2012). As mangrove peatlands store almost an order of magnitude more carbon per area (904 Mg/ha in this study, Fig. 5), this strengthens the case for prioritizing conservation of mangrove peatlands. Peat layer depths in tropical areas range from 0.5 m to >10 m (ASEAN Secretariat, 2014), although estuarine peat is typically 3 m thick (Donato et al., 2011). As such, the peat layer in PKWS is relatively thin with an average of 1.1 m, which is similar to values in oceanic systems (Donato et al., 2011). This may be due to the young age of mangrove forests in situ and their close proximity to open water, which may hinder formation of peat layers due to the disruptive nature of tides. The total carbon stock of peat soils in PKWS is 4.47 x 106 Mg, approximately 0.15 % of the total carbon stored in Cambodia’s terrestrial ecosystems (2.97 x 109 Mg) and about 0.007 % of the total carbon storage in Southeast Asian peatlands (58 Gt: Strack, 2008). Given the organic carbon to carbon dioxide emission factor of 3.67 (Agus et al., 2011), the PKWS peatlands could release 1.64 x 107 Mg of carbon dioxide emissions if burnt or otherwise destroyed. Organic carbon content Soil carbon estimation The organic carbon content of peat ranges from 18–58% when measured using the ‘loss on ignition’ (LOI) method (Agus et al., 2011), although Donato et al. (2011) obtained values of 7.9% and 14.6% for estuarine and oceanic systems respectively. Our values of 33.8–40.2% for PKWS are somewhat lower than those for U Minh Ha National Park in Vietnam (53.4–54.0%: Quoi, 2010). The higher values at the latter site may be due to its greater distance from the shoreline, and therefore reduced tidal influences compared to PKWS which is located in an estuary. Carbon density can be calculated from soil bulk density as a low cost option for estimating carbon stocks in tropical peat (Warren et al., 2012). To test the applicability of the regression equation developed by Warren et al. (2012) to PKWS, we calculated the theoretical carbon density and compared it with the value obtained from the LOI method. The former gave a value more than twice the latter, i.e. it over-estimated the amount of carbon in the peat by a factor of 2.2. However, Warren et al. (2012) recommend that the equation be used only for soils with © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 55–62 Carbon stock of peat soils climate change mitigation strategies and provide a basis for improving estimates of the potential of peatlands as carbon sinks in Southeast Asia. Fig. 4 A peat core showing living roots from Peam Krasaop Wildlife Sanctuary. Further work is required to characterize the peatlands of PKWS in detail. In particular, peat depths at the site should be validated because of the limited number of samples in our study. Further assessment of carbon for carbon credit programmes is also warranted to generate funds for conservation of PKWS through initiatives such as the Reduced Emissions from Deforestation and Degradation scheme. Other newly discovered peatlands in Cambodia, such as the Botum Sakor peatland, Koh Kong Province (Quoi et al., 2015) should also be characterized to understand their function, identify possible threats, and develop effective management practices. Ack now le dge m e nt s Fig. 5 Comparison of carbon stocks in Peam Krasaop Wildlife Sanctuary and forest soils (latter data taken from Toriyama et al., 2011; 2012). >40% carbon content. As this is higher than the organic content of peatlands in PKWS (33.8–40.2%), the linear relationship between BD and carbon density may not hold true due to physical properties of the soil affecting carbon content in low organic peat (Warren et al., 2012). In fact, BD and carbon density were negatively correlated in our samples. Conclusions The carbon stock of peat soils has received little attention in Cambodia to date. Our study adds to knowledge of tropical peatlands, highlights their importance, and can contribute to improving awareness of the value of peatlands with implications for their management and conservation. Sand mining, drainage, and deforestation in peatlands is likely to impact these ecosystems and release their sequestered carbon into the atmosphere. Our study can also inform further research in PKWS. In highlighting the importance of peatland and mangrove preservation, our results can support national Cambodian Journal of Natural History 2017 (1) 55–62 Dr Srey SunLeang, Dean of Faculty of Mathematics, Science and Engineering, Pannasastra University of Cambodia provided vital support for this study. We are grateful to Mangroves for the Future of IUCN for financial support and thank Dr Le Phat Quoi and Ms Julia Lo for their invaluable advice. We also thank Dr Top Neth for methodological assistance and Dr Uy Davin for his analytical support, including use of facilities. 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Cambodian Journal of Natural History 2017 (1) 55–62 Large mammals at Chhep Wildlife Sanctuary Ca m e ra t ra pping of la rge m a m m a ls in Chhe p Wildlife Sa nc t ua r y, nor t he r n Ca m bodia SUZUKI Ai1,*, THONG Sokha2, TAN Setha2 & IWATA Akihisa1 1 Ecology and Environment, Division of Southeast Asia, Graduate School of Asian and African Area Studies, Kyoto University, Yoshida-Honmachi, Sakyo-ku, Kyoto 606-8501, Japan. 2 Wildlife Conservation Society, No.21, Street 21, Tonle Bassac, Phnom Penh, 12000, Cambodia. * Corresponding author. Email suzuki.ai.47a@kyoto-u.jp Paper submitted 20 April 2017, revised manuscript accepted 21 June 2017. ɊɮɍɅʂɋɑɳȶſɆ ɴȼɅȹƙɊȲɑɁƛɵƙɈɴȸɆɑƏɩɁɳǷNJȴƴȶɳȹˊȶɵɅƙɆɳɃɑȲɊƕɭƺ ɑɊƓɮɌɳƽɋɵƙɈɌɳLJɹ (DDF) njɻ ȶɄɸɑɳɊƓˊɊʆ ƳɌɑɳȶžɁɳƽɋ Njɻ ɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩƙɁȪɎLJɅɳɄƛˊɳɓˊȶɳǷȲƒɭȶɴȼɅȹƙɊȲ ȲƒɭȶɌɋɺɳɈɍɈɪɌɌȼɮɎƙLJɸȶɆɅƎɆdžƐɆɽƵƒȴɬ ƹƒɸʒʐʑʒ ȼɍɽ ʒʐʑʓ Ʌɩȶ ʒʐʑʓ ȼɍɽ ʒʐʑʔʆ ɍɃƑɇɍLJɅɈɪƳɌƽȲɽNjɻɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩ ʗʔʘʓɋɆɽ ɃɃɯɍLJɅȲɸɀɁɽƙǂ ʓʗʖʗ ɵɅɂɅɩȲɑɁƛɄɸ ʓʐƙɆɳɉɃʆ ɍɃƑɇɍɌɆɑɽɳɋˊȶɆȦ ƅ ȲɽɈɪNJɈɆɅƎNjɅɵɅɂɅɩȲɑɁƛɄɸȲɭƒȶɵƙɈɌɳLJɹNjɅȼɮȷƺ ɌNjɸȶ(Rucervusȱ (Bosȱ eldii) ɃɳɅǜȶ javanicus) ɅɩȶƹƗɵƙɈ(Felisȱ chaus)ʆ ɑɸƴɅɽƺȶɳɅɹɳǵɳɃȢɁȴɬɎɁƎNjɅɑɸɳljȷɄɸ(Viverraȱ megaspila) ƺƙɆɳɉɃɌȶ ɳƙƵɹƺɑȲɍ ɴɁɌɮɆɂɁɌɆɑɽǏƙɁȪɎɳȵˊȻɳƙȷˊɅɍɸƽɆɽɃɪʔ ȲƒɭȶȷɸɳǁɊƙɆɳɉɃDŽɸȶɔɑɽɳǷȲƒɭȶɴȼɅȹƙɊȲʆ ɳɅɹɆƷƟȻɈɪǒɌɺ ɑɸƴɅɽƺɑȲɍɵɅɴȼɅȹƙɊȲɑɁƛɵƙɈɴȸɆ ɑƙNjɆɽƳɌɔɉɩɌȲƞƙȲȩɊɂɅɩȲɑɁƛȲɭƒȶɴȼɅɳɃɑNJɈɑɊƓɮɌɵƙɈɌɳLJɹɃɸdžɆ ɴɁƺƳɌɈɩɁ ɵƙɈɌɳLJɹɅɩȶɵƙɈɃɸdžɆɊɩɅƙɁȪɎƽȲɽƺɁɸɆɅɽƳɌljɌƙȴɆɽƙƵɅɽɳǷɳɓˊɋɳǷǕɑɭɪǕɳȴƒɋɿȼɪɳƵȲʆ Abst ra c t Chhep Wildlife Sanctuary in northern Cambodia comprises a large tract of deciduous dipterocarp forest (DDF). A camera trap survey was conducted in the wildlife sanctuary during two successive dry seasons, 2012–2013 and 2013– 2014. A total of 7,483 camera-trap-nights yielded 3,787 records of 30 large mammal species. Our results confirm the continued occurrence of DDF-associated large mammals such as Eld’s deer Rucervus eldii, banteng Bos javanicus, and jungle cat Felis chaus. Importantly, large-spotted civet Viverra megaspila, a globally Endangered species, was the fourthmost commonly photographed species in the wildlife sanctuary. This highlights the global significance of Chhep Wildlife Sanctuary for conservation of mammal assemblages in a lowland DDF-dominated landscape, given that DDF and lowland forests are under-represented by protected areas in mainland Southeast Asia. Ke yw ords Carnivores, deciduous dipterocarp forest, large-spotted civet, Preah Vihear Protected Forest, semi-evergreen forest, Viverra megaspila. CITATION: Suzuki A., Thong S., Tan S. & Iwata A. (2017) Camera trapping of large mammals in Chhep Wildlife Sanctuary, northern Cambodia. Cambodian Journal of Natural History, 2017, 63–75. Cambodian Journal of Natural History 2017 (1) 63–75 © Centre for Biodiversity Conservation, Phnom Penh 63 64 Suzuki A. et al. I nt roduc t ion Large areas of seasonally dry forest have disappeared across continental Southeast Asia and <10% of remaining deciduous dipterocarp forest (hereafter DDF) is in protected areas (McShea et al., 2005). In Thailand, DDF is under-represented in protected areas (Tantipisanuh & Gale 2013) and experienced 4.44% of annual tree cover losses between 2000 and 2012 (Johnson, 2015). Similarly, in Myanmar, Laos, and Vietnam, DDF has received limited protection (Wohlfart et al., 2014). In Cambodia, the large areas of DDF in the north and east of the country have received relatively better protection (Wohlfart et al., 2014). Chhep Wildlife Sanctuary, formerly Preah Vihear Protected Forest, is a protected area containing part of the largest contiguous tract of DDF in the northern plains of Cambodia. The wildlife sanctuary supports globally threatened species associated with DDF, such as Eld’s deer Rucervus eldii (McShea et al., 2005; Owen, 2009; McShea & Baker, 2011) and giant ibis Thaumatibis gigantea (BirdLife International, 2016). Despite its importance, ecological information on large mammals is limited in the landscape. This study aims to document records of large ground-dwelling mammals in Chhep Wildlife Sanctuary from intensive camera trapping surveys. M e t hods Fig. 1 Vegetation within Chhep Wildlife Sanctuary (Forestry Administration, 2010), and camera trap stations in the 2012– 2013 (upper) and 2013–2014 (lower) dry seasons. Upper right: the location of Chhep Wildlife Sanctuary (black area) and Preah Roka Wildlife Sanctuary (grey area) in Cambodia. Study area Chhep Wildlife Sanctuary is located in northern Preah Vihear Province, and borders Thailand and Laos (Fig. 1). The southwestern part of the wildlife sanctuary is contiguous with Preah Roka Wildlife Sanctuary which forms a corridor between Chhep Wildlife Sanctuary and Kulen Promtep Wildlife Sanctuary. Chhep Wildlife Sanctuary covers 1,900.27 km2 and comprises three main forest types: DDF (66.9%), evergreen forest (18.8%), and semievergreen forest (9.6%) (Forestry Administration, 2010) (Fig. 1). The wildlife sanctuary also contains smaller areas of seasonally-flooded grasslands, bamboo forests, seasonally-flooded riparian habitats and a network of temporary and permanent forest pools and streams. Data collection Our survey was conducted over two successive dry seasons from 2012 to 2014. Surveys during the wet season were not possible due to logistical and financial constraints. The purpose of the survey in the 2012–2013 dry season was to investigate the presence of carnivores as a part of feasibility study to determine future research © Centre for Biodiversity Conservation, Phnom Penh targets among four carnivores: large-spotted civet Viverra megaspila, large Indian civet Viverra zibetha, leopard cat Prionailurus bengalensis, and jungle cat Felis chaus. Passive infrared digital cameras were mainly set on animal trails, footpaths, motorbike tracks and waterways in semievergreen forest, evergreen forest, and DDF, under the assumption this would maximize detection of these species. At other sites, the two Viverra species were frequently recorded from cameras set in such areas (Gray et al., 2010) and leopard cats showed higher detectability along roads than off-trail locations (Sollomann et al., 2013). Compared with these species, records of jungle cat in Indochina are very limited, but scats have mostly been found on roads, trails, and dry river beds in India (Majumder et al., 2011), suggesting the species uses trails. Due to camera malfunctions, sampling efforts in DDF were limited and a total of 49 camera stations were set in the 2012–2013 dry season. Cameras were mounted approximately 30–50 cm above ground on trees at least 1 km apart and set to operate for 24 hours each day. No stations were baited. Cambodian Journal of Natural History 2017 (1) 63–75 Large mammals at Chhep Wildlife Sanctuary In the 2013–2014 dry season, the survey purpose was to understand occupancy patterns of civets of the subfamily Viverrinae. A total of 53 camera stations were set and as six of these were <50 m from stations set in the previous season, they were considered the same stations in analysis. Camera placement was similar to the 2012–2013 dry season, except that more sampling effort was undertaken in DDF. Both survey periods employed trail-based sampling, which biases measurements of relative abundance (Sollmann et al., 2013; Wearn et al., 2013) and probably fails to detect large mammals that use offtrail areas (Blake & Mosquera, 2014). In terms of species detected, however, trail-based sampling is less likely to differ from random sampling where sampling efforts exceed 1,400 camera-trap-nights, especially during the dry season (Cusack et al., 2015). Recognising the limitations of our non-random sampling approach, we consequently documented where species were detected and did not consider their relative abundances or compare levels of species richness with other sites. Species identification Species identifications employed the nomenclature of Wilson & Mittermeier (2009) for carnivores and Francis & Barrett (2008) for other species, and incorporated the taxonomic revisions of IUCN (2017). Because the survey targeted large non-volant mammals, small mammals such as treeshrews and most rodents were excluded (although porcupines were included). Species identifications for some mammals were unclear. For instance, though the presence of large-toothed ferret badger Melogale personata has been documented in Cambodia, it cannot be assumed that all ferret badgers recorded in the country represent this species (Schank et al., 2009). Second, Meijaard & Groves (2004) suggest chevrotains (Tragulus) in Cambodia could be T. kanchil affinis, but Gray et al. (2012) cautiously identified Tragulus only to genus level in the eastern plains, which is bordered by Vietnam and close to where the silver-backed chevrotain T. versicolor occurs (Meijaard & Groves, 2004). In this study, all photographs of Tragulus were assigned to T. kanchil. Third, hybrid individuals of crab-eating macaque Macaca fascicularis and rhesus macaque M. mulatta have been reported in northeastern Cambodia (Heng et al., 2010). The best feature for identifying these species is relative tail length, this being >90% for M. fascicularis, <60% for M. mulatta, and between these figures for hybrid individuals (Heng et al., 2010). Because relative tail length could not be determined confidently from photographs, however, all macaques with characters similar to M. fascicularis were assigned to this species. Cambodian Journal of Natural History 2017 (1) 63–75 Data analysis Because Yasuda (2004) found a large number of camera trap photographs of the same species occurred less than one minute after the first photograph and reached a plateau after 30 minutes, we defined a camera trap record as an independent record if it occurred at least 30 minutes after a photograph of the same species at a given station. Total sampling effort is expressed as the total number of camera-trap-nights, one camera-trap-night being defined as a continuous 24 hr period of normal camera operation. Records of large mammals in the two dry seasons are expressed as the proportion of camera stations where species were detected at least once (naïve occupancy), and encounter rates were calculated as the number of records / 1,000 camera-trap-nights. Detection of large mammals within each forest type was calculated as a percentage, namely the number of stations where a species was detected in the forest type divided by the total number of stations which detected the species. Forest types at each camera trap station were determined by calculating the proportion of dominant forest types within a 500 m buffer area around each camera station using ArcGIS. Classification of forest types followed Forestry Administration (2011). Although this classification has limited accuracy, observations of forest types at camera trap stations suggested it is suitable for indicative purposes. Of the 53 camera trap stations set during the 2013–2014 season, 42 stations matched DDF, semi-evergreen forest or evergreen forest. However, difficulties were experienced in distinguishing between evergreen and semi-evergreen forest because 21 of 23 stations identified as being located in evergreen forest by FA (2011) were observed to be semi-evergreen forest in the field. We therefore adopt “semi-evergreen and evergreen forest” (hereafter S/EGF) as a combined forest category. When a dominant forest type comprised >70% of a buffer zone, the station was defined as either deciduous dipterocarp forest (DDF) or semi-evergreen and evergreen forest (S/EGF). When both forest types occurred in the 500 m buffer zone, the location was defined as a mosaic of DDF and S/EGF. Another limitation of the FA classification was that nine stations in small areas of semi-evergreen forest were broadly identified as DDF. Buffer analysis helped to determine that two of these were not in DDF but in a mosaic of DDF and S/ EGF; however, seven camera stations remained in error. Activity patterns for four daily periods (dawn, day, dusk, and night) were examined for species with >20 records. Local sunset and sunrise times during the survey were obtained from the US Department of Commerce, National Oceanic and Atmospheric Administration (http://www.esri.noaa.gov/). During the survey period, © Centre for Biodiversity Conservation, Phnom Penh 65 66 Suzuki A. et al. sunrise varied from 05:55 to 06:30 hrs and sunset from 17:29 to 18:11 hrs. Dawn was defined as the period from 05:00 to 07:00 hrs and dusk from 17:00 to 19:00 hrs. Re sult s Thirty large mammal species were detected over the course of the two dry season surveys (Table 1). Asiatic black bears Ursus thibetanus, Sunda pangolins Manis javanica, and Indochinese silvered langurs Trachypithecus germaini were detected only in the 2012–2013 dry season while Eld’s deer were recorded only in the 2013–2014 dry season (Fig. 2). Survey effort during the 2012–2013 dry season was 2,370 camera-trap-nights and produced 1,198 records of confidently identified species (Table 1). Twentynine species representing seven orders were recorded, including, per IUCN (2017), one Critically Endangered species (Sunda pangolin Manis javanica) and six Endangered species including Asian elephant Elephas maximus (Fig. 2). The three most commonly photographed species were common palm civet Paradoxurus hermaphroditus, Eurasian wild pig Sus scrofa, and red muntjac Muntiacus muntjak. Survey effort during the 2013–2014 dry season was 5,113 camera-trap-nights and produced 2,589 records of 27 confidently identified species (Table 1). The three most commonly photographed species were golden jackal Canis aureus, Eurasian wild pig and common palm civet. Species occurrence and activity patterns Of the 21 mammal species detected at more than five camera trap stations, five (leopard Panthera pardus, lesser Oriental chevrotain Tragulus kanchil, gaur Bos gaurus, northern pig-tailed macaque Macaca leonina, and Asian elephant) were almost exclusively detected in S/EGF (Table 2). Large Indian civets were detected at 31 stations, only one of which was in DDF. Conversely, jungle cats were not detected in S/EGF, but only in DDF or a mosaic of DDF and S/EGF. Activity patterns of species with >20 records are shown in Fig. 3. Eleven species (comprising seven carnivores, two ungulates, Burmese hare Lepus peguensis and Malayan porcupine Hystrix brachyura) exhibited nocturnal patterns of activity, whereas five (two carnivores, one ungulate and two primates) exhibited diurnal patterns and lesser Oriental chevrotain showed crepuscular activity. © Centre for Biodiversity Conservation, Phnom Penh Disc ussion Our results provide preliminary information on large mammal communities during the dry season in Chhep Wildlife Sanctuary. Over two successive dry seasons, 30 large mammal species were detected including one Critically Endangered species and six Endangered taxa per IUCN (2017). This confirms the conservation importance of the sanctuary and our records, particularly those of two DDF-associated Endangered species—banteng Bos javanicus and Eld’s deer—highlight its significance in light of the under-representation of DDF in protected areas in Indochina. Tordoff et al. (2005) emphasized the importance of semi-evergreen forest in DDF landscapes in Indochina. Our findings corroborate this: 27 of the 30 species we detected were recorded by at least one station in S/EGF. Though we are unable to infer species habitat preferences due to sampling bias, these clearly use S/EGF despite its relatively small extent, at least in the dry season. In particular, leopards, lesser Oriental chevrotains, northern pig-tailed macaques, and Asian elephants were almost exclusively detected in S/EGF, with no records in DDF. Though our data are confined to the dry season and further work is required to determine seasonal movements of large mammals in the wildlife sanctuary, it is possible that some species may use S/EGF seasonally or much less during the wet season. In Thailand, banteng use dry evergreen forests in the day during the dry season, especially in the late dry season whereas they remain in DDF throughout the day in the wet season (Bhumpakphan & McShea, 2011; N. Bhumpakphan pers. comm.). Similarly, large Indian civets were observed shifting the centre of their home range from mixed deciduous forest to evergreen forest in the early dry season, and small Indian civets were observed shifting from DDF to evergreen forest in the late dry season (Rabinowitz, 1991). Carnivores Our data confirms the occurrence of 16 carnivore species at Chhep Wildlife Sanctuary. Bears and leopards were recorded less frequently and only in S/EGF. The latter is somewhat surprising as leopards were recorded at approximately 70% of the camera trap stations in DDF in the eastern plains of Cambodia (Gray et al., 2012) where un-baited camera trap pairs were spaced approximately 2–3 km apart along roads, trails, animal paths and ridgelines in mixed habitat types, with the highest proportion in DDF (R. Crouthers, per. comm.). Other studies in Indochina, where DDF persists, have also not found a strong association of leopards with evergreen forests and semi-evergreen forest (Simcharoen et al., 2008; Gray Cambodian Journal of Natural History 2017 (1) 63–75 Large mammals at Chhep Wildlife Sanctuary Table 1 Records of large mammals in Chhep Wildlife Sanctuary during the 2012–2013 and 2013–2014 dry seasons. IUCN status: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; LC = Least Concern. ND = Not detected. Species 1 IUCN status 2012–2013 dry season 2013–2014 dry season Naïve occupancy1 Encounter rate2 Naïve occupancy1 Encounter rate2 Golden jackal Canis aureus LC 0.26 23.63 0.61 84.88 Dhole Cuon alpinus EN 0.02 0.42 0.02 0.20 Jungle cat Felis chaus LC 0.06 1.69 0.11 2.74 Clouded leopard Neofelis nebulosa VU 0.02 0.42 0.03 0.39 Leopard Panthera pardus VU 0.06 2.95 0.06 1.56 Leopard cat Prionailurus bengalensis LC 0.22 9.28 0.35 9.97 Small Asian mongoose Herpestes javanicus LC 0.02 2.11 0.03 0.39 Crab-eating mongoose Herpestes urva LC 0.22 14.77 0.26 13.69 Yellow-throated marten Martes flavigula LC 0.10 2.11 0.19 4.11 Ferret badger Melogale sp. LC 0.12 6.75 0.13 6.45 Sun bear Helarctos malayanus VU 0.02 0.42 0.02 0.20 Asiatic black bear Ursus thibetanus VU 0.02 0.42 ND ND Common palm civet Paradoxurus hermaphroditus LC 0.66 81.01 0.69 61.22 Large-spotted civet Viverra megaspila EN 0.44 62.45 0.56 59.07 Large Indian civet Viverra zibetha LC 0.28 10.97 0.37 24.84 Small Indian civet Viverricula indica LC 0.24 14.77 0.37 39.31 Gaur Bos gaurus VU 0.08 1.69 0.05 1.76 Banteng Bos javanicus EN 0.04 0.84 0.02 0.20 Sambar Rusa unicolor VU 0.24 6.75 0.19 4.69 Eld's deer Rucervus eldii EN ND ND 0.02 0.20 Red muntjac Muntiacus muntjac LC 0.70 64.14 0.76 35.60 Eurasian wild pig Sus scrofa LC 0.80 80.59 0.87 68.06 Lesser Oriental chevrotain Tragulus kanchil LC 0.22 27.85 0.19 11.34 Burmese hare Lepus peguensis LC 0.10 7.59 0.23 35.79 Sunda pangolin Manis javanica CR 0.02 0.42 ND ND Crab-eating macaque Macaca fascicularis LC 0.38 20.68 0.29 15.84 Northern pig-tailed macaque Macaca leonina VU 0.26 18.99 0.26 9.00 Indochinese silvered langur Trachypithecus germaini EN 0.06 1.27 ND ND Asian elephant Elephas maximus EN 0.04 3.38 0.06 0.98 Malayan porcupine Hystrix brachyura LC 0.36 37.13 0.27 13.89 Proportion of stations a species was detected at least once; 2 Number of records / 1,000 camera-trap-nights. & Phan, 2011; Gray, 2012). Although sampling effort in DDF increased during the second survey period at Chhep Wildlife Sanctuary, leopards were not detected there or at stations in mosaics of DDF and S/EGF. Lower detectability in DDF is unlikely to be the only plausible explanation and it may be that leopards occur at relatively low densities in the wildlife sanctuary. Severe declines have occurred in leopard populations across Indochina Cambodian Journal of Natural History 2017 (1) 63–75 (Rostro-García et al., 2016), and Chhep is unlikely to be an exception. The reasons for this decline are many, but hunting for external markets has played a role, at least in the 1990s (Loucks et al., 2009). According to a hunter living near Chhep Wildlife Sanctuary, a leopard body was previously sold to a middleman for its skin and bones for about 200 USD. Bears were also were in demand. The gall bladder and bones of each Asiatic black bear sold for ca. © Centre for Biodiversity Conservation, Phnom Penh 67 68 Suzuki A. et al. Table 2 Percentage of camera trap stations in three forest types where mammal species were recorded in Chhep Wildlife Sanctuary during the 2012–2013 and 2013–2014 dry seasons. No. of Species stations detected Golden jackal Canis aureus 1 44 DDF S/EGF (n=28)1 (n=43)1 DDF & S/EGF mosaic (n=19)1 40.91 34.09 25.00 Dhole Cuon alpinus 2 50.00 50.00 0.00 Jungle cat Felis chaus 8 87.50 0.00 12.50 Clouded leopard Neofelis nebulosa 3 0.00 66.67 33.33 Leopard Panthera pardus 6 0.00 100.00 0.00 Leopard cat Prionailurus bengalensis 31 32.26 38.71 29.03 Small Asian mongoose Herpestes javanicus 3 66.67 33.33 0.00 Crab-eating mongoose Herpestes urva 24 25.00 42.00 33.00 Yellow-throated marten Martes flavigula 17 41.18 29.41 29.41 Ferret badger Melogale sp. 14 42.86 28.57 28.57 Sun bear Helarctos malayanus 2 0.00 100.00 0.00 Asiatic black bear Ursus thibetanus 1 0.00 100.00 0.00 Common palm civet Paradoxurus hermaphroditus 71 27.78 45.00 25.00 Large-spotted civet Viverra megaspila 52 42.31 25.00 30.77 Large Indian civet Viverra zibetha 31 3.13 74.00 23.00 Small Indian civet Viverricula indica 32 50.00 28.13 18.75 Gaur Bos gaurus 7 14.29 71.43 14.29 Banteng Bos javanicus 2 50.00 0.00 50.00 Sambar Rusa unicolor 23 21.74 52.17 26.09 Eld's deer Rucervus eldii 1 100.00 0.00 0.00 Red muntjac Muntiacus muntjac 76 31.58 46.05 21.05 Eurasian wild pig Sus scrofa 85 31.00 46.00 22.00 Lesser Oriental chevrotain Tragulus kanchil 20 0.00 90.00 10.00 Burmese hare Lepus peguensis 18 55.56 11.11 27.78 Sunda pangolin Manis javanica 1 0.00 100.00 0.00 Crab-eating macaque Macaca fascicularis 37 18.92 48.65 29.73 Northern pig-tailed macaque Macaca leonina 26 0.00 76.92 23.08 Indochinese silvered langur Trachypithecus germaini 3 33.33 33.33 33.33 Asian elephant Elephas maximus 6 0.00 83.33 16.67 Malayan porcupine Hystrix brachyura 32 12.12 60.61 27.27 n = Number of camera trap stations. Stations used in both dry seasons are counted as one. 100 USD and those of sun bear were sold for 30 USD in the 1990s. Near the Thailand border in 1994, prices for these species were 140 USD for a leopard skin, 3.20 USD/ kg for sun bear bones, and 80 USD for gall bladders from unidentified bears (Martin & Phipps, 1996). Besides the leopard, three medium or small cat species were recorded in Chhep Wildlife Sanctuary: © Centre for Biodiversity Conservation, Phnom Penh clouded leopard Neofelis nebulosa, jungle cat, and leopard cat. Clouded leopards were recorded at three stations including the edge of riverine forests and a juvenile was photographed in S/EGF. Leopard cats were recorded in all forest types, and a kitten with an adult was recorded in late December. The latter species (n=73) exhibited nocturnal (53.4%) and crepuscular (30.1%) behaviour in our study (Fig. 3) and its activity pattern varies from Cambodian Journal of Natural History 2017 (1) 63–75 Cambodian Journal of Natural History 2017 (1) 63–75 B C D E F G H I J K L Fig. 2 Mammal species recorded during the 2012–2013 and 2013–2014 dry seasons in Chhep Wildlife Sanctuary. A) Banteng Bos javanicus; B) Gaur Bos gaurus; C) Eld’s deer Rucervus eldii; D) Sambar Rusa unicolor; E) Asiatic black bear Ursus thibetanus; F) Leopard Panthera pardus; G) Clouded leopard Neofelis nebulosa; H) Dhole Cuon alpinus; I) Large-spotted civet Viverra megaspila; J) Indochinese silvered langur Trachypithecus germaini; K) Sunda pangolin Manis javanica; L) Asian elephant Elephas maximus. Large mammals at Chhep Wildlife Sanctuary © Centre for Biodiversity Conservation, Phnom Penh A 69 70 Suzuki A. et al. Fig. 3 Activity patterns of mammal species detected >20 times during the 2012–2013 and 2013–2014 dry seasons in Chhep Wildlife Sanctuary. Dawn (light grey): 05:00–7:00 hrs, day (horizontal lines): 07:01–16:59 hrs, dusk (dark grey): 17:00–19:00 hrs, night (black): 19:01–04:59 hrs. arrhythmic to nocturnal in Indochina (e.g., Rabinowitz, 1990; Grassman et al., 2005a; Austin et al., 2007; Kitamura et al., 2010; Gray et al., 2012). Jungle cats were almost exclusively detected in DDF and not in S/EGF despite greater sampling effort in the latter. This species was photographed 18 times at eight camera trap stations in total, including near a small pool, motorcycle trails, and a very small pocket of semi-evergreen forest in DDF. This is consistent with data from eastern Cambodia, where 96% of encounters were in DDF (Gray et al., 2012), and supports the idea that the species is strongly associated with DDF in Indochina (Duckworth et al., 2005). Jungle cats are likely to be naturally rare, or have become rare in Indochina (Duckworth et al., 2005) and recent studies suggest that the species is very rare in Vietnam (Willcox et al., 2014) and Thailand (Simcharoen et al., 2014; Tantipisanuh et al., 2014). Our records consequently highlight the importance of Chhep Wildlife Sanctuary for jungle cats in Indochina, together with the eastern plains (Gray et al., © Centre for Biodiversity Conservation, Phnom Penh 2014). Fishing cats Prionailurus viverrinus, Asiatic golden cats Catopuma temminckii, and marbled cats Pardofelis marmorata were not recorded in our study, although all three species have previously been recorded in other areas in the northern plains (Rainy & Kong 2010; Edwards & Demski 2012; Suzuki et al., 2015). Tigers Panthera tigris were also not recorded, although there are also historical records from the northern plains, including documentation that a minimum of 34 tigers was killed in 1998 (Sun, 2000). The golden jackal was the most commonly photographed carnivore while dholes Cuon alpinus were recorded only twice in two dry seasons. Golden jackals were detected in all forest types, however, naïve occupancy and encounter rates increased greatly in the 2013– 2014 survey period. Although comparisons between years in our dataset must be viewed with caution, this could be partially due to increased sampling effort in DDF in 2013–2014 (=22 camera trap stations vs. 8 in 2012– Cambodian Journal of Natural History 2017 (1) 63–75 Large mammals at Chhep Wildlife Sanctuary 2013). Like eastern Cambodia, where 98% of encounters were made in DDF (Gray et al., 2012), encounter rates of this species were probably high in DDF at Chhep Wildlife Sanctuary. Of six camera stations where jackals were photographed >20 times, five were in DDF and one in a small area of semi-evergreen forest approximately 20 m from DDF. In contrast to the golden jackal, dholes may occur in low densities at Chhep Wildlife Sanctuary. During our survey period, canine distemper virus (CDV) had possibly spread across Cambodia and could have lowered our detections of dholes which are susceptible to the disease (Kamler et al., 2015). Although dholes are less likely to occur in human-dominated landscapes than golden jackal (Jenks et al., 2015), they could be more susceptible to CDV (J. Kamler, pers. comm.) due to: 1) their requirement for larger group sizes to kill larger prey compared to jackals which hunt smaller prey (Johnsingh, 1982; Moehlman, 1983; Mukherjee et al., 2004; Jaeger et al., 2007); 2) amicable behaviour between within-group individuals (Fox, 1984). Further, when local people enter Chhep Wildlife Sanctuary they often bring dogs and dholes are often killed in snares in Cambodia (J. Kamler, pers. comm.). Two species of mongoose (Herpestidae) were confirmed during our survey: small Asian mongoose Herpestes javanicus and crab-eating mongooses H. urva. Encounter rates of small Asian mongoose were very low in both survey periods, but may not accurately reflect their status in the area; rather, they likely reflect sampling bias (see Duckworth et al., 2010). Villagers stated that the species is relatively common around villages (where no camera traps were set) and attacks poultry. In contrast, crab-eating mongooses were often photographed traveling in groups of up to four individuals during the day time in the wildlife sanctuary. The species was frequently recorded at three camera trap stations in particular. The first was at a river bed in DDF, and records began when the water became very shallow at the start of January. This station was set in both survey seasons, and produced the highest number of photographs of the species in both years, similar to the experience of Than Zaw et al. (2008) near a stream in the Hakaung Valley of Myanmar. The two other stations were in a small pocket of semi-evergreen forest near a dirt road in DDF and at a shallow water pool where water remains until February under the tangled branches of a shrub. Two species of Mustelidae were recorded, yellow throated-marten Martes flavigula and ferret badger. Hog badgers Arctonyx collaris were not recorded, although the species is thought to occur, or have occurred, in the wildlife sanctuary. Villagers reported using hog badger oil Cambodian Journal of Natural History 2017 (1) 63–75 Fig. 4 Camera trap stations where large-spotted civet (top), large Indian civet (middle), and small Indian civet (bottom) were surveyed at Chhep Wildlife Sanctuary. White squares represent detections and black points non-detections. for traditional medicinal use. Yellow-throated martens exhibited diurnal activity in line with previous studies (Grassman et al., 2005b; Johnson et al., 2009), and were recorded in both DDF and S/EGF. This species was mostly photographed once at each station where it was detected, but was photographed more than four times at two in particular. One of the latter stations was in a small dry stream (<5 m width) in DDF. The other was © Centre for Biodiversity Conservation, Phnom Penh 71 72 Suzuki A. et al. in a small patch of bamboo forest close to a large pond in DDF where a leaf litter fire was observed at the end of February in the 2013–2014 dry season. Local people were also frequently photographed at this station. Ferret badgers were photographed at 14 camera trap stations. The most frequent capture station was in a dry river bed at the edge of S/EGF with 21 records from December to March. In Cambodia, the presence of large-toothed ferret badger is confirmed and the presence of small-toothed ferret badger Melogale moschata also remains a possibility (Schank et al., 2009). Four species of civet (Viverridae) were recorded with high encounter rates. This is likely due in part to bias in camera trap placement as the original purpose of the second survey was to investigate the occupancy of the subfamily Viverrinae. Nevertheless, the high encounter rate of the Endangered large-spotted civet is significant. This species is rarely recorded in Myanmar (Than Zaw et al., 2008), southwestern Cambodia (Holden & Neang 2009), Malaysia (Hamirul et al., 2015), Thailand (except for Khao Ang Rue Nai Wildlife Sanctuary) (Chutipong et al., 2014), Laos, and Vietnam (W. Duckworth pers. comm.). In our study, the large-spotted civet was the second and third most commonly photographed carnivore in the 2012–2013 and the 2013–2014 dry seasons respectively, and the species was detected in all forest types (Fig. 4). This highlights the global conservation significance of Chhep Wildlife Sanctuary for the species and accords with records from Thailand (Chutipong et al., 2014) and across its range (Timmins et al., 2016). The camera trap stations with the top three highest encounter rates in our study were within or close to DDF, namely a shallow waterhole in a mosaic of DDF and S/EGF, a temporary pond in DDF, and a dry river bed in DDF. A high encounter rate was also reported near water sources in Thailand (Jenks et al., 2010) and southwestern Cambodia (Holden & Neang 2009). In contrast, large Indian civets were rarely detected in DDF (Fig. 4). This is consistent with Gray et al. (2010), but differs from Thailand where the species is common in DDF (Chutipong et al., 2014). The three stations with the highest encounter rates in our study comprised two at the intersection of animal trails in S/EGF and one on an animal trail close to a pond in S/EGF. The degree of spatial overlap between the sympatric large Indian civet and large-spotted civet is largely unknown (Gray et al., 2010), and likewise with small Indian civet Viverricula indica. In our study, the former two species were both photographed at 15 of the same camera stations. Eight of these were in S/EGF, five in a mosaic of DDF and S/EGF, and two in DDF (Fig. 4). Large-spotted civets were photographed with small Indian civets more often at the same station (20 stations) © Centre for Biodiversity Conservation, Phnom Penh than large Indian civets (3 stations). An occupancy study is currently underway to understand habitat use of these three Viverrinae in DDF-dominated landscapes at Chhep Wildlife Sanctuary. Large ungulates Large ungulates were detected at a minority of camera trap stations during our survey. However, gaur, banteng, Eld’s deer and sambar Rusa unicolor were all photographed at a single station which targeted a trail a few meters from a relatively small water hole in DDF, approximately 2 km from semi-evergreen forest. The water hole was surrounded by grass which was burnt in January and became dry in February. The area surrounding the waterhole was open, lacking tall grass or scrubs, allowing large ungulates easy access from many directions. This also increased the detection range of the camera beyond the targeted trail resulting in photographs of these ungulates travelling off-trail. Away from this seasonal water hole, gaur was detected only in an area of contiguous evergreen forest, stretching from southern Laos to Preah Roka Wildlife Sanctuary (Cambodia). These cameras were located at the intersections of animal trails and along a dry river bed in S/EGF. Gaur occur in a wide range of habitats (Bhumpakphan & McShea, 2011), and their use of different forest types varies according to season (Ahrestani et al., 2012), social class (Steinmetz et al., 2008), and their populations in relation to the availability of and competition over high-quality habitat (Steinmetz et al., 2010). Encounter rates of gaur, banteng, sambar and Eld’s deer were relatively low. This partly reflects bias in our placement of camera traps. Our surveys originally targeted small carnivores, and thus, if locations were deemed unsuitable for these or no signs of small carnivores were found, cameras were not set in locations even where salt licks or places where signs of large ungulates were present. This bias is evident when our results are compared with previous studies in Chhep Wildlife Sanctuary. For example, encounter rates of gaur and banteng were higher during small-scale camera trap surveys in 2010 and 2011 which targeted kouprey Bos sauveli (Wildlife Conservation Society, unpublished data). Although sampling bias must therefore be considered, large ungulates populations are likely to be decreasing in the wildlife sanctuary as well as many other places within their range. The status of banteng is especially of concern. Banteng almost exclusively uses DDF where plant species preferred by the species are common (Bhumpakphan & McShea, 2011), and the species was recorded in DDF during line transect surveys at Chhep Wildlife Sanctuary (Rainy et al., 2010). However, sightings of banteng were Cambodian Journal of Natural History 2017 (1) 63–75 Large mammals at Chhep Wildlife Sanctuary very rare during our monthly visits to camera traps in DDF over two successive dry seasons. Though the potential for sightings would be less if banteng used evergreen forests in the wildlife sanctuary more in the day time during the dry season, as in West Thailand (Bhumpakphan & McShea, 2011), sightings were very rare even in mornings and evenings. Conservation implications DDF-dominated landscapes are threatened and poorly represented in protected areas in mainland Southeast Asia (McShea et al., 2005; Tantipisanuh & Gale, 2013; Wohlfart et al., 2014; Johnson, 2015). Lowland forests are also poorly protected: >90% of protected areas created after 1965 are located above 200 m a.s.l. (Déry & Vanhooren, 2011). Given this situation, the confirmed occurrence of DDF-associated species—namely jungle cat and two globally Endangered species, Eld’s deer and banteng—highlights the conservation importance of Chhep Wildlife Sanctuary. In addition, the Endangered large-spotted civet, which likely prefers lowland areas, was commonly photographed in the wildlife sanctuary, suggesting potential for the site to provide a stronghold for the species. Consistent with Tordoff et al. (2005), small areas of S/EGF were used by 27 large mammal species, indicating that these areas are likely to be important components of DDF-dominated landscapes for some large mammals during the dry season. Further research on seasonal habitat use and movements would assist conservation management of large mammals in Chhep Wildlife Sanctuary. Ack now le dge m e nt s We thank the Forestry Administration of the Ministry of Agriculture, Forestry and Fisheries for permission to carry out the field survey. The survey was supported by the Wildlife Conservation Society, Cambodia Program. 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Et hnobot a nic a l k now le dge of t he Kuy a nd K hm e r pe ople in Pre y La ng, Ca m bodia Nerea TURREIRA-GARCIA1,*, Dimitrios ARGYRIOU1, CHHANG Phourin2, Prachaya SRISANGA3 & Ida THEILADE1,* 1 Department of Food and Resource Economics, University of Copenhagen, Rolighedsvej 25, 1958 Frederiksberg, Denmark. 2 Forest and Wildlife Research Institute, Forestry Administration, Hanoi Street 1019, Phum Rongchak, Sankat Phnom Penh Tmei, Khan Sen Sok, Phnom Penh, Cambodia. 3 Herbarium, Queen Sirikit Botanic Garden, P.O. Box 7, Maerim, Chiang Mai 50180, Thailand. * Corresponding authors. Email n.turreira@gmail.com, idat@ifro.ku.dk Paper submitted 30 September 2016, revised manuscript accepted 11 April 2017. ɊɮɍɅʂɋɑɳȶſɆ ȹɅƺɁɩɳȼˊɊNJȴɁɩȷ Ʌɩȶ ɑɒȴɊɅɿɴȼɍɈɫȶɴɇơȲɳɍˊɵƙɈɳȺˊ ƙɁȪɎLJɅɳȴȼɫȶǃNjɅȷɸɳɀɹȼɫȶɈɩɳɑɑ ɳɍˊɄɅDžɅɄɊƗƺɁɩɳǷȹɭɸ Ɏ ɩȻɁɸɆɅɽɈɯȲɳȴɌɑɽɳǷʆ ɳDŽɹƺnjɻ ȶǁ ƳɌȳɮȷɆɌ ɩǒƏɅLJɅɆɅƏɋȲƙɊɩɁɄɅDžɅɄɊƗƺɁɩɴȼɍDžƚɆɽNjɅ Ʌɩȶ ȴɸǍɊȲɸɴɒȶȼɍɽ NJɈɆɅƎNjɅɵɅȹɪɎɎɆƓɄɊɾɌɆɑɽȹɅƺɁɩɳȼˊɊɅɩȶƙɆƺȹɅȲƒɭȶɁɸɆɅɽɃɮDŽɸȶɈɩɉɈɳǎȲʆ ƳɌɑɩȲǜɳɅɹLJɅȷȶƙȲȶɡȲǒɌɌɭȲſƺɁɩ ɴȼɍɳƙɆˊƙLJɑɽɳƽɋƙɆƺȹɅɌɑɽɳǷȹɭɸɎ ɩȻɁɸɆɅɽɵƙɈɃɸdžɆɳɑɑɑɍɽȷɭȶɳƙƳɋɊɯɋȲƒɭȶƙɆɳɃɑȲɊƕɭƺʆ ƳɌƙɆɊɮɍɃɩɅƒɅʂɋLJɅ ɳɄƛˊɳǷȷɳdžƚɹƹƒɸʒʐʑʔ Ʌɩȶ ʒʐʑʖʆ ƳɌɳɄƛˊɴɇɅɃɪɑɩȲǜɳƽɋNjɅƳɌȷɮɍɌɯɊɈɪɑNjȹɩȲɑɒȴɊɅɿɅɩȶNjɅƳɌƙɆɊɮɍȴɸɅɩɁɳƽɋ ɳɑɌ ɪ (freeȬlisting) ƺɊɯɋƙɆƺȹɅȷɸɅɮɅʓʑdžȲɽ ɔƒȲƙɆɊɮɍɌɭȲƺ ſ ɁɩɅɩȶƳɌɈɩNJȲǜƺɊɯɋƙȲȩɊɳƵɍɳǮȷɸɅɯɅʑʒdžȲɽ ƙɁȪɎLJɅ ɳɄƛˊȲɭƒȶɉɮɊɩȷɸɅɯɅɆɪȲɭƒȶɳȳɁƎƙɈɹɎ ɩǓɌɅɩȶɑƐɫȶɴƙɁȶʆ ɑɌɭɆNjɅɌɭȲſƺɁɩɴȼɍɳȴɅɩɋɊɳƙɆˊȷɸɅɯɅʓʗʔƙɆɳɉɃ ɴȼɍƙɁȪɎLJɅȲɁɽƙǂ Ȳƒɭȶ ɳdžɹʙʐ% ƙɁȪɎLJɅƙɆɊɮɍ Ʌɩȶ ɳɄƛˊȷɸɴɀȲǃƒȲɽʆ ƙɆɳɉɃDŽɸȶɳɅɹNJȴɳƙȷˊɅƙɁȪɎLJɅɳƙɆˊƙLJɑɽƺɣɑɂ(ʖʗ%) ǕǓɌ(ʔʔ%)Ʌɩȶƺ ɑNjƖɌɺɳƙɆˊƙLJɑɽ(ʓʗ%) ɳɒˊɋNJȴɳƙȷˊɅƙɆɳɉɃɊɯɋƙɁȪɎLJɅɳƙɆˊƙLJɑɽɳƙȷˊɅnjɻ ȶʆ ɄɅDžɅɵƙɈɳȺˊ ɴȼɍɑɸƴɅɽƺȶɳȴɑƙNjɆɽ ȹɅƺɁɩȴɯɋȴɬƺƙɆɳɉɃɳȺˊ ɇƎɍɽȹʂɌɵɅɈɯȲDipterocarpus ɴȼɍƙɁȪɎLJɅƸɁɽȲɭƒȶƙɆɳɉɃɌȶɳƙƵɹɳƽɋɔȶƀƳɌIUCNʆ ɆɭɌɑɅɩȶ ȝɑƎɪLJɅǒƀɍɽƙɆɳɉɃɌɭȲƺ ſ ɁɩɴȼɍNjɅƙɆɳnjȹɅɿȲɭƒȶȷɸɅɯɅƙɆǓȲɽƙɆɴɒɍƵƒ ɳɒˊɋNjɅɌɳɆȢɆɵɅƳɌɳƙɆˊƙLJɑɽȳɭɑƵƒ (ɆɭɌɑƙɆ ɊɮɍƙɆɳɉɃɌɭȲſƺɁɩȼɮȷƵƒɳǵɅɫȶƙɆɳɉɃɴȼɍȝɑƎɪɳƙɆˊƙLJɑɽ)ʆ NjɅɌLJɋƳɌɀɿƺɳƙȷˊɅɑƎɪɈɪƙɆɳɉɃɌɭȲƺ ſ ɁɩNjɅƙɆɳnjȹɅɿɴȼɍ ɆƷƟȻɈɪƙɆɳɉɃɴȼɍɇƎɍɽǒɌɺɑɸƴɅɽɴɇƒȲɳɑȼƊȲɩȷƃɅɩȶɎɆƓɄɊɾ ƙɈɊDŽɸȶǒƏɅNJɈɌLJɋ Ʌɩȶ ɔɉɩɌȲƞʆ ƳɌɔɉɩɌȲƞɵƙɈɳȺˊ ɈɩɁƺ NjɅǒɌɺɑɸƴɅɽȲɭƒȶƳɌƙɃƙɃȶɽȹɪɎNJɈ Ʌɩȶ ȷɸɳɀɹȼɫȶljȲɽɈʂɅƑɅɫȶɌɭȲƺ ſ ɁɩɅɩȶɊɅɭɑƞɵɅƙɆƺȹɅȲƒɭȶɁɸɆɅɽ Ʌɩȶ ȹɅƺɁɩɳȼˊɊɳǷɵƙɈ ɓȶɽʆ Abst ra c t Indigenous peoples and forest-dependent communities are known to hold unique knowledge on natural resources in their surrounding environment. However, environmental degradation has diminished the availability of natural resources and threatens the bio-cultural survival of indigenous and local people world-wide. This study documented the plants used by people living in the vicinity of one of Cambodia’s last remaining lowland rainforests. Fieldwork took CITATION: Turreira-García, N., Argyriou, D., Chhang P., Srisanga, P. & Theilade, I. (2017) Ethnobotanical knowledge of the Kuy and Khmer people in Prey Lang, Cambodia. Cambodian Journal of Natural History, 2017, 76–101. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer place between 2014 and 2016. Participatory mapping exercises and ‘free-listings’ with 31 informants and participatory botanical collections and focus group discussions with 12 key informants were conducted across three villages in the Preah Vihear and Stung Treng provinces. A total of 374 useful ‘folk taxa’ were recorded, 90% of which were collected and identified. These species were mostly used as medicine (67%), food (44%) and/or materials (37%) with many species having multiple uses. The most important forest resources for the Kuy people were resin trees of the genus Dipterocarpus, some of which are listed as Endangered by IUCN. Men and women knew similar numbers of useful plants and played different roles in relation to these. Given the many useful plants reported, the indication of culturally and economically important species, and their distribution and conservation status, forest conservation appears to be essential to maintain the livelihoods and associated ethnobotanical knowledge of local and indigenous people in Prey Lang. Ke yw ords Bio-cultural diversity, knowledge loss, Kui, Kuoy, local ecological knowledge, participatory plant collection, Prey Long, traditional ecological knowledge. I nt roduc t ion Indigenous peoples and forest-dependent people in general hold a unique knowledge on natural resources in their surrounding environment (Martin, 2004). Among other things, their knowledge about plants useful for medicine, food, and construction improves their resilience to adversity. Worldwide, deforestation threatens the availability of natural resources useful for forest-dependent people, placing their bio-cultural survival under pressure. Ethnobotanical knowledge is directly related to the use of plant resources (Gadgil et al., 1993): if a plant is no longer available, it cannot be used and knowledge related to it may disappear. Under rapidly changing socio-economic, political and environmental conditions, knowledge related to the use of natural resources can be lost within a single generation (Reyes-García et al., 2013), especially given that ethnobotanical knowledge is usually orally transmitted and rarely documented (Case et al., 2005; Turreira-García et al., 2015). Documentation of ethnobotanical knowledge consequently provides an ancestal legacy for current and future generations. Ethnobotanical knowledge can also serve as an indicator of biodiversity (Salick et al., 1999) and as a measure of dependency upon the surrounding environment (Araújo & Lopes, 2011). There is a growing trend in employing local people as parataxonomists to provide biodiversity inventories (Janzen, 2004; Janzen & Hallwachs 2011; Zhao et al., 2016) and local knowledge is increasingly used in ecological and conservation research and monitoring. Local people are rarely actively involved in the research process, however (Brook & McLachlan, 2008). According to a recent review on the status of ethnobiology in Southeast Asia, Cambodia is one of the least researched countries (Hidayati et al., 2015) with only 13 ethnobiological publiCambodian Journal of Natural History 2017 (1) 76–101 cations between 1960 and 2014. Our reviews of recent ethnobotanical studies in Cambodia, Thailand, Vietnam, and Laos also reveal that most studies have been undertaken in Thailand and have mainly focussed on medicinal plants (Table 1). (Only studies that focused on ethnic groups and included (semi-)wild plants were taken into account. Studies that did not encompass local people’s knowledge, reviewed only one species or strictly inventoried homegardens were excluded). The only ethnobotanical studies involving the Kuy people in the literature were one Master’s thesis about materia medica employed by Kuy healers in Thailand, which documented the use of 333 medicinal plants (Virapongse, 2006), and a study of medicinal plants used for postpartum ailments (Grape et al., 2016). Prey Lang (‘our forest’ in Kuy language) covers 530,000 ha in the central plains of Cambodia and is considered the last intact lowland rainforest in mainland Indochina (MacDonald, 2004). In May 2016, 432,000 ha of Prey Lang were gazetted as a wildlife sanctuary. However, 4,700 ha of this area is affected by economic land concessions and mining concessions (Argyriou et al., 2016) and about 50,000 ha of forests bordering the sanctuary are impacted by 53 concessions for agro-businesses (LICADHO, 2016). Forest clearance within and nearby these concessions and rampant illegal logging throughout Prey Lang threaten its biodiversity and natural resources (Olsson & Emmett, 2007). An estimated 250,000 villagers also live in the vicinity of Prey Lang and depend on it for their livelihoods (Hüls Dyrmose et al., in press) and culture. The aims of our study were to: i) document the ethnobotanical knowledge of Kuy and Khmer people living nearby the Prey Lang forests (specifically regarding forest types, important natural resources, useful plants © Centre for Biodiversity Conservation, Phnom Penh 77 78 N. Turreira-García et al. Table 1 Previous ethnobotanical studies in Indochina based on searches made in Scopus, Web of Science and the Royal Library of Denmark and Copenhagen library services on 16 March 2017. UC = Plant use category (WEP = wild edible plants; MED = general medicine; DSD = digestive system disorder; CI= cognitive impairment; WH = women’s healthcare; REP = repellents and pesticides); Spp. = Number of species (not necessarily scientifically recognized species); Vill. = Number of villages; Inf. = Number of informants; n.s. = not stated; * = Includes cultivated species. Reference Ethnic group (Country) UC Spp. Vill. Inf. Grape et al. (2016) Kuy (Cambodia) MED, WH 68 4 50 Evergreen, semi-evergreen & deciduous dipterocarp forest Vegetation Chassagne et al. (2016) Buong (Cambodia) MED 214 28 202 Savanna, evergreen, semi-evergreen, deciduous dipterocarp & bamboo forest Whitney et al. (2016) Dao, Hmon, Kinh, Ma-Lieng, Sach, Tai, Tay, Xinh-Mun (Vietnam) n.s. 111 5 n.s. n.s. Cruz-Garcia & Struik (2015) Isaan (Thailand) WEP 20 1 7 Tangjitman et al. (2015) Karen (Thailand) MED, DSD 36 6 178 Neamsuvan et al. (2015) n.s. (Thailand) MED 95 7 7 Mangrove & swamp forest Offringa (2015) Khon Muang (Thailand) MED, CI n.s. n.s. 16 n.s. Elkington et al. (2014) Lao (Laos) MED 250 n.s. 12 Evergreen-mixed & deciduous forest Khuankaew et al. (2014) Tai Yai (Thailand) MED 141 4 126 n.s Junsongduang et al. (2014) Karen, Lawa (Thailand) MED 103 2 67 n.s Kosaka et al. (2013) Lao, Tai Leu, Tai Dam, Tai Deng, Khmu, Hmong (Laos) WEP 115 2 20 Paddy fields Tangjitman et al. (2013) Karen (Thailand) MED, WH 379* 14 458 Mixed deciduous, coniferous & hill evergreen forest Inta et al. (2013) Yuan (Thailand) MED 93 5 30 n.s. Srithi et al. (2012) Hmong (Thailand) MED, WH 79* 3 153 n.s. Cruz-Garcia & Price (2011) Isaan (Thailand) WEP 87 4 n.s. Dry monsoon forest (dipterocarp forest) Lamxay et al. (2011) Kry (Laos) MED, WH 49 3 20 n.s. de Boer et al. (2010) 17 groups (Laos) de Boer & Lamxay (2009) Brou, Saek, Kry (Laos) Dry monsoon forest (dipterocarp forest) Deciduous, tropical evergreen & dry dipterocarp forest REP 92 66 n.s. n.s. MED, WH 55 10 38 Secondary forest Inta et al. (2008) Akha (Thailand & China) MED 95 5 50 n.s. Libman et al. (2006) n.s. (Laos) MED 55 8 n.s. n.s. Johnson & Grivetti (2002) Karen (Thailand) WEP 47 2 32 Degraded secondary forest Van On et al. (2001) Dao (Vietnam) MED 200 n.s. n.s. Primary & secondary forest, bamboo thicket, grassland, plantation Anderson (1986a) Akha (Thailand) MED 121 n.s. n.s. Dry evergreen & lower montane (moist evergreen) forest Anderson (1986b) Lahu (Thailand) MED 68 n.s. n.s. Lower montane (moist evergreen) region © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer and forest-spirits); and, ii) investigate the ability of local and indigenous people to collect plant voucher specimens. We also compared the local names of plants and forest types to scientific classifications and assessed how much of their useful or culturally important flora was threatened. The study did not consider differences in knowledge between Kuy and Khmer people because of the cultural continuum between the two groups (Swift, 2013). Our findings will later be shared with the communities in the form of a book. M e t hods Study area and ethnicity The greater Prey Lang area extends over four provinces in the central plains of Cambodia: Preah Vihear, Stung Treng, Kratie, and Kampong Thom. The area contains seven vegetation types among its evergreen, semi-evergreen, and deciduous forests, which differ significantly in species composition, dominant tree species and plant community structure (McDonald, 2004; Olsson & Emmett, 2007; Theilade et al., 2011). Approximately 250,000 people live in the greater Prey Lang area and the dominant ethnic groups are Kuy (indigenous) and Khmer (Cambodian). The Kuy (also recorded as Kui, Kuoy, Kuay, Kouy, Suoy or Suay) occur in northeastern Thailand, southern Laos, and northern and northeastern Cambodia. Most of the Kuy people in Cambodia live in the Prey Lang area, with an unverified population estimate of 23,000 (Swift, 2013). Kuy and Khmer people are similar in terms of physical appearance, material culture, and religious practices: both groups are culturally and spiritually linked to the forest and practice of animism and Buddhism in Prey Lang (Swift, 2013). Lowland rice cultivation and swidden agriculture are widespread among both. The majority of inhabitants rely directly on the natural resources of Prey Lang for their livelihoods and resin tapping (extraction of oleoresin from dipterocarp trees) is the main source of cash income (Jiao et al., 2015; Hüls Dyrmose et al., in press). Differences between the Kuy and Khmer groups have become subtle since national integration and assimilation policies were adopted by the Cambodian Government following independence in 1953 (Baird, 2011). These policies were strengthened during the Pol Pot regime in the 1970s, when Kuy communities were resettled to lowland areas such as Prey Lang and those speaking Kuy language were punished. Interaction and inter-marriage between Kuy and Khmer is frequent and many Kuy Cambodian Journal of Natural History 2017 (1) 76–101 have adopted Khmer culture and traditions, although small differences still exist between the two groups. These include distinctive rituals (e.g., the Kuy practice communal fishing before the annual ceremony for the village spirit, perform rites for spirits before clearing new swiddens, or involve a certain species of turtle in weddings) and some characteristic crafts, foods, clothing and housing styles. While the two groups fomerly distinguished themselves through economic specialties such as iron production, their livelihood strategies of Kuy and rural Khmer are now very similar (Swift, 2013). In recent decades, the Kuy identity has been based upon language and/or family descent, whereby a person may identify themself as Kuy if they speak the language and/or have a Kuy parent. However, Kuy people sometimes deny their ethnicity because they may be perceived as being of lower status (Swift, 2013). The Kuy language also shares many terms with Khmer, which may be due to their shared roots (because both are MonKhmer languages) or borrowed from Khmer (Mann & Markowski, 2005). Three villages in Prey Lang were selected for the study: Thmea and Phneak Roluek in Preah Vihear Province and Spong in Stung Treng Province (Fig. 1). Thmea and Phneak Roluek were selected by representatives of the Prey Lang Community Network (PLCN) because they comprise traditional Kuy villages. The PLCN is a network of villagers within the Prey Lang area who advocate for forest protection through peaceful patrols and anti-logging interventions. Spong was selected by the authors due to its proximity to the core area of Prey Lang. This is the least disturbed area of the Prey Lang forests and is dominated by primary evergreen dipterocarp forest, with local residents reportedly being Khmer. At the time of the study, Thmea was the largest village (2,024 people), closest to a paved road (36 km), surrounded by disturbed and deciduous forest, and furthest from evergreen forest. Spong was the smallest village (497 people), furthest from paved roads (73 km) and markets (76 km), and mainly surrounded by primary evergreen dipterocarp forest. Phneak Roluek Village was intermediate in size (587 people), distance to a paved road (44 km) and distance to evergreen forest (CDB Online, 2010) (Fig. 1). Study formulation and methods The idea to conduct an ethnobotanical study was initially discussed by the authors and PLCN steering committee. The committee agreed that it would be useful to document their knowledge and agreed to co-design the study and participate in the research process. Fieldwork took © Centre for Biodiversity Conservation, Phnom Penh 79 80 N. Turreira-García et al. Fig. 1 Study sites in Prey Lang, Cambodia. Created using forest cover map (Open Development Cambodia, 2014) and natural earth data in QGIS. Fig. 2 Kuy woman carrying a handmade basket outside a traditional house. Phneak Roluek Village, Preah Vihear Province, September 2014 (© Nerea Turreira-García). © Centre for Biodiversity Conservation, Phnom Penh Fig. 3 Plant collector in Prey Lang, near Spong Village, Stung Treng Province, May 2015 (© Nerea Turreira-García). Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer place during September 2014, April–May 2015, and December 2016. Field methods included participatory mapping exercises, rankings, free-listings, forest walks, botanical collections and focus group discussions, and are explained in more detail below. To record local knowledge of plants used by the Kuy people at Prey Lang, the Thmea, Phneak Roluek, and Spong villages were visited three times. During the first visit, five to seven men and four to seven women participated in workshops led by the first author and an interpreter in each village. The men were 53 ±13 years old and women 52 ±3 years old on average. Workshop participants were decided by a PLCN representative from each village, based on participants’ knowledge of the area and its natural resources. Following the International Society of Ethnobiology guidelines (ISE, 2006), the study objectives were explained and participants gave their prior informed consent. Sharing and publication of results, and confidentiality were agreed upon. The workshops consisted of a participatory mapping exercise where participants drew their community boundaries, forest types, zones of use, and the most important sites and natural resources (Gilmore & Young, 2012). This helped the authors to become familiar with the area and local terms and aided the design of later forest walks and botanical collections with the participants. Participants were also asked to describe the defining characteristics of each-forest type and natural resources identified in the mapping exercise were ranked in order of importance. Men and women were separated into two genderbased groups to free-list useful plants, including those they did not use or only used infrequently. This allowed for smaller and more productive working groups, because men and women sometimes differ in their knowledge due to gender-based divisions of labour (Reyes-García et al., 2007). This was especially valuable for engaging the women who otherwise might have contributed less. Each group recorded the name, growth form, habitat, uses and flowering season of each plant (Martin, 2004; ReyesGarcía et al., 2006; Powell et al., 2014) and took about 90 minutes to complete their free-lists. During the second and third visits, plant species registered during the workshops (and others not included in the workshop lists) were collected during forest walks (Martin, 2004). Twelve people comprising two men and two women from each of the three villages assisted with the plant collection (Figs 2–3), seven of whom were Kuy and five Khmer. These were divided into male and female groups and trained in botanical specimen collection and note-taking. During the forest walks, local names for forest types were compared with the vegetation classifiCambodian Journal of Natural History 2017 (1) 76–101 cations and descriptions of McDonald (2004) and Rollet (1972). A total of 704 specimens were collected, after which the collectors were asked about the uses, parts used, preparation methods, and local (folk) names for each plant. Local names that referred to the same scientific species were regarded as synonyms and counted as one taxon in analysis. At the end of each visit, plant collectors cross-checked information recorded during focusgroup discussions. Information about forest spirits was collected through informal conversations with the plant collectors. Plant uses were later categorised following Cook (1995), who defined 12 use categories, plus two additional categories defined by Gruca et al. (2014), namely ‘cultural diseases and disorders’ and ‘ritual/magical’ uses, and two categories defined by the authors, namely ‘resin’ and ‘commerce’ (Table 2). Ailments treated using medicinal plants were translated verbatim. Plant voucher specimen were dried and pressed at the Forest and Wildlife Research Institute in Phnom Penh. These were identified by two of the authors (CP & PS) and a full set of specimens were deposited in the Queen Sirikit Botanic Garden in Thailand. Species names and family classifications were confirmed using The Plant List (2013), and IUCN (2017) was used to determine the conservation status of species. In our analysis and interpretation, we refer to folk taxa based on local names, and to scientific species. Terms given in italics are in Khmer language. Re sult s Forest types During the participatory mapping exercise and forest walks, all three communities claimed to distinguish four types of forest (prey in Khmer, also used by Kuy): 1. Prey robóh (‘sparse forest’, Fig. 4), described as a non-dense, deciduous forest that grows nearby their rice fields and chamkars (‘forest gardens’). Prey robóh corresponds to two forest types described by McDonald (2004), namely deciduous forest (<35 m tall) and short semi-evergreen forest (3–12 m tall). Local informants did not distinguish successional stages of the deciduous forest, whereas the short semi-evergreen forest is a combination of both deciduous and evergreen species. 2. Prey sralao’ (no English translation, Fig. 5) was described by the local communities as a tall evergreen forest at Prey Lang, denser than robóh but easily traversable, and characterised by dominance of the sralao’ trees (Lagerstroemia sp.). McDonald (2004) and Rollet (1972) classified this forest type with the same name. © Centre for Biodiversity Conservation, Phnom Penh 81 82 N. Turreira-García et al. Table 2 Description of plant use categories employed in this study for classifying plant use records (adapted from Cook (1995) and Gruca et al. (2014)). Use category Abbr. Description Food F Plants eaten by human beings, and plants used to make beverages Food additives Fa Processing additives and other additive ingredients used in food or beverages preparation Vertebrate food V Forage and fodder for domestic or wild vertebrates that are useful to humans Invertebrate food I Plants eaten by invertebrates that are useful to humans Apicolous A Plants that provide pollen, nectar or resins as sources for honey or propoleum production Fuel Fu Plants used to produce charcoal, or used as petroleum substitutes, alcohols, tinder or firewood Materials Ma Plants used for construction of houses, fences or bridges, or to elaborate handicrafts, music instruments, work tools, weapons, home objects, etc. This category includes fibres, waxes, oils, chemicals and their derived products (but not Resin), cosmetic products and dyes Social S Plants used for cultural purposes, which are not definable as food or medicines. This category includes stimulants, and plants used for games (modified according to local beliefs) Toxic to vertebrates Tv Plants that are poisonous to vertebrate animals, both accidentally and when deliberately applied, such as extracts and preparations used for fishing and hunting Toxic to nonvertebrates Tn Plants that are poisonous to non-vertebrates, both accidentally and when deliberately applied. This category includes insecticides and herbicides Medicinal M Plants used to cure human and animal sicknesses Environmental E Plants used to protect, improve, and fertilise soils; to provide shadow, as living fences, ornamentals or that form a structural part of agroforestry systems Cultural Diseases and Disorders CDD Plants used to treat disorders caused by spirits, such as mental illnesses and curses (modified according to local beliefs) Ritual/Magical Uses RMU Plants used during healing ceremonies, incantations, prayers, offerings and sacrifices made to deities, fetishes/amulets/charms, divination/oracles, black magic/bad medicines, incense Resin R This category is separated from ‘Materials’ due to its high importance in Cambodian livelihoods Commerce C Plants used for trade and are part of the household economy 3. Prey sdok (‘thick/narrow forest’), prey thom (‘tall forest’) and prey chas (‘old forest’) were Khmer synonyms for the ‘hard to penetrate’, tall forest at Prey Lang (Fig. 6). According to informants, this forest type is where more natural resources, expensive timber trees, resin trees, rattan and animals occur. It corresponds to the semi-evergreen and evergreen dipterocarp forests described by McDonald (2004) and the dense forest described by Rollet (1972). 4. Prey choam (in Kuy) or prey roneam (in Khmer, Fig. 7), was described as ‘the forest growing on land permanently covered by shallow water’. McDonald (2004) distinguished two types of swamp forest, deciduous swamp forest and evergreen swamp forest, and Theilade et al. (2011) provided a detailed account of the latter. Both types of swamp forest are rare and endemic to the region. Inhabitants of the three villages collect timber and nontimber forest products (NTFPs) in different areas of all four forest types. They usually follow rivers, trails which they create and maintain, and at Thmea Village, also a road built by a mining company. During their forest trips, © Centre for Biodiversity Conservation, Phnom Penh they hunt and collect wood for construction, medicinal plants, resin and rattan. Trip frequency, duration, transportation and distance travelled vary according to the purpose and needs of each trip. In the dry season for example, men usually travel in pairs to the forest by coyon (local tractor) to collect oleoresin from dipterocarp trees. These trips last about three days and the collectors sleep in hammocks in forest shelters. Women usually walk or are carried by coyon or motorbike to collect NTFPs in daily trips throughout the year. Importance of forest resources Our ranking exercise revealed that the most important resources for all three villages are the resin trees belonging to the Dipterocarpaceae, followed by pdao (Calamus viminalis Willd.), a rattan used to make furniture for sale and local use. The Prey Lang area was also reported to be important for medicinal plants, wild edible plants, other kinds of NTFPs, wild animals and timber. Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Fig. 4 Deciduous forest. Stung Treng Province, September 2014 (© Nerea Turreira-García). Fig. 7 Evergreen swamp forest. Stung Treng Province, December 2016. (© Nerea Turreira-García). Fig. 5 Sralao’ (Lagerstroemia sp.) forest. Preah Vihear Province, April 2015 (© Nerea Turreira-García). Fig. 6 Short semi-evergreen forest and evergreen dipterocarp forest. Preah Vihear Province, December 2016 (© Nerea Turreira-García). Cambodian Journal of Natural History 2017 (1) 76–101 Fig. 8 Spirit house near Phneak Roluek. Preah Vihear, September 2014 (© Nerea Turreira-García). © Centre for Biodiversity Conservation, Phnom Penh 83 84 N. Turreira-García et al. Folk taxa Our free-listing exercises and plant collections yielded 374 folk taxa, 337 (90%) of which were collected and five photographed. Of the 337 folk taxa collected, eight were identified to family, 31 to genus and 288 to species (Appendix 1). Ten were not identified to species level. Thirty-two plants were not collected or photographed, either because they were locally extinct, occurred too far away or because (in two cases) our local plant collectors did not know them. Informants claimed to use at least 11 species of fungi, of which four belong to the Basidiomycota phyla. These are not considered further in our analysis. The folk taxa recorded belonged to 83 families and the families most frequently listed were Leguminosae (10%), Rubiaceae (8%), Annonaceae (4%), Apocynaceae (4%), Malvaceae (4%), and Dipterocarpaceae (3%). Species known by most informants included highly valuable timber species such as Hopea odorata Roxb. (korki), Afzelia xylocarpa (Kurz) Craib (beng), Heritiera javanica (Blume) Kosterm. (doungchem), Dalbergia oliveri Prain (neanghoun), Pterocarpus macrocarpus Kurz (thnong), Shorea roxburghii G. Don (porpael), Sindora siamensis Miq. (korkoh), and Terminalia mucronata Craib & Hutch. (bramdomleng); Lagerstroemia speciosa (L.) Pers. (kraol), a medicinal plant with abundant and flashy purple flowers at the time of the collection; several resin-yielding species including Dipterocarpus alatus Roxb. & G.Don (chhertheal) and D. intricatus Dyer (trach); and finally, edible species and species with medicinal properties: Azadirachta indica A. Juss. (sdao), Hymenodictyon orixense (Roxb.) Mabb. (aolaok), and Syzygium zeylanicum (L.) DC. (smarch). Most of the plants used were trees and shrubs (70%), followed by vines, including woody and non-woody lianas and climbers (24%), although herbaceous plants (5%) and palms (1%) were also registered. A total of 630 uses were recorded for the 374 folk taxa (Fig. 9) and each taxon had 2 ±1 (mean ± SD) uses on average. Most were used for medicine (n=249, 67%), food (n=165, 44%) or as material (n=138, 37%), especially for construction of houses, fences and huts. Most medicinal folk taxa were used for a single ailment (51% of all medicinal folk taxa), 32% for two ailments, and 17% for more than two ailments. For instance, Lagerstroemia speciosa was reported to cure seven different illnesses. Almost 30% of the medicinal folk taxa were used to treat postpartum ailments, usually to stimulate appetite, milk production, blood circulation or uterus contraction. This was followed by plants that cured fever (20%), skin problems (17%) and stomach problems (10%). Informants often agreed on the uses of folk taxa, although they sometimes used the same taxon for different ailments. For example, women from Phneak Roluek Village usually grind the leaves of Drynaria sparsisora (Desv.) T. Moore for boils, whereas men from Spong Village claimed that chewing the root of this plant cured urine infection. In addition, different parts of the same folk taxon were sometimes used for the same ailment. In Spong for example, the bark of Terminalia mucronata is Fig. 9 Percentage of folk taxa (n=374) per plant-use category in Prey Lang, Cambodia. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer boiled to treat diarrhoea, whereas women from Phneak Roluek Village boil the root for the same purpose. Men from Thmea Village claimed that the bark of Ficus benjamina should be boiled for skin infections, while men from Spong and women from Phneak Roluek prepare cold infusions of the root and/or leaves for the same purpose. Men often knew the medicinal plants for postpartum ailments, but seldom knew their specific uses. The plant collectors from all three villages explained that men often collect the these plants for their wives and so recognize them, but that women usually prepare the medicines. Women consequently provided more information on the preparation of medicinal plants for postpartum ailments, although they did not always agree on what the postpartum plants were specifically used for. For example, women from Phneak Roluek boiled or made a tincture from the bark of Hymenocardia punctata Wall. ex Lindl. to improve postpartum blood circulation, whereas women from Spong boiled the root of the species to stimulate appetite, milk production and postpartum health. Men collected and free-listed 237 species and women 235 species in total. Men knew 65 folk taxa that women did not free-list or collect, and women knew 81 folk taxa that men did not. Of the folk taxa known only to men, 47 were medicinal (19% of all medicinal folk taxa), 20 were materials (14%) and 12 were food (7%). Of the folk taxa only known to women, 46 were medicinal (18%, of all medicinal folk taxa), 40 were food (24%) and 18 were materials (13%). Informal conversations with the plant collectors on the differences between plants known and used by women and men revealed that they do not perceive knowledge as being influenced by gender. In their view, both men and women know the same plants. Forest and village spirits Informants explained during the plant collection that many spirits occur in the Prey Lang forest. Multiple forest spirits or village spirits exist, such that each community takes care of a particular forest-spirit, or group of spirits, and sometimes different communities take care of the same spirit. In addition, some trees have their own spirit. For example, when Hopea odorata (korki) and Dipterocarpus alatus (chhertheal) are large, these trees are inhabited by a spirit. Other large trees that possess their own spirit include Irvingia malayana Oliv. ex A.W.Benn. (chombork), Sindora siamensis (korkoh), Lagerstroemia calyculata Kurz (sralao’) and all resin-yielding trees. Ficus pubilimba Merr. (chhrey) trees also have a spirit, irrespective of size. Spirit trees are not supposed to be cut, and villagers must ask permission from the spirit if they wish to do so. In general, people pray to the forest-spirit of the area in Cambodian Journal of Natural History 2017 (1) 76–101 spirit houses (Fig. 8) and sacred sites before entering the forest. In their prayers they ask for permission to take its natural resources, and believe that if they fail to do so, the spirit will take revenge and harm them. They also make an offering to spirits before eating or drinking rice wine. Some people reported being angry at the spirits because they do not harm illegal loggers and companies that clearcut forest areas. However, they continue to praise the spirits out of respect (and possibly also fear). A given spirit can either be male or female. The male spirit is usually called neak ta or lok ta, and the female spirit yeay in Khmer and yeak in Kuy. These names change according to the community. The culture of respect for the spirits is passed on through the generations. The forest and tree spirits can also have family members such as parents, spouse and/or children. Conservation status Thirty-five of the species recorded have been assessed by IUCN (2017) and a quarter of these belong to the Dipterocarpaceae, notably Shorea guiso Blume (chorchong, Critically Endangered), Dipterocarpus alatus (chhertheal, Endangered), Shorea roxburghii (porpael, Endangered) and Anisoptera costata Korth. (pdeak, Endangered). The dipterocarps are used for resin extraction and construction. Pinus merkusii Jungh. & de Vriese (srorl, Vulnerable) is also used for resin tapping. Some of the luxury wood species are also globally threatened, such as Afzelia xylocarpa (beng, Endangered), Dalbergia oliveri (neanghoun, Endangered), Hopea odorata (korki, Vulnerable), although Sindora siamensis is not (korkoh, Least Concern). A number of plant species used for food and medicine also occur on the IUCN list (though not necessarily in a threatened category), including Curcuma sparganiifolia Gagnep. (kra chork anderk, Near Threatened), Aglaia edulis (Roxb.) Wall. (bang kau, Lower Risk/Near Threatened) and Irvingia malayana Oliv. ex A.W.Benn. (chombork, Least Concern), as do species used for black magic such as Xylopia pierrei Hance (kray sor, Vulnerable). Disc ussion Prey Lang is a mosaic of forest types (McDonald, 2004; Theilade et al., 2011) and its inhabitants are tightly linked to this area culturally, spiritually and economically. This forest-dependency has created a great body of ethnobotanical knowledge. The study participants, who were mainly middle-aged and older people, demonstrated extensive knowledge of useful flora in Prey Lang. Participants explained that some young people know less about the forest and do not show interest in such knowledge. © Centre for Biodiversity Conservation, Phnom Penh 85 86 N. Turreira-García et al. The youth would need time to accumulate ethnobotanical knowledge if they were interested to do so, if the resources were still available, and if their socio-political conditions were unchanged when they became adults (Reyes-García et al., 2013). The congruence between local and scientific forest classifications in our study supports the notion that local people can play a role in classification of forest types (Halme & Bodmer, 2007) and ecological conservation and research (Janzen, 2004). Most of the ethnobotanical terminology used by the participants was in Khmer, which suggests that use of the Kuy language for plantrelated matters may be vanishing. As noted previously, the Kuy culture has largely been assimilated into Khmer culture in Cambodia (Swift, 2013). Study participants also reported that many children were separated from their parents during the Khmer Rouge (1963−1997) and lost the ability to speak Kuy. This contrasts with the culture of forest knowledge and respect for spirits, which has clearly survived. The ethnobotanical knowledge of the inhabitants of Prey Lang encompasses mainly trees and shrubs, which may reflect the abundance and distribution of vegetation here. Most of the plants used were used for medicine, food and construction, similar to patterns of plant use by Kuy healers in Thailand (Virapongse, 2006). Compared with other studies in similar vegetation in Indochina, the numbers of medicinal plants used in Prey Lang (n=249) were similar or greater than those reportedly used by the Lao (n=250; Elkington et al., 2013), Dao (n=200; Van On et al., 2001), Akha (n=121; Anderson, 1986a) and Lahu (n=68; Anderson, 1986b) ethnic groups. Somewhat higher figures have been reported for Kuy healers (n=333; Virapongse, 2006) and the Karen ethnic group in Thailand, however (n=379; Tangjitman et al., 2013), possibly due to greater survey coverage or because these studies included more cultivated species. The Kuy people also appear to know more wild edible plants (n=165) than the Isaan (n=87; Cruz-Garcia & Price, 2011) and Karen (n=47; Johnson & Grivetti, 2002) ethnic groups in Thailand. Previous studies suggest postpartum ailments are the most frequent conditions treated with medicinal plants by the Kuy in Cambodia (Grape et al., 2016). On revisiting the research sites of Grape et al. (2016), we found 11 new plants used for postpartum ailments, which suggests that potential remains to find additional useful plants in Prey Lang. This contrasts with other studies that have found that fever and digestive problems are the most frequently treated ailments in the region (Virapongse 2006; Tangjitman et al., 2013; Elkington et al., 2014; Neamsuvan et al., 2015) and world-wide (e.g., Hanazaki et al., 2000; Casagrande, 2002; Ayodele, 2005; Liu et al., 2009). © Centre for Biodiversity Conservation, Phnom Penh Resin trees, the main source of income to local households (Jiao et al., 2015; Hüls Dyrmose et al., in press), were ranked in our study as the most important resources of Prey Lang, together with trees used for construction. Many of these trees were also considered spirit trees and thus constitute a strong bio-cultural and economic connection to the forest. Unfortunately, many of these trees are also luxury timber trees which have been logged illegally for decades, and are now endangered locally and globally. Illegal logging consequently threatens the bio-cultural life of the Kuy and Khmer people at Prey Lang. Other studies have found gender-based differences in ethnobotanical knowledge across most use-categories (Nesheim et al., 2006; Araújo & Lopes, 2011; Müller et al., 2014). These are usually represented as differences in number of species known, and/or that men and women know different species because of their different roles in society. Our results suggest the reverse: that many plants are known by both men and women but their use is gendered (i.e., men collect the species whereas women oversee their use). Conversations with plant collectors on the differences between plants known and used by women and men revealed that they did not perceive plant knowledge as gendered: in their view, men and women know the same plants. Further studies are consequently warranted to determine if gender-specific plant knowledge exists in Prey Lang or not. The participatory nature of our study encouraged local people to gain ownership of the research. As it was made clear from the onset that the results would be shared with the communities in the form of an ethnobotanical book, this motivatived study participants to extensively collect useful plants and explain their uses in detail. The plant collectors also felt that a book might motivate younger generations to take interest in the subject, and subsequently pass on their knowledge to future generations. The plant list generated in this study was used to create a database to support community-based biodiversity monitoring and our study demonstrates that local experts can effectively contribute to forest categorisation and voucher specimen collection. As indigenous knowledge is constantly changing, being produced as well as reproduced, discovered as well as lost (Ellen et al., 2000) and is also site-specific (Mutchnick & McCarthy, 1997), we acknowledge that additional plants may have been used in the past or in other regions of Prey Lang. Nevertheless, this study serves as an indicator of the biocultural diversity and importance of Prey Lang and it points to the need to conserve this ecosystem to sustain the livelihoods of its inhabitants. Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Ack now le dge m e nt s The authors are grateful to the inhabitants of Prey Lang who shared their time and knowledge and to the Prey Lang Community Network for their brave work protecting the forest. Thanks are due to University of Copenhagen, Danmission, Oticon Fonden and O.H.F og A.J.-E Heilmanns Fond for financing the research. Special thanks go to Narith Nou for helping with the logistics and to Victoria Helene Grape and Anne-Mette Hüls Dyrmose for their assistance during the fieldwork. We also wish to thank our interpreters Vathana, Sokhan, Kim, Vuthy, Raksmey and Sokchea for their work. 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About t he Aut hors NEREA TURREIRA-GARCÍA is a PhD student at the University of Copenhagen, Denmark. She studied environmental sciences at the University of the Basque Country (Spain) and forest and nature management at the University of Copenhagen. Her main line of research concerns local ecological knowledge and forest monitoring and she has worked in Cambodia, Vietnam, Guatemala, Spain and the Netherlands. DIMITRIOS ARGYRIOU is a consultant and data manager in the project “It’s our forest too”, concerning Prey Lang forest in Cambodia. He studied Agricultural engineering in Democritus University of Thrace, Greece and an MSc in Food Safety in Wageningen University, Netherlands. He has collaborated with research projects in Cambodia, Guatemala and the Netherlands. PHOURIN CHHANG is the Deputy Director of the Forest and Wildlife Research Institute in Phnom Penh, Cambodia. He has extensive experience of botanical surveys in the central lowlands of Cambodia and Bokor National Park. PRACHAYA SRISANGA is a curator at Queen Sirikit Botanic Garden Herbarium, Chiang Mai, Thailand. His main research is on species diversity of plants in mainland Southeast Asia (Laos, Myanmar and Thailand), especially within the Juglandaceae and Violaceae. He also has an interest in ethnobotanical studies of ethnic groups in Southeast Asia. IDA THEILADE is a senior researcher at the University of Copenhagen. Her current research concerns community monitoring of forest biodiversity, carbon stocks and resources, and the role of local knowledge and institutions in conservation and management of tropical forests. © Centre for Biodiversity Conservation, Phnom Penh 89 90 N. Turreira-García et al. Appe ndix 1 I nfor m at ion on spe c ie s fre e -list e d a nd c olle c t e d in nor t hw e st e r n Pre y La ng, Ca m bodia . Use categories: C = Commerce, CDD = Cultural diseases and disorders, E = Environmental, F = Food, FA = Food additives, Fu = Fuel, Ma = Materials, M = Medicinal, R = Resin, RMU = Ritual/Magical Uses, S = Social, TV = Toxic to vertebrates. Ethnospecies names in italics are in Kuy, otherwise Khmer. Vouchers are deposited at Queen Sirikit Botanic Garden, Chiang Mai, Thailand. Scientific name Family Ethnospecies name Life form Voucher No. Use(s) Acacia harmandiana (Pierre) Gagnep. Leguminosae Thmea Tree 76, 387PR, 904 M, Ma, CDD Acacia pennata (L.) Willd. Leguminosae Mchoo Som Bour Shrub 948 F Acacia pennata subsp. insuavis (Lace) I.C. Nielsen Leguminosae Vor Em Vine 580, 585 M, F Acacia sp. Leguminosae Vor Torleng Vine 92 M, TV Acronychia pedunculata (L.) Miq. Rutaceae Tromel Tree 480 M Afzelia xylocarpa (Kurz) Craib Leguminosae Beng Tree 227, 669, 676, 35, 43 M, Ma, F Aganonerion polymorphum Spire Apocynaceae Vor Tneng Vine 347 F Aglaia edulis (Roxb.) Wall. Meliaceae Bang Kau Tree 921 F Aglaia lawii (Wight) C.J. Saldanha Meliaceae Bang Kau Sva Tree 222, 664 M, F Albizia lebbeck (L.) Benth. Leguminosae Chres Tree 78 F, Ma Allophylus cobbe (L.) Raeusch. Sapindaceae Sleuk Bei Shrub 775 M Alpinia galanga (L.) Willd. Zingiberaceae Rom Deng (Prey) Herb 197, 246 M, F, Ma Amaranthus spinosus L. Amaranthaceae Ptebanla Herb 243 M, F Amorphophallus sp. Araceae Teal Shrub 934 M, F Amphineurion marginatum (Roxb.) D.J.Middleton Apocynaceae Sralao' Ompae Vine 515, 488, 46 M Ancistrocladus tectorius (Lour.) Merr. Ancistrocladaceae Khanma, Ktong Vine 147, 132, 651 M, Ma Anisoptera costata Korth. Dipterocarpaceae Pdeak Tree 193, 645, 668 Ma Anisoptera sp. Dipterocarpaceae Stearng Tree 196, 639, 666 Ma, R Antidesma ghaesembilla Gaertn. Euphorbiaceae Dongkeabkdam Tree 278, 489, 462 M, F, RMU, Fu Antidesma japonicum Siebold & Zucc. Euphorbiaceae Trormouch, Mchoo Trormouch Shrub 165, 172, 483, 915 M, F, RMU Aporosa ficifolia Baill. Phyllanthaceae Krong Tree 413, 526, 759 M Aporosa planchoniana Baill. ex Müll. Arg. Phyllanthaceae Propech Chongva Tree 565, 570 M Ardisia crenata Sims Primulaceae Kandetmean Shrub 158.2, 581, 574 M, F Areca triandra Roxb. ex Buch.-Ham. Arecaceae Chnarb Palmlike 463, 426 S, Ma, F Argyreia mollis (Burm. f.) Choisy Convolvulaceae Vor Tror Jeark Tun Sai Vine 527 Ma Artocarpus chama Buch.-Ham. Moraceae Knorprey Tree 266, 289 M, Ma, F Artocarpus nitidus subsp. lingnanensis Moraceae (Merr.) F.M.Jarrett Sombour Tree 359, 473, 690 M, S, Ma, F Azadirachta indica A. Juss. Meliaceae Sdao Tree 315, 286, 667, 678, 83 M, F, Ma Baccaurea ramiflora Lour. Phyllanthaceae Pnheav Tree 213, 118, 613, 618 F © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Appe ndix 1 Cont ’d Scientific name Family Ethnospecies name Life form Voucher No. Use(s) Baeckea frutescens L. Myrtaceae Mrichtonsay Tree 637, 670 M, F Barringtonia acutangula (L.) Gaertn. Lecythidaceae Reang Tree 99, 661, 624 M, F, Ma, TV Bauhinia bracteata (Benth.) Baker Leguminosae Jerngkow, Klaenpor Tree 603, 552, 510, 123 M, S, Ma Bauhinia hirsuta Weinm. Leguminosae Cheungkhu Tree 327, 174 M, F Beaumontia murtonii Craib Apocynaceae Vor Thlork Vine 785, 999 Ma Berrya mollis Wall. ex Kurz Malvaceae Sor Seurm, Trorserm Tree 373PR, 36, 907 M, Ma, Fu Blumea balsamifera (L.) DC. Compositae Baymart Shrub 457 M Bombax anceps Pierre Malvaceae Rorkar Tree 323, 302, 435, 608, 31 M, Ma Breynia vitis-idaea (Burm.f.) C.E.C. Fisch. Phyllanthaceae Muntrei, Miat Kar Vine 837 M, F Bridelia ovata Decne. Euphorbiaceae Pnektrey Tree 2 F Bridelia sp. Phyllanthaceae Chhlikpork Tree 62 M Brucea javanica (L.) Merr. Simaroubaceae Bromatmunus, Damley Smang Shrub 333, 38 M Buchanania cochinchinensis (Lour.) M.R. Almeida Anacardiaceae Laingchey, Romchey Tree 433, 450 M, F Butea superba Roxb. Leguminosae Vor Char Vine 326 M, Ma Caesalpinia digyna Rottler Leguminosae Vor Kvav Vine 912 M Caesalpinia sappan L. Leguminosae Kvav Banla Tree 6 M Calamus palustris Griff. Arecaceae Pdao Chvang Vine 229, 228, 13 M, Ma, F Calamus rudentum Lour. Arecaceae Vor Dombong Vine 320, 951 F, C, Ma Calamus tetradactylus Hance Arecaceae Vor Seung Vine 198 Ma, C, F Calamus viminalis Willd. Arecaceae Chongpdao, Pdao Vine 111, 455, 789, 21 M, Ma, F Calophyllum calaba var. bracteatum (Wight) P.F.Stevens Clusiaceae Paong Tree 395, 390, 199 Ma, F, Ma Cananga latifolia (Hook.f. & Thomson) Finet & Gagnep. Annonaceae Chkaesraeng Tree 295, 308, 595, 592, 39, 66 M Capparis micracantha DC. Capparaceae Kounh Chur Beay Dach Shrub 152 M Careya arborea Roxb. Lecythidaceae Kondaul Tree 385PR, 379SP, 492, 49 M, Ma Caryota mitis Lour. Arecaceae Tunsae, Ansae, Chongsae Tree 139, 116, 497, 620 M, Ma, F Cassia javanica L. Leguminosae Kal Tree 445 S Cassytha filiformis L. Lauraceae Vor Rom saysork Vine 449 M Catunaregam tomentosa (Blume ex DC.) Tirveng. Rubiaceae Rorveang, Rveang Sor Tree 382, 572 M, S Ceiba pentandra (L.) Gaertn. Malvaceae Kor Tree 698 F, Ma Celastrus sp. Celastraceae Vor Kolab Vine 622 M Chionanthus ramiflorus Roxb. Oleaceae Spet, Marey Tree 547, 505, 476 M, S Chionanthus sp. Oleaceae Archdaek Tree 7 Ma Cambodian Journal of Natural History 2017 (1) 76–101 © Centre for Biodiversity Conservation, Phnom Penh 91 92 N. Turreira-García et al. Appe ndix 1 Cont ’d Ethnospecies name Life form Voucher No. Use(s) Compositae Pka’Sor, Kon Traeng Kaet Shrub 900 M, E, F Cinnamomum bejolghota (Buch.Ham.) Sweet Lauraceae Teppiroo Tree 179 M Cinnamomum cambodianum Lecomte Lauraceae Tepproo Tree 821 M Cinnamomum polyadelphum (Lour.) Kosterm. Lauraceae Slapok Tree 101, 423, 563, 536 M, S Citrus lucida (Scheff.) Mabb. Rutaceae Kror Sang Tree 818 Clausena excavata Burm. f. Rutaceae Kanhchrok Shrub 241, 96, 484, 459 CDD, RMU Cleistanthus sp. Phyllanthaceae Neang Leav Tree 762 M, F, Ma Scientific name Family Chromolaena odorata (L.) R.M.King & H.Rob. Colona auriculata (Desf.) Craib Malvaceae Preal Shrub 437, 652, 168 M, Fu, Ma Colona sp. Malvaceae Tangek Tree 309 Ma Combretum latifolium Blume Combretaceae Vor Rormeat Vine 647, 314 M, F Combretum micranthum G. Don Combretaceae Vor Khnos Vine 230, 516, 471 M, F Combretum quadrangulare Kurz Combretaceae Sangkae Tree 363, 589 M, Ma, Fu Connarus cochinchinensis (Baill.) Pierre Connaraceae Vor Lompoh Vine 521, 496, 825 M Coptosapelta flavescens Korth. Rubiaceae Vor Tonling Plerng Vine 235 M Costus speciosus (J.Koenig) C.D.Specht Costaceae Tar Thok Herb 812 M, F Cratoxylum formosum (Jacq.) Benth. & Hook.f. ex Dyer Hypericaceae Lngeang Tree 45, 159 M, F, Ma, Fu Crotalaria pallida Aiton Leguminosae Chongkrong Sva Shrub 453, 810 M, F Croton sp. Euphorbiaceae Montek Tree 257, 764 M Curculigo sp. Hypoxidaceae Tnoutley Herb 322 Ma Curcuma alismatifolia Gagnep. Zingiberaceae Chahouy Herb 318, 930 F Curcuma longa L. Zingiberaceae Rormeat Herb 952 M, F, Ma Curcuma sparganiifolia Gagnep. Zingiberaceae Kra Chork Anderk Herb 593, 143, 914 F Cyclea barbata Miers Menispermaceae Vor Phraskrong Vine 905, 195, 44 M, F Daemonorops jenkinsiana (Griff.) Mart. Arecaceae Saom Vine 774 F, Ma Dalbergia cochinchinensis Pierre Leguminosae Krornhong Tree 573, 560, 84 Ma Dalbergia oliveri Prain Leguminosae Neanghoun Tree 290, 551, 502, 53 Ma Dalbergia sp. Leguminosae Vor Chas Vine 916 Dalbergia thorelii Gagnep. Leguminosae Vor Ampil Vine 523, 494 M, Ma Dalbergia lanceolaria subsp. paniculata (Roxb.) Thoth. Leguminosae Snoul Tree 303, 300, 441, 458, 25 M, F, Fu Dasymaschalon macrocalyx Finet & Gagnep. Annonaceae Cheungchab Shrub 110, 479 M, F Dendrolobium lanceolatum (Dunn) Schinedl. Leguminosae Tronoumbangkhuy Shrub 247, 310, 553, 460 M, F, RMU Dialium cochinchinense Pierre Leguminosae Vor Kralarnh Vine 770 Ma © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Appe ndix 1 Cont ’d Family Ethnospecies name Dianella ensifolia (L.) DC. Xanthorrhoeaceae Dillenia hookeri Pierre Dilleniaceae Scientific name Life form Voucher No. Use(s) Kontoykrorper Herb 409 M Ploosbart Shrub 381PR, 87 M, F Dillenia indica L. Dilleniaceae Plou Tree 98, 411 M, F, Ma Dillenia pentagyna Roxb. Dilleniaceae Rovey Tree 75, 82 M, Fu, F Dimocarpus longan Lour. Sapindaceae Meanprey Tree 183, 102, 493, 582, 468, 16 M, F, Fu Dioscorea brevipetiolata Prain & Burkill Dioscoreaceae Domlong Tean Vine 203, 927 F Dioscorea esculenta (Lour.) Burkill Dioscoreaceae Domlong Shar Vine 935 F Dioscorea pentaphylla L. Dioscoreaceae Vor Dom Loung Teuk Vine 800 F Dioscorea poilanei Prain & Burkill Dioscoreaceae Domlong Kour Vine 926 F Dioscorea polyclados Hook. f. Dioscoreaceae Domlong Romeat Vine 950 F Diospyros ehretioides Wall. ex G. Don Ebenaceae Mormeang Tree 307, 288 M, TV Diospyros filipendula Pierre ex Lecomte Ebenaceae Ambengprah Tree 769, 917, 58 M, Ma, F, Fu Diospyros lobata Lour. Ebenaceae Chherkmao Tree 56 Ma, Fu Diospyros pendula Hasselt ex Hassk. Ebenaceae Khchas Tree 910 F, Ma Diospyros sp. Ebenaceae Chaas, Ches Tree 906 F, Fu Diospyros sylvatica Roxb. Ebenaceae Khanhchas, Krorchas Tree 100, 814 M, Fu, F, Ma Diospyros undulata Wall. ex G. Don var. cratericalyx (Craib) Bakh. Ebenaceae Chi Plerng Tree 287, 561, 422, 771 TV, F Diospyros venosa Wall. ex A.DC. Ebenaceae Chherkmao II Tree 415, 520 Ma, Fu Dipterocarpus alatus Roxb. & G.Don Dipterocarpaceae Chhertheal Tree 107, 164, 621, 456 M, R, Ma Dipterocarpus intricatus Dyer Dipterocarpaceae Trach Tree 217, 208, 375SP, 376, 29 M, R, Ma Dipterocarpus obtusifolius Teijsm. ex Miq. Dipterocarpaceae Tbaeng Tree 383PR, 47 M, Ma Dipterocarpus tuberculatus Roxb. Dipterocarpaceae Khlong Tree 50 Ma Dischidia major (Vahl) Merr. Apocynaceae Vor Bampong sromouch Vine 569 M Donax canniformis (G. Forst.) K.Schum Marantaceae Ron Herb 777, 623, 627 M, Ma Dracaena elliptica Thunb. & Dalm. Asparagaceae Tbaldaek Shrub 542 M Dracaena angustifolia (Medik.) Roxb. Asparagaceae Angraedaek Shrub 482, 188, 173, 501 M, F, Ma Drynaria sparsisora (Desv.) T. Moore Polypodiaceae Borbrok Herb 270 M Elephantopus scaber L. Compositae Chen Veal Herb 758 F Ellipanthus tomentosus Kurz Connaraceae Kdor Komprok Shrub 267 M, F Entada rheedii Spreng. Leguminosae Vor Ang Kunh Vine 774 M, Ma Erythrophleum teysmannii (Kurz) Craib Leguminosae Kreul Tree 932 Ma Cambodian Journal of Natural History 2017 (1) 76–101 © Centre for Biodiversity Conservation, Phnom Penh 93 94 N. Turreira-García et al. Appe ndix 1 Cont ’d Ethnospecies name Life form Voucher No. Use(s) Erythroxylaceae Chompussek, Changkung sek Shrub 412, 406 M Euonymus cochinchinensis Pierre Celastraceae Koomouy Tree 519, 448 M Eurycoma longifolia Jack Simaroubaceae Angtongsor Shrub 321, 316, 567, 388 M, S Scientific name Family Erythroxylum cambodianum Pierre Fagraea fragrans Roxb. Gentianaceae Tatrav Tree Photo Ma Fagraea racemosa Jack Gentianaceae Changka Trong Tree 786 Ma Ficus annulata Blume Moraceae Chrey Vor, Vor Chrey Vine 248 M Ficus benjamina L. Moraceae Chhreykruem Tree 686, 64 M Ficus callophylla Blume Moraceae Chrey Klaok Tree 628 M Ficus hirta Vahl Moraceae Lavadey Tree 533 F Ficus hispida L.f. Moraceae Roveadey Tree 242 M Ficus pubilimba Merr. Moraceae Chhrey Tree 537 F, Ma Ficus pumila var. awkeotsang (Makino) Corner Moraceae Vor Krorbeytraos Vine 830, 260 M Ficus racemosa L. Moraceae Lovear Tree 665, 674, 251 M, F Firmiana simplex (L.) W.Wight Malvaceae Samroung Tree 325 Ma Flacourtia indica (Burm.f.) Merr. Salicaceae Krorkob (Prey) Tree 607, 15 M, F Garcinia celebica L. Clusiaceae Proos Tree 129, 451, 9, 442 Ma, F Garcinia cochinchinensis (Lour.) Choisy Clusiaceae Mchhoosandan, Sandan Tree 834, 692 F Garcinia merguensis Wight Clusiaceae Kres, Yeam Tree 578, 220 M, F, S Garcinia oliveri Pierre Clusiaceae Trormoong, Mchoo Trormoong, Tronoumseik, Tromongchea Tree 421, 24 M, F, Fu Garcinia vilersiana Pierre Clusiaceae Prorhoot Tree 633, 656, 10 Ma, F Gardenia angkorensis Pit. Rubiaceae Daiklar Tree 375PR M, C Gardenia sootepensis Hutch. Rubiaceae Barkdong Tree 293 M, F, Ma Garuga sp. Burseraceae Sdavkhmoch Tree 5 Ma Getonia floribunda Roxb. Combretaceae Kor Nhours Vine 813 M Glochidion kerrii Craib Phyllanthaceae Sesach Tree 486 M Gmelina asiatica L. Lamiaceae Anhcharnh Tree 93, 507 M Gnetum montanum Markgr. Gnetaceae Khlout Vine 233, 124, 465, 658 F, Ma Gomphia serrata (Gaertn.) Kanis Ochnaceae Pesles Tree 175, 112, 391, 380 M Goniothalamus repevensis Pierre ex Fin. & Gagnep. Annonaceae Vor Krovan Vine 138 M, Ma Goniothalamus tamirensis Pierre ex Finet & Gagnep. Annonaceae Moom Shrub 629 M, TV Grewia sp. Malvaceae Jeay moa Tree 336 M Haldina cordifolia (Roxb.) Rids. Rubiaceae Kvav Tree 606 M, Ma © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Appe ndix 1 Cont ’d Ethnospecies name Life form Simaroubaceae Klentea Vine 254, 598,793 M, F Rubiaceae Slabbrang Tree 838 M Helicteres hirsuta Lour. Malvaceae Phrealphnom, Preal Momis Shrub 40, 454 M Helicteres sp. Malvaceae Neang Moa Shrub 753 M Scientific name Family Harrisonia perforata (Blanco) Merr. Hedyotis sp. Voucher No. Use(s) Heliotropium indicum L. Boraginaceae Bromony Domrey Herb 158, 68 M Heritiera javanica (Blume) Kosterm. Malvaceae Doungchem Tree 268, 625, 648, 700 Ma Holarrhena curtisii King & Gamble Apocynaceae Tekdors, Vor Chhuy, Tuekdoh Veal Vine 341, 279 M Hopea odorata Roxb. Dipterocarpaceae Korki Tree 640, 14 Ma Hoya sp. Apocynaceae Vor Krobay Vine 416 E Hydnocarpus anthelminthicus Pierre ex Laness. Achariaceae Krorbao Tree 335, 642 M, F Hydnocarpus ilicifolia King (unresolved name) Achariaceae Chambokkaek Tree 922 Ma Hymenocardia punctata Wall. ex Lindl. Phyllanthaceae Komkhneang Tree 185, 120, 619, 614 M, F, Fu Ovlok Tree 299, 284, 587, 77, 72 M, F M, Ma Hymenodictyon orixense (Roxb.) Mabb. Rubiaceae Imperata cylindrica (L.) Raeusch. Poaceae Sbaupleang Herb 600, 778 Indigofera tinctoria L. Leguminosae Trom Prey Shrub 121 M Irvingia malayana Oliv. ex A.W.Benn. Irvingiaceae Chombork Tree 161, 671, 474, 22, 170 M, F, Ma, Fu Ixora javanica (Blume) DC. Rubiaceae Pkakroham Shrub 549, 440, 91 M, F Ixora nigricans R.Br. ex Wight & Arn. Rubiaceae Pkamuchol Shrub 87PR M Ixora sp. Rubiaceae Chhongkonghing Shrub 103 M Jasminum scandens (Retz.) Vahl Oleaceae Vor Chuengpoh Vine 485B M Lagerstroemia calyculata Kurz Lythraceae Sralao' Tree 317, 584, 34 M, Ma, F, Fu Lagerstroemia floribunda Jack (unresolved name) Lythraceae Trobekprey Tree 125 M Lagerstroemia ovalifolia Teijsm. & Binn. (unresolved name) Lythraceae Sralao' Trobek Tree 503, 663 F, Ma Lagerstroemia speciosa (L.) Pers. Lythraceae Kraol Tree 345, 330, 447, 500, 634 M, Ma, E Lasianthus hirsutus (Roxb.) Merr. Rubiaceae Skun Shrub 150, 649, 650 M Leea indica (Burm. f.) Merr. Vitaceae Baykdaing, Kandan Bay Shrub 564 M, S Leea thorelii Gagnep. Vitaceae Lounglang Tree 361.2 M Lepisanthes rubiginosa (Roxb.) Leenh. Sapindaceae Chunlous, Tumlos Tree 166, 4, 938, 949 M, F Licuala spinosa Wurmb Arecaceae Paav Palm 903, 169, 146, 397, 399 F, Ma Limnophila geoffrayi Bonati (unresolved name) Plantaginaceae Ma Orm Herb 833 M, F Cambodian Journal of Natural History 2017 (1) 76–101 © Centre for Biodiversity Conservation, Phnom Penh 95 96 N. Turreira-García et al. Appe ndix 1 Cont ’d Scientific name Family Ethnospecies name Life form Voucher No. Use(s) Limnophila sp. Plantaginaceae Bror Mae Herb 836 F Loeseneriella pauciflora (DC.) A.C. Sm. (unresolved name) Celastraceae Vor Angtong Vine 660 M, Ma Lygodium flexuosum (L.) Sw. Lygodiaceae Vor Trom, Vor Ovlor Vine 176, 673, 680, 11, 779 M, Ma, RMU RMU Machilus thunbergii Siebold & Zucc. Lauraceae Yeangboung Shrub 831 Macroptilium atropurpureum (DC.) Urb. Leguminosae Vor Sangdek bangkuoy Vine 791 Madhuca butyrospermoides A.Chev. Sapotaceae Srorkom Tree 475, 428 F, Ma, Fu Mallotus glabriusculus (Kurz) Pax & K.Hoffm. Euphorbiaceae Kansamta oa Shrub 89PR M, F, Ma Mallotus nanus Airy Shaw Euphorbiaceae Konsomthao Tree 576 M Mammea siamensis T.Anderson (unresolved name) Calophyllaceae Sophi Tree 282 TV Mangifera longipetiolata King (unresolved name) Anacardiaceae Svay Prey Tree 274, 472 M, F, Ma Markhamia stipulata (Wall.) Seem. Bignoniaceae Dakpor Tree 137, 216 M, F Melastoma malabathricum L. Melastomataceae Baynhenh Shrub 119, 90, 119B M, F Melastoma saigonense (Kuntze) Merr. Melastomataceae Baynhenh (fem) Shrub 399A M Melastoma sanguineum Sims Melastomataceae Baynhenh (male) Shrub 401, 446, 410 M Melientha suavis Pierre Opiliaceae Prech, Prechprey Tree 945 F, Ma Melodorum fruticosum Lour. Annonaceae Romduol Tree 108, 477, 611 M, F, Fu, Ma Memecylon caeruleum Jack Melastomataceae Phlorng Tree 495, 8,211 Ma Microcos tomentosa Sm. Malvaceae Porplear Tree 225, 407 Ma, F, Fu Mimosa pudica L. Leguminosae Phrasklob Herb 756 M Mischocarpus sp. Sapindaceae Promarksan Shrub 181 M Mitragyna hirsuta Hav. Rubiaceae Ktom, Ktomtom Tree 355, 602 M, Ma Mitragyna speciosa (Korth.) Havil. Rubiaceae Ktumphnom Tree 294 M Momordica cissoides Planch. ex Benth. Cucurbitaceae Vor M’reas Prey Vine 832 F Morinda coreia Buch.-Ham. Rubiaceae Nhio (Prey) Tree 343, 334, 51, 931 M Murraya siamensis Craib (unresolved name) Rutaceae Brohoungarkas Shrub 232, 617, 797 M, RMU Myrialepis paradoxa (Kurz) J. Dransf. Arecaceae Chnuo Vine 283 Ma Myristica iners Blume Myristicaceae Kuok Tree 944 F, Ma Nauclea orientalis (L.) L. Rubiaceae Kdol Tree 601, 594, 513, 632 M, Ma Ochna integerrima (Lour.) Merr. Ochnaceae Angkea Sel Tree 312, 439, 384, 429B, 189A M Ocimum tenuiflorum L. Lamiaceae M’reas Prov Shrub 177 F Ocotea lancifolia (Schott) Mez Lauraceae Krolor Tree 201 M, F Olax scandens Roxb. (unresolved name) Olacaceae Orkktong Vine 511 M, F © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Appe ndix 1 Cont ’d Ethnospecies name Life form Voucher No. Use(s) Rubiaceae Thnungkanhchos, Vor Sneng kropey Shrub 250, 772, 182 M Paederia foetida L. Rubiaceae Vor Phorm Vine 269 F Pandanus humilis Lour. Pandanaceae Romchekprey Screw 171, 467, 530 -pine M, Ma Pandanus sp. Pandanaceae Chak Screw 761 -pine F Parinari anamensis Hance Chrysobalanaceae Thlork Tree 223, 204, 373SP, 374, 80 M, F Peliosanthes teta Andrews Asparagaceae Tbaldaek, Tbaltark Herb 828, 659 M, RMU Peltophorum dasyrrhachis (Miq.) Kurz Leguminosae Trorsek Tree 94, 381SP, 432, 133, 19 Ma, F Scientific name Family Oxyceros horridus Lour. Pentacme siamensis (Miq.) Kurz Dipterocarpaceae Reangphnom Tree 294, 682, 60 Ma Phyllanthus emblica L. Phyllanthaceae Kontoutprey Tree 349, 324, 630, 583 M, F, Fu Phyllodium pulchellum (L) Desv. Leguminosae Kom Prum Bae Kroy Shrub 937 M Physalis angulata L. Solanaceae Pengposprey Herb 271 M, F Pinus merkusii Jungh. & de Vriese Pinaceae Srorl Tree 535, 504 M, R Piper sarmentosum Roxb. Piperaceae Chhiplou Herb 259 F Ploiarium alternifolium (Vahl) Melch. Bonnetiaceae Sreurng Tree 631, 544 Ma M, F, Ma Polyalthia cerasoides (Roxb.) Bedd. Annonaceae Knaydael, Snaydel Tree 329, 615, 57 Polyalthia evecta Finet & Gagnep. (unresolved name) Annonaceae Sanghasbart Tree 920 Premna herbacea Roxb. Lamiaceae Ruschin Shrub 371 M Prismatomeris filamentosa Craib Rubiaceae Romdenhmeas Shrub 210, 402, 189 M Prismatomeris memecyloides Craib Rubiaceae Romdenh Shrub 417 M, F Prismatomeris sessiliflora Pierre ex Pit. Rubiaceae Romdenhmeas II Shrub 55 M Psychotria asiatica L. Rutaceae Sraomdav Shrub 393 M, Ma Psychotria sp. Rubiaceae Slerkreum Shrub 531 M Psychotria sp.1 Rubiaceae Reum Shrub 438 M Psydrax dicoccos Gaertn. Rubiaceae Bongkorng Tree 641 Ma Psydrax pergracilis (Bourd.) Ridsdale Rubiaceae Mekorng Tree 41 Ma Pternandra caerulescens Jack Melastomataceae Changketbrak Tree 559 F Pterocarpus macrocarpus Kurz Leguminosae Thnong Tree 205, 192, 487, 466 M, Ma Rhodamnia dumetorum (DC.) Merr. & L.M.Perry Myrtaceae Plorng (Uol) Shrub 539, 815 F Rhodomyrtus tomentosa (Aiton) Hassk. Myrtaceae Pouch Uol, Trobekprey Shrub 541, 508 F Rinorea anguifera Kuntze (unresolved name) Violaceae Dom Nek Pro Ma Tree 136 M Salacia chinensis L. Celastraceae Pengphorng, Vorveay Vine 32, 400, 543, 313 M, F Cambodian Journal of Natural History 2017 (1) 76–101 © Centre for Biodiversity Conservation, Phnom Penh 97 98 N. Turreira-García et al. Appe ndix 1 Cont ’d Ethnospecies name Life form Voucher No. Use(s) Celastraceae Vor Kondabchongae Vine 256 M, F Salacia typhina Pierre (unresolved name) Celastraceae Kon Darb Jong Ae Vine 924 M, F Sandoricum koetjape (Burm. f.) Merr. Meliaceae Kompinhreach Tree 823 M, Ma, F Scientific name Family Salacia cochinchinensis Lour. Sauropus sp. Phyllanthaceae Thmehntrey Shrub 249 M Schleichera oleosa (Lour.) Merr. Sapindaceae Pongror, Tomroos, Ta Tok Tree 902, 37, 291, 913 M, F, Fu, Ma Scindapsus officinalis (Roxb.) Schott Araceae Vor Chum Vine 272 M Scleropyrum pentandrum (Denn.) Mabb. Santalaceae Rlokkeo, Aolaokkao Tree 529, 524, 141 M Senna alata (L.) Roxb. Leguminosae Donghet Shrub 88, 763 M, F Senna garrettiana (Craib) H.S.Irwin & Barneby Leguminosae Haisan Tree 67 M Shorea guiso Blume Dipterocarpaceae Chorchong, Pchuek Aodom Tree 215, 154, 635, 662, 829 R, Ma Pchek Tree 361.1, 328, 59 M, Ma Tree 219, 296, 377SP, 386, 48 Ma, F Shorea obtusa Wall. ex Bl. (unresolved Dipterocarpaceae name) Shorea roxburghii G. Don Dipterocarpaceae Porpael Sindora siamensis Miq. Leguminosae Korkoh Tree 298, 682, 60 M, Ma, F Smilax lanceifolia Roxb. Smilacaceae Porpreus, Vor Porpeay Vine 130 M Smilax megacarpa A. DC. Smilacaceae Porpreus, Vor Rombers Vine 131, 525, 550, 663V, 672, 817 M, F Smilax sp. Smilacaceae Vor Thnamchin Vine 825 M Spatholobus acuminatus Benth. Leguminosae Vor Tar Arn Vine 236, 942 M, Ma, F Spirolobium cambodianum Baill. Apocynaceae Chhertheal trang (young), Preay Kbalbromboy (old) Tree 644, 827, 532 M, CDD, F Spondias pinnata (L. f.) Kurz Anacardiaceae Mkark prey, Phloch Tree 157, 234, 684, 754, 909, 754 M, F, FA, Ma Stemona sp. Stemonaceae Kbeas Shrub 263, 114 M Stenochlaena palustris (Burm. f.) Bedd. Blechnaceae Vor Thnanh Vine 127, 577, 777 M, F, Ma Sterculia sp. Malvaceae Prorlob Tree 688 Ma Streblus asper Lour. Moraceae Snay Tree 609, 604 M Streptocaulon juventas (Lour.) Merr. Apocynaceae Vor Chuy, Vor Joch Vine 339, 509, 396 M Strychnos nux-blanda A.W. Hill Loganiaceae Kompolvek Tree 389PR M Strychnos nux-vomica L. Loganiaceae Sleng Tree 306 M Strychnos polyantha Pierre ex Dop Loganiaceae Vor Sleng Vine 518, 281 M Suregada multiflora (A.Juss.) Baill. Euphorbiaceae Markdaok Tree 490 M, F Syzygium fruticosum DC. Myrtaceae Pring Angkam Tree 953 M, F, Fu Syzygium grande (Wight) Walp. Myrtaceae Pring Som Bork Krars Tree 153, 387SP, 816 M, F © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Appe ndix 1 Cont ’d Scientific name Family Ethnospecies name Life form Voucher No. Use(s) Syzygium siamense (Craib) Chantaran. & J.Parn. Myrtaceae Pring Kbal Nakta Tree 943 M, Ma Syzygium sp. Myrtaceae Smarch Tuk Tree 557 F Syzygium syzygioides (Miq.) Merr. & L.M.Perry Myrtaceae Pring Bay Tree 81, 811 M, Ma, F Syzygium zeylanicum (L.) DC. Myrtaceae Smarch Tree 163, 190, 419, 404, 30 M, F, Ma, Fu Tabernaemontana bufalina Lour. Apocynaceae Matesprey Tree 534, 766 M Tadehagi triquetrum (L.) H.Ohashi Leguminosae Angkrorng, Chang Kes Angkrong Shrub 167, 126, 332 M Tamarindus indica L. Leguminosae Ampil Tree VS2 M Tamilnadia uliginosa (Retz.) Tirv. & Sastre Rubiaceae Rompok Tree 292 M Tarenna hoaensis Pit. Rubiaceae Chantornear Shrub 527 M, Ma Terminalia alata Roth (unresolved name) Combretaceae Chhlik Tree 61 M, Ma Terminalia bialata (Roxb.) Steud. Combretaceae Pealkhe, Porpaelkae Tree 1, 596 M Terminalia chebula Retz. Combretaceae Sramor, Srormor Lau Tree 351, 933 M, F, Ma Terminalia mucronata Craib & Hutch. (unresolved name) Combretaceae Bramdomleng Tree 357, 431, 498, 52 M, Ma Terminalia nigrovenulosa Pierre Combretaceae Bayarm Tree 599, 646, 28 M, F, Ma, Fu Terminalia pierrei Gagnep. (unresolved name) Combretaceae Sev Tree 751 M Tetracera loureiri (Finet & Gagnep.) Pierre ex W.G. Craib Dilleniaceae Vor Dakun Vine 113, 206, 128, 443, 378 M Thunbergia sp. Acanthaceae Vor Dakpor Vine 63 M Thyrsanthera suborbicularis Pierre ex Gagnep. Euphorbiaceae Rus Bong Ki, Vongsa Preahatit Vine 929, 752 M Tiliacora triandra Diels (unresolved name) Menispermaceae Vor Yeav Vine 568 Ma, F, Fu Tinospora crispa (L.) Hook. f. & Thomson Menispermaceae Bondolpich Vine 86 M Tristaniopsis merguensis (Griff.) Peter G.Wilson & J.T.Waterh. Myrtaceae Srorngam Tree 506 Ma Urceola rosea (Hook. & Arn.) Midd. Apocynaceae Mchoo Tneng, Vor Tneng Vine 936 F Uvaria fauveliana Pierre ex Ast (unresolved name) Annonaceae Saomaoprey Vine 186, 575 M, F, Ma Uvaria hahnii (Finet & Gagnep.) J.Sinclair (unresolved name) Annonaceae Songkhouch Vine 261, 258, 548 M, F Uvaria rufa Blume Annonaceae Treal Sva Vine 97, 750 F Uvaria sp. Annonaceae Vor Doskrobey, Vor Treal, Teu Doh Krobai Vine 262, 792 M, F Cambodian Journal of Natural History 2017 (1) 76–101 © Centre for Biodiversity Conservation, Phnom Penh 99 100 N. Turreira-García et al. Appe ndix 1 Cont ’d Scientific name Family Ethnospecies name Life form Voucher No. Use(s) Uvaria littoralis (Blume) Blume Annonaceae Vor Chekprey Vine 765 M, F Vatica odorata (Griff.) Symington Dipterocarpaceae Chrormas Tree 385SP, 558, 27 Ma, F Ventilago cristata Pierre (unresolved name) Rhamnaceae Vor Tonlueng Vine 638 M, Ma Vitex pinnata L. Lamiaceae Porpool Tree 304, 427, 522, 54 M Vitex sp. Lamiaceae Protespray Shrub 224 M, RMU Walsura villosa Wall. ex Hiern Meliaceae Sdok Sdao Tree 928 M Waltheria indica L. Malvaceae Preash Proa Veal Shrub 89 M Willughbeia edulis Roxb. Apocynaceae Koy Vine 155, 389SP, 408 M, F Wrightia arborea (Dennst.) Mabb. Apocynaceae Klengkong Tree 3 M Xanthophyllum colubrinum Gagnep. Polygalaceae Trop Tum Tree 514, 545, 776 F, Ma Xerospermum noronhianum (Blume) Blume Sapindaceae Mean Angkarm, Seman Tree 135, 106, 657, 420, 917 F, Fu Xylia xylocarpa (Roxb.) Taub. Leguminosae Sokrom Tree 301, 280, 591, 17 M, Ma Xylopia pierrei Hance (unresolved name) Annonaceae Kray Sor Tree 212, 403, 394, 911 M, RMU, Ma, Fu Xylopia vielana Pierre Annonaceae Kray Krahorm Tree 901 M, Fu Zanthoxylum nitidum (Roxb.) DC. Rutaceae Preah Kom Jart Tree 605, 276, 586, 760 M, F, CDD Zingiber zerumbet (L.) Roscoe ex Sm. Zingiberaceae Phtue Herb 908 F Ziziphus cambodianus Pierre (unresolved name) Rhamnaceae Vor Angkrong Vine 20, 616 M, S, Ma Ziziphus oenopolia (L.) Mill. Rhamnaceae Vor Sangkher Vine 566, 555, 187, 178, 33 M, F - Acanthaceae Bromatksan Tree 180 M - Apocynaceae Vor Preah Trorheng Vine 696 M - Araceae Vor Prork Vine 767 Ma - Asclepiadaceae Vor Chlous Vine 554 Ma - Leguminosae Sombour II Tree 839 RMU - Primulaceae Vor Preah Samkong Vine 925 M - Rubiaceae Lout Tree 540 Ma - Scrophulariaceae S’mao Kreung Herb 820 F - - Derm Kon Tuy Mian Herb Photo M - - Dermprus Tree - Ma - - K’Cheay Shrub - F - - K’Dourch Vine - F - - Kachdek Tree - Ma - - Khchaeng, Krorcheng Tree Photo Ma, TV, F - - Kom Pong Tro aoh Tree - F - - Korkithmor Tree - Ma - - Kramuon Tree - M, F © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 76–101 Ethnobotanical knowledge of the Kuy and Khmer Appe ndix 1 Cont ’d Scientific name Family Ethnospecies name Life form Voucher No. Use(s) - - Krasaeang Tree - M, F - - Krolanh Tree - M, Ma - - Krorlunch Tree - F - - Lovear dei Shrub Photo M - - Lumpoung Tree - M - - Mermchin - M - - Ploo Tree 923 M, F, Fu - - Pouk Shrub - Ma - - Preah Oproveal Tree - M - - Preah Trorheng Tree - M, RMU, F - - Proteng Herb 379PR M, F, Ma - - Ptheark Tree - Ma - - Ro Ngoung Tree - RMU - - Rodong Tree - M - - Rompukrorhorm Tree - M, Ma, F - - Rumduol Sbart Shrub - M - - Russey Shrub - F, Ma - - Russlar - - M, S - - Sluekprich - - F - - Smarkrorbey Tree - M, F - - Spong Tree - M, Ma - - Sro Kum Bay Tree 819 F - - Svarkhom Tree - M - - Tha’Kao Tree - Fu - - Thnenn Vine 577, 127 F - - Trameng Tree - M - - Treal Var/ Kon Treal Var Vine - F - - Trouyprich Tree - F - - Tuntreankhet Shrub - M, E - - Vor K’morng Vine Photo TV - - Vor Lanchoeung Vine 929 Ma - - Vor Pouh Vien Mean Vine 252 M - - Vor Tasan Vine - Ma Cambodian Journal of Natural History 2017 (1) 76–101 © Centre for Biodiversity Conservation, Phnom Penh 101 102 Eam S. et al. M ove m e nt of c a pt ive -re a re d Sia m e se c roc odile s Croc odylus sia m e nsis re le a se d in t he Sout he r n Ca rda m om N at iona l Pa rk , Ca m bodia EAM Sam Un1,*, SAM Han1,2, HOR Leng1,2, Me’ira MIZRAHI1 & Jackson L. FRECHETTE1 1 Fauna & Flora International - Cambodia Programme, No.19, Street 360, Boeung Keng Kang 1, PO Box 1380, Phnom Penh, Cambodia. 2 Forestry Administration, Phnom Penh, No. 40 Preah Norodom Boulevard, Phsar Kandal 2, Khann Daun Penh, Phnom Penh, Cambodia. * Corresponding authors. Email samun.eam@fauna-flora.org Paper submitted 11 April 2017, revised manuscript accepted 8 May 2017. ɊɮɍɅʂɋɑɳȶſɆ DžƚɆɽɴɁNjɅɌLJɋɄɸɃɮǎɋɳǷǕɑɭɪǕɳȴƒɋɿ ƙȲɳɈˊɉɸ(ƒ Crocodylusȱsiamensis) ɆȷƃɭɆƓɅƒƺƙɆɳɉɃȹɩɁɇɭɁɈɮȹɄƂɅɽɄƂɌɆɸɇɭɁƺ ɑȲɍɳǷȲƒɭȶȷɸɳǁɊɈɈɯȲƙȲɳɈˊDŽɸȶɔɑɽʆ ɳȴɳȹȟƺȲɽǃƙɆɳɃɑȲɊƕɭƺƺȹƙɊȲȼʁɄɸƺȶɳȴɆɸɇɭɁɵɅɆɻɮɈɭɋǔɑƘɭȶɵƙɈɴȼɍɳǷ ɳɑɑɑɍɽɳɅɹ ȼɮɳȷƒɹƳɌɈƙȶɫȶƳɌɔɉɩɌȲƞƙȲɳɈˊɉɸȲ ƒ ɭƒȶƙɆɳɃɑȲɊƕɭƺ ƺƳɌƷɌǕɃɩNJɈɆɸɇɭɁʆ ƳɌɑɩȲǜɳɅɹNjɅɳƵɍɆɸɀȶ ɴɑƛȶɋɍɽɈɪɆɸǎɑɽɃɪɅɩȶƳɌɌɑɽɳǷɌɆɑɽƙȲɳɈˊɉɸƒɴȼɍLJɅɴɍȶɳǵȲƒɭȶɵƙɈɄɊƗƺɁɩ ɴȼɍƺȲɊƗɎ ɩɄɪǃƒȲɽƺɁɩɑɪɈ Ǝ ɪƳɌǒƎɌƙȲɳɈˊɉɸƒ ɳɓˊȶɎ ɩȻʆ ɳǷȲƒɭȶƳɌɑɩȲǜɳɅɹ ƙȲɳɈˊɉɸNj ƒ ɅȷɸɅɯɅʑʕȲǙɍƙɁȪɎLJɅɆɸljȲɽəɆȲɌɀɿɎ ɩɃƘɭDŽȲɽɃȶɅɩȶǂɊƽɅɌɋɺɳɈɍʑʘɴȳ ɆdžƐɆɽɈɪLJɅɴɍȶɳǵȲƒɭȶəɃǚɅƺɁɩȹɯɌɉƒɸƙȲǏȻƴȶɁƓɮȶ ɴɆɻȲɅɩɌɁɪɵɅƙɆɳɃɑȲɊƕɭƺʆ ƙȲɳɈˊɉɸȷ ƒ ɸɅɯɅʑʓȲǙɍLJɅLjƚɑɽɃɪƙɆNjɀ ʗʐʐɊɈɪɃɪȲɴɅƚȶɴɍȶ ȲƒɭȶɔɸɓɭȶɳɈɍǂɊƽɅ (ƳɌǂɊƽɅƙɆƙɈɫɁƎɳǵɳǷɌȼɮɎƙLJɸȶ)ʆ ɍɃƑɇɍɳɅɹ ɆƷƟȻǃǕȲɆƓȲɩɌ ɩnjɌɆɑɽ ƙȲɳɈˊɉɸNJ ƒ ȴɳƙȷˊɅɊɩɅLjƚɑɽɃɪƹƂɋɈɪȲɴɅƚȶɴɍȶɳɃ ɴȼɍƙɁȪɎƵƒɳǵɅɫȶƳɌɑɩȲǜȲɅƚȶɊȲɳɍˊƙȲɳɈˊɉɸƒɄɊƗƺɁɩɳǷȲƒɭȶɁɸɆɅɽȹɯɌɉƒɸƙȲ ǏȻʆ ƳɌɌɑɽɳǷȹɩɁȲɴɅƚȶɴɍȶǕȷƺƙɆɳnjȹɅɿȷɸɳljɹȳƚɯɅǏ ɤƘɈɯȲǏNjɅȹɳNjƚɹƺɊɯɋɊɅɭɑƞʆ ɳƙljɹǏɆɅƏɋɣƳɑȲƒɭȶƳɌȳɩɁȹɩɁƺɋȹƙɊȲɞɁɸɆɅɽɴȼɍɆȶž Abst ra c t Once widely distributed throughout Southeast Asia, the Siamese crocodile Crocodylus siamensis is currently one of the world’s most Critically Endangered crocodilian species. Because Cambodia is home to the largest remaining wild population, conservation efforts within the country should be considered of upmost importance. This study was aimed to understand the movement and survival of captive-reared Siamese crocodiles released as a part of a national reintroduction and reinforcement programme. In the study, 15 juvenile and sub-adult crocodiles fitted with VHF radio transmitters were monitored for up to 18 months after release in the Southern Cardamom National Park, southwestern Cambodia. Thirteen of the crocodiles were detected within 700 m of the release site during monitoring, which occurred mostly during the dry season. Their sedentary behaviour was consistent with previous studies of young Siamese crocodiles in the Cardamom Mountains. Remaining close to release sites may be beneficial for crocodiles by reducing chances of their moving to more marginal habitats or areas where conflict could potentially occur with people. Ke yw ords Movement patterns, radio-tracking, reintroduction, Siamese crocodile. CITATION: Eam S.U., Sam H., Hor L., Mizrahi, M. & Frechette, J.L. (2017) Movement of captive-reared Siamese crocodiles Crocodylus siamensis released in the Southern Cardamom National Park, Cambodia. Cambodian Journal of Natural History, 2017, 102–108. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 102–108 Movement of Siamese crocodiles I nt roduc t ion The Siamese crocodile Crocodylus siamensis is considered one of the world’s most Critically Endangered crocodilian species (IUCN, 2015), and is one of the least studied in the wild. The species was once widely distributed throughout Southeast Asia, specifically in Indonesia, Malaysia, Thailand, Laos, Cambodia, Vietnam and possibly Myanmar and Brunei (Ross, 1998). By the early 1990s, Siamese crocodile was reported to be effectively extinct in the wild, but scattered populations have since been discovered in Cambodia, Laos and Indonesia (Kalimantan) (Daltry & Chheang, 2000; Kurniati et al., 2005; Platt et al., 2006; Simpson et al., 2006). Cambodia has the largest remaining wild population, but this is widely fragmented across the country and very few nests are recorded annually (Bezuijen et al., 2006; Sam et al., 2015). Populations of the Siamese crocodile have been greatly depleted and fragmented in Cambodia by illegal poaching for the skin trade, live capture for stock farms, egg-collection, and hunting for meat (Nao & Tana, 1994; Thorbjarnarson, 1999). Additionally, crocodile farms frequently hybridize Siamese crocodile with Cuban crocodile C. rhombifer and saltwater crocodile C. porosus, and there is at risk of such hybrids escaping or being released into the wild (Simpson & Bezuijen, 2010; Starr et al., 2010; Daltry et al., 2016). Although the threat of hunting and wild capture has reduced in Cambodia, recovery of wild populations is still impeded by poor reproductive rates, habitat conversion and degradation, and incidental deaths from drowning in fishing gear (Simpson & Sam, 2004). Furthermore, the chances of unaided population recovery are reduced by the small and scattered nature of existing populations in the wild. Although Siamese crocodile was originally described over 200 years ago, very little is known about ecology and biology of the species in the wild (Simpson & Sam, 2004). The few studies published indicate that the species inhabits a wide range of freshwater habitats including slow-moving rivers, streams, oxbow lakes, seasonal lakes, marshes, and swamps up to 730 m a.s.l. (Daltry et al., 2003; Simpson et al., 2006). Studies conducted during the dry season (December–April) in the Cardamom Mountains of southwestern Cambodia have found wild crocodiles to be highly sedentary, typically remaining within a lake or short length of river (Simpson et al., 2006). Conversely, during the wet season (May–November) adult Siamese crocodiles have been recorded dispersing up to 25 km before returning to a dry season site (Simpson et al., 2006). ment via the release of captive-bred animals into secure natural habitats is an important strategy to recover wild populations (Thorbjarnarson, 1992; National Crocodile Conservation Network, 2012). Understanding how released animals adapt to their new environment is critical to the success of any reintroduction or reinforcement programme. We present the results of radio-tracking conducted on 15 captive-reared Siamese crocodiles that were released in the Southern Cardamom National Park in 2012 (n=10) and 2014 (n=5). The aim of our study was to understand the movements and survival of crocodiles taken from captivity and released into the wild. M e t hods Study site The crocodiles were released in one of eight sites identified by the Cambodia Crocodile Conservation Programme for reintroduction and reinforcement of Siamese crocodile in the Cardamom Mountains (National Crocodile Conservation Network, 2012) (Fig. 1). The site is located in Srae Ambel District, Koh Kong Province within the Southern Cardamom National Park and comprises hill evergreen forest, semi-evergreen forest and open forest and grassland, with an elevation range of 10–600 m a.s.l. Names of localities and landscape features at the release site are not provided in this paper to protect the crocodiles. The release site comprised a section of river located at an elevation of 70 m a.s.l., approximately 5 km from its headwaters. The nearest village was located 12 km downstream. The width of the river ranged from 30 to 50 m, and riverine habitats included a mixture of rocks, sandbars, and vegetation in the water, with rapids separating deepwater sections (anlong in Khmer) which have an average minimum dry season depth of 4.39 m (± 1.95m SD) and length of 300–1,000 m. Release procedures Because remaining populations of the species are small and fragmented, reintroduction and/or reinforce- Release of the crocodiles followed protocols given in the Siamese Crocodile Reintroduction and Reinforcement Strategy and Action Plan (National Crocodile Conservation Network, 2012). Prior to release, all of the crocodiles in this study were cared for at the Phnom Tamao Wildlife Rescue Centre (PTWRC), and originated from confiscations made by law enforcement officials and donations from other captive facilities. All of the animals were identified as ‘purebred’ Siamese crocodiles through DNA testing (Starr et al., 2010). They were also medically cleared by veterinarians belonging to the Wildlife Cambodian Journal of Natural History 2017 (1) 102–108 © Centre for Biodiversity Conservation, Phnom Penh 103 104 Eam S. et al. Fig. 1 General location of Siamese crocodile release area in Cambodia and last recorded locations of crocodiles. Conservation Society to ensure they were in good health before release. Data collection and analysis A temporary enclosure (5 m x 8 m) was constructed on the river bank at the release site which extended four metres into the river. The crocodiles were kept in the enclosure for about one week to allow them to recover from the journey from PTWRC and to monitor their condition and ensure they were ready for release. Eighteen crocodiles were released in the project site in 2012 and eight in 2014. Ten and five of these, respectively, were fitted with VHF radio-transmitters (150 MHz, Sirtrack Ltd, New Zealand) (Table 1). Small radiotransmitters (weighing ca. 40 g) with expected battery life of 13 months were attached to the dorsal base of the tail of four juvenile animals (ranging from 132 to 149 cm in total length). Larger radio-transmitters (weighing ca. 50 g) with expected battery life of 18 months were attached to the same part of 11 sub-adult crocodiles (ranging from 155 to 183 cm in total length). Radio-telemetry of the 15 crocodiles was undertaken for a total of 282 days from December 2012 to June 2015. Because the release site and surrounding area were inaccessible in July–October due to the wet season, all data collection was conducted during the dry season. Monitoring was conducted for 5–7 days every month when the site was accessible (specifically November–June). Observations were carried out on boat or on foot using a portable 3-element Yagi antenna attached to a 16-channel receiver (Advanced Telemetry Systems, USA). Signals could usually be detected from distances up to 500 m and 5–10 m in depth and crocodile locations were registered using hand-held GPS receivers. A 9-km stretch of river was surveyed (comprising 2 km upstream and 7 km downstream of the release site) for five days during each survey. If crocodiles were not located during this period, one to two additional days were devoted to surveying areas beyond the aforementioned stretch of river. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 102–108 Movement of Siamese crocodiles The mean minimum daily distance travelled by individual crocodiles was calculated using location data from successive days and calculating Euclidean distances between points. The release site was used as the reference point to determine average distances that crocodiles travelled from the release site. ArcGIS (vers. 10.2.2) and MS Excel were used for calculations. Re sult s The number of times each crocodile was detected after release varied greatly. One individual (M11) was detected only once, three days after release; five (M1, F2, M8, M12 & F13) were detected less than six months after release; and five (M3, F4, F6, M9 & F14) were found six to seven months after release. Four crocodiles (M5, M7, F10 & F15) were detected after one year and the latest detection occurred 18 months after release (Table 1). None of crocodiles released were found or reported dead. Thirteen of the 15 crocodiles remained within 700 m of the release site during the survey period. The two exceptions were sub-adult females (F6 and F15; Table 1) whose final detections were in the same anlong ca. 10 km downstream of the release site and ca. 3 km upstream of the nearest village (Fig. 5). The mean minimum distance travelled by crocodiles per day was 280.91 m (SD = 189.87) and the mean minimum distance crocodiles were located from the release site was 741.12 m (SD = 1,095.28) (Table 1). There was little difference between the distances travelled by male and female crocodiles. On average, females (n=7) travelled 349.56 m/day (SD = 250.86), whereas males (n=8) travelled 220.84 m/day (SD = 96.55 m/day) (Fig. 2). Mean distances from the release site recorded for female crocodiles were 1,138 m (SD = 1561.71) and 393.28 m (SD = 111.37) for males (Fig. 3). However, the figures for females are likely distorted by the two individuals that travelled 10 km downstream of the release site, while the remaining five females travelled a minimum distance of 435 m (SD = 234.09) from the release site. Juvenile crocodiles (n=4) appeared to move less than sub-adult crocodiles (n=11), travelling 171.68 m/day (SD = 32.12) and 320.63 m/day (SD = 208.93) on average, respectively (Fig. 4). Juveniles were located 368.87 m (SD = 169.85) from the release site on average, whereas the equivalent figure for sub-adults was 876.49 m (SD = 1,263.03) (Fig. 5). Again however, the latter figures are likely distorted by the two individuals that travelled 10 km downstream from the release site. Table 1 Siamese crocodiles released and monitored in southwestern Cambodia from 2012 to 2015. Code Status name Total length (cm) Start date End date No. of times recorded Mean minimum distance (m) travelled per day Mean distance from release site (m) M1 Juvenile 132 13 December 2012 27 February 2013 19 203.33 388.64 F2 Sub-adult 156 13 December 2012 30 January 2013 11 260.36 161.44 M3 Sub-adult 176 13 December 2012 12 June 2014 20 327.66 609.98 F4 Juvenile 144 13 December 2012 19 January 2014 31 134.38 596.12 M5 Sub-adult 183 13 December 2012 10 May 2014 40 138.68 282.33 F6 Sub-adult 160 13 December 2012 21 May 2013 28 330.46 1,201.67 M7 Sub-adult 163 13 December 2012 12 June 2014 43 175.71 430.39 M8 Sub-adult 168 13 December 2012 26 February 2013 6 319.98 329.07 M9 Juvenile 148 13 December 2012 12 June 2014 17 193.03 290.15 F10 Juvenile 149 13 December 2012 12 June 2014 39 155.98 200.58 M11 Sub-adult 173 17 January 2014 20 January 2014 1 333.79 333.79 M12 Sub-adult 172 26 February 2014 20 March 2014 5 74.56 481.94 F13 Sub-adult 173 27 February 2014 12 June 2014 9 732.12 648.28 F14 Sub-adult 155 26 February 2014 12 June 2014 10 671.88 569.05 F15 Sub-adult 155 19 January 2014 25 April 2015 4 161.77 4,593.49 Cambodian Journal of Natural History 2017 (1) 102–108 © Centre for Biodiversity Conservation, Phnom Penh 105 106 Eam S. et al. Fig. 2 Mean minimum daily distances (m) travelled by eight male and seven female Siamese crocodiles in southwestern Cambodia. Bold lines represent mean values, whiskers represent minimum and maximum values and boxes respresent lower and upper quartiles. Fig. 3 Mean minimum distances (m) travelled from release site by eight male and seven female Siamese crocodiles in southwestern Cambodia. Bold lines represent mean values, whiskers represent minimum and maximum values and boxes respresent lower and upper quartiles. Fig. 4 Mean minimum daily distances (m) travelled by four juvenile and eleven sub-adult Siamese crocodiles in southwestern Cambodia. Bold lines represent mean values, whiskers represent minimum and maximum values and boxes respresent lower and upper quartiles. Fig. 5 Mean minimum distances (m) travelled from release site by four juvenile and eleven sub-adult Siamese crocodiles in southwestern Cambodia. Bold lines represent mean values, whiskers represent minimum and maximum values and boxes respresent lower and upper quartiles. Disc ussion Few studies have monitored the movements of captivereared Siamese crocodiles released as part of a reintroduction programme, although similar releases have been undertaken e.g., Cat Tien National Park in Vietnam (Polet et al., 2002). Our study suggests that captive-reared juvenile and sub-adult Siamese crocodiles released into novel environments show site fidelity during the dry season, at least for the first year after release. This suggests that such animals may have good prospects for surviving at suitable release sites. Because visual sightings of the released crocodiles were rare due to their wariness, the fate of those not observed could not be confirmed. More than half of our study animals could not be located after six months, but no dead crocodiles were found or reported by local communities. The lack of observations in these cases could be due to other factors, such as equipment failure. For instance, Strauss et al. (2008) found that only 50% of batteries in VHF transmitters affixed to Nile crocodiles lasted longer than six months. Other crocodile release programmes have also experienced similarly high rates © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 102–108 Movement of Siamese crocodiles of ‘disappearance’ (e.g., Ballouard et al., 2010), but without direct evidence of mortality, it is difficult to know if this might be due to batteries expiring, device failure, or dispersal beyond detection range and/or death of crocodiles outside of the study area. Because no evidence of mortality was observed or reported during the study period, we suspect that most, if not all, of our study crocodiles survived. Previous studies of Siamese crocodile movements during the wet season reveal that the species can travel further than our study animals did during the dry season. Simpson et al. (2006) recorded a juvenile female moving 2–4 km across flooded forests and between the Areng River (southwestern Cambodia) and oxbow lakes from March to September. They also monitored an adult male which moved up to 11.9 km along the same river over three and a half months during the dry season. Studies on saltwater crocodiles Crocodylus porosus in Australia have demonstrated that the linear range of the species varies from 1.3 ± 0.9 km during the dry season to 62 km in the wet season (Kay, 2005). On release, captive-bred gharials Gavialis gangeticus also varied greatly in their movements, with some remaining close to the release site and others settling up to 40 km away (Ballouard et al., 2010). Because habitat quality is likely a driver of site fidelity among released crocodiles (van Weerd et al., 2011), the fact that most of our crocodiles stayed near the release site could indicate its habitat quality was good. However, the real measure of success for reintroduction and reinforcement programmes is breeding success. As the first animals we released in 2012 will take at least five years to reach sexual maturity, further surveys should be conducted to search for evidence of nests and juveniles. the communities to garner support for the recovery of an animal that has the potential for conflict with humans. Ack now le dge m e nt s The authors are grateful to the Phnom Tamao Wildlife Rescue Centre and the Cambodian Forestry Administration for their support. Many other individuals and organizations provided valuable technical assistance and support including: Adam Starr, Boyd Simpson, Hang Chandaravuth, Jenny Daltry, Lonnie McCaskill, Sarah Brook, Sorn Piseth and community crocodile wardens for the local area. This project would not be possible without the support of the following donors: Association of Zoos and Aquariums, Disney Conservation Fund, Mohammed bin Zayed Species Conservation Fund, Ocean Park-Hong Kong, Oren Taylor, SOS-Save Our Species, U.S. Fish and Wildlife Service Critically Endangered Animals Conservation Fund, and other private donors who choose to remain anonymous. Re fe re nc e s Ballouard, J.M., Priol, P., Oison, J., Ciliberti, A. & Aawely, A.C. (2010) Does reintroduction stabilize the population of the critically endangered gharial (Gavialis gangeticus, Gavialidae) in Chitwan National Park, Nepal? Aquatic Conservation, Marine and Freshwater Ecosystems, 20, 756–761. Bezuijen, M.R., Phothitay, C., Hedemark, M. & Chanrya, S. (2006) Preliminary Status Review of the Siamese crocodile (Crocodylus siamensis Schneider, 1801) (Reptilia: Crocodylia) in the Lao People’s Democratic Republic. Mekong Wetlands Biodiversity Conservation and Sustainable Use Programme, Vientiane, Laos. With fewer than three wild nests reported each year for Siamese crocodiles in Cambodia, existing populations are too small and fragmented to recover in the absence of reintroduction and reinforcement efforts. This study is part of ongoing efforts to establish viable populations of the species in the country, and our discovery that young crocodiles typically remain near release sites during the dry season is encouraging because it suggests the risk of crocodiles straying from release sites into areas with less suitable habitat or greater risk from people may be low. Our release of captive-reared animals follows a decade of conservation work to determine the distribution and natural history of Siamese crocodiles (Sam et al., 2015) and secure key areas as sanctuaries. It was of course critical to understand the historical distribution and habitat needs of this species before any sort of reintroduction, and to eliminate the reasons it was extirpated in the first place. Another critical step was to engage with Daltry, J.C. & Chheang D. (2000) Siamese crocodiles discovered in the Cardmom Mountains. Crocodile Specialist Group Newsletter, 19, 7–8. Cambodian Journal of Natural History 2017 (1) 102–108 © Centre for Biodiversity Conservation, Phnom Penh Daltry, J.C., Chheang D., Em P., Poeung M., Sam H., Tan T. & Simpson, B.K. (2003) Status of the Siamese Crocodile in the Central Cardamom Mountains, Cambodia. Fauna & Flora International Cambodia Programme, and Department of Forestry and Wildlife, Phnom Penh. Daltry, J.C., Langelet, E., Solmu, G.C., van der Ploeg, J., van Weerd, M. & Whitaker, R. (2016) Successes and failures of crocodile harvesting strategies in the Asia Pacific. In Tropical Conservation: Perspectives on Local and Global Priorities (eds A.A. Aguirre & R. Sukumar), pp. 345–362. Oxford University Press, Oxford, UK. IUCN (2015) The IUCN Red List of Threatened Species. Http:// www.iucnredlist.org [accessed 10 November 2015]. Kay, W.R. (2005) Movements and home ranges of radio-tracked Crocodylus porosus in the Cambridge Gulf region of Western Australia. Wildlife Research, 31, 495–508. 107 108 Eam S. et al. Kurniati, H., Widodo, T. & Manolis, C. (2005) Surveys of Siamese Crocodile (Crocodylus siamensis) Habitat in the Mahakam River, East Kalimantan, Indonesia. Indonesian Institute of Sciences, Bogor, Indonesia. Nao T. & Tang T.S. (1994) Country report on crocodile conservation in Cambodia. In Crocodiles: Proceedings of the 12th Working Meeting of the IUCN/SSC Crocodile Specialist Group, pp. 3–15. IUCN, Gland, Switzerland. National Crocodile Conservation Network (2012) Siamese Crocodile Reintroduction and Reinforcement Strategy and Action Plan for the Royal Kingdom of Cambodia: 2012–2031. Ministry of Agriculture, Forestry, and Fisheries, Fauna & Flora International Cambodia Programme and Wildlife Conservation Society, Phnom Penh, Cambodia. Simpson, B.K. & Bezuijen, M.R. (2010) Siamese crocodile Crocodylus siamensis. In Crocodiles: Status Survey and Conservation Action Plan. Third Edition (eds S.C. Manolis & C. Stevenson), pp. 120–126. IUCN/SSC Crocodile Specialist Group, Darwin, Australia. Simpson, B.K. & Sam H. (2004) Siamese crocodile (Crocodylus siamensis) surveys in Cambodia. In Crocodiles. Proceedings of the 17th Working Meeting of the IUCN/SSG Crocodile Specialist Group, pp. 110–120. IUCN, Gland, Switzerland. Simpson, B.K., Sorn P., Pheng S., Pok S., Sok P. & Prumsoeun W. (2006) Habitat use and movement of wild Siamese crocodiles in Cambodia. In Crocodiles. Proceedings of the 18th Working Meeting of the IUCN/SSC Crocodile Specialist Group, pp. 345. IUCN, Gland, Switzerland. Platt, S.G., Sovannara H., Kheng L., Stuart, B.L. & Walston, L. (2006) Crocodylus siamensis along the Sre Ambel River, southern Cambodia: habitat, nesting, and conservation. Herpetological Natural History, 9, 183–188. Starr, A., Daltry, J.C. & Nhek R. (2010) DNA study reveals Siamese crocodiles at the Phnom Tamao Wildlife Rescue Centre, Cambodia. Crocodile Specialist Group Newsletter, 28, 5–7. Polet, G., David, J. Murphy, Phan V.L. & Tran V.M. (2002) Crocodile conservation at work in Vietnam, re-establishing Crocodylus siamensis in Cat Tien National Park. In Crocodiles. Proceedings of the 16th Working Meeting of the IUCN/SSC Crocodile Specialist Group, pp. 86–95. IUCN, Gland, Switzerland. Strauss, M., Botha, H. & Van Hoven, W. (2008) Nile crocodile Crocodylus niloticus telemetry: observations on transmitter attachment and longevity. South African Journal of Wildlife Research, 38, 189–192. Ross, J.P. (1998) Crocodiles: Status Survey and Action Plan. Second Edition. IUCN/SSC Crocodile Specialist Group, Gland, Switzerland. Sam H., Hor L., Nhek R., Sorn P., Heng S., Simpson, B., Starr, A., Brook, S., Frechette, J.L. & Daltry, J.C. (2015) Status, distribution and ecology of the Siamese crocodile, Crocodylus siamensis, in Cambodia. Cambodian Journal of Natural History, 2015, 153–164. © Centre for Biodiversity Conservation, Phnom Penh Thorbjarnarson, J.B. (1992) Crocodiles: An Action Plan for Their Conservation. IUCN/SSC Crocodile Specialist Group, Gland, Switzerland. Thorbjarnarson, J.B. (1999) Crocodile tears and skins: international trade, economic constraints, and limits to the sustainable use of crocodilians. Conservation Biology, 13, 465–70. van Weerd, M., Balbas, M., Telan, S., Rodriguez, D., Guerrero, J. & van de Ven, W. (2011) Philippine crocodile reintroduction workshop. Crocodile Specialist Group Newsletter, 30, 10–12. Cambodian Journal of Natural History 2017 (1) 102–108 Carbon dynamics in dry dipterocarp forest St a nd c a rbon dyna m ic s in a dr y Ca m bodia n dipt e roc a rp fore st w it h se a sona lly fl oode d sa ndy soils ITO Eriko1, FURUYA Naoyuki1, TORIYAMA Jumpei2, OHNUKI Yasuhiro2, KIYONO Yoshiyuki2, ARAKI Makoto2, SOKH Heng3, CHANN Sophal3, KHORN Saret4, SAMRETH Vanna4, SO Thea3, TITH Bora3, KETH Samkol3, LY Chandararity3, OP Phallaphearaoth3, MONDA Yukako5 & KANZAKI Mamoru5 1 Hokkaido Research Center, Forestry and Forest Products Research Institute, No. 7 Hitsujigaoka, Toyohira, Sapporo, Hokkaido, 062-8516 Japan. 2 Forestry and Forest Products Research Institute, No. 1 Matsunosato, Tsukuba, Ibaraki, 305-8687 Japan. 3 Institute of Forest and Wildlife Research and Development, Forestry Administration, Street 1019, Phum Rongchak, Sankat Phnom Penh Thmei, Khan Sen Sok, Phnom Penh, Cambodia. 4 Department of Forestry and Community Forestry, Forestry Administration, No. 40 Preah Norodom Boulevard, Phnom Penh, Cambodia. 5 Graduate School of Agriculture, Kyoto University, Kyoto City, Kyoto, 606-8502 Japan. * Corresponding author. Email iter@affrc.go.jp Paper submitted 6 December 2016, revised manuscript accepted 12 June 2017. ɊɮɍɅʂɋɑɳȶſɆ ɳȼˊɊƓɪɔɅɭɎɁƎɎ ɩDžɅƳɌ “ȲɸɳɀˊɅ-LJɁɽɆȶɽ” ɵɅȴɳƙNjȶɇƎɯȷɳɇƎˊɊ ”ƳɌƳɁɽɆɅƏɋƳɌǒɋNJɋƳɆɮɅ Ɇǁ Ǝ ɍɊȲɈɪƳɌƳɆɽɅɩȶƳɌɆɸ LjƚȻɵƙɈɳȺˊ ” ɳɋˊȶɇƎɍɽƳɌLJɻ ɅɽǒƗɅȲɸɳɀˊɅƳɆɮɅ ɈɪƳɌɳȲˊɅɳɓˊȶɵɅȹɪɎNjɻ ɑɳȼˊɊɳȺˊ ƙɆƸɸƹƒɸ Ʌɩȶ ƳɌLJɁɽɆȶɽȹɪɎNjɻ ɑNjɅȹɪɎ ɩɁ Ɇǁ Ǝ ɍɊȲɈɪɄɊƗƺɁɩɅɩȶɑȲɊƗNJɈɊɅɭɑƞ ɳǷȲƒɭȶɵƙɈɌɳLJɹȼɪɍǙɋȳǜȷɽɵɅƙɆɳɃɑȲɊƕɭƺʆ ɳɋˊȶLJɅƙɁȫɁɈɩɅɩɁƘɵƙɈɌɳLJɹȼɪ ɍǙɋȳǜȷɽ ƺɵƙɈɴȼɍɑɊƓɮɌɳƽɋɳȼˊɊɴɁƓȶ (Dipterocarpusȱobtusifolius) ɴȼɍȼɭɹɳǷɁɸɆɅɽȼɪɍǙɋȳǜȷɽʆ ɳɋˊȶLJɅɎ ɩNJȴ ɈɪɔƙǂɅɭȲɮɍǒȝɑƎ Ʌɩȶ ȲɸɳɀˊɅɔȶžɁɽɇɩƃɁȲɊƕɑɽƙɃȪȶ ɴȼɍNjɅɔȶžɁɽɇɩƃɁ>ʕɑɊ ɳƽɋɴɇơȲɳɍˊƳɌɳɄƛˊȹɸɳɌȟɅɳȼˊɊɳȺˊ ɔɑɽɌɋɺ ɳɈɍʑʑƹƒɸ(ʒʐʐʓ ȼɍɽ ʒʐʑʔ)ȲɅƚȶɊȲʆ ɃɪǂɸȶɑɩȲǜLJɅɆƷƟȻɈɪȲƙɊɩɁDŽɆɵɅȲɸɳɀˊɅƳɆɮɅ(ʐ,ʑʘɊƙȲ ƳɆɮɅ/ʐ,ʒʔɒɩȷ ǂ/ƹƒɸ) ȷɸɳljɹƳɌɑƎɭȲƳɆɮɅȼɸɆɮȶNjɅ(ʑʑ,ʓɊƙȲ ƳɆɮɅ/ʐ,ʒʔɒɩȷǂ) ɳɄȢɆɳǵɅɫȶƙɆɳɉɃɵƙɈɳȺˊ ȲɊƕɭƺȼɵɃɳɃȢɁɴȼɍƙɁȪɎLJɅ ɑɩȲǜȲɅƚȶɊȲʆ ɔƙǂƷɆɽɵɅɳȼˊɊɳȺˊ ɴȼɍNjɅɔȶžɁɽɇɩƃɁȲɊƕɑɽƙɃȪȶ ǃʑʐɑɊ NjɅȲƙɊɩɁDŽɆ(ʑ,ʐʓ%/ƹƒɸ) ƺɳɒɁɭdžɸɤƘNjɅƳɌ LJɁɽɆȶɽƳɆɮɅɁɩȷɁɯȷ(ʐ,ʑʒɊƙȲ ƳɆɮɅ/ʐ,ʒʔɒɩȷǂ) ɈɪɳȼˊɊɳȺˊ ƷɆɽɳƽɋɄɊƗƺɁɩ ȲƒɭȶɔɸɓɭȶɳɈɍɊɭɅƳɆɽ(ʒʐʐʓ ȼɍɽ ʒʐʑʑ)ʆ ƳɌƳɆɽɳȺˊ ɳƽɋȹɅɔdžɊɩȲLJɅƳɁɽɆɅƏɋƳɌɑƎɭȲƳɆɮɅƙɆNjɀʕ,ʓʗɊƙȲ ƳɆɮɅ/ʐ,ʒʔɒɩȷǂ (ʔʒ,ʗ% ɵɅɁɵɊƚ ɊɭɅɳɈɍƳɆɽ) ƺɊɯɋɅɫȶƳɌȳɮȷƴɁɆdžƐɆɽɆɅƞɸ (ʐ,ʐʐʖɊƙȲ ƳɆɮɅ/ʐ,ʒʔɒɩȷǂ) ɴȼɍɑNjNjƙɁɳǵɅɫȶȲɸɳɀˊɅƳɆɮɅȲƒɭȶɌɋɺ ɳɈɍ ʓʐƹƒɸȲɅƚȶɊȲʆ ȲɸɳɀˊɅƳɆɮɅƽȷɽƴɁ ȲɸɳɀˊɅƳɆɮɅDŽɆ Ʌɩȶ ƳɌƳɆɽɳȺˊ ȲɭƒȶƙɃȶɽƙDŽɋɄɸɴȼɍLJɅɔɳȶžɁɳǷȲƒɭȶɃɪǂɸȶ ɑɩȲǜ ɆƷƟȻǃɵƙɈɌɳLJɹȼɭɹɳǷɁɸɆɅɽȼɪɍǙɋȳǜȷɽ ƸɸLJȷɽƙɁȪɎɴɁNjɅƳɌɴɆȶɴȷȲƺƙȲȩɊɌȶɵɅɵƙɈƙȹȩɹɑƚɫȲɳɅɹ ɳǷɳɈɍɔɅɭɎɁƎ Ɏ ɩDžɅƳɌ “ȲɸɳɀˊɅ-LJɁɽɆȶɽ” ɳɅɹʆ CITATION: Ito E., Furuya N., Toriyama J., Ohnuki Y., Kiyono Y., Araki M., Sokh H., Chann S., Khorn S., Samreth V., So T., Tith B., Keth S., Ly C., Op P., Mona Y. & Kanzaki M. (2016) Stand carbon dynamics in a dry Cambodian dipterocarp forest with seasonally flooded sandy soils. Cambodian Journal of Natural History, 2017, 109–127. Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh 109 110 Ito E. et al. Abst ra c t To implement the gain-loss approach from the ‘Reducing Emissions from Deforestation and Forest Degradation’ initiative, we provide estimates of gains from the annual increase in tree biomass and losses in live biomass caused by natural and anthropogenic processes in a sandy dry dipterocarp forest in Cambodia. We examined a sandy dry dipterocarp forest—a form of forest characterised by the strong dominance of Dipterocarpus obtusifolius—which was distributed on sites with sandy soils. We analysed the demography and diameter increment of trees with diameters at breast height (DBH) >5 cm based on an 11 year tree census (2003–2014). The study plot showed a low carbon (C) increment (0.18 Mg C 0.24 ha−1 y−1) for the initial C stock (11.3 Mg C 0.24 ha−1) compared to other Cambodian forest types that have been studied in the past. The low mortality of trees with DBH ≥10 cm (1.03% y–1) resulted in a small C loss from naturally dying trees (0.12 Mg C 0.24 ha−1) in the pre-cutting period (2003–2011). Logging by unknown parties decreased C stocks by 5.37 Mg C 0.24 ha−1 (42.7% of the pre-logging value) with relatively less collateral damage (0.006 Mg C 0.24 ha−1); this was equivalent to C increments that had accrued over 30 years. The low absolute C increment, the relatively low C increment, and the intensive logging observed in the study plot suggest that sandy dipterocarp forests need to be stratified into subdivisions of deciduous forests when implementing the gain-loss approach. Ke yw ords Carbon stock, Dipterocarpus obtusifolius, forest degradation, mortality, recruitment, Reducing Emissions from Deforestation and Forest Degradation (REDD), tree increment. I nt roduc t ion Reducing Emissions from Deforestation and Forest Degradation and the role of conservation, sustainable management of forests and enhancement of forest carbon stocks in developing countries (REDD+) is an effort to create financial incentives for developing countries to reduce carbon (C) emissions from forested lands (UNFCCC, 2009). To implement the global climate change mitigation programme, IPCC (2006) presented two methods for calculating changes in C stocks: the gain-loss method and the stock-difference method. The choice of method largely depends on a country’s available data, domestic capacity, and forest transition stage (GOFC-GOLD, 2008; Murdiyarso et al., 2008). Cambodia has attracted the attention of the REDD+ programme because it is a hotspot for deforestation and forest degradation (FAO, 2010; 2011). In recent years, a several studies have attempted to prepare a full implementation programme for REDD+ activities in Cambodia (Sasaki et al., 2006, 2011, 2012, 2013, 2016; Sasaki & Yoshimoto, 2010). To reduce uncertainties in C estimation for countries experiencing significant forest degradation (Sasaki, 2006), application of the gain-loss method was recently examined in Cambodia (Sasaki et al., 2006, 2012, 2013, 2016; Kiyono et al., 2017). The gain-loss method is built around the gains from annual increases in biomass and the losses in live biomass caused by natural and anthropogenic processes (Murdiyarso et al., 2008). Those estimates must be obtained for each forest type, appropriately stratified by eco-types and degradation © Centre for Biodiversity Conservation, Phnom Penh processes (Murdiyarso et al. 2008; Pearson et al., 2014; GOFC-GOLD, 2016). Cambodia has a diversity of forest types (Rollet, 1972; Rundel, 1999) and the extent of their degradation varies, adding a layer of complexity to the task of data collection. The Cambodian Forestry Administration (FA) classified forest cover into four types: evergreen, semi-evergreen, deciduous, and other forests including different types such as forest re-growth, inundated forests, stunted forests, mangrove forests, and forest plantations (FA, 2011). Specific information for each forest-type has been obtained from ground-based studies in Cambodia, and includes data on forest structure (Kimphat et al., 2000, 2002a; Kao & Iida, 2006; Ouk, 2006; Pin et al., 2013; Toyama et al., 2015; Chheng et al., 2016), biomass (Top et al., 2004a; Kiyono et al., 2010; Khun et al., 2012; Samreth et al., 2012; Chheng et al., 2016; Monda et al., 2016), and biomass increment (FA, 2004; Top et al., 2004a; Kiyono et al., 2017). Despite these, data collection is still fragmented. For example, data on annual increment are relatively limited, possibly because neither the continuous maintenance of permanent sample plots nor frequent tree measurements for forest biomass increment are necessarily straightforward in countries with frequent deforestation and forest degradation (Kiyono et al., 2017). Positive relationships have been reported between annual above-ground biomass increment and aboveground biomass for Cambodian forests without stratifying forest types (Top et al., 2004a; Kiyono et al., 2017). The coefficients of determination for this relationship Cambodian Journal of Natural History 2017 (1) 109–127 Carbon dynamics in dry dipterocarp forest in both previous studies were relatively low. Top et al.A(2004a) suggested that stand age, soil types, microtopography, or species composition might explain the residuals, whereas Kiyono et al. (2017) could not explain these by differing mean annual precipitation or the soil fertility index. The gain-loss method should be built on an ecological understanding of how forests grow (Murdiyarso et al. 2008). Additional data on C increment is still required, in addition to data on fundamental forest structure and dynamics for the various forest types. Deciduous forests comprising dry mixed deciduous forests and dry dipterocarp forests predominate in Cambodia, and cover 24.68% of its land area (FA, 2011). Dry dipterocarp forests are described as forêt claire (Rollet, 1972), deciduous dipterocarp forests or woodlands (Rundel, 1999). They exist throughout IndoBurma (Ashton, 2014) and are scattered among lowland forest areas in Cambodia (JICA, 2002). The name of this community is a result of dominance by a small number of deciduous species of Dipterocarpaceae, such as Shorea siamensis, S. obtusa, Dipterocarpus tuberculatus, D. intricatus and D. obtusifolius (Rundel, 1999). Dry dipterocarp forests have been further subdivided into four forms, with different combinations of soil type and dominant dipterocarp species (Rollet, 1972). is unclear whether sandy dipterocarp forests achieve C increments to the same degree as other deciduous forests (Top et al., 2004a; Kiyono et al., 2017). There is also a dearth of data related to C emissions due to forest degradation, which is a significant contributor to C emissions in the atmosphere (GOFC-GOLD 2008). Considerable international variation exists in C emissions from tropical forest degradation caused by selective logging (Pearson et al., 2014). Forest degradation from uncontrolled tree cutting has resulted in significant losses of C stock in Cambodian evergreen forests (Sasaki & Putz, 2009). As such, further data on deciduous forest degradation would contribute to implementation of the gain-loss approach in Cambodia. Destablished a permanent sample plot (30 × 80 m) We to investigate stand structure and dynamics in a sandy dipterocarp forest in the Kampong Thom Province of Cambodia. During a chronological tree census (2003– 2014), logging by unknown parties occurred in the plot. The plot, while small and isolated, had the potential to provide useful information about sandy dipterocarp forests. The objectives of our study were to clarify the stand dynamics and annual increment in sandy dipterocarp forest based on 11 years of tree census data, and to assess C emission from cutting events and related collateral damage. One form of dry dipterocarp forest is characterised by the strong dominance of D. obtusifolius (Tbeng in Khmer), which favours sandy soil, gravelly soil or B (Smitinand et al., 1980). This forest (hereafter laterite sandy dipterocarp forest) has been referred to as forêt claire à Dipterocarpus obtusifolius (Vidal, 1960), D. obtusifolius on sand or grey soil (à D. obtusifolius, sur sable ou terre grise: Rollet, 1972), D. obtusifolius community (Baltzer et al., 2001) and D. obtusifolius stand type (Hiramatsu et al., 2007). Sandy dipterocarp forests are most characteristic of areas east of the Mekong River in Cambodia at sites with thin sandy soils over laterites (Rundel, 1999) and are F scattered as forest patches among evergreen forests often at sites with deep sandy soils subject to seasonal flooding in Kampong Thom Province, northeast of the Tonle Sap Lake (Hiramatsu et al., 2007). We conducted our study in the Kampong Thom Province in central Cambodia (12.8°N, 105.5°E; elevation: 70 m a.s.l., Fig. 1). The site was positioned on quaternary sedimentary rock and located on a largely flat, slightly undulating alluvial plain (Toriyama et al., 2011). The tropical Gis seasonal, and the months between November climate and April are dry. The mean annual temperature was 27°C, and the annual rainfall (mean ± SD) was 1,542 ± 248 mm (2000–2010; NIS, 2012). Sandy dipterocarp forests are often low in species richness (Hiramatsu et al., 2007) and have open structures with 40–70% canopy cover (Rundel, 1999; Hiramatsu et al., 2007; FA, 2011). They are associated with ground fires (Rundel, 1999; Hiramatsu et al., 2007), have nutrientpoor sandy soils (Rollet, 1972; Toriyama et al., 2007a, b), and experience seasonal flood and drought conditions (Rollet, 1972; Rundel, 1999; Baltzer et al., 2001; Araki et al., 2007). Nevertheless the predominant D. obtusifolius is ecologically plastic and stress tolerant (Rundel, 1999). Its annual growth is still unknown; in other words, it We established a permanent sample plot (30 × 80 m) for vegetation in a sandy dipterocarp forest with a canopy openness of 50% (Hiramatsu et al., 2007, Fig. 2). The forest includes D. obtusifolius (ca. 50% of stand basal area and 60% of stand tree numbers) and Gluta laccifera (35% and 6%, respectively) as the dominant species (Hiramatsu et al., 2007). These are the major species in seasonal tropical forests in Asian monsoon areas and are indicative of a dry forest type (Rundel & Boonpragob, 1995; Eiadthong, 2011). The forest lacked auxiliary deciduous species such as Shorea obtusa, Pterocarpus macrocarpus, and Xylia Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh M e tE hods Study site 111 112 Ito E. et al. (DBH) ≥5 cm were enumerated and identified to species (except for one specimen which was identified to genus, Appendix 1). We measured stem girth to the nearest 1 mm (1.3 m above ground level) during the tree censuses except in the 2010 census. The height of each tree was measured in 2010 using either a Vertex III clinometer (Haglöf, Långsele, Sweden) or a telescopic height-measuring pole. We spatially mapped stems with an accuracy <1 m and measured crown diameters of major and minor axes in 2012 (Fig. 2). Analysis Fig. 1 Location of the study site (solid circle) in Kampong Thom Province, Cambodia. xylocarpa, which usually co-occur in dry dipterocarp or deciduous dipterocarp forests (Royal Forest Department, 1962; Tani et al., 2007; Hiramatsu et al., 2007; Eiadthong, 2011; Pin et al., 2013). Edaphic limitation is a potential factor limiting species richness (Hiramatsu et al., 2007). The forest was classified as a deciduous forest using FA (2011); however, the component tree species showed irregular and incomplete leaf shedding (Ito et al., 2007). For example, leaf longevity of D. obtusifolius often exceeded one year (Ito et al., unpublished data), a trend which has also been observed in the Thai highlands (Elliott et al., 2006). The soil of the study area was generally sandy and nutrient-poor, similar to other G. laccifera habitats (Eiadthong, 2011). The soils have been classified as acrisols (WRB), but with albic and arenic features that suggest a closer relationship with arenosols (WRB) (Toriyama et al., 2007a). The ground surface was waterlogged several times in the middle of the rainy season (August through September: Araki et al., 2007) and the ground vegetation includes Xyris complanata R.Br. and insectivorous plants (Drosera sp. and Nepenthes sp.), which suggest low-nutrient edaphic conditions (Hiramatsu et al., 2007). Annual mortality and recruitment rate were calculated using commonly used logarithmic models (e.g., Swaine et al., 1987; Phillips & Gentry, 1994; Lewis et al., 2004). More specificially, mortality rate (λ) and recruitment rate (k) were calculated as follows: λ = (ln(n0)−ln(n0−nd))/t (Eq. 1), k = (ln(n0+nr )−ln(n0))/t (Eq. 2), Field surveys were conducted in 2003, 2008, 2009, 2010, 2011, 2012 (pre-logging), and 2014 (post-logging) to investigate tree growth and demography. Censuses in 2003, 2011, and 2014 were conducted in February each year. Censuses in 2009 and 2010 were conducted in December each year. Censuses in 2008 and 2012 were conducted in June and November, respectively. All standing woody stems with a diameter at breast height where n0 is the number of trees present at the beginning of the census interval, and nd and nr are the number of trees that died of natural causes and the recruited trees during time (t) of the census interval. We used a 2 × 2 contingency table to assess differences in mortality and recruitment rate between individuals with a DBH of 5–10 cm and ≥10 cm. Given the stems present during the initial 2003 inventory, the numbers of surviving and dying stems at the end of the census interval (2014) provided the values for mortality analysis. Recruited stems were omitted from this analysis. Using the stems present at the end of the 2014 inventory, numbers of existing stems during the initial 2013 inventory and recruited stems during the 11-year census, we were able to identify values for recruitment rate analysis. Dying stems were omitted from this analysis. Hypothesis tests were performed using one-tailed probabilities based on Fisher’s exact test. The observed number of dying or recruited trees with DBH ≥5 cm was compared to the expected number of dying or recruited trees. The expected number was calculated by assuming that there were no mortality or recruitment rate differences during the 11-year census. Significance was tested by the chi-squared test for goodness of fit. To avoid periods with fewer than five expected dying or recruited trees, we divided our observations from all 11 years into three periods: 2003–2008, 2008–2011, and 2011–2014. Numbers of observed and predicted dying or recruited trees for each period were compared to a significance level of p=0.05. Diameter distributions of the predominant Dipterocarpus obtusifolius were evaluated for skewness (s) and kurtosis (κ), and were compared to a normal distribution using the pre-logging 2011 census © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 109–127 Data collection Carbon dynamics in dry dipterocarp forest Fig. 2 Individual tree locations and crown sizes within a 30 × 80 m study plot in Kampong Thom Province, Cambodia. Symbol sizes are canopy areas calculated from crown diameters and assume each crown was circular. Letters indicate status changes between 2012 and 2014: L = logged for timber (n=5); I = collateral damage from logging (n=2); U = cut for unknown reasons (n=5); N = natural death (n=6). data and the Shapiro-Wilk test. We used a log-normal distribution to best describe the diameter distribution of D. obtusifolius, where we estimated two parameters, μ (scale) and σ (shape). A goodness-of-fit test was examined using the Kolmogorov-Smirnov test. We assessed whether the log-normal distribution fitted for D. obtusifolius was significantly different from those of associated species using the Kolmogorov-Smirnov test. Four of the tested species were selected based on their basal area in the 2011 census (G. laccifera, Parinari anamensis, Syzygium oblatum, and Memecylon scutellatum). Diameter increments per year were compared between the 2003–2011 census and the 2011–2014 census for trees that had survived since the 2003 (first) census to the 2014 post-logging census (n=69) using a Wilcoxon signed rank test. We estimated the total biomass (TB) of the trees using the latest published allometric equations developed in a deciduous dipterocarp forest in Kratie Province, Cambodia (Model 4: Monda et al., 2016): ln (AGB) = −2.438 + 2.518 ln (DBH) sum of AGB and BGB. Field-measured DBH’s of each tree in the 2003, 2011, and 2014 censuses were applied to the equation for evaluating chronological changes in stand C storage, C emissions, and C increment because these three censuses were conducted in the same month of the dry season (February), which avoids DBH swelling in the rainy season. Allometry equations with tree height parameters were not applicable because tree height was measured only during the 2010 census. To evaluate the difference in biomass estimates, we estimated tree biomass for the corresponding DBH dataset from the 2011 census and tree height from the 2010 census using an allometry equation with parameters for tree height (H, m) (Model 2: Monda et al., 2016): ln (AGB) = −2.710 + 0.924 ln (DBH2 × H) ln (BGB) = −4.030 + 0.928 ln (DBH2 × H) (Eq. 4). The difference (%) in the stand TB results between Equations 3 and 4 was calculated as: (stand TB by Eq. 3 − stand TB by Eq. 4) / (stand TB by Eq. 3) × 100. where AGB and BGB represent the dry above-ground biomass and dry below-ground biomass of each tree (kg), respectively, and DBH (cm) is the diameter of the stem 1.3 m above-ground. TB was obtained by calculating the We found a difference of 6.8% in the study plot, which was considered small, although it suggests relatively low tree height in the study plot compared to the deciduous dipterocarp forest in Kratie where the equations were developed. Besides the TB estimates found using Equation 3, we obtained tree biomass by applying Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh ln (BGB) = −3.734 + 2.521 ln (DBH) (Eq. 3), 113 114 Ito E. et al. various equations identical to several previous studies to maintain consistency in the biomass estimates among comparative studies. Top et al. (2004a) estimated the AGB of trees with DBH >10 cm using the equation developed by Brown (1997): AGB = 42.69–12.800DBH + 1.242DBH2 (Eq. 5). Kiyono et al. (2017) estimated the ABG of trees with DBH >5 cm using the equation developed by Kiyono et al. (2006): AGB = 11545 BA1.24 (Eq. 6), where BA is the basal area of a stem at a height of 1.3 m (m2). Samreth et al. (2012) estimated the TB of trees with DBH >7.5 cm using the equation developed by Kiyono et al. (2011): TB = 4.08 × BA1.25 × WD1.33 (Eq. 7), where WD is the basic density (kg m–3) of the stem wood. The species-specific WD data obtained from neighbouring countries used for Equation 7 were selected from a global WD database provided by Chave et al. (2009) and Zanne et al. (2009). When species-specific data were not available in the database, we used 570, which is the default value for tropical Asian wood (Brown, 1997). The values of WD used for each species are listed in Appendix 1. Kimphat et al. (2002b) and Khun et al. (2012) presented biomass as standing volume (SV, m3). We converted AGB to SV similar to Sasaki et al. (2016) using the equation developed by Brown (1997): AGB = SV × 0.001 WD × BEF (Eq. 8), were calculated as the difference between their estimated biomass at the beginning and end of the interval. For stand C increment, biomass increments were summed for all trees that had survived, were recruited, and were dying. To compare our results with previous studies, the indices of stand structure, biomass, and increment were rescaled per hectare, given that the number of individuals increased linearly with area. A combination of the plotwise annual AGB increment and the initial AGB in the study plot was compared to previous studies (Top et al., 2004a; Kiyono et al., 2017). These studies were conducted for Cambodian forests without stratifying forest types, which derived positive relationships between annual AGB increment and initial AGB. A logarithmic relationship was obtained for trees with DBH >10 cm using data from 32 plots (four deciduous forest plots, where the details of the forest type were unknown) located in Kampong Thom (Top et al., 2004a). A linear relationship was obtained for trees with DBH >5 cm using data from 49 plots (24 deciduous forest plots, not including sandy dipterocarp forests, Kiyono, pers. comm.) in nine provinces (Kiyono et al., 2017). To maintain consistency in the biomass estimates between the present and previous studies, we used Equations 5 and 6 for biomass estimation and set tree size criteria identical to these particular studies. We recorded the presence of logged trees and collaterally damaged trees from the 2014 census. C loss associated with the cutting procedure was estimated based on DBH from the 2011 census. Tree C emissions by natural and anthropogenic causes were compared during the 2011–2014 censuses. Plot canopy coverages from the 2011 and 2014 censuses were estimated based on crown diameters measured in 2012. We assumed that each crown was circular, and plot canopy coverage was therefore provisionary and calculated as the sum of the crown area within the rectangular plot, with overlapping crown areas excluded. where BEF is the biomass expansion factor (1.74). Stand tree C stock in the study plot was obtained for the 2003, 2011, and 2014 censuses. The C stored in the trees was obtained by multiplying the C content in the dry wood biomass (0.5: IPCC, 2003) by the estimated biomass of each tree. Plot-specific values of tree C emissions by natural causes and C increment without anthropogenic effects were estimated for the 2003–2011 censuses. For stand C emissions estimation, the biomass of all the trees that died during the interval were summed, and the biomass of each tree was estimated from their last measured diameters during the interval. Stand C increment was calculated using a revised method of Clark et al. (2001). Increments for surviving trees during the interval were calculated as the difference between their estimated biomass at the beginning (2003) and end of the interval (2011). Increments for recruited trees during the interval were calculated as the difference between their estimated biomass at the end of the interval (2011) and the biomass of a tree of the minimum measured diameter (DBH of 5 or 10 cm). Increments for dying trees during the interval Stand-level tree density ranged 91 to 104 stems 0.24 ha–1 and 56 to 61 stems 0.24 ha–1, for individuals with DBH ≥5 cm and ≥10 cm across five pre-logging censuses (2003, 2008, 2009, 2011, 2012) respectively (Fig. 3). During the 11-year census, 121 stems with DBH ≥5 cm were enumerated, of which 23 died from natural causes and 23 were recruited (Table 1). Sixty-three stems with DBH ≥10 cm were enumerated, of which six died from natural causes and six were recruited (Table 1). Individuals in the 5–10 cm DBH class that died comprised M. scutellatum (n=10), © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 109–127 Re sult s Stand structure and dynamics Carbon dynamics in dry dipterocarp forest Fig. 3 Chronological changes in stem density, mortality, and recruitment rate. This is a bar-plot of the number of stems in the study plot (0.24 ha) across five pre-logging censuses (2003, 2008, 2009, 2011, 2012) and one post-logging census (2014). Black and white bars show stems with DBH of 5–10 cm and ≥10 cm, respectively. Open circles and closed squares indicate mortality and recruitment rates for stems with DBH ≥5 cm, respectively. Rates were calculated for the periods 2003–2008, 2008–2011, 2011–2014, and 2003–2014 (dotted lines). Artocarpus sp. (n=5), and D. obtusifolius (n=2). Individuals with DBH ≥10 cm that died comprised D. obtusifolius (n=3), Syzygium oblatum (n=1), Artocarpus sp. (n=1), and Anneslea fragrans (n=1). Recruited individuals in the 5–10 cm DBH class were M. scutellatum (n=7), D. obtusifolius (n=6), Xylopia vielana (n=4), Parinari anamensis (n=3), and Calophyllum calaba var. bracteatum (n=3). Recruited individuals in the DBH ≥10 cm class were D. obtusifolius (n=3), P. anamensis (n=3), and Symplocos cochinchinensis (n=1). Mortality (λ) for stems with DBH ≥5 cm during 2003–2014 was 2.08% y–1 (Fig. 3). Significantly higher λ was found for stems of 5–10 cm DBH (3.69% y–1) than for stems with DBH ≥10 cm (1.03% y–1) (one tailed Fisher’s exact test, p=0.0031). The chi-squared test for goodness of fit indicated that there was a significant difference in the occurrence of dying trees among the different census periods (2003–2008, 2008–2011, 2011–2014) (χ2=7.242, df=2, p<0.05) and only the number of dying trees between the 2011 and 2014 censuses were larger than expected values (Fig. 3). Fig. 4 Frequency distribution of DBH for predominant and associated tree species in the study forest based on pre-logging 2011 census data. Hatched patterns in the bars indicate tree status during the pre-logging 2011 census. A) Dipterocarpus obtusifolius; B) Gluta laccifera; C) Parinari anamensis; D) Syzygium oblatum; E) Memecylon scutellatum. The recruitment rate (k) for stems with DBH ≥5 cm in 2003–2014 was 1.92% y–1 (Fig. 3). The recruitment rate for 5–10 cm DBH stems (3.98% y–1) significantly exceeded ≥10 cm DBH stems (1.07% y–1) (one-tailed Fisher’s exact test, p=0.0010). Chi-squared test for goodness of fit showed that the expected number of trees recruited in each period between censuses significantly differed from the occurrence of period categories within the study plot (χ2=18.447, df=2, p<0.05); where the expected figure was greater than their actual occurrence during the 2003– 2008 censuses (Fig. 3). Size distributions according to basal area in the 2011 census are shown for five representative tree species in the study plot in Fig. 4. The diameter distribution of D. obtusifolius was significantly different from a normal distribution (Shapiro-Wilk test, n=59, W=0.87, p<0.0001). Dipterocarpus obtusifolius showed significant positive skewness (coefficient s=1.52, p<0.05) and large kurtosis (coefficient κ=2.93, p<0.05), with a peak in the 10–15 cm DBH class and a long tail indicating several rare adults (Fig. 4a). The diameter distribution of D. obtusifolius did not differ significantly from a log-normal distribution (Kolmogorov-Smirnov test, n=59, D=0.07, p=0.150). The best fitting parameters with a 95% CI were 2.63 (2.50–2.76) and 0.51 (0.43–0.61) for μ and σ, respectively. The species-specific diameter Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh 115 116 Ito E. et al. Table 1 Transition of DBH size classes and conditions over the 11-year study period (2003–2014) for stems in study plot, Kampong Thom Province, Cambodia. Stem size class in 2003 Stem size class in 2014 Condition in 2014 5–10 cm DBH Alive b ≥10cm DBH 5–10 cm DBH 16 20 Dieback 0 1 Dead 3 14 Cut 3 0 Collateral damage 1 0 ≥10 cm DBH Alive 6 43 0 0 Dead 0 6 Cut 1 6 Collateral damage 0 1 42 56 23 Total 58 Dieback Total a <5 cm DBHa 63 121 Not enumerated in 2003; b Dieback of primary stems reduced the height of the stem by less than 1.3 m. distributions of associated species were significantly different from D. obtusifolius (G. laccifera, n=6, D=0.85, p=0.010, Fig. 4b; S. oblatum, n=4, D=0.42, p=0.015, Fig. 3d; M. scutellatum, n=11, D=0.93, p=0.010, Fig. 3e), except for P. anamensis (n=8, D=0.23, p=0.150, Fig. 4c). Visual inspection suggested that G. laccifera had a distinct distribution skewed to the larger size class (Fig. 4b), and M. scutellatum had a distinct distribution limited to the 5–10 cm DBH class (Fig. 4e). Detailed structure data for each species are given in Appendix 2. Annual diameter increments during the 2003–2014 censuses averaged 0.14 ± 0.12 cm y–1 and 0.11 ± 0.08 cm y–1 for all of the specimens (n=69) and the D. obtusifolius (n=48) trees that survived over the course of the censuses. Diameter increments for trees that survived the 2003– 2014 censuses were significantly higher between the 2011 and 2014 censuses (0.18 ± 0.16 cm y–1) than between the 2003 and 2011 censuses (0.12 ± 0.12 cm y–1) (Wilcoxon signed rank test, p<0.0001). Plotwise and species-specific diameter increments for each size class are described in Appendix 3. Visual inspection suggested that P. anamensis and C. calaba var. bracteatum showed relatively high increments. The bark of M. scutellatum was often peeling, which resulted in highly variable increments. Basal areas of stems with DBH ≥5 cm and ≥10 cm increased from 11.24 to 12.31 m2 ha−1 and from 10.54 to 11.74 m2 ha−1 between the 2003 and 2011 pre-logging censuses, respectively (Appendix 2). Emissions of C for dying individuals with DBH ≥5 cm and ≥10 cm during the 2003–2014 census were 0.36 Mg C 0.24ha–1 and 0.25 Mg C 0.24ha–1, respectively (Table 2). The DBH classes for dying trees of five representative tree species in the study plot during the 2011–2014 censuses are described in Fig. 4. Tree C increments were larger than C emissions from dead trees from 2003 to 2011; thus, tree C stock in the stand slightly increased (from 11.26 to 12.60 MgC 0.24 ha–1 for trees with DBH ≥5 cm, Table 2). Aboveground tree C stock estimates derived from several equations are also shown in Table 2. Annual above-ground biomass increments in the study plot were lower than estimated values from previously reported regression relationships between the annual above-ground biomass increment and the above-ground biomass (Fig. 5). The annual above-ground biomass increment in this study plot (1.61 Mg ha-1 y-1, estimated using Eq. 5) was 48% of the expected value derived from the logarithmic relationship developed by Top et al. (2004a) (3.38 Mg ha-1 y-1) given an initial above-ground biomass of 113.4 Mg ha-1 (estimated using Eq. 5). The annual above-ground biomass increment (1.08 Mg ha-1 y-1, estimated using Eq. 6) was 32% of the expected value derived from the linear relationship reported by Kiyono et al. (2017) (3.34 Mg ha-1 y-1), given an above-ground biomass of 69.2 Mg ha-1 (estimated using Eq. 6). Estimates of stand structure and tree biomass are summarised and compared to previous studies of Cambodian forest in Table 3. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 109–127 Size increment and C emissions Carbon dynamics in dry dipterocarp forest Table 2 Plot-wise tree carbon (C) dynamics in study plot, Kampong Thom Province, Cambodia. Item (MgC 0.24 ha–1) C stock Census year DBH size criteria (cm) 2003 ≥5 2011 2014 C emissions from dead trees 2003–2011 2011–2014 a C emissions from logging and collateral damage 2011–2014 C increment 2003–2011 Total biomass Above-ground biomass Eqs. 3a Eqs. 3a Eq. 5b Eq. 6c 11.26 8.8 14.0 8.3 ≥10 10.93 8.6 13.6 8.0 ≥5 12.60 9.87 15.53 9.27 ≥10 12.32 9.65 15.24 9.05 ≥5 7.56 5.9 9.6 5.6 ≥10 7.28 5.7 9.4 5.4 ≥5 0.12 0.10 0.14 0.09 ≥10 0.06 0.05 0.07 0.05 ≥5 0.24 0.19 0.29 0.18 ≥10 0.19 0.15 0.24 0.14 ≥5 5.37 4.21 6.33 3.90 ≥10 5.36 4.19 6.30 3.88 ≥5 1.42 1.12 1.66 1.04 ≥10 1.31 1.02 1.54 0.95 Monda et al. (2016); b Brown (1997) used in Top et al. (2004a); c Kiyono et al. (2011) used in Kiyono et al. (2017). Cutting operations in sandy dipterocarp forests ≥10 cm declined to 67.3% (from 12.31 to 8.29 m2 ha−1) and 65.7% (from 11.74 to 7.71 m2 ha−1), respectively (Appendix 2). Basal area depletion as a result of cutting and collateral damage was 34.4% of the initial area (4.23 of 12.31 m2 ha−1). Total tree C stock for trees with DBH ≥5 cm was 12.60 Mg C 0.24 ha–1 in the 2011 census, where 5.37 Mg C 0.24 ha–1, 0.006 Mg C 0.24 ha–1 and 0.24 Mg C 0.24 ha–1 were lost to logging, damage and natural mortality until 2014, respectively (Table 2). Logging intensity and collateral damage associated with cutting are summarised and compared to previous tropical forest studies in Table 4. Logging by unknown parties occurred at the study site between November 2012 and February 2014, likely in late 2013 or early 2014, judging from the freshness of stumps. Ten logged trees were scattered within the 0.24 ha plot (Fig. 2). Five large logged stems were either removed from the plot or left behind partially sawn. Another five small fallen trunks were left behind; their locations were not related to visible skid trails, which indicated normal removal operations. The average DBH of the former was 50.1 cm (35.9 and 46.9 cm for D. obtusifolius, n=2; 54.3– 57.8 cm for G. laccifera, n=3) in the 2011 census (Fig. 4a, b). The average DBH of the unlogged trees of these two species was 16.4 cm (range: 5.3–39.0 cm for D. obtusifolius, n=54; 23.4–47.1 cm for G. laccifera, n=3). The average DBH of the latter was 8.2 cm (range: 5.5–13.8 cm for D. obtusifolius, n=3; 5.9 cm for C. calaba var. bracteatum, n=1; 4.9 cm for P. anamensis, n=1) in the 2011 census (Fig. 4a, c; C. calaba not shown). In addition to the logged trees (n=10), logging operations caused collateral damage to remaining individuals (Figs 2 & 4). Broken-stemmed trees (n=2) occurred in the vicinity of logged tree stumps; a form of damage directly attributable to cutting operations. From 2011 to 2014, densities of trees with DBH ≥5 cm and ≥10 cm fell 87.6% (from 97 to 85 stems 0.24 ha−1) and 80.3% (from 61 to 49 stems 0.24 ha−1), respectively (Fig. 3). Basal areas of individuals with DBH ≥5 cm and Sandy dipterocarp forests are characterised by the strong dominance of D. obtusifolius, with poor associated species richness (Appendix 1; Hiramatsu et al., 2007; Tani et al., 2007). Generally, numbers of dying and recruited trees of D. obtusifolius were similar in our study, suggesting a stable dynamic, resulting in a positively skewed lognormal distribution (Fig. 4a). These results are not inconsistent with previous reports of the species adapting well to edaphic and meteorological conditions specific to sandy dipterocarp forests (Norisada & Kojima, 2005a, 2007; Miyazawa et al., 2014a, 2014b). However, it remains Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh Disc ussion Stand structure and dynamics 117 118 Ito E. et al. Table 3 Stand structure and tree biomass estimates reported in previous studies of Cambodian deciduous forests. This study Index (unit) Tree density (trees ha–1) 2 –1 (pre-logging) Previous studies 379–433 626 Forests and measurement details in previous studiesa References Mixed deciduous forest in Kampong Thom- Sandan; trees with DBH >5 cm; tree components lacking Do, Di, Dt, Ss, and Ta Kimphat et al. (2002a) Basal area (m ha ) 11.2–12.3 32.0 Standing volume (m3 ha–1) 74.1–83.0 179.2 113.4–127.0 189 Deciduous forest in Kampong Thom; trees with DBH >10 cm; forest details unknown Top et al. (2004) Deciduous forest in Mondulkiri-Seima; trees with DBH >5 cm; dominated by Dt Khun et al. (2012), Sasaki et al. (2016) Deciduous forest in Preah Vihear, Kratie-Snoul, Kratie-Sandan; trees with DBH >5 cm; dominated by Dt, Di, So, and Ta Kiyono et al. (2010) Aboveground biomass (Mg ha–1) Basal area (m2 ha–1) 11.2–12.3 24.8 Standing volume (m3 ha–1) 74.1–83.0 192.1 Above-ground C stock (Mg C ha–1) 36.8–41.1 95.1 Total (above-ground + belowground) biomass (Mg ha–1) 131.7–147.6 257.8 ± 92.0b Total tree C stock (Mg C ha–1) 66.7–74.8 55.2 ± 6.9c, 56.2 ± 6.7d Deciduous forest in Ratanakiri, Siem Reap, Kampong Thom, Kratie; trees with DBH >7.5 cm; common species are Do, Dt, Ss, So, and Ta Samreth et al. (2012) Total biomass (Mg ha–1) 93.9–105.0 32.2–158.9 Deciduous dipterocarp forest in Kratie; trees with DBH >5 cm; forest details unknown Monda et al. (2016) Annual diameter increment (cm y–1) 0.14 ± 0.12e 0.12 ± 0.08f 0.19 ± 0.15g 0.32 Deciduous forest in Kampong Thom; trees with DBH > 3 cm; commercial species; forest details unknown FA (2014) 0.45 Same; in Kratie 0.56 Same; in Siem Reap 0.14 Same; in Ratanakiri Do = Dipterocarpus obtusifolius; Di = D. intricatus; Dt = D. tuberculatus; Ss = Shorea siamensis; So = S. obtusa; Ta = Terminalia alata. b Mean ± SD. c 1998 census, Mean ± SE. d 2000–2001 census, Mean ± SE. e All species, 2003–2014, Mean ± SD. f D. obtusifolius, 2003–2014, Mean ± SD. g Trees with DBH >30 cm, D. obtusifolius and G. laccifera, 2003–2011, Mean ± SD. a unclear why D. obtusifolius did not expand to occupy ground spaces which lacked tree canopy occupants (Fig. 2) and this is unlikely to be related to low-light conditions or a lack of flowering trees. Flowering was observed in much smaller trees (minimum diameter for flowering 11.8 cm; Ito et al., 2016) than other dipterocarps (≥50 cm, Sist et al., 2003b). Mycorrhizal limitation is a potential factor limiting recruitment, as symbiotic fungi promote the survival and growth of juveniles (Alexander et al., 1992). As adult dipterocarps are a source of mycorrhizal colonisers that infect juvenile plants, logging of adult trees reduces the size of the fungal source pool. The average mortality of trees with DBH ≥10 cm in the tropics is well documented (1.81 ± 0.16% y–1, mean ± 95% CI; Lewis et al., 2004). Our study site had very low tree mortality for these individuals over the 11-year census (1.03% y–1). High mortality for trees in the 5–10 cm DBH class (3.69% y–1) was mainly due to high turnover of some shrubby species (e.g., M. scutellatum; Fig. 4e). Numbers of recruited trees were balanced with the number of dying trees for trees with DBH ≥5 cm and ≥10 cm. Plotwise stem density was condequently relatively stable until the 2012 census (Fig. 3). This implies that the study forest is likely to be sustained in the absence of anthropogenic disturbance. Over the course of the study, we found that plot-wise mortality and recruitment rates were relatively high between the 2011–2014 censuses, followed by the 2008–2011 censuses and the 2003–2008 censuses (Fig. 3). Diameter increments for trees that survived during the course of the study were also higher during 2011–2014 than 2003–2011 (Appendix 3). The casual relationship underlying these trends in stand © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 109–127 Carbon dynamics in dry dipterocarp forest Table 4 Collateral damage associated with forest logging reported in previous studies. This study Previous studies No. of collaterally damaged trees per felling of a single tree (tree tree–1) <1 ca. 10 Ratio of carbon stock, collateral trees to logged trees (%) 3.2 Carbon stock loss to collateral tree damage (Mg C ha–1) Index of collateral damage (unit) Conditions of forests and logging operations in previous studies References Closed-canopy neotropical forests with DBH 60–140 cm harvested Jackson et al. (2002), Feldpausch et al. (2005) 110 Amazonian humid tropical forest Feldpausch et al. (2005) 0.7 3.2 Amazonian humid tropical forest Feldpausch et al. (2005) Cutting intensity normalised by proportion of damaged trees (2.5% of original tree population) (trees ha–1) 20.8 <4 East Kalimantan forest Sist et al. (1998) Proportion of canopy lost at cutting intensity of 50 m3 ha–1 (%) 20.8 39.1 Ratio of skidding damage of original stand area (%) Not measured 22.7 25 dynamics are unknown, but they may reflect changing climatic or edaphic conditions. Regression from 11 previous studies Pereira et al. (2002) including a Brazilian moist tropical forest, conventional cutting operations Same, reduced-impact logging operations Pereira et al. (2002) Dense evergreen forest in Indonesian Borneo Sist et al. (2003a) Stand C storage and increments of the expected values for initial aboveground biomass using regression relationships (Fig. 5). The absolute and relative low C increment in our study plot suggests that sandy dipterocarp forests might need to be stratified into Sandy dipterocarp forests are characterised by an open structure (Rundel, 1999; Hiramatsu et al., 2007; FA, 2011). This is consistent with lower plot-wise C storage and its relating forest structure indices found in the study plot compared with various Cambodian deciduous forests (Table 3). Given the positive relationship between annual above-ground biomass increment and above-ground biomass (Top et al., 2004a; Kiyono et al., 2017), the biomass increment in sandy dipterocarp forests is expected to be lower than other types of deciduous forests. The mean increment for above-ground stand biomass for the average initial above-ground stand biomass has been estimated to be 4.59 Mg ha-1 y-1 for deciduous forests (Top et al., 2004a), and 4.79 Mg ha-1 y-1 for unstratified forest categories (Kiyono et al., 2017). These estimates are fairly high compared to the values estimated in our study (1.61 Mg ha-1 y-1 and 1.08 Mg ha-1 y-1, using Eqs. 5 and 6, respectively). The latest application of the gain–loss method nationally in Cambodia assumed that a more conservative mean annual increment would be identical to all of the forest categories (1.5 MgC ha–1 y–1, Sasaki et al., 2016), which was twice our study estimate (0.74 MgC ha-1 y-1, using Eq. 3). It is also noteworthy that the above-ground biomass increments in our study plot were 48% and 32% Fig. 5 Relationship between plot-wise annual above-ground biomass increment and initial above-ground biomass. Dotted and solid lines represent regression lines of the relationships reported by Top et al. (2004a) and Kiyono et al. (2017), respectively. Circles indicate a combination of annual above-ground biomass increment and initial aboveground biomass in study plot. Open and closed circles indicate biomass estimation using equations identical to Top et al. (2004a) and Kiyono et al. (2017), respectively. Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh 119 120 Ito E. et al. subdivisions of deciduous forests when implementing the gain–loss approach. Given the low mortality and balanced recruitment of our study plot, the relatively low increment suggests that tree growth was limited. Though seemingly lower diameter increments were found at our study site than those reported for Cambodian deciduous forests (FA, 2004), lack of detailed information for the latter prevent meaningful comparisons (Table 3; Appendix 3). We still do not know why sandy dipterocarp forest has a relatively low increment, although this is not related to light availability. Dipterocarpus obtusifolius physiologically adapts to seasonal flood and drought conditions (Miyazawa et al., 2014a, 2014b), high irradiance levels (Norisada & Kojima, 2005a), and high temperatures (Norisada & Kojima, 2007). The nutritional responses of the species are also similar to other canopydominating dipterocarp species in sandy soils (Norisada & Kojima, 2005b). Acrisols underlying sandy dipterocarp forests are generally nutrient-poor soils compared to more nutrient-rich plinthite in dry dipterocarp forests on the east bank of Mekong (Toriyama et al., 2007a, 2010). Thus, growth of D. obtusifolius might be limited by the nutrient-poor edaphic conditions of our study site. Logging and its impact on C stock The four logged and fallen tree species in our study are often targets for both timber and fuel extraction (Top et al., 2004b; San et al., 2012). The wood of D. obtusifolius is graded as durable (Grade II: MAFF, 2005) and subject to commercial exploitation (Kim Phat et al., 1999). Large logged trees of D. obtusifolius and G. laccifera in the studied plot were possibly used for timber, while the small trunks (<30 cm DBH) left behind may have been felled for fuel (Top et al., 2004b) or used to test chainsaw performance. Over-exploitation of forest resources has been reported in the region (KimPhat et al., 2001; Top et al., 2004c). Basal area depletion from logging in our study plot (34.4%) was above the indicative criteria for sustainable logging operations (<15%) proposed in East Kalimantan (Sist & Nguyen-Thé, 2002). For Cambodian mixed forests, Kimphat et al. (2002a) tentatively proposed a sustainable harvest potential as selective felling of 30% of the stand volume during a 30-year cutting cycle. Logging in the study plot removed substantial C stocks (5.37 MgC 0.24 ha–1, 42.6% of the pre-logging value), which is equivalent to a C increment of 30 years (accrual rate 1.42 Mg C 0.24 ha–1 8y–1; Table 2). Notably, the annual tree C stock increment would decline after cutting operations. A sustainable cutting cycle prediction should be based on the remaining basal area after felled trees are deducted (Sist et al., 2003c; Kimphat et al., 2004). © Centre for Biodiversity Conservation, Phnom Penh Selective logging, particularly in operations without a sustainable management plan, is frequently associated with serious collateral forest damage (Pereira et al., 2002; Sasaki & Putz, 2009). The extent of collateral damage in our study plot, however, appeared to be less than that reported for logging operations in other tropical forests (Table 4). This is obviously due to the low tree density of our study plot. Tree density and basal area for individuals with <10 cm DBH values in previous studies were ca. 525 trees ha-1 and 57 m2 ha–1, respectively (Sist et al., 1998; Pereira et al., 2002; Feldpausch et al., 2005; Table 4). The respective values for our study plot were rescaled to 254 trees ha–1 and 11.7 m2 ha–1, respectively. Such low tree densities and basal areas are found in other sandy dipterocarp forests (e.g., 235 trees ha–1 and 5.5 m2 ha–1: FA, unpublished data). Directional felling without collateral damage to other trees is greatly simplified in sparse forests (Feldpausch et al., 2005). Skid damage could also be minimised with basic requirements for logging roads in sparse forests compared to dense forests (Sist et al., 2003a). How intensive logging alters stand dynamics in sandy dipterocarp forests is still unknown. Logging has the potential to accelerate the growth of surviving trees (Putz et al., 2001), as light conditions improve when a dense forest is thinned (Kao et al., 2011). However, our opencanopied sandy dipterocarp forest likely had unlimited light availability. Along increasing environmental stress gradients, sandy dipterocarp forests become increasingly open in structure and lower in stature, grading eventually into savanna woodlands with decreasing woody cover (Rundel, 1999). Further studies are necessary to clarify whether over-exploitation increases environmental stress and leads to further forest degradation. Ack now le dge m e nt s This paper reports results obtained by the emergency project to develop the structure of promoting REDD action supported by the Forestry Agency Japan. The authors are deeply indebted to H.E. Dr Ty Sokhun, Secretariat of State and to H.E. Dr Chheng Kimsun, Head of Forestry Administration at the Ministry of Agriculture, Forestry, and Fisheries for permission to use permanent sample plot data and undertake field research. The authors also thank Dr T. Toma for helpful comments, and Dr Y. M. Ito for statistical advice. We deeply thank two anonymous reviewers for constructive comments and corrections on the manuscript. 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Family (APG III) Species a Khmer name a Anacardiaceae Gluta laccifera (Pierre) Ding Hou Krouel Basic Data density b c (kg m−3) source 689 Cambodia Annonaceae Xylopia vielana Pierre Kray Krahom 570 Default Calophyllaceae Calophyllum calaba L. var. bracteatum (Wight) P.F.Stevens Phaong 570 Default Chrysobalanaceae Parinari anamensis Hance Thlok 641 Cambodia Dipterocarpaceae Dipterocarpus obtusifolius Teijsm. ex Miq. Tbeng 650 Vietnam Melastomataceae Memecylon scutellatum (Lour.) Hook. & Arn. Phlorng 570 Default Moraceae Artocarpus sp. Kroung 570 Default Myrtaceae Syzygium oblatum (Roxb.) Wall. ex A.M.Cowan & Cowan Pring 570 Default Pentaphylacaceae Anneslea fragrans Wall. Sorphy 634 Myanmar Rubiaceae Catunaregam tomentosa (Blume ex DC.) Tirveng. Ror Veang 570 Default Symplocaceae Symplocos cochinchinensis subsp. laurina (Retz.) Nooteboom Luot 538 Cambodia Scientific names and Khmer names refer to Toyama et al. (2013); Chave et al. (2009) and Zanne et al. (2009) were used for data selection; Country name indicates place where original data were obtained, while default indicates the default value for tropical Asia (Brown, 1997). a © Centre for Biodiversity Conservation, Phnom Penh b c Cambodian Journal of Natural History 2017 (1) 109–127 Carbon dynamics in dry dipterocarp forest Appe ndix 2 Spe c ie s-spe c ifi c ba sa l a re a a nd st e m de nsit y pe r DBH cla ss in t he fi rst (2 0 0 3 ), pre -logging (2 0 1 1 ), a nd post -logging (2 0 1 4 ) c e nsuse s at st udy plot , K a m pong T hom Provinc e , Ca m bodia . 2003 (first) census. Basal area is given as m2 ha-1 and stem density (in parentheses) is given as trees 0.24 ha-1. Only genus names are shown (see Appendix 1 for full scientific names). DBH class (cm) Genus 5–10 10–20 20–30 30–40 40–50 50–60 Total Gluta – (–) – (–) 0.15 (1) – (–) 2.07 (3) 1.81 (2) 4.03 (6) Xylopia – (–) – (–) – (–) – (–) – (–) – (–) – (–) Calophyllum – (–) – (–) – (–) – (–) – (–) – (–) – (–) Parinari 0.07 (3) 0.06 (1) 0.44 (2) – Dipterocarpus 0.34 (18) 2.06 (28) 1.31 (8) 1.36 Memecylon 0.16 (13) – (–) – (–) – Artocarpus 0.07 (5) 0.03 (1) – (–) – Syzygium – (–) 0.33 (4) 0.17 (1) Anneslea – (–) 0.06 (1) – (–) Catunaregam 0.03 (2) – (–) – Symplocos 0.03 (1) – (–) – 0.70 (42) 2.54 (35) 2.07 Total (–) – (–) – (–) 0.57 (6) 0.69 (1) – (–) 5.76 (58) (–) – (–) – (–) 0.16 (13) (–) – (–) – (–) 0.10 (6) – (–) – (–) – (–) 0.50 (5) – (–) – (–) – (–) 0.06 (1) (–) – (–) – (–) – (–) 0.03 (2) (–) – (–) – (–) – (–) 0.03 (1) (12) 1.36 (3) 2.76 (4) 1.81 (2) 11.24 (98) (3) 2011 (pre-logging) census. Basal area is given as m2 ha-1 and stem density (in parentheses) is given as trees 0.24 ha-1. DBH class (cm) Genus 5–10 10–20 20–30 30–40 40–50 50–60 Total Gluta – (–) – (–) 0.18 (1) – (–) 1.38 (2) 2.90 (3) 4.46 (6) Xylopia – (–) – (–) – (–) – (–) – (–) – (–) – (–) Calophyllum 0.04 (3) – (–) – (–) – (–) – (–) – (–) 0.04 (3) Parinari 0.03 (2) 0.20 (4) 0.20 (1) 0.41 (1) – (–) – (–) 0.84 (8) Dipterocarpus 0.30 (16) 2.30 (30) 1.52 (9) 1.39 (3) 0.71 (1) – (–) 6.22 (59) Memecylon 0.14 (12) – (–) – (–) – (–) – (–) – (–) 0.14 (12) Artocarpus 0.03 (2) – (–) – (–) – (–) – (–) – (–) 0.03 (2) – (–) 0.27 (3) 0.18 (1) – (–) – (–) – (–) 0.46 (4) Syzygium Anneslea Catunaregam Symplocos Total – (–) 0.06 (1) – (–) – (–) – (–) – (–) 0.06 (1) 0.03 (2) – (–) – (–) – (–) – (–) – (–) 0.03 (2) – (–) 0.04 (–) – (–) – (–) – (–) – (–) 0.04 (1) 0.57 (36) 2.87 (39) 2.08 (12) 1.8 (4) 2.09 (3) 2.9 (3) 12.31 (97) Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh 125 126 Ito E. et al. Appe ndix 2 Cont ’d 2014 (post-logging) census. Basal area is given as m2 ha-1 and stem density (in parentheses) is given as trees 0.24 ha-1. DBH class (cm) Genus 5–10 20–30 30–40 40–50 50–60 Total – (–) – (–) 0.19 (1) – (–) 1.45 (2) – (–) 1.64 Xylopia 0.04 (4) – (–) – (–) – (–) – (–) – (–) 0.04 (4) Calophyllum 0.06 (3) – (–) – (–) – (–) – (–) – (–) 0.06 (3)a Parinari 0.02 (1) 0.24 (4) 0.21 (1) 0.44 (1) – (–) – (–) 0.91 (6) Dipterocarpus 0.31 (16) 1.92 (23) 1.74 (10) 0.98 (2) – (–) – (–) 4.95 (51) Memecylon 0.12 (10) – (–) – (–) – (–) – (–) – (–) 0.12 (10) Artocarpus – (–) – (–) – (–) – (–) – (–) – (–) – (–) Syzygium – (–) 0.16 (2) 0.33 (2) – (–) – (–) – (–) 0.49 (4) Anneslea – (–) – (–) – (–) – (–) – (–) – (–) – (–) 0.03 (2) – (–) – (–) – (–) – (–) – (–) 0.03 (2) – (–) 0.05 (1) – (–) – (–) – (–) – (–) 0.05 (1) 0.58 (36) 2.37 (30) 2.47 (14) 1.42 (3) 1.45 (2) – (–) 8.29 (85) Gluta Catunaregam Symplocos Total a 10–20 (3) One tree was cut higher than 1.3 m, but still alive. Appe ndix 3 Spe c ie s-spe c ifi c dia m e t e r inc re m e nt s pe r DBH cla ss for t he fi rst (2 0 0 3 ), pre -logging (2 0 1 1 ), a nd post -logging(2 0 1 4 ) c e nsuse s in st udy plot , K a m pong T hom Provinc e , Ca m bodia . 2003–2011 (first to pre-logging) census. Diameter increments are given as mean ±SD, (cm year-1). Sample sizes are given in parenthesis. Only genus names are shown (see Appendix 1 for full scientific names). DBH class (cm) Genus 5–10 10–20 20–30 30–40 40–50 50–60 Total Gluta – (–) – (–) 0.22 (1) – (–) 0.29 ±0.09 (3) 0.33 ±0.04 (2) 0.29 ±0.07 (6) Xylopia – (–) – (–) – (–) – (–) – (–) – (–) – (–) Calophyllum – (–) – (–) – (–) – (–) – (–) – (–) – (–) Parinari 0.33 ±0.14 (3) 0.38 (1) 0.52 ±0.27 (2) – (–) – (–) – (–) 0.40 ±0.18 (6) Dipterocarpus 0.10 ±0.09 (17) 0.11 ±0.07 (28) 0.08 ±0.05 (8) 0.05 ±0.08 (3) 0.07 (1) – (–) 0.10 ±0.07 (57) Memecylon 0.04 ±0.12 (7) – (–) – (–) – (–) – (–) – (–) 0.04 ±0.12 (7) Artocarpus -0.01 (1) – (–) – (–) – (–) – (–) – (–) -0.01 (1) – (–) 0.01 ±0.02 (3) 0.13 (1) – (–) – (–) – (–) 0.04 ±0.06 (4) Syzygium – (–) -0.01 (1) – (–) – (–) – (–) – (–) -0.01 (1) Catunaregam 0.02 ±0.00 (2) – (–) – (–) – (–) – (–) – (–) 0.02 ±0.00 (2) Symplocos 0.24 (1) – (–) – (–) – (–) – (–) – (–) 0.24 (1) Total 0.11 ±0.13 (31) 0.11 ±0.09 (33) 0.17 ±0.19 (12) 0.05 ±0.08 (3) 0.24 ±0.13 (4) 0.33 ±0.04 (2) 0.12 ±0.13 (85) Anneslea © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 109–127 Carbon dynamics in dry dipterocarp forest Appe ndix 3 Cont ’d 2011–2014 (pre-logging to post-logging) census. DBH class (cm) Genus Gluta 5–10 – 10–20 (–) – (–) 20–30 0.24 (1) 30–40 – (–) 40–50 0.38 ±0.22 (2) 50–60 – (–) Total 0.33 ±0.17 (3) – (–) – (–) – (–) – (–) – (–) – (–) – (–) Calophyllum 0.48 ±0.13 (3) – (–) – (–) – (–) – (–) – (–) 0.48 ±0.13 (3) Parinari 0.24 ±0.23 (2) 0.39 ±0.24 (4) 0.34 (1) 0.40 (1) – (–) – (–) 0.35 ±0.19 (8) Dipterocarpus 0.10 ±0.17 (14) 0.20 ±0.14 (24) 0.17 ±0.15 (9) 0.06 ±0.08 (2) – (–) – (–) 0.16 ±0.15 (49) Memecylon 0.10 ±0.15 (9) – (–) – (–) – (–) – (–) – (–) 0.10 ±0.15 (9) Artocarpus – (–) – (–) – (–) – (–) – (–) – (–) – (–) Syzygium – (–) 0.15 ±0.06 (3) 0.39 (1) – (–) – (–) – (–) 0.21 ±0.13 (4) Anneslea – (–) – (–) – (–) – (–) – (–) – (–) – (–) 0.10 ±0.06 (2) – (–) – (–) – (–) – (–) – (–) 0.10 ±0.06 (2) – (–) 0.32 (1) – (–) – (–) – (–) – (–) 0.32 (1) 0.15 ±0.19 (30) 0.23 ±0.16 (32) 0.21 ±0.15 (12) 0.17 ±0.21 (3) 0.38 ±0.22 (2) – (–) 0.20 ±0.17 (79) Xylopia Catunaregam Symplocos Total 2003–2014 (first to post-logging) census. DBH class (cm) Genus 5–10 10–20 20–30 30–40 40–50 50–60 Total Gluta – (–) – (–) 0.22 (1) – (–) 0.29 ±0.01 (2) – (–) 0.26 ±0.04 (3) Xylopia – (–) – (–) – (–) – (–) – (–) – (–) – (–) – (–) – (–) – (–) – (–) – (–) – (–) – (–) Parinari 0.34 ±0.17 (3) 0.39 (1) 0.48 ±0.21 (2) – (–) – (–) – (–) 0.39 ±0.16 (6) Dipterocarpus 0.10 ±0.09 (15) 0.14 ±0.08 (23) 0.10 ±0.05 (8) 0.02 ±0.00 (2) – (–) – (–) 0.12 ±0.08 (48) Memecylon 0.07 ±0.06 (5) – (–) – (–) – (–) – (–) – (–) 0.07 ±0.06 (5) Artocarpus – (–) – (–) – (–) – (–) – (–) – (–) – (–) Syzygium – (–) 0.05 ±0.03 (3) 0.20 (1) – (–) – (–) – (–) 0.09 ±0.08 (4) Anneslea – (–) – (–) – (–) – (–) – (–) – (–) – (–) 0.04 ±0.01 (2) – (–) – (–) – (–) – (–) – (–) 0.04 ±0.01 (2) Calophyllum Catunaregam Symplocos Total – (–) 0.26 (1) – (–) – (–) – (–) – (–) 0.26 (1) 0.12 ±0.12 (26) 0.14 ±0.09 (27) 0.18 ±0.16 (12) 0.02 ±0.00 (2) 0.29 ±0.01 (2) – (–) 0.14 ±0.12 (69) Cambodian Journal of Natural History 2017 (1) 109–127 © Centre for Biodiversity Conservation, Phnom Penh 127 128 Thesis abstracts Re c e nt M a st e r’s T he se s This section presents the abstracts of research theses produced by Royal University of Phnom Penh graduates recently awarded the degree of Masters of Science in Biodiversity Conservation. The abstracts have been edited for English. Dive rsit y a nd populat ion dyna m ic s of m osquit o ve c t ors of Ja pa ne se e nc e pha lit is virus in a pe ri-urba n a nd rura l pig fa r m se t t ing in Ca m bodia PENG Borin ɊɮɍɅʂɋɑɳȶſɆ Ɏ ɪɌɭɑɆȶžȹɸȶɬɌǎȲȳɯɌȲǙɍȹɆɻɭɅ (Japaneseȱ encephalitisȱ virusȱ(JEV)) ȴɬƺɆɭɈƛɳɒɁɭɆȶžȹɸȶɬɌǎȲȳɯɌȲǙɍɳǷǕɑɭɪʆ ɆȲƞɪ ɃɫȲƺɳƙȷˊɅƙɆɳɉɃNjɅɇƐɭȲɎ ɪɌɭɑɳɅɹ ȷɸɴɀȲɡɑɁƛƙȹȪȲȴɬƺɄƗɯɍɎ ɪɌɭɑ ɳɅɹʆ ɳƽɋƳɌɋɍɽȼɫȶɈɪɔɅƎɌȲɊƗɌǏȶɄƗɯɍ Ʌɩȶ NJƒȲɽƷɌȷɊƚȶ JEV ɳɒˊɋɊɮɑɔƷžɊ (Culex) ƺNJƒȲɽƷɌɑɸƴɅɽȷɊƚȶȹɸȶɬ JEV ɳǷNjɅȲƙɊɩɁȲƒɭȶɆɌ ɩɆɃƳɌɌ ɪȲǍɍƽɍɵɅɅȴɌɮɆɅɪɋȲɊƗ ɳɋˊȶ LJɅɔɳȶžɁɈɪdždžNJɈ Ʌɩȶ ɆɴƙɊɆƙɊȫɍ ɆɻɮɈɭɋǔɑƘɭȶɵɅNJƒȲɽƷɌɆȶžȹɸȶɬ JEV ɳǷȲɑɩƽƊɅȷɩȥƃɫɊƙȹȪȲȲƒɭȶɁɸɆɅɽƺɋƙȲȩȶ ɅɩȶȹɅ ɆɃȲƒɭȶƙɆɳɃɑȲɊƕɭƺʆ ɊɮɑƙɁȪɎLJɅƸɆɽȲɭƒȶɌɋɺɳɈɍʑʒɴȳ ȴɬƸɆɽɈɪɴȳȲȲžƽ ƹƒɸʒʐʑʕ ȼɍɽɴȳȲȲžƽ ƹƒɸʒʐʑʖ ɳƽɋ ɳƙɆˊɔdžƐȲɽɳƙȷˊɅɋɆɽƺɆɽʉ ɑɌɭɆNjɅɊɮɑʘʓ.ʕʓʑ ȲǙɍƙɁȪɎLJɅƸɆɽʆ ɊɮɑʑʗƙɆɳɉɃƙɁȪɎLJɅȲɸɀɁɽ ƙǂɈɪɁɸɆɅɽƺɋƙȲȩȶ Ʌɩȶ ʑʕƙɆɳɉɃɳɃȢɁɈɪȹɅɆɃʆ Culexȱgelidus ƺƙɆɳɉɃƸɆɽLJɅɳƙȷˊɅƺȶɳȴɳǷɁɸ ɆɅɽƺȶƙȲȩȶ ɁɸǁȶɤƘʓʖ.ʗʘ% ɵɅɡȲɁƎɑɌɭɆ ɅɩȶɳƙȷˊɅɆdžƐɆɽȴɬɑNjȹɩȲɵɅƙȲȩɊɌȶ C.ȱ vishnui (ʒʘ%) C.ȱ tritaeniorhynchus (ʑʙ.ʘ%) Culexȱ sp. (ʑʑ.ʕ%) Anophelesȱ sp. (ʑ.ʕ%) Ʌɩȶȱ C.ȱ quinquefasciatus (ʑ.ʔ%)ʆ ɑNjȹɩȲɵɅƙȲȩɊɌȶ C.ȱ vishnui (ʕʑ.ʖ%) ȴɬƺƙȲȩɊɴȼɍɑɊƓɮɌƺȶɳȴɳǷɃɪ ȹɅɆɃ ɆdžƐɆɽɊȲȴɬ Ʌɩȶ Culexȱsp. (ʑʗ%) C.ȱgelidus (ʑʔ.ʙ%)ȱC.ȱtritaeniorhynchus (ʑʐ.ʙ%) C.ȱquinquefasciatus (ʑ.ʘ%) Anophelesȱ sp. (ʑ.ʘ%)ʆ ȷɸɅɯɅɊɮɍɴȼɍLJɅƙɆɊɮɍȲƒɭȶɴȳɅɪɊɯɋʉLJɅɳȲˊɅɳɓˊȶȲƒɭȶɴȳɄƒɮ Ʌɩȶ ɴȳȲȲžƽ ɳɒˊɋɃɸɅȶƺɇƎɍɽ ɇɍɆɻɹljɍɽȼɍɽɑɭȳNJɈɑɁƛƙȹȪȲ ɑɁƛɳƵɅɩȶƙȲɆɪ ɳǷȲƒɭȶɁɸɆɅɽɑɩȲǜ (ƺɈɩɳɑɑɳǷ ɁɸɆɅɽƺɋƙȲȩȶ) ɳǷȲƒɭȶȲɸɓɭȶɳɈɍɳdžɹʆ dždžNJɈ Ʌɩȶ ɆɴƙɊɆƙɊȫɍɆɻɮɈɭɋǔɑƘɭȶɵɅNJƒȲɽƷɌɆȶžȹɸȶɬ ɳǷȲƒɭȶƙɆɳɃɑȲɊƕɭƺ ɈɩɁǁɑɽDŽɊDŽɌɤƘNjɅƳɌɑɩȲǜɆɴɅƏɊ ɳɃȢɁʆ Abst ra c t Japanese encephalitis virus (JEV) is a leading cause of encephalitis in Asia. Several water bird species are reservoirs for the virus, whereas pigs act as hosts for JEV and several Culex mosquitoes are important vectors for its transmission. As understanding of interactions between JEV hosts and vectors remains limited in the context of expanding urbanization, we investigated the diversity and population dynamics of potential JEV vectors in a peri-urban and a rural pig-farming setting in Cambodia. Mosquitoes were sampled for 12 months from July 2015 to July 2016 using consecutive night traps which captured a total of 83,531 mosquitoes. Seventeen mosquito species were recorded in the peri-urban study site and 15 mosquito species in the rural study site. Culex gelidus was the most abundant species in the former, representing 36.7% of individuals, followed by members of the C. vishnui subgroup (28%), C. tritaeniorhynchus (19.8%), Culex sp. (11.5%), Anopheles sp. (1.5%), and C. quinquefasciatus (1.4%). Members of the C. vishnui subgroup (51.6%) were most abundant at the latter study site, followed by Culex sp. (17%), C. gelidus (14.9%), C. tritaeniorhynchus (10.9%), C. quinquefasciatus (1.8%) and Anopheles sp. (1.8%). Numbers of mosquitoes sampled each month increased in December and July and likely impacted pig and/or cattle livestock in the study sites (particularly the peri-urban site) during these periods. The diversity and population dynamics of mosquito vectors in Cambodia warrant further study. Citation: Peng B. (2017) Diversity and population dynamics of mosquito vectors of Japanese encephalitis virus in a peri-urban and rural pig farm setting in Cambodia. Cambodian Journal of Natural History, 2017, 128. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 128–133 Thesis abstracts T he c urre nt a nd hist oric a l im pa c t of c om m unit y fi she rie s on se a t ur t le s in Koh Sda ch a nd Chrouy Sva y, sout hw e st e r n Ca m bodia SAN Satya ɊɮɍɅʂɋɑɳȶſɆ ɔɳɀˊ Ǝ ȲɑɊɭƙɃʕƙɆɳɉɃƙɁȪɎLJɅɳȴǒƀɍɽƺƙɆɎɁƎɩǒȝɑƎɳǷƙɆɳɃɑȲɊƕɭƺ ɳDŽɹɆɪƺNjɅɴɁɈɪɌƙɆɳɉɃ(ɔɳɀˊ Ǝ ȲɍƗɩȷ Chelonia mydasɅɩȶɔɳɀˊƎ ȲƙƳɑɽ Eretmochelys imbricata) ƙɁȪɎLJɅɳȴɳɄƛˊȲɸɀɁɽƙǂȲƒɭȶɔɸɓɭȶʑʐƹƒɸȷɭȶɳƙƳɋȲʁɳƽɋʆ ƙɆɳɉɃƴȶ ɳȼˊɊƙɁȪɎLJɅɳȴƸɁɽɃɭȲǃƺƙɆɳɉɃɌȶɳƙƵɹ ɳɒˊɋƙɆɳɉɃɆdžƐɆɽƺƙɆɳɉɃɌȶɳƙƵɹɄƂɅɽɄƂɌʆ ɆɻɮɈɭɋǔɑƘɭȶɌɆɑɽƙɆɳɉɃDŽɸȶɈɪɌ ƙɁȪɎ LJɅɳȴɳȹȟƺȲɽǃNjɅƳɌɂɋȷɭɹɳƽɋǒɌƳɌƺɆɽɳƽɋɵȷȼɅƘɅɩȶƷɆɽɳǷȲƒɭȶəɆȲɌɀɿɳɅǒɃƙɁɪʆ ɳɓˊɋɈɪȲƙɊɩɁɵɅƳɌƷɆɽ ɳƙljɹǏƺƳɌƺɆɽɳƽɋɵȷȼɅƘȲƒɭȶȼɸɳɀˊɌƳɌɳɅǒɃɑɊɭƙɃ NjɅƳɌȼɫȶɁɩȷɁɯȷɳǷ ƳɌɑɩȲǜɌɆɑɽȳɭɸƇNjɅɆɸɀȶɆɳȶžˊɅƳɌ ɋɍɽȼɫȶɈɪƙɆɳɉɃəɆȲɌɀɿɳɅǒɃɴȼɍLJɅɳƙɆˊȲɭƒȶƙɆɳɃɑȲɊƕɭƺ Ʌɩȶ ȲƙɊɩɁɆɻɹljɍɽɳɇƞȶʉɳǵɳɍˊɆɻɮɈɭɋǔɑƘɭȶɌɆɑɽɔɳɀˊ Ǝ Ȳ ɑɊɭƙɃʆ ɃɩɅƒɅʂɋƙɁȪɎLJɅƙɆɊɮɍǂɊɌɋɺȲƙɊȶɑɸɀɯɌɑNjƖɑɿɔȲ ƒ ɳɅǒɃȷɸɅɯɅʒʒʔdžȲɽ ɳǷȲƒɭȶɉɮɊɩȷɸɅɯɅʔ ɳǷȯɑȩȲɳƳɹɳɑƎȷ ɅɩȶȯɑȩȲɳƙƺɋǒƛɋ ɳȳɁƎɳƳɹȲɭȶ NJȴɅɩɌɁɪɵɅƙɆɳɃɑȲɊƕɭƺ ɈɪɴȳȲɭɊƖɺȼɍɽɴȳɊɪdž ƹƒɸʒʐʑʖʆ ɳDŽɹƺƙɆɳɉɃəɆȲɌɀɿɳɅǒɃ ɴȼɍLJɅɳƙɆˊNjɅƳɌLjƚɑɽɆɮɌȲɅƚ Ǝ ȶɊȲ ɳɒˊɋɔƒȲɳɅǒɃȳƚɹɳƙɆˊƙLJɑɽɳƙȷˊɅƺȶɊɯɋƙɆɳɉɃ ɑɸǁȻɽɆƷƀɅɩȶɴɇƚɑɅƐɮȷɆɴɆɍ ƙɁȪɎLJɅɳȴɳƙɆˊƙLJɑɽɳƙȷˊɅƺȶɳȴȲƒɭȶȷɸɳǁɊəɆȲɌɀɿɴȼɍLJɅɳƙɆˊɳƽɋɔƒȲɳɅǒɃ ȴɬƙɆɴɒɍƺʔʔ%Ʌɩȶʒʘ% ǂɊɍɸƽɆɽʆ ɳnjȶǂɊȷɳɊƚˊɋɑNjƖɑɅɿ ɑɅƐɮȷɆɴɆɍɇƎɍɽƳɌȴɸǍɊȲɸɴɒȶȼɍɽɔɳɀˊ ž ȲɑɊɭƙɃƴƚɸȶƺȶɳȴ LJɅȷɸɅɯɅʔʑȲǙɍɳƽɋɵȷȼɅƘ ɳɄȢɆɅɫȶɔɯɅLJɅȷɸɅɯɅʙȲǙɍ Ʌɩȶ ɴȼɍǏɊɯɋɊɭȳƸɆɽLJɅɔɳɀˊ Ǝ Ȳ ɁɩȷƺȶɳɅɹȷɸɳljɹƙɆɳɉɃəɆȲɌɀɿɳɅǒɃɳɇƞȶɳɃȢɁʆ ɍɃƊɇɍɵɅƳɌɑɩȲǜɌɆɑɽȳɭɸƇLJɅɆƷƟȻǃ ƳɌǂɊƽɅɳɍˊɔɳɀˊ ž ȲɑɊɭƙɃɴȼɍƸɆɽɳƽɋɵȷȼɅƘǂɊɌɋɺƳɌɳɅǒɃɑɊɭƙɃ Ʌɩȶ ƳɁɽɆɅƏɋȲƙɊɩɁɵɅƳɌƸɆɽɳɅɹ ȴɬƺƳɌƷɌɆdžƐɅɽʆ ƳɌƷɌɳɅɹȴɯɌǍɆɽɆȥɮƃ ɍDŽɸȶƳɌɆɳȶžˊɅƳɌƙɁȫɁɈɩɅɩɁƘƳɌɳɅǒɃȳɭɑȷǙɆɽʆ Abst ra c t Five species of marine turtle are historically known in Cambodia, although only two—green turtle Chelonia mydas and hawksbill turtle Eretmochelys imbricata—have been recorded in the last 10 years. The former species is Endangered and the latter Critically Endangered and populations of both are believed to be decreasing due to incidental capture and subsequent mortality in fishing gear. As little is known about the extent of mortality due to by-catch in marine fisheries, my study aimed to improve understanding of the types of fishing gear used in Cambodia and their relative impacts on sea turtle populations. Data were collected through questionnaire-based interviews of 224 fishermen in four villages in the Koh Sdach and Chrouy Svay districts of Koh Kong Province, southwestern Cambodia, in February–March 2016. Though types of fishing gear used have changed over time and some fishermen use more than one type, shrimp gill nets and ray hooks were most common gear types used, being employed by 44% and 28% of fishermen respectively. According to respondents, ray hooks posed the greatest threat to sea turtles with these alone resulting in a by-catch of 41 turtles, compared to nine reported for single trawling and lesser numbers for other types of fishing gear. My results suggest that monitoring of turtle by-catch in marine fisheries, and measures to reduce this are urgently needed. These should include improved control of illegal fishing. Citation: San S. (2017) The current and historical impact of community fisheries on sea turtles in Koh Sdach and Chrouy Svay, southwestern Cambodia. Cambodian Journal of Natural History, 2017, 129. Cambodian Journal of Natural History 2017 (1) 128–133 © Centre for Biodiversity Conservation, Phnom Penh 129 130 Thesis abstracts An a sse ssm e nt of re m ot e sa m pling m e t hodologie s for e st im at ing be a r populat ions in t ropic a l fore st SIM Sovannarun ɊɮɍɅʂɋɑɳȶſɆ NjɅƴƚȵƗɸɭɈɪɌƙɆɳɉɃɳǷƙɆɳɃɑȲɊƕɭƺȴɬƴƚȵƗɸɭɁɮȷ (Helarctosȱ malayanus) ɅɩȶƴƚȵƗɸɭɄɸ (Ursusȱ thibetanus)ʆ ɈɯȲǏƺƙɆɳɉɃ ɑɁƛɌɑɽɳǷȲƒɭȶɵƙɈ Ʌɩȶ ƙɆɃɹɳȵˊȻɳǷǂɊɁɸɆɅɽƳɌljɌƺɳƙȷˊɅ ɳDŽɹɆɪƺNjɅƳɌȼɫȶɁɩȷɁɯȷɳǷɳɓˊɋɔɸɈɪɆɻɮɈɭɋǔɑƘɭȶɌɆɑɽǏ ɴȼɍLJɅɅɫȶȲɸɈɭȶɂɋȷɭɹɳƽɋǒɌƳɌɆɌLJȻɽɳɄƛˊljɀɩȹƅȲɊƗ ƳɌLJɁɽɆȶɽɃɪȹƙɊȲ Ʌɩȶ ƳɌɳɆˊȲɑɩƽƊɅȷɩȥƃɫɊƴƚȵƗɭɸʆ Ɏ ɪɄɪǒȝɑƎƙɆ ɊɮɍɃɩɅƒɅʂɋ Remoteȱ sampling ȼɮȷƺƳɌȲɁɽƙǂƽɅɞǒƒɊ ɑɸǁȲɴɑɅ Ʌɩȶ Njɻ ɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩ NjɅǒɌɺɑɸƴɅɽ ǁɑɽɑƙNjɆɽLJɻɅɽƙɆNjɀɆɻɮɈɭɋǔɑƘɭȶɑɁƛɵƙɈ ƺɈɩɳɑɑƙɆɳɉɃɌȶƳɌȴɸǍɊȲɸɴɒȶ ɳƙljɹɎ ɩɄɪǒȝɑƎɳɅɹɊɩɅɁƙɊȪɎɤƘɔƒȲȯǒɎ ƙƺɎƸɆɽɑɁƛɳɃʆ ƳɌɑɩȲǜɌɆɑɽȳɭɸƇNjɅɳƵɍɆɸɀȶǏɋɁɵɊƚɈɪƳɌɳƙɆˊƙLJɑɽɎ ɪɄɪǒȝɑƎɳɅɹ ɑƙNjɆɽLJɻɅɽƙɆNjɀɆɻɮɈɭɋǔɑƘɭȶ ƴƚȵƗɸɭȲɭȶɁɸ ƒ ɆɅɽɵƙɈdždžʆ ƳɌƙɆɊɮɍɑɸǁȲƙɁȪɎLJɅɳɄƛˊɳɓˊȶɳǷɃɪǂɸȶɈɪɌɃɩɑɅɩɌɁɪɵɅƙɆɳɃɑȲɊƕɭƺ ƸɆɽɈɪɴȳɊɪdž ȼɍɽɴȳəɑNJ ƹƒɸʒʐʑʖʈ ʑ) əɃǚɅƺɁɩƙȲǏȻɁƓɮȶ ȵɭɸȹɪLjȶ ɳȳɁƎɳƳɹȲɭȶ ʒ) əɃǚɅƺɁɩƙȲǏȻȲǁ Ǝ ɍ ȯɑȩȲɔɮɌɳǒɊ ɳȳɁƎɳljɄȨ ǒɁɽʆ ƳɌƽȲɽNjɻɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩ Ʌɩȶ ƳɌƙɆɊɮɍɑɸǁȲɳǍɊ(ǂɊɌɋɺəɆȲɌɀɿDŽȲɽɋȲɳǍɊ) ɑƙNjɆɽƳɌɎ ɩNJȴDNA ƙɁȪɎLJɅɳɄƛˊ ɳɓˊȶɳǷɃɪǂɸȶNjɅƽȲɽȷɸɀɪɳǷǂɊɃɪǂɸȶDŽɸȶɈɪɌʆ ƴƚȵƗɭɸɄɸɊɩɅƙɁȪɎLJɅɳɄˊƛȲɸɀɁɽƙǂɳɃȲƒɭȶɔɸɓɭȶɳɈɍɑɩȲǜɳDŽɹɆɪƺƴƚȵƗɭɸɄɸ NjɅȷɸɅɮɅɆɯɅȲǙɍ ɆƷƟȻǃƴƚȵƗɸɭɁɮȷNjɅɳƙȷˊɅƺȶɳǷȲƒɭȶɁɸɆɅɽʆ ƙɆɳɉɃɌȶƳɌȴɸǍɊȲɸɴɒȶ Ʌɩȶ ƙɆȺɊƳɌȴɸǍɊȲɸɴɒȶɊɯɋ ȷɸɅɯɅɳɃȢɁȲʁƙɁȪɎLJɅɳɄƛˊȲɸɀɁɽƙǂɇȶɴȼɌ(ǍɆɽɆȥɮƃ ɍDŽɸȶ ȼɸɌ ɪǕɑɭɪ ɴȸžɵƙɈ ȳƐɪȶ ɌNjɸȶ ƴƚɈɈȲ ƴƚɂƇɪɂƗɴȲɎ Ʌɩȶ ƴƚɳɍȟȶNjɑ) ɴȼɍ ɳɅɹƺɉɑƎɭǂȶȼʁɑɸƴɅɽɆƷƟȻɤƘNjɅƳɌɔɉɩɌȲƞɳǷȲƒɭȶɁɸɆɅɽʆ ɔɁƎɑȦƈɀɡȲɁƎƴƚȵƗɭɸɁɮȷɊɩɅǕȷɳɄƛˊLJɅɳɓˊɋ ɈɪɳƙljɹǒƒɊɳǷ ɳɍˊƙɃȪȶɊɩɅƙɁȪɎLJɅɂɁɳƽɋNjɻ ɑɭɪɅɂɁɑƛʂɋƙɆɎɁƎɩɳɃ ɳɒˊɋɑɸǁȲɳǍɊȲʁƙɆɊɮɍɊɩɅLJɅȲƒɭȶɔɸɓɭȶɳɈɍɑɩȲǜʆ ȲɌɀɪɌɮɆɂɁƙɆ ɴɒɍǕȷɆǁ Ǝ ɍɊȲɈɪȷɸɀɪƙɁȪɎLJɅɑɭɪɆɞɋȲɳȷȻɳƽɋɑɁƛɳɇƞȶɳɃȢɁ ƳɌƽȲɽȷɸɀɪɊɩɅƙɁȪɎɃɪǂɸȶɞɑɁƛɌɁɽɳƽɋǒɌɈɅƚɬ ɳɉƚˊȶNjɻ ɑɭɪɅɂɁʆ Ɏ ɪɄɪǒȝɑƎDŽɸȶɈɪɌɁƙɊȪɎɤƘNjɅƳɌɴȲɍɊơɌɳȼˊɊƓɪɆɳȶžˊɅɍɃƊNJɈɳɄƛˊɔɁƎɑȦƈɀɡȲɁƎ Ʌɩȶ LJɻ ɅɽƙɆNjɀɆɻɮɈɭɋǔ ɑƘɭȶƴƚȵƗɭɸʆ Abst ra c t Two bear species occur in Cambodia: sun bear Helarctos malayanus and Asiatic black bear Ursus thibetanus. Both are forest dwelling and occur in many protected areas. Little is known about their populations, which are decreasing due to commercial hunting, habitat loss and bear farming. Remote sampling methods, such as sign-based surveys, genetic sampling and camera trapping, are useful for estimating populations of wildlife and threatened species in particular because they do not require animal handling. My study aimed to assess the utility of these methods for estimating bear populations in forest areas. Sampling was conducted in two sites in southwestern Cambodia from March to June 2016: 1) Kravanh Khang Tbong National Park, Chi Phat Commune, Koh Kong Province; 2) Central Cardamoms National Park, Ou Saom Commune, Pursat Province. Camera trapping and collection of hair samples (through hair-snag traps) for DNA analysis were undertaken at baited locations at both sites. Asiatic black bear was not recorded during the study, although four sun bears were, suggesting the latter species is more abundant in the region. Several other globally threatened or near-threatened species were also recorded (including Asian elephant, dhole, gaur, sambar, clouded leopard, marbled cat and Asiatic golden cat): proof of their conservation importance. Recognition of individual sun bears was not possible because chest markings were not captured in camera trap images and because hair samples were not obtained during the study. Failure to record the bears’ chest markings may have been due to baits being removed by other species, inappropriate positioning of baits or animal movement due to camera flash. Both survey methods require refinement to improve their prospects for facilitating recognition of individual bears and estimation of their populations. Citation: Sim S. (2017) An assessment of remote sampling methodologies for estimating bear populations in tropical forest. Cambodian Journal of Natural History, 2017, 130. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 128–133 Thesis abstracts Est im at ing Asia n e le pha nt Ele pha s m a x im us dist ribut ion pat t e r ns during t he dr y se a son in Pre y La ng Wildlife Sa nc t ua r y, Ca m bodia : im plic at ions for c onse r vat ion a nd fut ure re c ove r y SOUN Visal ɊɮɍɅʂɋɑɳȶſɆ ɴȼɅȹƙɊȲɑɁƛɵƙɈɵƙɈɓȶɽ (IndoȬBurmaȱ ȴɬƺɁɸɆɅɽɵƙɈɳȯǒȶɃɸdžɆɴȼɍɳǷɳɑɑɑɍɽȷɭȶɳƙƳɋ ɑƏɩɁȲƒɭȶɃɪǂɸȶȹɪɎɺȷƙɊȩɹȳƕɑɽɗɀɮ Ƌ ɉɮNj hotspot) ɳɒˊɋɂƗɪʉɳɅɹǏƳɅɽɴɁɌȶȲɸɳɀˊɅƳɌƽȷɽƺɆɸɴɀȲʉ ɳƽɋǒɌƳɌɂɋȷɭɹɵɅȴƙɊɆɵƙɈʆ NjɅƳɌ ɑɩȲǜȯǒɎƙƺɎɊɯɋȷɸɅɯɅɁɮȷDŽȲɽɃȶɅɫȶȹɪɎɺȷƙɊȩɹɳǷɁɸɆɅɽ ɴȼɍɳɈɍɳɎǎɅɩȶɎ ɩǒɍNJɈƳɌƷɌȯǒɎƙƺɎDŽɸȶɳdžɹɑɭɃɴƑ ɁɳǷ NjɅȲƙɊɩɁʆ ƳɌɑɩȲǜɌɆɑɽȳɭɸƇLJɅɳƙɆˊƙLJɑɽƳɌɎ ɩNJȴǂɊɴɆɆ Remoteȱ sensing ȲƒɭȶƳɌǂɊƽɅɌLJɋȼɸɌ ɪǕɑɭɪ (Elephasȱ maximus) ȲƒɭȶɌȼɮɎƙLJɸȶɳǷɁɸɆɅɽɵƙɈɓȶɽ ɳƽɋNjɅɳƵɍɆɸɀȶȲɸɀɁɽɃɪǂɸȶƸɸLJȷɽɑƙNjɆɽƳɌɔɉɩɌȲƞ ɅɩȶǒƎɌȷɸɅɯɅȼɸɌ ɪɳɓˊȶ Ɏ ɩȻȲƒɭȶɁɸɆɅɽʆ ɳƽɋɴɇơȲɳɍˊȼɸɳɀˊɌƳɌɈǚȲɌɀɿǂɊ ȷɸɅɯɅʔ ƙɁȪɎLJɅƽȲɽɆȥɮƃ ɍƵƒɳǷȲƒɭȶȲɊƗɎ ɩɄɪ ArcMap ȯɑDŽɆɽɃɩɅƒɅʂɋɁɸɆɅɽɉɮɊɩǒȝɑƎȷɸɅɯɅʓ Ʌɩȶ ɃɩɅƒɅʂɋɆɌ ɩǒƏɅ MaximumȱEntropy(MaxEnt) ɴȼɍɁɵɊƚɈǚȲɌɀɿɌɆɑɽǏ (replicatedȱruns) ƺɊɄƘɊ ȴɬƺȶʑʕȼȶʆ ƺɄɊƗǂ ɔɳɂɌǁɊɯɋȷɮɍɌɯɊȷɸɴɀȲƳɅɽɴɁƴƚɸȶɳǷȲƒɭȶɊɻɮɴȼɍ ƺɔɳɂɌɴȼɍNjɅɗɃƑɩɈɍƴƚɸȶȷɸɳljɹ ƳɌɈǚȲɌɀɿɈɪɎɁƎNjɅɵɅȼɸɌ ɪǕɑɭɪȲɭƒȶɁɸɆɅɽɳdžɹʆ ȲƙɊɩɁɃɫȲɳɉƚȣȶȲƒɭȶɔɸɓɭȶɳɈɍƙɁƺȲɽɆɸɇɭɁɵɅƙɁɪNjɑƙɆƸɸƹƒɸ LJɅȷɮɍɌɯɊƳɌ ɈǚȲɌɀɿɍơƺȶɳȴȴɬɌɒɮɁȼɍɽʔʔ.ʕ% ȲƒɭȶɳdžɹɴȼɌ ƙɆɉɈɃɫȲɌɯɊȷɸɴɀȲʔʑ.ʘ%ɳǷȲƒɭȶɊɻɮɴȼɍ ɳɒˊɋɔɳɂɌɳɇƞȶʉɳɃȢɁLJɅȷɮɍ ɌɯɊȷɸɴɀȲɁɩȷƺȶʒʐ%ʆ ɊɻɮɴȼɍNjɅɁɵɊƚ ACU ʐ.ʘʕʔ ɆƷƟȻǃƳɌɈǚȲɌNjɅNJɈƙɁɫɊƙɁȪɎʆ ɳƽɋɳƙɆˊɍɃƑɇɍDŽɸȶɳɅɹ ȯɑDŽɆɽraster ƙɁȪɎLJɅɆɴɊƚȶɳǷȲƒɭȶȲɊƗɎ ɩɄɪ ArcMap ɳȼˊɊƓɪɆɳȶžˊɁɴɇɅɃɪȹƙɊȲɑɊȯɑɆɅɩȶɁɸɆɅɽɔɉɩɌȲƞɑɸƴɅɽʆ ɍɃƑɇɍƳɌ ɑɩȲǜɌɆɑɽȳɭɸɆ Ƈ ƷƟȻɈɪNJɈƸɸLJȷɽȲɭȶƳɌȲɸ ƒ ɀɁɽƳɌɌ ɸƴɅɳƽɋɑȲɊƗNJɈɊɅɭɑƞɳǵɵɂƂɔdžȴɁɳǷɴȼɅȹƙɊȲɑɁƛɵƙɈɵƙɈɓȶɽ Ʌɩȶ ɆƷƟȻɈɪɁɯdžɃɪɑɸƴɅɽ ɵɅɍȲſȳʂɀƋǕƳɑDžɁɭ ƙɆɉɈɃɫȲ Ʌɩȶ ȴƙɊɆɵƙɈ ɑƙNjɆɽƳɌɌɑɽǍɅNjɅȹɪɎ ɩɁ Ʌɩȶ ɳȲˊɅɳɓˊȶɎ ɩȻɵɅȷɸɅɯɅ ȼɸɌ ɪǕɑɭɪɳǷɁɸɆɅɽɳɅɹʆ Abst ra c t Prey Lang Wildlife Sanctuary is one of the last remaining areas of lowland evergreen forest in the Indo-Burma hotspot and has become increasingly isolated with reduced forest cover in recent years. Relatively few biodiversity surveys have been conducted at the site and these were limited in duration and scope. My study employed remote sensing analysis to investigate the distribution patterns of Asian elephant Elephas maximus during the dry season in Prey Lang and aimed to identify areas required to conserve and recover remaining populations of the species there. Following a projection process performed in ArcMap, three geographical and four environmental data layers were combined in a Maximum Entropy (MaxEnt) software model whose predictive values were averaged over 15 replicated runs. Not surprisingly, the more a variable contributed to the model, the greater the impact it had in predicting the occurrence of E. maximus at the site. Precipitation during the coldest yearly quarter made the greatest predictive contribution of 44.5%, whereas water sources contributed 41.8% to the model, and the remaining variables collectively contributed <20%. The model had an ACU value of 0.854, implying that it was robust. Using these results, a raster layer was converted in ArcMap to produce a map of habitat suitability and conservation hotspots at the site. The results of my study highlight the necessity of limiting future anthropogenic disturbance at Prey Lang Wildlife Sanctuary and indicate the important roles played by climatic conditions, availability of water and forest cover for survival and recovery of Asian elephant populations at the site. Citation: Soun V. (2017) Estimating Asian elephant (Elephas maximus) distribution patterns during the dry season in Prey Lang Wildlife Sanctuary, Cambodia: implications for conservation and future recovery. Cambodian Journal of Natural History, 2017, 131. Cambodian Journal of Natural History 2017 (1) 128–133 © Centre for Biodiversity Conservation, Phnom Penh 131 132 Thesis abstracts Com pa rat ive de nsit y of gre e n le a fhoppe r N e phot e t t ix vire sc e ns a nd brow n pla nt hoppe r N ila pa r vat a luge ns in ric e fi e lds a round t he Tonle Sa p La ke , Ca m bodia SUOR Kimhuor ɊɮɍɅʂɋɑɳȶſɆ ɑɁƛɊNjȷɳǂƒɁ(Nilaparvataȱlugens)ɅɩȶɊNjȷɵɆɁȶ(Nephotettixȱvirescens) ȴɬƺɑɁƛȷɵƙȶȼʁȷɊƓȶɳɍˊȼɸǁɸȯɑȪɎɳǷɁɸɆɅɽ ǕɑɭɪǕɳȴƒɋɿ ɴȼɍɳɄƛˊɤƘȯɑȪɎɍɮɁǎɑɽɋɬɁȼɮȷƺ ɴɇƒȲȲɊƕɑɽ ɵɇƐɑɫȲ ƚ ȲƙɊɩɁɵɅƳɌɳɄƛˊɌɑƗɪɑɸɳnjȴɅɩȶǕɑɮɁʆ ɊNjȷɳǂƒɁƺɔƒȲ Ɇȥɮƅ ɅɎ ɪɌɭɑɆȶžȹɸȶɬɳɁȟɌɯȻɑƚɫȲ(RaggedȱStuntȱVirus)ɅɩȶɎ ɪɌɭɑȹɸȶɬɳɁȟɃɭɸɑɫȲ ƚ (riceȱgrassyȱstuntȱvirus) ɳɒˊɋɊNjȷɵɆɁȶȴɬ ƺNJƒȲɽƷɌȷɊƚȶɎ ɪɌɭɑɌɯɊNjɅɎ ɪɌɭɑɳɄƛˊɳǕɋɍɮɁǎɑɽɋɬɁ(tungroȱ virus) Ɏ ɪɌɭɑȹɸȶɬɳɁȟɳɍȟȶ(yellowȱ dwarfȱ virus) Ɏ ɪɌɭɑȹɸȶɬ ɳɍȟȶɃɭɸɑɫȲ ƚ (yellowȬorangeȱleafȱvirus) ɅɩȶɎ ɪɌɭɑȹɸȶɬɳɍȟȶɊɩɅƺɆɽǎɆɽ(transitoryȱyellowingȱvirus)ʆ ƳɌɑɩȲǜɌɆɑɽ ȳƇɭɸNjɅɳƵɍɆɸɀȶɳƙɆȣɆɳɄȢɆȼȶɽɑɭɪɳɁɵɅN.ȱlugens Ʌɩȶ N.ȱvirescens ɳǷȲƒɭȶɴȯɑ Ʌɩȶ ȲɸɀɁɽȲǂƎɴȼɍǕȷNjɅɗɃƑɩɈɍȼɍɽ ȼȶɽɑɭɪɳɁɌɆɑɽǏʆ ƳɌƙɆɊɮɍɃɩɅƒɅʂɋƙɁȪɎLJɅɳɄƛˊɳɓˊȶɳǷʕʐɃɪǂɸȶ ȲƒɭȶȯɑȩȲȷɸɅɯɅɈɪɌɵɅɳȳɁƎɳljɄȨǒɁɽ ɅɩȶȯɑȩȲȷɸɅɯɅɈɪɌɵɅɳȳɁƎ LJɁɽȼɸɆȶ ȴɬƸɆɽɈɪɵɂƂɃɪʖ ȼɍɽ ʒʕ ɴȳɎ ɩȷƄɩƳ ƹƒɸʒʐʑʕʆ ȲƒɭȶɃɪǂɸȶɅɪɊɯɋʉ ɑɸǁȲɑɁƛƙɁȪɎLJɅƙɆɊɮɍɈɪȲɭƒȶɆƚȶɽƙɁȶɽ (Ɇɳǁ Ǝ ɋʑʐʐɊ Ʌɩȶ ɃɃɫȶʒɊ) ȷɸɅɯɅʓɴȳƞ ɳƽɋɳƙɆˊȲȜɅƎȶɵɑƓǏɁɽƸɆɽ Ʌɩȶ Njɻ ɑɭɪɅɆɮɊƸɆɽɑɁƛɍɩɁ ơ ɳɒˊɋȲɑɩȲɌȲʁƙɁȪɎLJɅ ɑNjƖɑɇȶɴȼɌɳǷɃɪǂɸȶDŽɸȶɳdžɹʆ ƺɍɃƑɇɍ ȼȶɽɑɭɪɳɁɵɅɊNjȷɵɆɁȶɅɩȶɊNjȷɳǂƒɁNjɅɍȲſɀɺȯɑɳȼȢȶƵƒɌǏȶɴȯɑȲƒɭȶɳȳɁƎ DŽɸȶɈɪɌ ɳDŽɹN.ȱ virescens ǓȲɽNjɅȼȶɽɑɭɪɳɁȳƕɑɽƺȶȲƒɭȶɳȳɁƎDŽɸȶɈɪɌʆ ȲǂƎɊɯɋȷɸɅɯɅȼɮȷƺɑɪɁɭɀƟNJɈɑɸɳɀˊɊ ȼɸǁȲɽƳɍ ɍɮɁǎɑɽɅɩȶȲɊƕɑɽȯɑȪɎ ɊɩɅNjɅɗɃƑɩɈɍȼɍɽȼȶɽɑɭɪɳɁɌɆɑɽǏDŽɸȶɈɪɌɳɃʆ ɳDŽɹƺɴɆɆɳɅɹȲƎɪ ǏǕȷɆǁ Ǝ ɍɊȲɈɪɌɋɺɳɈɍƙɆ ɊɮɍɃɩɅƒɅʂɋȳƚɪ ɳƽɋɑɪɁɭɀƟNJɈɅɩȶɑɸɳɀˊɊNjɅNJɈȳɭɑƵƒɁɩȷɁɯȷȲƒɭȶɔɸɓɭȶɳɈɍɑɩȲǜʆ ɍɃƑɇɍɳɅɹLJɅɳɄƛˊɳǕɋɳȵˊȻɅɮɎ NJɈȷɸLJȷɽɵɅƳɌƙɆɊɮɍɑɸǁȲ ɳƽɋɳƙɆˊƙLJɑɽɎ ɩɄɪǒȝɑƎɳɍˊɑɈɪɊɯɋ Ʌɩȶ NjɅɌɋɺɳɈɍȯǒɎƙƺɎɋɮɌ ɳȼˊɊƓɪȲɸɀɁɽȲǂƎɴȼɍ ȹɹɗɃƑɩɈɍȼɍɽȼȶɽɑɭɪɳɁɵɅɑɁƛɍɩɁ ơ ɆɸLjƚȻȯɑȪɎɳǷȲɊƕɭƺʆ Abst ra c t The brown planthopper Nilaparvata lugens and green leafhopper Nephotettix virescens are major pests of rice crops in Southeast Asia, stunting their growth, height, leaf area, photosynthetic rate and nitrogen content. Brown planthopper transmits the ragged stunt virus and rice grassy stunt virus, whereas green leafhopper is a vector for rice diseases including tungro virus, yellow dwarf virus, yellow-orange leaf virus and transitory yellowing virus. My study aimed to compare the density of N. lugens and N. virescens in rice fields and identify possible factors influencing this. Fieldwork was undertaken at 50 sites in two districts in Pursat Province and two districts in Battambang Province from 6–25 November 2015. At each site, samples were collected from three transect lines (each measuring 100 m in length and 2 m in width) using sweep nets and a vacuum aspirator. Farmers were also interviewed at each site. My results suggest that densities of N. virescens and N. lugens are similar between the rice fields of the two provinces and that N. virescens appears to occur at higher densities in both. Factors such as temperature, humidity and the growth stage and height of rice plants did not influence the density of either species. However, this may have been due to the relatively short sampling duration, as temperature and humidity varied little during the study period. My results highlight the importance of using more than one sampling method and sampling for longer periods in research aiming to identify factors influencing the density of rice insect pests in Cambodia. Citation: Sour K. (2017) Comparative density of green leafhopper Nephotettix virescens and brown planthopper Nilaparvata lugens in rice fields around the Tonle Sap Lake, Cambodia. Cambodian Journal of Natural History, 2017, 132. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2017 (1) 128–133 Thesis abstracts De nsit y e st im at ion of gre e n pe a fow l Pavo m ut ic us in Sre pok Wildlife Sa nc t ua r y, Ca m bodia TAK Chandara ɊɮɍɅʂɋɑɳȶſɆ ɆɻɮɈɭɋǔɑƘɭȶɌɆɑɽɑɁƛɳƳƂȲɵɆɁȶ ɴȼɍƺɑɁƛɃɃɯɍɌȶɳƙƵɹȹɩɁɇɭɁɈɮȹɳǷɳɍˊɈɩɉɈɳǎȲ ƙɁȪɎLJɅɂɋȷɭɹɃɮDŽɸȶɁɸɆɅɽǕɑɭɪǕ ɳȴƒɋɿ ɳƽɋǒɌɴɁƳɌLJɁɽɆȶɽɃɪȹƙɊȲ Ʌɩȶ ƳɌɆɌLJȻɽʆ ɁɸɆɅɽNJȴɘǒɅɵɅƙɆɳɃɑȲɊƕɭƺ ƺȲɴɅƚȶɴȼɍƙɃƙɃȶɽɆɻɮɈɭɋǔɑƘɭȶ ƙɆɳɉɃɳɅɹɄɸƺȶɳȴ ɴȼɍɳɄƛˊɳǕɋǏƳƚɋƺɁɸɆɅɽǕɃɩNJɈɆƙNjɆɽƳɌɔɉɩɌȲƞʆ ɳƵɍɆɸɀȶɵɅƳɌɑɩȲǜɳɅɹȴɬɳȼˊɊƓɪLJɻɅɽƙɆNjɀ ȼȶɽɑɭɪɳɁɵɅɆɻɮɈɭɋǔɑƘɭȶɑɁƛɳƳƂȲɵɆɁȶɳǷɴȼɅȹƙɊȲɑɁƛɵƙɈɴȯɑɈȲʆ ʒ ʒ ƳɌɑɩȲǜLJɅɳɄƛˊɳɓˊȶɳǷȲƒɭȶɁɸɆɅɽɑɮɍ ƒ (ɵɇƐƙȲǔ ʑ.ʓʙʘ ȴɊ ) ɅɩȶɁɸɆɅɽȹɭɸɎ ɩȻɁɸɆɅɽɑɮɍ ƒ (ɵɇƐƙȲǔ ʑ.ʙʖʗ ȴɊ ) ɵɅɴȼɅȹƙɊȲ ɈɪɴȳɄƒɮ ƹƒɸʒʐʑʕ ȼɍɽ ɴȳəɑNJ ƹƒɸʒʐʑʖʆ ƳɌ LJɻ ɅɽƙɆNjɀȼȶɽɑɭɪɳɁɑɁƛɳƳƂȲȴɬ ǂɊɌɋɺƳɌǍɆɽɑɸɳɓȶɋɸɳǷǂɊȷɸɀɭȷǒƎɆɽDŽɸȶʘʐɃɪǂɸȶɳǷȲƒɭȶɁɸɆɅɽ ɴȼɍʔʘɃɪǂɸȶɑƏɩɁ ƒ ȲƒɭȶɃɪǂɸȶɅɪɊɯɋʉ NjɅȷɸɀɭȷǒƎɆɽȷɸɅɯɅɈɪɌ ɴȼɍȷɸɀɭȷɊɯɋǒƎɆɽɳǷ ȲƒɭȶɁɸɆɅɽɑɮɍ ƒ Ʌɩȶ ʓʒɃɪǂɸȶɑƏɩɁɳǷȹɭɸɎ ɩȻɁɸɆɅɽɑɮɍʆ ɳɈɍƙɈɫȲ Ʌɩȶ ȷɸɅɭȷɊɯɋɳɃȢɁǒƎɆɽɳǷɳɈɍǎƂȷʆ ɑɸɳɓȶɌɆɑɽɑɁƛɳƳƂȲɳƻƗɍȷɸɅɯɅʖʕȲǙɍƙɁȪɎLJɅɠɳǷȲƒɭȶɁɸɆɅɽɑɮɍ ƒ Ʌɩȶ ƒ ʗʑɳǷȹɭɸɎ ɩȻɁɸɆɅɽɑɮɍ ƒ ƺƳɌLJɻ ɅɽǒƗɅȼȶɽɑɭɪɳɁȴɬ NjɅɳƳƂȲɳƻƗɍȷɸɅɯɅ ʐ,ʕʕ ȲǙɍ/ȴɊʒ ɳɑƗˊɅɫȶʗʖʘȲǙɍȲƒɭȶɁɸɆɅɽɑɮɍ ɅɩȶɳǷȹɭɸɎ ɩȻɁɸɆɅɽɑɮɍNjɅȼȶɽ ƒ ɑɭɪɳɁʑ,ʐʗȲǙɍ/ȴɊʒ ȲǙɍ/ȴɊ ʒ ɴȼɍɳɑƗˊɅɫȶʒ.ʑʐʔȲǙɍʆ ȼȶɽɑɭɪɳɁɌɯɊɵɅɁɸɆɅɽDŽɸȶɈɪɌȷɮɍƵƒȴɬʐ,ʗʗ ɴȼɍɳɑƗˊɅɫȶʒʕʙʑȲǙɍ ɴȼɍɆƷƟȻǃɵƙɈɳǷɁɸɆɅɽNJȴɘǒɅɵɅƙɆɳɃɑȲɊƕɭƺ ƺɁɸɆɅɽɴȼɍNjɅǒɌɺɑɸƴɅɽ ƴƚɸȶȼɍɽƙɆɳɉɃɑɁƛɳƳƂȲɵɆɁȶʆ ɳǷȹɭɸɎ ɩȻɁɸɆɅɽɑɮɍɵɅɴȼɅȹƙɊȲɑɁƛ ƒ ɵƙɈɴȯɑɈȲNjɅȼȶɽɑɭɪɳɁɑɁƛɳƳƂȲɵɆɁȶȳƕɑɽ Ǖȷɳƽɋ ǒɌɴɁNjɅƳɌɌ ɸƴɅɈɪɊɅɭɑƞƴƚɸȶɳǷȲƒɭȶɁɸɆɅɽɑɮɍʆ ƒ NjɅȲǂƎȷɸɅɯɅɈɪɌɴȼɍNjɅɗɃƑɩɈɍȼɍɽȲƙɊɩɁɵɅƳɌȲɸɀɁɽɎɁƎNjɅɌɆɑɽ ɳƳƂȲʈ ʑ) ȷNjƂɋɈɪɃɳɅƚ ʒ) ȷNjƂɋɈɪɉɮɊɩdždžʆ DŽȲɽɃȶɅɫȶȲǂƎɃɪʑ ɳȴǕȷǒƗɅɈɪɎɁƎNjɅɵɅɑɁƛɳƳƂȲLJɅɳƽɋǒɌǏȷɮɍȷɩɁƎ ɳǷɴȲƓɌȲɴɅƚȶNjɅɃɫȲƺɔȷɩɵȜɅƎɋɿʆ ȲƒɭȶɑȲɊƗNJɈɔɉɩɌȲƞɑɁƛƙɆɳɉɃɳɅɹ ȴɯɌɤƘǕɃɩNJɈɳɍˊƳɌȷɮɍɌɯɊɈɪɑɒȴɊɅɿɴȼɍNjɅƙɆ ƺȹɅɳƙȷˊɅ ɳȼˊɊƓɪɳɍˊȲȲɊƕɑɽƳɌɌɑɽɳƽɋɑɅƎɩNJɈƺɊɯɋƵƒɌǏȶɊɅɭɑƞɅɩȶɑɁƛɵƙɈ ɴȼɍǕȷɃɃɯɍLJɅǂɊɌɋɺƳɌɔɆɽɌ ɸ ɋɭɃƑ džƳɌɆɳȶžˊɅƳɌɋɍɽȼɫȶ Ʌɩȶ ƳɌɳɍˊȲɃɫȲȷɩɁƎɴɇƒȲɳɑȼƊȲɩȷƃȼɮȷƺƳ ɌɃɑƞdžɑɁƛǒƚɆ Ʌɩȶ ɃƙɊȶɽɳɇƞȶʉɳɃȢɁɵɅɳɔȲɮɳɃɑ ȷɌɀɿʆ Abst ra c t Populations of the globally Endangered green peafowl have declined across Southeast Asia due to habitat loss and hunting. Northeast Cambodia likely supports the largest remaining populations of the species, making it a priority area for its conservation. The aim of my study was to estimate the population density of green peafowl in Srepok Wildlife Sanctuary. The study was conducted in the core zone (1,398 km2) and outer zone (1,967 km2) of the sanctuary between December 2015 and May 2016. Density estimates were derived from point counts at 80 locations at the site, 48 of which were situated in the core zone and 32 within the outer zone. Two point counts were conducted at each location, one in the morning and one in the evening. Sixty-five calling males were detected in the core zone and 71 in the outer zone, providing a density estimate of 0.55 calling males / km2 and a total estimate of 768 calling males in the former, and 1.07 calling males / km2 and 2,104 calling males in the latter. These provided a combined estimate of 0.77 calling males / km2 and 2,591 calling males for the site, which supports the notion that forests in northeastern Cambodia are an important stronghold for the species. The higher density of birds recorded in the outer zone of Srepok Wildlife Sanctuary may be due to greater human disturbance within the core zone. Two factors were found to affect detection rate: 1) distance from rivers; 2) distance from villages. The former was expected because the green peafowl is thought to prefer areas near permanent water. Conservation actions for the species should prioritize community engagement where larger human populations occur to promote the peaceful coexistence of people and wildlife. This could include education, awareness campaigns and economic incentives such as bird-watching and other forms of ecotourism. Citation: Tak C. (2017) Density estimation of green peafowl Pavo muticus in Srepok Wildlife Sanctuary, Cambodia. Cambodian Journal of Natural History, 2017, 133. Cambodian Journal of Natural History 2017 (1) 128–133 © Centre for Biodiversity Conservation, Phnom Penh 133 134 Instructions for Authors I nst ruc t ions for Aut hors Purpose and Scope The Cambodian Journal of Natural History (ISSN 2226– 969X) is an open access, peer-review journal published biannually by the Centre for Biodiversity Conservation at the Royal University of Phnom Penh. The Centre for Biodiversity Conservation is a non-profit making unit, dedicated to training Cambodian biologists and the study and conservation of Cambodia’s biodiversity. references) are welcomed on topics relevant to the Journal’s focus, including: • Research on the status, ecology or behaviour of wild species. • Research on the status or ecology of habitats. • Checklists of species, whether nationally or for a specific area. The Cambodian Journal of Natural History publishes original work by: • Discoveries of new species records or range extensions. • Cambodian or foreign scientists on any aspect of Cambodian natural history, including fauna, flora, habitats, management policy and use of natural resources. • Reviews of conservation policy and legislation in Cambodia. • Cambodian scientists on studies of natural history in any part of the world. The Journal especially welcomes material that enhances understanding of conservation needs and has the potential to improve conservation management in Cambodia. The primary language of the Journal is English. For full papers, however, authors are encouraged to provide a Khmer translation of their abstract. • Conservation management plans for species, habitats or areas. • The nature and results of conservation initiatives, including case studies. • Research on the sustainable use of wild species. The Journal does not normally accept formal descriptions of new species, new subspecies or other new taxa. If you wish to submit original taxonomic descriptions, please contact the editors in advance. Readership News The Journal’s readers include conservation professionals, academics, government departments, non-governmental organisations, students and interested members of the public, both in Cambodia and overseas. In addition to printed copies distributed in Cambodia, the Journal is freely available online from: http://www.fauna-flora.org/ publications/cambodian-journal-of-natural-history/ Concise reports (<300 words) on news of general interest to the study and management of Cambodia’s biodiversity. News items may include, for example: Manuscripts Accepted The following types of manucripts are accepted: • Announcements of new initiatives; for example, the launch of new projects, conferences or funding opportunities. • Summaries of important news from an authoritative published source; for example, a new research technique, or a recent development in conservation. • Full papers (2,000–7,000 words, excluding references) Letters to the Editors • Short communications (300–2,000 words, excluding references) Informative contributions (<650 words), usually in response to material published in the Journal. • News (<300 words) • Letters to the editor (<650 words) Recent Literature Full Papers (2,000–7,000 words, excluding references) and Short Communications (300–2,000 words, excluding Copies or links to recent (<18 months) scientific publications concerning Cambodian biodiversity and the management of natural resources. These may include journal papers, project technical reports, conference posters and student theses. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2016 (2) 134–136 Full Papers and Short Communications Instructions for Authors How to Submit a Manuscript Manuscripts are accepted on a rolling basis each year and should be submitted by email to the editors (Editor. CJNH@gmail.com). In the covering email, the lead (corresponding) author should provide the names and contact details of at least three suitably qualified reviewers (whom the editors may or may not contact at their discretion) and confirm that: • The submitted manuscript has not been published elsewhere, • All of the authors have read the submitted manuscript and agreed to its submission, and • All research was conducted with the necessary approval and permit from the appropriate authorities. Authors are welcome to contact the editors at any time if questions arise before or after submitting a manuscript. Preparation of Manuscripts Authors should consult previous issues of the journal for general style, and early-career authors are encouraged to consider guidance provided by: Fisher, M. (2012) Editorial – To shed light on dark corners. Cambodian Journal of Natural History, 2012, 1–2. Daltry, J., Fisher, M. & Furey, N.M. (2012) Editorial – How to write a winning paper. Cambodian Journal of Natural History, 2012, 97–100. Manuscripts should be in English and use UK English spelling (if in doubt, Microsoft Word and similar software should be set to check spelling and grammar for ‘English (UK)’ language). Lines should be double-spaced. Submissions can be in ‘doc’, ‘docx’ or ‘rtf’ format, preferably as a single file attached to one covering email. The order of sections in the manuscript should be: cover page, main text, references, short biography of each author, tables and figures (including photographs). All pages should be numbered consecutively. Cover page: This should contain the institutions and full mailing addresses of all authors and the email address of the corresponding author. Title: A succinct description of the work, in no more than 20 words. Cambodian authors are strongly encouraged to submit a Khmer translation of the English abstract. Keywords: (Full papers only). Up to eight pertinent words, in alphabetical order. Main text: (Short communications). This should avoid the use of headed sections or subsections. Main text: (Full papers). This should comprise the following sections in order: Introduction, Methods, Results, Discussion and Acknowledgements. Subsections may be included in the Methods, Results and Discussion sections if necessary. Conclusions and recomendations should be included in the Discussion. References: These should be cited in the text in the form of Stuart & Emmett (2006) or (Lay, 2000). For three or more authors, use the first author’s surname followed by et al.; for example, Rab et al. (2006) or (Khou et al., 2005). Multiple references should be in chronological order, for example, Holloway & Browne (2004); Kry & Chea (2004); Phan (2005); Farrow (2006). The reference list should be presented in alphabetical order. Cambodian, Vietnamese and other authors who typically write their family name first are presented in the form <surname> <initials> without a comma (thus, Sin Sisamouth becomes Sin S.). Western author names are presented in the form <surname> <comma> <initials> (thus Charles Robert Darwin becomes Darwin, C.R.). The titles of articles and journals should be written in full. The following are examples of house style: Papers: Berzins, B. (1973) Some rotifers from Cambodia. Hydrobiologia, 41, 453–459. Neang T. (2009) Liquid resin tapping by local people in Phnom Samkos Wildlife Sanctuary, Cambodia. Cambodian Journal of Natural History, 2009, 16–25. Tanaka S. & Ohtaka A. (2010) Freshwater Cladocera (Crustacea, Branchiopoda) in Lake Tonle Sap and its adjacent waters in Cambodia. Limnology, 11, 171–178. Books and chapters: Khou E.H. (2010) A Field Guide to the Rattans of Cambodia. WWF Greater Mekong Cambodia Country Programme, Phnom Penh, Cambodia. MacArthur, R.H. & Wilson, E.O. (1967) The Theory of Island Biogeography. Princeton University Press, Princeton, USA. Abstract: (Full papers only). This should describe, in no more than 250 words, the aims, methods, major findings and conclusions. The abstract should be informative and intelligible without reference to the text, and should not contain any references or undefined abbreviations. Rawson, B. (2010) The status of Cambodia’s primates. In Conservation of Primates in Indochina (eds T. Nadler, B. Rawson & Van N.T.), pp. 17–25. Frankfurt Zoological Society, Frankfurt, Germany, and Conservation International, Hanoi, Vietnam. Cambodian Journal of Natural History 2016 (2) 134–136 © Centre for Biodiversity Conservation, Phnom Penh 135 136 Instructions for Authors Reports: Lic V., Sun H., Hing C. & Dioli, M. (1995) A Brief Field Visit to Mondolkiri Province to Collect Data on Kouprey (Bos sauveli), Rare Wildlife and for Field Training. Unpublished report to Canada Fund and IUCN, Phnom Penh, Cambodia. Theses: Yeang D. (2010) Tenure rights and benefit sharing arrangements for REDD: a case study of two REDD pilot projects in Cambodia. MSc thesis, Wageningen University, Wageningen, The Netherlands. tuation: e.g., Asian elephant Elephas maximus. English names should be in lower case throughout except where they incorporate a proper name (e.g., Asian flycatcher, Swinhoe’s minivet, long-billed vulture). Abbreviations: Full expansion should be given at first mention in the text. Units of measurement: Use metric units for measurements of area, mass, height, etc. Websites: Review and Editing IUCN (2010) 2010 IUCN Red List of Threatened Species. Http:// www.redlist.org [accessed 1 December 2010]. All authors are strongly advised to ensure that their spelling and grammar is checked by a native English speaker before the manuscript is submitted to the journal. The editorial team reserves the right to reject manuscripts that need extensive editing for spelling and grammar. About the Author(s): This section is optional for Full Papers and Short Communications. It should describe the main research interests of each author (<150 words each), apart from what is obvious from the subject of the manuscript and the authors’ affiliations. Tables and figures (including plates): All tables and figures should be cited in the text and placed at the end of the manuscript. These should be self-explanatory, have an appropriate caption and be placed on separate pages. Figures, including maps, should ideally be in black and white. Plates (photographs) should be included only if they are of good quality and form part of evidence that is integral to the study (e.g. a camera trap photograph of a rare species). Appendices: Long tables and other supporting materials, such as questionnaires, should be placed in Appendices. All manuscripts are subject to rigorous peer review by a minimum of two qualified reviewers. Proofs will be sent to authors as a portable document format (PDF) file attached to an email note. Acrobat Reader can be downloaded free of charge from <www. adobe.com> to view the PDF files. Corrected proofs should be returned to the Editor within three working days of receipt. Minor corrections can be communicated by email. Authors are permitted to post their papers on their personal and institutional webpages on condition that access is free and no changes are made to the content. Species names: The first time a species is mentioned, its scientific name should follow without intervening punc- Publisher: Centre for Biodiversity Conservation, Room 415, Main Campus, Faculty of Science, Royal University of Phnom Penh, Confederation of Russian Boulevard, Phnom Penh, Cambodia. © Centre for Biodiversity Conservation, Phnom Penh Cambodian Journal of Natural History 2016 (2) 134–136 Cambodian Journal of Natural History The preparation and printing of this volume was generously supported by: Royal University of Phnom Penh—Centre for Biodiversity Conservation RUPP is Cambodia’s oldest university, with over 9,000 students and over 400 teachers. The Department of Biology founded the Centre for Biodiversity Conservation to provide training and support for national scientists. The Centre delivers a Masters of Science curriculum in Biodiversity Conservation and has established a library, classrooms, herbarium and zoological reference collection for use by students and scholars of Cambodian natural science. Website: www.rupp.edu.kh/master/biodiversity/?page=CBC Fauna & Flora International FFI protects threatened species and ecosystems worldwide, choosing solutions that are sustainable, are based on sound science and take account of human needs. Operating in more than 40 developing countries worldwide, FFI saves species from extinction and habitats from destruction, while improving the livelihoods of local people. Founded in 1903, FFI is the world’s longest established international conservation body. FFI has been active in Cambodia since 1996. Website: www.fauna-flora.org The present issue was also supported by a major foundation that chooses to remain anonymous. The Cambodian Journal of Natural History does not charge subscription fees. The journal depends upon the generosity of its partner organisations and sponsors to be published and distributed free of charge to readers throughout Cambodia and worldwide. If you or your organisation are interested in supporting the Cambodian Journal of Natural History or the Centre for Biodiversity Conservation, kindly contact the editors (Editor.CJNH@gmail.com) or the Centre for Biodiversity Conservation (mbiodiversity. info@rupp.edu.kh). The names and logos of all supporters will be published in the journal unless they wish to remain anonymous. The Editors are grateful to our reviewers and to Chantha Nasak, Chhin Sophea, Eam SamUn, Hun Seiha, Kheam Sokha, Srey Saovina, Thi Sothearen and Regine Weckauf for their kind assistance with the production of this issue. Cambodian Journal of Natural History Volume 2017, Number 1 Cont e nt s 1 Editorial— The future of Payments for Ecosystem Services in Cambodia, Virginia Simpson & Nicholas Souter. 4 Short Communication— New records of Orchidaceae from Cambodia IV, André Schuiteman, Rudolf Jenny, Khou Eang Hourt, Nay Sikhoeun & Att Sreynak. 10 Short Communication— Gastrodia exilis (Orchidaceae), a newly recorded mycoheterotrophic genus and species in Cambodia, Suetsugu Kenji, Hsu Tian-Chuan, Tagane Shuichiro, Chhang Phourin & Yahara Tetsukazu. 14 Short Communication—Revision of Cambodia’s National Biodiversity Strategy & Action Plan to integrate conservation and sustainable use of biological resources into national decision making, Matthew Maltby, Chan Somaly & Jady Smith. 17 Full Paper— The flora of the Bokor Plateau, southeastern Cambodia: a homage to Pauline Dy Phon, Philip Rundel & David Middleton. 38 Full Paper— Status and conservation significance of ground-dwelling mammals in the Cardamom Rainforest Landscape, southwestern Cambodia, Thomas Gray, Andrew Billingsley, Brian Crudge, Jackson Frechette, Romica Grosu, Vanessa Herranz-Munoz, Jeremy Holden, Keo Omaliss, Kong Kimsreng, David Madonald, Neang Thy, Ou Ratanak, Phan Channa & Sim Sovannarun. 49 Full Paper— The impact of shrimp farming on water quality in Anlung Pring, a protected landscape in Cambodia, Yav Net, Seng Kimhout, Nhim Sophea, Chea Vannara, Bou Vorsak & Tomos Avent. 55 Full Paper— Carbon stock of peat soils in mangrove forest in Peam Krasaop Wildlife Sanctuary, Koh Kong Province, southwestern Cambodia, Taing Porchhay, Eang Phallis, Tann Sotha & Chakraborty Irina. 63 Full Paper— Camera trapping of large mammals in Chhep Wildlife Sanctuary, northern Cambodia, Suzuki Ai, Thong Sokha, Tan Setha & Iwata Akihisa. 76 Full Paper— Ethnobotanical knowledge of the Kuy and Khmer people in Prey Lang, Cambodia, Nerea Turreira-Garcia, Dimitrios Argyriou, Chhang Phourin, Prachaya Srisanga & Ida Theilade. 102 Full Paper— Movement of captive-reared Siamese crocodiles Crocodylus siamensis released in the Southern Cardamom National Park, Cambodia, Eam Sam Un, Sam Han, Hor Leng, Me’ira Mizrahi & Jackson Frechette. 109 Full Paper— Stand carbon dynamics in a dry Cambodian dipterocarp forest with seasonally flooded sandy soils, Ito Eriko, Furuya Naoyuki, Toriyama Jumpei, Ohnuki Yasuhiro, Kiyono Yoshiyuki, Araki Makoto, Sokh Heng, Chann Sophal, Khorn Saret, Samreth Vanna, So Thea, Tith Bora, Keth Samkol, Ly Chandararity, Op Phallaphearaoth, Monda Yukako & Kanzaki Mamoru. 128 Recent Master’s Theses— Peng Borin, San Satya, Sim Sovannarun, Soun Visal, Suor Kimhuor & Tak Chandara. 134 Instructions for Authors.