The Biodiversity of the Tonle Sap Biosphere Reserve 2005 status review

March 2006

Peter J.A. Davidson (compiler)

This publication is a technical output of the UNDP/GEF-funded Tonle Sap Conservation Project. Acknowledgements

The following provided valuable discussion, unpublished data and/or useful background information: David Ashwell, Eric Baran (World Fish Centre), Kristen Davies (Sam Veasna Centre for Wildlife Conservation, Siem Reap), Paul Dolman (University of East Anglia, U.K.), Tom Evans (WCS), Tom Gray (University of East Anglia, U.K.), Heng Sovannara (WCS), Hong Chamnan (WCS), Rohan Holloway (Australia National University), Dani Jump, Long Kheng (TSBR Secretariat), Ro Borey (WCS), Sin Saenglay (WCS), Sun Visal (WCS), Richard Salter (UNDP TSCP), Joe Walston (WCS). Steve Platt (Oglala Lakota College, South Dakota, U.S.A.) and Bryan Stuart (Field Museum of Natural History, Chicago, U.S.A.) provided references on reptiles and reviewed the her- petology section. Angela Frost (University of Queensland, Australia) provided reference material on rodents. Tom Clements (WCS) provided data and advised on large waterbird monitoring in the Prek Toal Core Area. Zeb Hogan provided a detailed account of the Mekong Giant Catfish. Mr. Cheng Phen (Department of Fisheries) provided useful dis- cussion on the fish sanctuaries. Isabel Beasley and James Cook University provided references and unpublished results on Irrawaddy Dolphin.

Nhem Nyta greatly assisted with the reference search, and summarily reviewed many documents in both English and Khmer. Kuoch Kunthea Voreak assisted with the reference search and compiled species lists from various sources. Contents

LITERATURE REVIEW...... 2 Information Sources...... 2 Synopsis of Biodiversity Study in the TSBR...... 2

BACKGROUND...... 4 Physical Geography and Climate...... 4 Drainage and Hydrology...... 4 Demography and Land-use...... 5 Conservation Status of the Tonle Sap Floodplain...... 5

BIODIVERSITY...... 7

FUNGI & FLORA...... 10

MACROPHYTE FLORA...... 11

FAUNA...... 18 Mammals...... 18 Birds...... 22 Reptiles...... 29 Amphibians...... 34 Fish...... 34

INVERTEBRATES...... 42

KEY THREATS TO BIODIVERSITY IN THE TSBR...... 44

INTRODUCED SPECIES...... 47

BIODIVERSITY IN THE TONLE SAP BIOSPHERE RESERVE: WHAT WE KNOW, AND WHAT WE NEED TO KNOW...... 49

REPTILES AND AMPHIBIANS...... 51

REFERENCES...... 53 ANNEX 1: PLANT SPECIES RECORDED IN THE TSBR...... 63

ANNEX 2: FISH SPECIES RECORDED IN THE TSBR...... 68

ANNEX 3: BIRD SPECIES RECORDED IN THE TSBR...... 72

ANNEX 4: MAMMAL SPECIES RECORDED IN THE TSBR...... 76 List of Tables

ECOLOGICAL PROCESSES/EVENTS BY HABITAT TYPE...... 8

List of Figures

FIG. 1 CROSS SECTION OF FLOOD PLAIN HABITATS...... 11

FIG. 2 DOMINANT PLANT SPECIES IN THE TONLE SAP FLOODPLAIN...... 16

List of Maps

TONLE SAP LAKE...... 1 1 INFORMATION SOURCES SYNOPSIS OF BIODIVERSITY TUDY IN THE This study is in essence a literature review. All S TSBR traceable documents relating to biodiversity in the Tonle Sap floodplain have been viewed by The exceptional biodiversity of the Tonle Sap the editor, including published journal papers was first documented as early as the 13th (relatively few), occasional book chapters, the Century by Zhou Daguan, a Chinese visitor to bi-annual journal Cambodia Bird News, pub- Cambodia, who noted the abundance of fish, lished reports (numerous, notably from the frogs, tortoises, turtles, lizards, large croco- Wildlife Conservation Society and the Mekong diles and molluscs on and around the lake River Commission/WUP-FIN, also the United (Zhou 2002). The bas-reliefs of the Angkorian Nations Food and Agriculture Organisation – Bayon Temple depict several key elements of FAO), identification and species reference the Great Lake’s fauna, in particular fish, croc- guides, unpublished reports (very numerous, odiles, turtles and large waterbirds (Roberts including many “trip reports” from field sur- 2002, Bailleux 2003). veys, often in Khmer) and even posters. In cases where references could not be viewed French explorers reported on the rich fisheries directly this is indicated in the text by the of the Tonle Sap (e.g. Garnier 1996), and by phrase “cited in” (e.g. Anon. 2003 cited in the early 20th Century the exceptional produc- Anon. 2005). tivity of Tonle Sap fisheries for export was well known (Petillot 1911, Chevey 1935 cited Reference material was gathered from the in Hortle et al. 2004). Fish studies conducted libraries of the following organisations and during the French colonial period, synthesised institutions: Tonle Sap Biosphere Reserve in Chevey and Le Poulain (1940) and Blache Secretariat (Ministry of Environment - MoE), and Goosen (1954) are still the most compre- Department of Fisheries and the Forest hensive reviews of fish status, distribution and Administration (Ministry of Agriculture, Forests diversity conducted on the Tonle Sap (E. and Fisheries – MAFF), Wildlife Conservation Baran, World Fish Centre, verbally September Society (WCS), World Wide Fund for Nature 2005). French naturalists resident in Cambodia (WWF), Conservation International (CI), Flora during the first half of the 20th Century pub- and Fauna International (FFI), Cambodia lished the first significant information from the Development Resource Institute (CDRI), Tonle Sap region, on birds (Delacour 1929, Cambodia National Mekong Committee Delacour and Jabouille 1928, 1931, Engelbach (CNMC), Mekong River Commission (MRC, 1940). now based in Vientiane, Lao PDR) including the Water Utilisation Programme – Finnish During the post-colonial era (ca.1953-1970), Environment Institute (WUP-FIN), United the French synthesised information on the Nations Development Programme (UNDP), the flora of selected parts of Cambodia, including World Fish Centre, the Royal University of the Tonle Sap (e.g. Dy Phon 1970, 1982, Agriculture (RUA) and the Royal University of Legris and Blasco 1972, Rollet 1972). Bill Phnom Penh (RUPP). Thomas made further ornithological observa- tions and collated these with the earlier stud- Additional material was gathered through per- ies of birds (Thomas and Poole 2003), and sonal correspondence and discussion between Bardach (1959) conducted a study of the fish- the editor and many people working on differ- eries. No significant biodiversity work was ent aspects of biodiversity in the TSBR (see conducted between 1970 and the early Acknowledgements). 1990s.

Immediately after the civil war period, the Tonle Sap formed a central focus for biological exploration. This began with short aerial and 2 boat-based surveys for birds (Archibald 1992, birds in parts of the outer floodplain in Banteay Scott 1992, Carr 1994), followed by more Meanchey, Siem Reap and Kompong Thom comprehensive aerial, ground and interview provinces during 2001-2002, as part of the surveys during the dry seasons of 1994 Important Bird Area identification process (Mundkur et al. 1995) and 1996 (Parr et al. (Seng et al. 2003, BirdLife International 1996). This work established that breeding 2004a). Similarly, widespread interview and colonies of several globally threatened large follow-up field surveys were conducted for waterbirds persisted in the forested habitats crocodiles in 2001-2002 (Platt et al. 2004). along the north-western shores of the Tonle First attempts to quantify the watersnake were Sap, south-west of the floating village of Prek conducted in 2000 (Stuart et al. 2000), and Toal, but that they were being heavily exploit- have since been followed up with more ed for their eggs and chicks. Surveys investi- detailed investigations (Brookes et al. 2005). gating the socio-economic drivers behind the waterbird harvest, conducted in 1997, includ- Fish catches from the Tonle Sap have been ed conservation awareness work, which led to reported on annually and in some detail since a substantial reduction in collection that year the mid-1990s (e.g. van Zalinge and Touch (Ear-Dupuy et al. 1998). 1996, Nao 1997, van Zalinge et al. 1999, Lieng and van Zalinge 2001), but very few In February-March 1997, a detailed study of studies of fish diversity and distribution have the inundated forest-shrubland flora was con- been conducted (Lim et al. 1999, Lamberts ducted as a contribution to the nomination of 2001). Since 2002, a series of documents the Tonle Sap as a UNESCO Biosphere summarising knowledge of fish and fisheries in Reserve (McDonald et al. 1997). This remains the Lower Mekong Basin has been published the most comprehensive (indeed, the only by the Mekong River Commission (e.g. major field) study of the vegetation of the Mattson et al. 2002, Poulsen et al. 2002 & Tonle Sap floodplain. 2004, Coates et al. 2003, Hortle et al. 2004), and modelling of the physical processes of the Since 2000, Prek Toal Core Area’s waterbird Tonle Sap has been addressed with reference colonies have been the focus of annual to biodiversity by WUP-FIN (e.g. Kummu et al. counts, and with the establishment of the Prek 2005, in press a & b). Toal Conservation Team in 2001, the site has had almost year-round field coverage (Goes 1998, Goes and Hong 2002, Goes 2005, Cambodia Bird News nos.4-13). Conservation outreach and Bengal Florican monitoring research activities were initiated in the middle- outer floodplain grasslands in Kompong Thom in 2000 (Goes et al. 2001, Davidson 2004, Davidson et al. in prep.). Widespread interview and follow-up field surveys were conducted for

3 HYSICAL EOGRAPHY AND Cambodia and the northern slopes of the P G Cardamom Mountain coastal ranges. CLIMATE The Tonle Sap Lake expands and contracts seasonally in response to monsoonal rainfall The Tonle Sap Great Lake (hereafter simply and the flow of the Mekong River, in effect referred to as Tonle Sap) is the largest perma- acting as a storage reservoir which regulates nent freshwater lake in Southeast Asia (Scott flooding in the Mekong Delta. When the depth 1989). It is situated in the centre of the low-lying of the Mekong reaches approximately 9m Cambodian plain, which has an average eleva- (Lamberts 2001) in the early wet season, tion of 10-30 m asl (Lamberts 2001). water is forced north-west ‘up’ the Tonle Sap Quaternary sediments of 15-30 m cover large into the lake, which results in extensive inun- portions of this plain (Rundel 2000). The lake dation of the surrounding floodplain. The lake itself was formed by subsidence of the plain can increase in depth from its dry-season approximately 5,700 years ago (Carbonnel (Dec-June) depth of 1-2 m up to 8-10 m at 1963, Tsukawaki 1997 cited in Hortle et al. the peak of the flood, and its surface area 2004). The Tonle Sap floodplain (defined as 2 the area at maximum flood encompassed by expands from 2,500-3,000 km in the dry sea- 2 the 10-m asl contour line, including the open son up to 15,000 km during the flood season, lake; Giesen 1998) extends 250-300 km when at maximum flood level it covers about along a north-west to south-east axis, and is 6-8% of the land area of Cambodia (Nao up to 100 km wide (Scott 1989, Lamberts 1997, MRC 2003, Kummu et al. in press a). 2001). During the wet season, the volume of the lake increases from about 1.3 km3 up to 60-70 3 The region experiences a tropical monsoon cli- km , depending of the flood intensity. The mate, characterised by a pronounced May to reverse Mekong flow phenomenon results in November rainy season determined by the peak water levels in Tonle Sap lagging 1-2 south-west monsoon, with mean monthly max- months behind the peak of seasonal rainfall. imum rainfall reaching ca. 270mm in Floodwaters begin to recede in the early dry September and October (Giesen 1998, season in November, with the lowest water Lamberts 2001). Nonetheless, the Tonle Sap levels occurring at the end of the dry season basin is one of the driest areas in Cambodia in early May (Scott 1989). (together with lowland areas to the west and the lower Mekong basin south-east of Phnom The Mekong River yields approximately 51% Penh), receiving 1,000-1,500 mm of annual of this inflow of water, but is responsible for rainfall (Catling 2001), with a mean of 1,477 the majority (ca.75%) of the sediment deposit- mm at Siem Reap and 1,350 mm at ed in the Tonle Sap Lake (Kummu et al. in Battambang (Crocker 1962), and 1,550 mm press a). It is estimated that more than 80 % (2000-2003 four-year mean) at Kompong of the sediment the Tonle Sap system receives Thom (Kompong Thom Provincial Department from the Mekong River and its tributaries is of Forestry data). stored in the lake and its floodplain. The dynamic nature of sediment movement into and out of Tonle Sap has been the subject of much speculation, i.e. that the lake is rapidly DRAINAGE AND HYDROLOGY filling up with sediment due increasing sedi- ment loads coming from its catchment. The overall drainage basin of the Tonle Sap However, recent studies have shown that net sediment accumulation within the Tonle Sap covers 67,600 km2 (Pantulu 1986; see map in Lamberts 2001). Major tributaries include the Lake itself is low, and not accelerating with Stung Sen and the Stung Chinit, which drain respect to the long-term sediment dynamics of much of northern Cambodia, and the Stung the system (Kummu et al. 2005). The majority Sangke and Stung Poutisat that drain western of the sediments settle in the inundated forest

4 and shrubland areas of the floodplain (Sarkulla Fishing lots are enclosed with bamboo fences et al. 2003). However, there are local prob- that extend for many kilometers and lot opera- lems associated with high sedimentation and tors strictly control access for the duration of erosion rates in the TSBR (Heinonen 2005 the fishing season (November – May inclu- cited in Kummu et al. 2005). sive). Rice cultivation has traditionally been almost exclusively of deepwater (floating) rice; historically about half (200,000 ha) of EMOGRAPHY AND AND USE Cambodia’s deepwater rice was grown in the D L - Tonle Sap inundation zone, but by 2000 only ca.70,000 ha remained, of which 38% was in The human population on and around the Kompong Thom province (Catling 2001). Dam Tonle Sap was estimated to have at least construction for more intensively cultivated dry tripled between the 1940s and 1995 (Baran et season rice has become prevalent in the flood- al. 2001), when it was thought that the num- plain since 2000 (Evans et al. 2005). ber of people in fishing dependent communes in the six provinces bordering the lake num- bered almost 3 million (Nao et al. 1996 cited in Baran 2004). Giesen (1998) reports one CONSERVATION STATUS OF hundred and sixty communes inhabited by an THE ONLE AP FLOODPLAIN estimated 1.02 million people are located on T S the periphery of Tonle Sap, and about 170 floating villages ranging in size from two to The Tonle Sap Lake and most of its floodplain over 100 households occur on the lake itself was approved as the Tonle Sap Biosphere and move relative to the seasonally fluctuating Reserve (TSBR) under UNESCO’s Man and the water levels. The most recent and accurate Biosphere Program (MAB) in October 1997 estimate is of approximately 470,000 people (UNESCO 2005). This approval was recog- living within the 10 m inundation zone (up to nised within Cambodia by the Royal Decree for NR-5 and NR-6), and approximately 1.19 mil- the Establishment of the Tonle Sap Biosphere lion in all zones including urban centres Reserve passed in April 2001. The UNESCO (Keskinen 2003). A small out-migration of peo- Biosphere Reserve conceptual design consists ple from all provinces surrounding the lake but of strictly protected core areas surrounded by for Kompong Chhnang has recently been buffer and transitional zones, where sustain- reported (Haapala 2003 cited in Baran 2004), able resource extraction and human occupancy due to decreasing fish catches, droughts and is permitted. In contrast to this standard, sub- unpredictable flooding which is impacting on sistence and commercial fishing are permitted rice yields, and reduction in river water quality in the TSBR core areas. Three core areas were due to increased sediment loads. proposed: Prek Toal (31,282 ha), Moat Khla - Boeng Chhma (32,969 ha), and Stoeng Sen Rice cultivation, fuelwood collection, aquacul- (6,586 ha) (Giesen 1998), each considered to ture (chiefly fish), and subsistence and com- represent unique ecosystems demarcated for mercial fishing are the principal economic long-term protection and conservation (Neou activities of the communities living on the lake 2001). By the time the TSBR was legally and surrounding the floodplain (Nesbitt 1997, established in April 2001, however, the Prek Giesen 1998, Gum 1998). Fishing is the main- Toal Core Area had been reduced to 21,342 stay of communities situated closer to open ha, and the Moat Khla – Boeng Chhma Core water and major waterways; rice cultivation Area was reduced to 14,560 ha (encompass- and other agriculture are economically more ing Boeng Chhma only) (e.g. Goes 2005). The significant to communities in the outer flood- three core areas are surrounded by a buffer plain (Keskinen 2003). Commercial fishing is zone of just under 500,000 ha, which in turn concentrated in 57 administrative fishing lots, is encompassed within a transition zone of which encompass extensive areas of the lake approximately 900,000 ha, bordered to the and surrounding wetlands (Giesen 1998). south-west by National Route 5 (NR-5) and to 5 the north-east by NR-6 (Giesen 1998, Gum habitats in Stung, Stung Sen and Santuk dis- 1998). The transition zone does not cover the tricts of Kompong Thom province, and adja- entire Tonle Sap floodplain. A substantial por- cent Chikreng district in Siem Reap province tion (approximately 90,000 ha) of the outer (e.g. Goes et al. 2001). BirdLife International floodplain in Kompong Thom province lies out- has been conducting conservation outreach in side the transition zone, comprising a grass- two IBAs within the TSBR, in Siem Reap and land/agro-ecosystem mosaic of considerable Banteay Meanchey provinces. biodiversity conservation importance (e.g Seng et al. 2003).

Eight designated fish sanctuaries are physically demarcated on the Tonle Sap Lake itself. These reportedly became operational between 1980 and 1982 (Cheng Phen verbally 2005), although it is not clear on what basis they were designated (E. Baran and Cheng Phen verbally 2005). They are manned by armed personnel for 4-6 months of each dry season, and essentially function as dry season no-fish- ing and boat exclusion zones. Experimental protection from fishing of the lower water col- umn has begun in two fish sanctuaries using concrete tubes (Cheng Phen verbally 2005).

The Boeung Chhma Core Area was designated as a 28,000 ha site of international signifi- cance under the Ramsar Convention (Boeng Chhmar and Associated River System and Floodplain) in June 1999 (Wetlands International 2005). It is one of only three Ramsar sites in Cambodia. The other two are the Mekong River north of Stung Treng (the “Stung Treng Ramsar site”) and Koh Kapik and associated islets on the Gulf of Thailand coast in Koh Kong province.

The TSBR wholly encompasses or overlaps nine Important Bird Areas (IBAs), which are internationally recognised sites of conservation importance because they regularly support populations of globally threatened birds and/or regionally significant populations of congrega- tory waterbirds (Seng et al. 2003, BirdLife International 2004b).

Since 2001, the WCS Cambodia programme’s Tonle Sap Project has been conducting wide- ranging conservation action in the Prek Toal Core Area (e.g. Goes and Hong 2002, Goes 2005) and in grassland and agro-ecosystem

6 The biodiversity of the Tonle Sap Lake and its Seasonality of key processes and their implica- floodplain is relatively species poor in compari- tions for conservation son to other tropical lowland ecosystems, pri- marily due to the extreme stresses imposed on The wet-season flood-pulse is the primary the ecosystem by the annual climatic and physical process driving a sequence of signifi- hydrological fluctuations. Unique floral and fau- cant ecological events that are the mainstay of nal assemblages have evolved to cope with the Tonle Sap ecosystem’s annual cycle. The these harsh environmental conditions, howev- annual flood-pulse from the Mekong transfers er, and the annual flood-pulse and nutrient the floodplain’s terrestrial primary products replenishment cycle supports abundant popula- into an aquatic phase (e.g. Lamberts, 2001). tions of this adaptive flora and fauna. These Table 1 presents the major physical and include significant populations of many species land/human-use processes, and the major of global conservation concern. The TSBR is annual ecological events as they occur chrono- one of the most important wetlands for biodi- logically in the Tonle Sap ecosystem’s highly versity conservation in Southeast Asia, particu- seasonal annual cycle. larly for birds, reptiles and plant assemblages. Because of the extreme environmental changes The Tonle Sap is widely believed to be one of experienced in the TSBR through the course of the most productive inland waters in the world a year, many species are only able to use the (e.g. Lamberts 2001, Lieng and van Zalinge Tonle Sap on a seasonal basis. Their conserva- 2001, Baran 2004, and references therein). tion therefore depends not only on measures Key factors behind this exceptional productivi- taken within the TSBR, but also on factors ty include i) the annual inundation cycle with operating in other parts of these species’ its extensive, long-lasting monsoon floods, ii) annual ranges. For many, there is currently lit- the floodplain vegetation, in particular the tle or no conservation regulation implemented “flooded forest” and “flooded shrub”, and iii) in other parts of their range; indeed, the other high rates of nutrient cycling (Lamberts 2001). parts of many of these species’ ranges are Another highly plausible factor is the sustained often rather poorly known. high exploitation of fish over the past century (Petillot 1911, Chevey 1934, both cited in Baran 2004), which may have resulted in selection for dominant fast growing and short life span species, mostly Cyprinids (Baran 2004). The precise mechanisms of this pro- ductivity remain rather poorly understood and require detailed further research.

7 Season Month(s) Key Physical, Land- and Ecological Processes/Events by Habitat Type EcologicalPr Human-use Processes Open Lake Swamp forests Short-tree shrublands Grasslands/agro-ecosystems Reversal of Tonle Sap River flow In-migration of "white Tree and shrub flowering and Tree and shrub flowering and Out-migration of terrestrial into Tonle Sap Lake - flood- fish" from Mekong via fruit production fruit production breeding species (grassland

pulse begins Tonle Sap River birds and larger mammals) as flood rises Deepwater rice crops In-migration of "white fish" In-migration of "white fish" In-migration of post-breeding ocesses /Eventsb germinating in outer floodplain for feeding, nesting and for feeding, nesting and large waterbird congregations spawning spawning (storks, ibises, herons and Early egrets, pelicans) May-July monsoon Watersnake harvest begins Lateral migrations of "black Lateral migrations of "black In-migration of wet-season fish" to nesting and spawning fish" to nesting and spawning breeding waterbirds (e.g. rails, habitats habitats crakes, bitterns) Departure of large waterbird In-migrations of some black and breeding colonies (storks, white fish species for nesting,

spawning and juvenile growth. y Habitat pelicans, ibises, herons and egrets)

8 Expansion of Tonle Sap Lake to Tree and shrub flowering and Tree and shrub flowering and

max. inundation through reverse fruit production fruit production flow mechanism and catchment

rainfall T Largest sediment inputs Deciduous tree leaf fall Deciduous tree leaf fall ype (underwater) (underwater) Fishing (outside lot system) in Fish nesting, spawning, Fish nesting, spawning, Fish nesting, spawning, feeding Mid- August- outer floodplain agro-ecosystems feeding and juvenile growth feeding and juvenile growth and juvenile growth period monsoon October period period Deepwater rice crop main growth Cormorants and Darters return Main phase of deepwater rice phase to nest colonies and growth in the outer floodplain commence breeding agro-ecosystems Watersnake harvest peaks Watersnake breeding period Watersnake breeding period Fish perform function of natural pest regulators in traditional rice agricultural systems Season Month(s) Key Physical, Land- and Ecological Processes/Events by Habitat Type Human-use Processes

Open Lake Swamp forests Short-tree shrublands Grasslands/agro-ecosystems Flood water begins to recede as Out-migration of Leaf flush in all deciduous Leaf flush in all deciduous Continued growth of "floating Mekong River level drops and "white fish" to and evergreen tree species and evergreen tree species grasses" Tonle Sap River reverses flow Mekong River via Late October- again and begins draining the Tonle Sap River and monsoon November lake floodplains Large-scale commercial fishing Watersnake breeding period Watersnake breeding period begins in all fishing lots and main waterways draining Tonle Sap lake Deepwater rice ripening phase Large-scale commercial fishing Out-migration of Cormorants and Darters In-migration of terrestrial continues in all fishing lots and "white fish" to complete breeding and fledge breeding species begins main waterways draining Tonle Mekong River via (grassland birds including Sap lake Tonle Sap River and Bengal Florican) as land re- floodplains continues exposed on flood recession November- Early dry JanuaryFlood waters recede increasingly Large feeding Large waterbirds (storks, Large in-migration of Palearctic 9 rapidly aggregations of terns ibises, pelicans, herons and "winter visitor" bird populations and gulls egrets) return to nesting (raptors, chats, hirundines, colonies warblers, pipits, wagtails)

Second watersnake harvest peak

Deepwater rice harvest Main breeding period for large Black fish concentrated in Main breeding period for Large-scale commercial fishing ?Concentration of waterbrids (storks, ibises, remaining waterbodies and terrestrial grassland birds (e.g. begins in all fishing lots and black fish as pelicans, herons and egrets) performing "overland" Bengal Florican, quails and main waterways draining Tonle surroundingplain dries out migrations buttonquails) SapIncreasingly lake drought like floodLarge feeding High concentrations of fish Estimated main breeding Black fish concentrated in conditions develop in outer aggregations of terns behind fish traps in fishing period for turtles and pythons remaining waterbodies and flooplain and gulls lots performing "overland" Mid-late January- migrations dry May Extensive burning of grassland Estimated main breeding Local movements of Palearctic and shrubland Livestock grazing period for turtles and pythons birds in response to availability (January-early April) of key food resources

Preparation of land for Domestic livestock may partly deepwater rice cultivation fulfil grazing function of extirpated herbivore populations FUNGI FLORA

The only information traced on fungi is PHYTOPLANKTON from McDonald et al. (1997). Several fruit- ing Basidiomycetes were observed on The phytoplankton of the Tonle Sap itself large trunks drifting in the water in Moat has not received specific study, but during Khla in late February, one of which resem- the late 1980s, a Vietnamese study bled Pholiotus morphologically. Two poly- (Nguyen and Nguyen 1991 cited in pores (tough, leathery poroid mushrooms, Lamberts 2001) identified 197 species of similar to boletes) were also observed in phytoplankton in “the Mekong, the Tonle large colonies on floating logs, one bright Sap channel and the floodplains”, with the orange the other dull whitish in colour largest number of species found in the (McDonald et al.1997). Mekong mainstream. Green algae and diatoms account for 64% of the plankton Aquatic fungi were not observed by species; one quarter comprise blue-green McDonald et al. (1997), but evidence for algae and the remainder are composed of their occurrence was apparent in the boles Euglenophyta, Xantophyta, Pyrrophyta and of Barringtonia and Diospyros tree spp., Chrysophyta. Approximately 10% of the which were frequently “scarred through- diatoms are specific to the river or to the out with exposed patches of black, car- floodplain. Phytoplankton densities are bonised heartwood”, creating an effect generally higher in the flooded areas than reminiscent of a lightning strike. in the rivers, where they vary consider- ably, being about 40 times higher in the dry season than during the flood period (Nguyen and Nguyen 1991 cited in Lamberts 2001). McDonald et al. (1997) report an alga occurring in submerged green masses, apparently Hydrodictyon sp., usually found interlacing populations of Utricularia aurea Lour.

10 DISTRIBUTION OF HABITATS agriculture, much of which is probably the result of hundreds or thousands of years of human modification (e.g. Chandler 2003). In The floodplain vegetation can be crudely differ- places, scrub, grassland and agriculture form a entiated in irregular concentric bands radiating complex mosaic landscape (Davidson 2004). out from the dry season low water mark of Widespread and numerous permanent and sea- the Tonle Sap Lake (Fig. 1). Forest (up to 20 sonal wetlands are found beyond the dry sea- m tall) and scrub (up to 5 m in height) forms son low water mark. Large areas of the sea- a semi-continuous belt of vegetation, 7–40 km sonal inundation zone are effectively “common wide, surrounding the open water surface. land” and heavily utilised by the human com- Trees and scrub become lower in stature with munities surrounding the floodplain (e.g. increasing distance from the lakeshore and as Davidson 2004, Evans et al. 2005). duration of flooding decreases, and patchily distributed grasslands occur within this vegeta- tion belt (McDonald et al. 1997). The scrub is encircled by an outer band of grassland and

Fig. 1. Cross section of floodplain habitats in Kampong Thom province during (a) October/November (above) and (b) May/June (below); reproduced from Balzer et al. (2002).

FLOODLAND OF THE TONLE SAP & KOMPONG THOM DURING OCTOBER/NOVEMBER

FLOODLAND OF THE TONLE SAP & KOMPONG THOM DURING MAY/JUN

Open Lake Swamp Forest Short-tree Grasslands and Settlements Deciduous shrublands Floating Rice Dipterocarp Forest

DIVERSITY, FLORAL AFFINITIES AND ENDEMISM The Tonle Sap floodplain supports a low overall diversity of plant species, but is rich in species that are unique to the Indochinese floristic region (Rundel 2000), some of which are entirely restricted to the Tonle Sap floodplain (McDonald et al. 1997). The Cambodian National Mekong Committee (1998 cited in Rundel 2000) published a preliminary checklist

11 of approximately 200 vascular plant THE TERM “FLOODED FOREST” species present in the Tonle Sap flood- The dominant wooded habitat of the plain; this list needs to be evaluated to floodplain is most often referred to as determine its correctness and complete- “flooded forest”. However, the more ness (Rundel 2000). Based on four main descriptive “seasonally flooded freshwater sources (Rollet 1972, McDonald et al. swamp forest” is technically a more accu- 1997, Lamberts 2001 and Davidson rate term for these formations (e.g. Rundel 2004), 206 species are identified from the 2000). Seasonally flooded freshwater Tonle Sap floodplain, a further 19 addi- swamp forest differs markedly from clas- tional species that occur are unidentified, sic South-east Asian swamp forests in and six more species are listed but with- both structure and species composition, out a primary source for the data (plant because it is flooded seasonally, and not species list provided by TSBR Secretariat permanently. For example, palms, often a 2005). characteristic component of typical swamp forests, are almost entirely absent The swamp forest communities of the from the Tonle Sap floodplain but for two Tonle Sap floodplain appear to be unique or three rattan species (Rundel 2000), and both in structure and floristic composition, pneumatophores, aerial roots and vascular and contain a number of narrow endemics, epiphytes are all lacking (Rollet 1972). e.g. Samandura harmandii (B. Rollet) (Simaroubaceae), which occurs in the The Tonle Sap’s seasonally flooded fresh- Stung Sen Core Zone (McDonald et al. water swamp forests are adapted to with- 1997). Some elements of the freshwater stand twice-annual fluctuations in water floodplain’s flora show affinities with level of up to 10 m, and to cope with this coastal mangrove habitats (Rundel 2000). stress, have developed some very distinc- For example, Terminalia cambodiana tive physiological characteristics Gagnep. (Combretaceae) is endemic to the (McDonald et al. 1997, Rundel 2000): Tonle Sap and the southern coastal zone of Cambodia and adjacent Thailand; • Most seasonally flooded swamp forest species relationships in the genera plants are “flood-deciduous”, i.e. leaf Diospyros and Barringtonia show similar fall occurs underwater, when the flood patterns (Rundel 2000). McDonald et al. submerges the branches (leaf fall in (1997) reported the collection of what most tropical forests coincides with appears to be an undescribed taxon of the dry season periods of insufficient mangrove Lumnitzera cf. racemosa water stress); the flush of new leaves (Combretaceae) in the outer Tonle Sap occurs as receding waters (in floodplain. Mekong Basin endemics includ- November–December) expose the ing Coccoceras anisopodum Gagnep. trees and shrubs to the air again; (Euphorbiaceae), Diospyros bejaudii • Flowering and fruit production is Lecomte and D. cambodiana H. Lec. delayed for several months after the (Ebenaceae), Garcinia loureiri Pierre leaf flush, occurring during the late (Guttiferae), Acacia thailandica Nielsen dry and early wet season, suggesting (Mimosoideae) and Hydnocarpus saigonen- the possibility that fish may play a key sis Pierre (Flacourtiaceae) are well repre- role in seed dispersal. sented in the TSBR (Dy Phon 1982, McDonald et al. 1997, Rundel 2000). Most are locally distributed along river FOREST ASSOCIATIONS channels in relatively small and fragment- Two major forest associations are ed populations (Rundel 2000), for which described for the extensive floodplain area the lower Steung Sen catchment is an of Tonle Sap: i) a variously termed gallery important area (McDonald et al. 1997). (e.g. McDonald et al. 1997) or swamp for- est (e.g. Rundel 2000) formation around A complete list of plant species document- the lake itself, and along some of the ed from the TSBR is presented in Annex1. lake’s major tributaries (e.g. the Stung Sen

12 and Stung Sangkhe), and ii) a short-tree Coccoceras anisopodum, Crataeva rox- shrubland in the middle-outer floodplain burghii R.Br., C. nurvala Ham. (McDonald et al. 1997, Rundel 2000). In (Capparidaceae), and Terminalia cambodi- addition, an evergreen forest association ana), which tend to occur as co- or sub- with tropical rainforest affinities occurs dominant constituents with Crudia locally in riverine locations (McDonald et chrysantha, Caesalpiniaceae, Diospyros al. 1997). and Ficus spp., Garcinia loureiri, Elaeocarpus griffithii, Hydnocarpus Swamp or gallery forest anthelmintica and Samandura harmandii, This forest association receives inundation forming a more stratified vegetation cover (of 4-6 m, locally up to 8 m) for up to including trees with trunks reaching 1-3 m eight months each year, and occurs in diameter and 15 m in height (McDonald around the dry-season shoreline of the et al. 1997). Also present in the stratified lake itself and along the major river chan- canopy of this forest are large lianas of nels, as a mosaic of stands of large trees the rattan Calamus palustris Griff. and open areas with floating aquatic vege- (Palmeae), as well as a diverse assem- tation typical of the lake itself (McDonald blage of typical swamp forest lianas et al. 1997, Rundel 2000). The primary including Acacia thailandica, Breynia rham- dominants are two rather stunted (7- 15 noides, Cissus hexangularis Thorel m tall) tree species, Barringtonia acutangu- (Vitaceae), and Combretum trifoliatum. la (L) Gaertn. (Lecythidaceae) and The under storey of this community is Diospyros cambodiana, which typically occupied by a mixture of shrubs typical of occur with a variety of woody lianas, the short-tree shrubland association includ- including Combretum trifoliatum Vent. ing Brownlowia paludosa Kost. (Tiliaceae), (Combretaceae), Breynia rhamnoides Arg. Cudrania cambodiana Gagnep. (Moraceae), (Euphorbiaceae), Tetracera sarmentosa Gmelina asiatica L. (Verbenaceae), Vahl (Dilleniaceae) and Acacia thailandica. Mallotus cochinchinensis Lour. Barringtonia acutangula and Combretum (Euphorbiaceae), Phyllanthus sp, Vitex trifoliatum both retain their leaves year- holaedenon D.Dop (Verbenaceae) and four round (but flush new leaves contempora- locally unique, unidentified species, and neously with the deciduous species). A the under storey herb Schummanianthus similar gallery forest association formerly dichotomus (Marantaceae) was only occurred along channels of many major encountered in this habitat (McDonald et rivers in southern Cambodia (Rundel al. 1997). 2000). Other common tree species in this forest type noted by Rollet (1972) include This forest association contains a few Barringtonia micrantha (Lecythidaceae), species that do not delay flowering and Coccoceras anisopodum, Elaeocarpus grif- fruit production: e.g. Combretum trifolia- fithii Mast. and E. madropetalus Pierre tum, Crataeva nurvala and C. roxburghii, (Elaeocarpaceae), Hydnocarpus and Gmelina asiatica. anthelmintica Pierre and H. saigonensis (Flacourtiaceae), Albizia lebbekoides (A.P. Short-tree shrublands DC.) Benth. (Mimosaciae), and Cynometra This vegetation type consists of dense, sp. (cf. dongnaiensis) (Caesalpiniaceae) fairly homogeneous stands of 2–4 m tall and Stephegyne cf. diversifolia Rubiaceae trees and scrub. The flora of the short-tree (Nauclaeceae). shrublands is dominated by species of Euphorbiaceae, Fabaceae and Evergreen riverine forest Combretaceae (McDonald et al. 1997). An evergreen riverine forest habitat identi- Widespread common and/or dominant fiable by a tall and species-rich gallery for- species include Acacia spiralis est is described along the Stung Sen (Mimosaceae), Barringtonia acutangula embankments (McDonald et al. 1997). In (Lecythidaceae), Bridelia cambodiana addition to the typical swamp forest Gagnep. (Euphorbiaceae), Brownlowia species (e.g. Barringtonia acutangula, paludosa, Capparis micrantha, Cissus 13 hexangularis, Croton mekongensis geneity of soil moisture conditions and Gagnep. and Croton krabas Gagnep. seasonal flood dynamics as human distur- (Euphorbiaceae), Dalbergia pinnata Pierre bance (Rundel 2000). (Papilionoaceae), Gardenia cambodiana Pitard (Rubiaceae), Gmelina asiatica, Phyllanthus taxodiifolius Beille AQUATIC HERBACEOUS COMMUNITIES (Euphorbiaceae), Popowia diospyrifolia The shallow shoreline of Tonle Sap lake Pierre (Annonaceae), Quisqualis indica L. and patches within the gallery/swamp (Combretaceae), Stenocaulon kleinii W. & forests support dense mats of herbaceous Arn. (Asclepiadaceae), Terminalia cambodi- vegetation, 1-3 m tall, that may be emer- ana and Vitex holoadenon. gent from shallow water but are more typ- ically floating (McDonald et al. 1997). Several species (e.g. Combretum trifolia- True submerged species are lacking, but tum), which form woody lianas under wet- surface creepers capable of increasing ter conditions, form extensive areas of their stem length with rising water are scrub in the outer floodplain under the very common. These large, often clonal seasonal cycle of flooding and drought assemblages float freely over the lake, and disturbance. Herbaceous cover is colonising large openings and gaps within largely lacking in these short-tree shrub- the swamp forest. Extensive mats of the lands, but returns rapidly following clear- floating legume Sesbania javanica Miq. ance. Several alien woody species have (Papilionaceae), the grass Brachiaria muti- now invaded this habitat and are greatly cus Stapf., Polygonum barbatum L. expanding their cover and dominance (Polygonaceae) and the introduced (McDonald et al. 1997). The giant Eichhornia crassipes (Martius) Solms mimosa, Mimosa pigra L. (Mimosaceae), (Pontederiaceae) occur, and along the currently presents the most serious prob- Stung Sangkhe, Achyranthes aquatica lems (Jantunen 2003, Hellsten et al. 2003). R.Br. (Amaranthaceae) co-dominates with these species. Occasional widespread The low stature shrubland which domi- species include Commelina salicifolia nates much of the Tonle Sap floodplain Roxb. (Commelinaceae), Ipomoea aquatica has led some authors to describe this veg- Forsk. and I. chrysioides Ker.-Gawl. etation formation as forest degraded by (Convolvulaceae), Ludwigia adscendens logging and other human activities (see (L.) Hara. (Onagraceae), Pistia stratiotes L. Rundel 2000). The shrublands and swamp (Araceae) and Salvinia cucullata Roxb. forests of the Tonle Sap floodplain have (Salviniaceae) (McDonald et al.1997). been modified by centuries of human Given the long history of human activities activities (e.g. Chandler 2003), an impact around the lake, the structure and diversi- that has certainly accelerated over the ty of the aquatic flora has likely altered past decade. Although extensive areas from its original composition (Rundel now comprise short-tree shrublands, it is 2000). unclear whether or not these are second- ary formations. McDonald et al. (1997) A notable floristic character of the aquatic provide a compelling argument that these flora of Tonle Sap is the paucity of fami- formations are essentially a climax vegeta- lies of rooted monocots and dicots that tion, while others argue that it is more characterise similar habitats elsewhere in logically a result of repeated disturbance, Asia (Rundel 2000). These families especially burning, supported by the include the Hydrocharitaceae, observation that gallery forest exists only Marantaceae, Najadaceae, Nympaeaceae, on the wettest substrates, i.e. those Pontederiaceae and Potamogetonaceae where trees are safe from fire. While (McDonald et al. 1997). The large sea- much disturbance has clearly taken place, sonal change in water level of the lake the structure and composition of woody and its strong turbidity are the likely caus- vegetation on the floodplain appears to be es for the absence of these groups. One as much a function of the microhetero- rooted monocot, the grass Phragmites

14 karka Trin. (Poaceae), is a local dominant rice) is the dominant grassland type within at the mouth of Tonle Sap and at other the inundated forest-scrub of the inner and river inlets and exits. middle floodplain (e.g. Veal Srangai IBA A distinctive aquatic community occurs in where it is rather extensive), occurring in isolated ponds in the outer floodplain, in homogeneous dense mats mixed with which grasses and sedges are important, Leersia hexandra Sw. (Poaceae) (Davidson including Cyperus pilosus, Rhynchospora 2004). Burning promotes species diversity sp. and Actinoscirpus grossus (L.f.) in these grasslands: Oryza dominated Goetgh. & D.A. Simpson (Cyperaceae), as swards burnt during the previous dry sea- well as a group of dicot emergents, includ- son often support a wider range of ing Aeschynomene indica L. species, including the sedges Cyperus digi- (Papilionaceae), Impatiens sp. tatus Roxb. and C. iria L. (Cyperaceae), (Balsaminaceae) and lotus Nelumbo Aeschynomene indica, and tall stands of nucifera (L) Gaertn. (Nelumbonaceae). Panicum luzonense J. Presl. (Poaceae) Lamberts (2001) and Hellsten et al. (Davidson 2004). Further from the (2003) note that Nelumbo nucifera is lakeshore, Oryza rufipogon and Leersia missing from the inner floodplain. hexandra become less common, merging into Paspalum-type grasslands including co-dominants Paspalum scrobiculatum L. GRASSLANDS AND AGRO-ECOSYSTEMS (Poaceae) and other, unidentified rhizome- A wide variety of grassland types occur in forming species. Gmelina asiatica scrub is the middle-outer floodplain. Their structure a widespread dominant throughout these and species composition has largely been areas. Waterlogged substrates in the inner determined by a combination of physical and middle floodplain support stands of factors (the inter-related seasonal inunda- Phragmites sp./spp. (especially in the tion gradient, soil type and fertility) and Tonle Sap ‘delta, e.g. McDonald et al. land-use processes, deepwater (chiefly 1997), and (often extensive) Actinoscirpus floating) rice cultivation, burning and graz- grossus sedge beds and stands of ing (Davidson 2004). Deepwater rice culti- Sesbania javanica (Davidson 2004). vation—in which seed is hand-broadcast onto areas that flood naturally, and grown Large homogeneous areas of apparent fire- with minimal chemical inputs (Catling climax communities, including a distinctive 1992)—has modified the middle and outer Imperata-Arundinella-Eragrostis-Themeda floodplain areas for centuries, but has grass community, comprising dominant declined in extent by over 60% since the Imperata cylindrica (L.) P. Beauv. late 1960s (Catling 2001). (Poaceae) and Rhynchospora sedge sp./spp. (including Rhynchospora submar- Examination of MRC/GTZ (1999) land- ginata Kuk. (Cyperaceae)) and, patchily cover data and 2002 Landsat Thematic co-dominant Arundinella setosa Trin., Mapper (TM) raw satellite imagery suggest Eragrostis atrovirens (Desf.) Trin. and E. that grassland areas within and around the brownii (Kunth) Nees. (Poaceae), occur on Tonle Sap floodplain may cover more than impoverished soils subject to annual burn- 2,000 km2 (Davidson 2004), quite possi- ing. These areas also support numerous bly the largest extent of seasonally inun- tall stands of the Saccharum-like dated grassland habitat anywhere in Sclerostachya fusca (Roxb.) A. Camus. South-East Asia (Davidson et al. in prep.). and more patchy Cymbopogon cambogien- Grasslands occur throughout the flood- sis (Balansa) E.G & A. Camus, Themeda plain, especially in the middle and outer arundinacea (Roxb.) Ridl. (Poaceae) and reaches which receive shorter duration the sedge Fimbristylis dura (Zoll. ex inundations, and exhibit considerable Maritz.) Merr. (Cyperaceae) (Davidson diversity in both structure and community 2004). composition (Davidson 2004). In recent fallows, Panicum repens L. Oryza rufigpogon Griff. (Poaceae) (wild (Poaceae), Eragrostis atrovirens and other,

15 unidentified Eragrostis spp., dominate, with patchy Desmostachya bipinnata (L.) Stapf. (Poaceae), and short (15-20 cm) Fimbristylis dichotoma (L.) Vahl and F. umbellaris (Lam.) Vahl sedge swards. Tall Chrysopogon festucoides tussocks are widespread across drier and less recently cultivated substrates. On the outer edge of the floodplain, on very poor sandy soils, structurally simple, extensive sparse short grasslands occur, including Fimbristylis swards, with occasional tiny isolated clumps of the rattan Calamus salicifolius Becc. (Davidson 2004).

The wet-season floating ricefield agro-ecosystem near Roluos in Kompong Thom supports a community very similar to the floating aquatic vegetation mats of the open lake and gallery/swamp forest clearings described by McDonald et al. (1997), including Sesbania javanica, Eichhornia crassipes (introduced), Ipomoea aquatica, Ludwigia adscendens and Pistia stratiotes, together with Polygonum tomentosum Willd. (Polygonaceae), Nelumbo nucifera, Limnophila geoffrayi Bonati. (Scrophulariaceae), Nymphae sp., and Trapa natans L. (Trapaceae), the latter not noted by other authors, and perhaps a cultivar (Balzer et al. 2002).

Fig. 2. Vertical distribution of dominant plant species in the Tonle Sap floodplain (to 6m elevation), reproduced from Hellsten et al. (2003).

16 SITES WITH DISTINCTIVE VEGETATION were unique to the site in the McDonald ASSEMBLAGES et al. (1997) survey. Much of the seasonally inundated swamp forest and short-tree shrubland in the GLOBALLY THREATENED SPECIES TSBR comprises a characteristic mix of Only two of the plant species recorded the same dominant species. Particularly thus far (Annex 1) are afforded a global large tracts of this are found in the Prek threat status. However, the flora of the Toal Core Area, which is notable for its Tonle Sap floodplain is still rather poorly common and large (10+ m tall) known, and its importance in a regional Terminalia cambodiana trees, and in Boeng context is inadequately described. Chhma Core Area where there is a diversi- Research into the status if the substantial ty of vegetation heights and structures, number of endemics reported from the interspersed with patches of open water Tonle Sap floodplain and adjacent areas supporting extensive herbaceous (McDonald et al. 1997, Rundel 2000) may hydrophytic flora (McDonald et al. 1997). reveal that more species meet the global threat status criteria. Stung Sen Core Area and river- mouth Cynometra (cf.) inaequifolia IUCN Vulnerable Gallery forest along the Stung Sangkhe is Cynometra cf. inaequifolia was noted as a the most species rich, tall forest encoun- constituent species of the seasonally tered in the TSBR by McDonald et al. flooded freshwater swamp forest commu- (1997), and includes the largest number of nity of the Tonle Sap by Rollet (1972) but rare species. Common elements of the not by subsequent authors. It is unclear swamp or gallery forest formations typical whether this is due to a misidentification of many parts of the floodplain are co- or by Rollet (1972), or it it perhaps indicates subdominant to populations of a decline, natural scarcity or a localised Hydnocarpus anthelminthica, Ficus spp. distribution. This small-stature tree is (some very large examples), Elaeaocarpus known from Thailand, peninsular Malaysia, griffithii, Garcinia loureiri, Crudia chrysan- Malaysian Borneo and the Philippines, and tha and Samandura harmandii. has undergone a rapid decline due to ongoing clearance for agriculture and Moat Khla area extraction for timber (IUCN 2004). Large and dense gallery forests developed along landlocked waterways between Dalbergia entadoides IUCN Data Peam Bang and Moat Khla are a mature Deficient and relatively undisturbed example of This shrubby liana occurs commonly in the closed canopy inundated swamp forest. swamp or gallery forest formations of the Generally very typical in overall species Tonle Sap’s seasonally flooded freshwater composition, there are examples of swamp forest community (Rollet, 1972, endemics in this formation, including the McDonald et al. 1997). Its known range narrow endemic Homalium brevidens covers Cambodia, Lao, Thailand and Gagnep. (Flacourtiaceae). Vietnam (IUCN 2004). Outer floodplain north of Bak Prea This is an exemplary area of “upland” veg- etation on the Tonle Sap floodplain in terms of physiognomy and species compo- sition. The relatively short stature vegeta- tion supports emergent trees of up to 12m, and species diversity is higher than in formations associated with the lakeshore and riverways McDonald et al. 1997). Eight unidentified shrub species

17 MAMMALS

OVERVIEW internationally significant numbers. It The mammals of the Tonle Sap have not seems likely that one large mammal, the yet been well studied, and no specific sur- Irrawaddy Dolphin Orcaella brevirostris, veys have been conducted for any mam- was extirpated only very recently. mal species. Most field records are of inci- dental observations made during waterbird GLOBALLY THREATENED SPECIES and reptile surveys, and by the Prek Toal Conservation Team rangers. The other Loris sp. Nycticebus sp. main source of records is market speci- One loris was confiscated from a hunter mens and captive animals. within the Prek Toal Core Area in November 2000. The animal superficially Despite the almost total lack of specific resembled Northern Slow Loris Nycticebus historical information on mammals from bengalensis (IUCN Data Deficient), but the the Tonle Sap floodplain, it is highly likely unlikely possibility of it being Pygmy Slow that it was seasonally important for a vari- Loris Nycticebus pygmaeus (IUCN ety of large herbivores, when floodplain Vulnerable) could not be discounted (Goes forests were connected to other major for- and Hong 2002). Fishermen and hunters est blocks to the north, east and south- report that the species occurs in the Prek west (e.g. McDonald et al. 1997). Kol Toal Core Area (Sun Visal, WCS, verbally (1987 cited in MoE 1994) mentions that 2005). the Tonle Sap forests “feature a range of wildlife including troupes of elephants Long-tailed Macaque Macaca fascicu- (200-300 per troupe) wild buffalo and laris IUCN Near Threatened deer at certain times”. These may have This primate is still widespread and locally included Eld’s Deer Cervus eldi, which still common in taller inundated forest-scrub occurs in small populations just beyond within the TSBR, particularly around the the inundation zone (McShea et al 2005 lakeshore itself (e.g. Goes and Hong Seng et al. 2003) and the regionally extir- 2002). Large numbers (up to 170 individu- pated Hog Deer Axis porcinus. Whatever als in one consignment) have been seized their former status, however, wild large from hunters and traders, especially during herbivores have now been extirpated from the wet season, e.g. in Prek Toal (Goes the TSBR. These extirpations may have and Hong 2002) and in Kompong Thom occurred as recently as 10-20 years ago town (P.J.A.D. unpublished data). The (McDonald et al. 1997, P.J.A.D. unpub- population is likely to have declined as a lished interview data). The most recent result of hunting to supply captive breed- report of elephants was of a group seen ing facilities in Cambodia, Vietnam and crossing NR-6 in 1994 from the Tonle Sap China (e.g. Goes 2002b). Captive individu- floodplain travelling towards Boeng Per als are frequently seen in many of the Wildlife Sanctuary (N. Thayer pers. comm. floating villages around the lake (Goes and to D. Ashwell, 1994). Hong 2002).

The TSBR does not support a particularly It is common in Prek Toal Core Area (e.g. diverse mammal fauna, but several species Goes and Hong 2002), where it is encoun- that do still occur are listed as globally tered on a estimated 70% of visits during threatened (IUCN 2004), and may occur in

18 the wet season, when animals are more Recent surveys across Cambodia have visible than during the dry season in the revealed that T. v. germaini is relatively reduced area of available habitat (Sun widespread, with post-2000 records from Visal, WCS, verbally 2005). It also occurs the Cardamom Mountains, the Northern in the Boeung Chhma Core Area (Goes Plains (Preah Vihear) and the north-east and Hong 2002), the Stung Sen Core Area (Mondulkiri and Ratanakiri). The paucity of (Seng et al. 2003), and the Veal Srangai, records from elsewhere within its range Stung Sen-Santuk-Baray and Boeung underscore the regional conservation Chhma Moat Khla IBAs (Seng et al. importance of the Cambodian population. 2003). However, the relative significance of the Tonle Sap population within a national Germain’s Silver Leaf Monkey context cannot be assessed objectively at Trachypithecus villosus germaini IUCN present. Data Deficient This is the species referred to as Silvered Otters Langur Semnopithecus cristatus by many Most otter records in Cambodia originate previous authors. Recent taxonomic work from markets or captivity, and specific first considered T. v. germaini a full locality data only exists for two species, species in its own right, Indochinese Hairy-nosed Otter Lutra sumatrana and Lutung Trachypithecus germaini (Groves Smooth Otter Lutrogale perspicillata, both 2001), but this has since been rejected of which occur in areas of inundated for- (Brandon-Jones et al. 2004). Nonetheless, est and scrub within the TSBR (Poole given the uncertainties that remain over 2003). Otters in Cambodia are trapped for this species (and other primate)’s tax- their skins, which are used in Traditional onomies, Brandon-Jones et al. (2004) urge Khmer Medicine to assist women during that conservation measures be implement- pregnancy and childbirth, and can fetch up ed for as many subspecies as possible. to $45/kg (Walston 2005). The relative There are very few recent records of T. v. lack of otter records from habitats where germaini from a large part of its range, they would be expected to occur, com- which encompasses continental Thailand bined with the demand for skins, make and parts of Myanmar, Lao PDR, Vietnam otters a high conservation priority in and Cambodia, prompting concern that it Cambodia. The seasonally inundated for- may qualify for the IUCN threat category est and scrub in the TSBR, which may hold Vulnerable, or possibly Endangered (J. globally significant populations of Hairy- Walston in litt. 2005). nosed Otter, is the single most important priority area for otter surveys in Cambodia Germain’s Silver Leaf Monkey occurs in (Poole 2003). the Prek Toal and Boeung Chhma Core Zones (e.g. Seng et al. 2003), and Veal Hairy-nosed Otter Lutra sumatrana Srangai IBA (Goes et al. 2001). In Prek IUCN Data Deficient Toal Core Zone it is most regularly Two of the four confirmed records of this observed during the wet season, when species from Cambodia come from the animals become confined to discrete Tonle Sap: two captive individuals, which groups of trees by the flood waters (Sun had reportedly been caught along the Prek Visal, WCS, verbally September 2005). Kantiel within the Prek Toal Core Area, Field observations suggest the species were photographed in Prek Toal village in may be less well adapted to swimming June 1998, and a young captive animal long distances than Long-tailed Macaque was photographed in Peam Bang village Macaca fascicularis (Sun Visal, WCS, near the Boeng Chhma Core Area in pers. obs.). December 1998 (Poole 2003). Villagers in Prek Toal considered the species may be 19 close to extinction in the Prek Toal area cycles and motorboats (Baird and Beasley (Goes and Hong 2002). Hairy-nosed Otter 2005). As of 1997, occasional dolphin was also reported to occur at Boeung sightings have been reported from the lake Chhma (Doroshenko et al. 1998, Long (Baird and Beasley 2005), but there have 2003). been no confirmed or published sightings since 1969 (Lloze 1973). In March 2004, Smooth Otter Lutrogale perspicillata 567 kilometres of boat-based surveys con- IUCN Vulnerable ducted over 40 hours by the Mekong The only confirmed record of this species Dolphin Conservation Project yielded no from the TSBR is of an individual caught sightings. Interview surveys with 156 pro- by hunters in the inundated forest-scrub fessional fishers confirmed that dolphins near Kbal Taol village south of the Prek had not been seen in the lake since 1990 Toal Core Area in mid 2002 (Poole 2003). (I. Beasley unpublished data). This is the first time two otter species have been confirmed to co-exist at one Additional notes on mammal site (Prek Toal) in Cambodia. species Giant Flying Squirrel Petaurista sp. Unidentified otters Reported from the Prek Toal Core Area, An unidentified otter was reported where it is considered to be very rare (Sun between Phnom Kraom and Prek Toal in Visal, WCS, verbally September 2005). October 2001 (Goes and Hong 2002). A group of four otters was seen along the Asiatic Jackal Canis aureus Prek Preah Daem Chheu in March 2003 The only records reported by biologists are (Sun Visal, WCS, verbally September from the Kruos Kraom area within the 2005). Prek Toal Conservation Team Stung Sen-Santuk-Baray IBA, where it has rangers report seeing otters annually, in been observed twice, in July 2000 (Goes groups of 4-8 individuals, and usually in et al. 2001) and March 2002 (P.D. unpub- the dry season (per Sun Visal, WCS, lished data) in the outer floodplain grass- September 2005). Some stream owners land-scrub mosaic and deepwater rice complain that otters destroy fish traps (per agro-ecosystem. Farmers in the area Sun Visal, WCS, September 2005). report that jackals were formerly common, but have declined as a result of hunting Fishing Cat Prionailurus viverrinus for food (Goes et al. 2001). IUCN Vulnerable Frequently reported by local people, this Leopard Cat Prionailurus bengalensis elusive cat’s occurrence in the TSBR was One captive individual, photographed in supported when tracks were measured the Fisheries Office at Phet Sanday village, and photographed by dried up pools in the at the mouth of the Stung Sen in outer floodplain part of the Prek Toal Core Kompong Thom province on 2 December Zone in March/April 2002. Fresh tracks 1998 (Goes and Hong 2002), was the with measurements and a general impres- only record traced, and the only evidence sion matching that of Fishing Cat were that the species may occur within the seen in February 2002 in western Kruos TSBR. Kraom (P.D. unpublished data). Small Asian Mongoose Herpestes Irrawaddy Dolphin Orcaella brevirostris javanicus IUCN Data Deficient The few scattered records of this species Irrawaddy Dolphin formerly occurred in the (including from Prek Toal Core Area and Tonle Sap Lake, but may now be extirpat- the Siem Reap-Kompong Thom border ed. The population was decimated in the region), suggest a widespread distribution 1970s by the Khmer Rouge, who hunted (Goes and Hong 2002). This mammal is them for their oil, which was reportedly widespread in Cambodia, and seemingly used as a petroleum substitute for motor- relatively resilient to human disturbance in

20 the variety of landscapes it occurs in; as favourable. The Lesser Bandicoot Rat B. such it is not (currently) of conservation savilei has a more patchy distribution concern. (within the TSBR, it has thus far only been recorded from Kompong Chhnang Variable Squirrel Callosciurus fin- province), and is much less common. Field laysonii Rat Rattus rattus, which may comprise The distinctive deep rufous-coloured two distinct taxa, is widespread in most morph of this squirrel is widespread in the habitats and common in ricefields. forested areas of the floodplain (e.g. Ricefield Rat R. argentiventer only occurs Bailleux 2003, P.J.A.D. unpublished data). in the southern part of the TSBR, in Kompong Chhnang province; its distribu- Bats tion in Cambodia is restricted to the south- Lyle’s or Large Flying-Fox Pteropus east of the country. Lesser Ricefield Rat R. losea is less common than R. argen- lylei/vampyrus tiventer, but more widespread in the TSBR A roost of several thousand was pho- and locally common in Siem Reap, occur- tographed 8km south-west of Phnom ring in ricefield habitats in both the south- Kraom, Siem Reap province, in August east and north-west. Field Mouse Mus cer- 1997 (Sam 1997 cited in Goes and Hong vicolor occurs in the north-west of the 2002). Approximately 1,000 roosted in TSBR, and favours grassland habitats, trees at Dey Roneath in 2002 (month not including ricefields (Frost et al. in prep.). given), but were disturbed regularly by fishermen and hunters, and did not return Farmers interviewed by Frost et al. (in in 2003 (Sun Visal, WCS, verbally 2005). prep.) describe a strong link between the Roosts are occasionally reported in the presence of seasonally inundated shrub- Prek Toal Core Area, and residents of Kbal land and high rodent rice crop damage: in Taol village reported that there was a areas where scrub and forest have been roost near the Pursat/Battambang border cleared, crop damage is significantly in 2004. reduced, clearly indicating that short-tree shrubland is a key habitat for rodent popu- Small numbers of microchiropteran bats lations, probably as shelter and nesting are regularly observed (perhaps more in habitat during the inundation period. In the wet season) feeding over the lake sur- Kompong Cham, interviewees report that face, and roosts occur in pagodas around the fields closest to the floodwaters gener- the fringes of the floodplain (Goes and ally have higher levels of crop damage Hong 2002, P.D. unpublished data). from rodents than those further away Frost et al. (in prep.). These factors infer Rodents that the Tonle Sap inundation zone sup- In 2003-4, studies of rats in ricefield habi- ports significant populations and/or high tats have been conducted in four densities of several rodent species. Casual provinces around the Tonle Sap, Kompong dry-season field observations in Kompong Chhnang, Siem Reap, Banteay Meanchey Thom confirm this, where the abundant and Battambang. At least six species have rodent populations associated with deep- been trapped within ricefield habitats, i.e. water rice cultivation provide an important outside villages and the immediate com- prey base for substantial populations of mensal environments shared with human birds of prey (P.J.A.D. unpublished data). populations, and identified through molec- ular analysis of DNA and comparison with other specimens from the region (Frost et al. in prep.).

Bandicoot rat Bandicota indica is wide- spread in lowland rice growing areas, and is very common where conditions are

21 birds, Greater Adjutant Leptoptilos dubius BIRDS (Endangered), Lesser Adjutant Leptoptilos Birds are the best-studied faunal group javanicus, Spot-billed Pelican Pelecanus within the TSBR, and consequently their philippensis and Milky Stork Mycteria status is better known than for any other cinerea (all Vulnerable), and the Near faunal group. Two bird communities of Threatened Painted Stork Mycteria leuco- great global significance have been the cephala, Black-headed Ibis Threskiornis focus of WCS conservation and monitor- melanocephalus and Darter Anhinga ing projects since 2001. Counts and pro- melanogaster (BirdLife International 2001, tection of the large waterbird colonies 2003, Goes and Hong 2002, Goes in have been conducted in Prek Toal Core prep.). The site supports 10-30% of the Area since the late 1990s, and since global population of each of these species 2003 have been the subject of an increas- (except Milky Stork), totaling up to 5,000 ingly sophisticated monitoring system breeding pairs. Prek Toal is the only site in (Goes et al. 1998, Hong 1999, Goes and South-east Asia where Black-necked Stork Hong 2002, Goes 2005). Bengal Florican Ephippiorhynchus asiaticus (Near Houbaropsis bengalensis and other species Threatened)—a solitary nester—has been using the grasslands and agro-ecosystems confirmed to breed (Sun 2005). In addi- of the floodplain in Kompong Thom and tion, the Masked Finfoot Heliopais person- Siem Reap provinces have been the focus ata (Vulnerable) is an uncommon wet sea- of monitoring and research since 2001 son breeding visitor to the inundated (Davidson 2004, Hong and Ro 2005, swamp forest of Prek Toal Core Area Davidson et al. in prep.). In addition, large (Goes 2001, Goes and Hong 2002, Seng areas of the Tonle Sap floodplain, espe- et al. 2003). cially north and east of the lake, were covered during the Important Bird Area Internationally significant populations identification process in 2000-2002 (Seng (>1% of the Asian biogeographic popula- et al. 2002, 2003), and various ad hoc tion) of several other large waterbirds visits have been made to potentially inter- breed within the Prek Toal Cora Area: esting sites for birds at different times of Little Cormorant Phalacrocorax niger, year (Carr 1993, 1994, Mundkur et al. Indian Cormorant P. fuscicollis, Great 1995, Parr et al. 1996, Goes and Hong Egret Casmerodius albus and Asian 2002, Long and Sun 2002). Openbill Anastomus oscitans (Seng et al. 2003). An additional suite of large water- By June 2005, 210 bird species had been bird species, each of which is threatened recorded in the TSBR (Annex 3), of which as a breeding species, at least, in the adja- 17 are globally threatened or Near cent countries of Thailand (Round 2000), Threatened (BirdLife International 2004). Laos (Duckworth et al. 1999) and Vietnam (Safford and Buckton 2004), also occur in Breeding colonies of large water- substantial numbers within the TSBR: Woolly-necked Stork Ciconia episcopus, birds Glossy Ibis Plegadis falcinellus, Grey Heron Ardea cinerea, Purple Heron Ardea pur- Prek Toal Core Area purea and Great Cormorant Phalacrocorax The TSBR supports the largest remaining carbo. fragment of a bird megafauna that was once widespread across much of South- It is important to note that each large east Asia (Davidson et al. 2001). The vast waterbird species only uses Prek Toal dur- majority of these large waterbirds breed in ing its breeding season, and as such Prek the inundated forest and scrub of Prek Toal cannot be viewed in isolation when Toal Core Area. Prek Toal represents considering the conservation of large either the largest or the only remaining waterbirds in Indochina. After breeding, nesting site in South-east Asia for seven most individuals of all species disperse Globally Threatened colonial large water- widely across Cambodia, using other parts

22 of the Tonle Sap floodplain (e.g. Boeng This inland delta at the mouth of the lake, Tonle Chhma/Moat Khla and the Kompong and area of extensive Phragmites karka Thom grasslands), and areas north-west of beds and shallow water, where the Stung the TSBR (e.g. Ang Trapeang Thmor Sarus Sen River enters and the Tonle Sap River Crane Reserve) in particular, as well as the begins, seasonally supports 6,000+ Mekong delta and Cambodia’s northern Indian Cormorants (see above), 1,500 plains (e.g. Davidson et al. 2001, Seng Purple Swamphens Porphyrio porphyrio Kimhout et al. 2003). Small numbers (part of a regionally important population range farther, into north-east and central estimated at 10,000 birds in the TSBR), Thailand, southern Lao PDR and southern 500 Black-winged Stilts Himantopus Vietnam (Duckworth et al. 1999, BirdLife himantopus (1-10% of the Asian biogeogr- International 2004a, Buckton and Safford pahic population) and large numbers of 2004). migratory shorebirds (March - May) that utilise the mudflats that are only exposed Dei Roneath IBA at the height of the dry season (Carr The seasonally inundated swamp forest 1993, Rose and Scott 1997, Goes 2001). along the shores of the lake in Pursat province may support a significant colony Tonle Sap Lake of Lesser Adjutants (100+ pairs reported The open lake is an important feeding area in 2001) and Darters (1,000+ pairs for an estimated 2,000 Brown-headed reported in 2001), in addition to the Gulls Larus brunnicephalus (2% of the largest known colony of Purple Heron global population) and 10,000+ Ardea purpurea in Cambodia, if not South- Whiskered Terns Chlidonias hybridus (per- east Asia, comprising 1-10% of the Asian haps the highest known concentration in biogeographic population (Rose and Scott South-east Asia) (Goes 2001, Davidson 1997, Goes 2001, Goes and Hong 2002). 2005). The current status of these colonies is unknown because they are not monitored Bird fauna of grassland and agro- annually. ecosystems A suite of globally threatened species and Other significant swamp forest and internationally significant populations sea- permanent wetland sites for water- sonally utilise the grasslands and agricul- birds tural habitats of the middle and outer Tonle Sap floodplain. These habitats sup- Boeng Chhma Core Area and Moat Khla port the largest known breeding popula- The seasonally inundated swamp forests tion of Bengal Floricans Houbaropsis ben- and short-tree shrublands of the Boeng galensis (Endangered) in South-east Asia, Chhma Core Area are less well studied, and perhaps the world (Davidson et al. in but do support important post-breeding prep.). The floricans breed during the dry feeding concentrations (May-July) of and early wet seasons, from January to Greater Adjutant and Spot-billed Pelican July, when the rising flood forces them to and the highest count of Black-necked migrate across the agricultural belt of land Storks (6 in late June 1998) from the flanking NR-6 to patchy grasslands within TSBR (Goes 2001, Seng et al. 2003), open deciduous dipterocarp woodlands along with 3,500 Indian Cormorants and outside the inundation zone (Davidson 200+ Great Cormorants Phalacrocorax 2004). The largest known breeding con- carbo. The total population of Indian centrations are in Veal Srangai, Stung Cormorant in the TSBR is equivalent to Sen-Santuk-Baray and Stung-Chikreng- 30% of the global population, with key Kompong Svay IBAs in Kompong Thom concentrations occurring seasonally at and Siem Reap provinces, but large areas Chhnuk Tru, Prek Toal and Boeng Chhma of grassland and agro-ecosystem have not (Carr 1993, Rose and Scott 1997, Goes been surveyed (Seng et al. 2003, 2001). Davidson 2004).

23 The many small wetlands scattered (Near Threatened) nests in riparian larger throughout the floodplain grassland and bushes and trees flanking some of the agricultural habitats are important feeding Tonle Sap’s tributaries (Seng et al. 2003). areas for large waterbirds, chiefly when Red Avadavat Amandava amandava the flood recedes during November- January and when the flood rises again in Grassland habitats in the outer floodplain June-July, but also during periods of (especially in Stung-Chikreng-Kompong heavy rainfall mid dry season (P.D. unpub- Svay and Stung Sen-Santuk-Baray IBAs) lished data). Most important among these support healthy dry season populations of are the White-shouldered Ibis Pseudibis several grassland/agro-ecosystem special- davisoni (Critically Endangered), which is ists that have undergone substantial infrequently recorded in very small num- declines in other parts of South-east Asia bers (Sam 1999, P.J.A.D. unpublished (e.g. Duckworth et al. 1999, Round 2000, data), and large flocks of Greater Adjutant 2002). These include Small Buttonquail and Lesser Adjutant (maximum national Turnix sylvatica, Blue-breasted Quail counts of both come from the Tonle Sap Coturnix chinensis, Red Avadavat floodplain grasslands). In addition, large Amandava amandava, Striated Grassbird flocks of Black-headed Ibis, Painted Stork Megalurus palutris, Australasian Bushlark and Asian Openbill, and smaller numbers Mirafra javanica and Oriental Skylark Woolly-necked and Black-necked Storks Alauda gulgula. use these grasslands seasonally. Deepwater rice stubbles provide important Flocks of Sarus Crane Grus antigone food sources for large flocks of both (Vulnerable) use the drier grasslands as granivorous species, especially Yellow- migration staging posts, and in recent breasted Bunting Emberiza aureola (Near years a flock of up to 25 has spent the Threatened), which has undergone wide- non-breeding season (December-April) in spread declines at wintering sites across Stung-Chikreng-Kompong Svay IBA much of its range (BirdLife International (Davidson 2005). 2004a), and insectivores, including Oriental Pratincole Glareola maldivarum, The abundant rodent populations associat- Black Drongo Dicrurus macrocercus, ed with deepwater rice agro-ecosystems Bluethroat Luscinia svecica, Red-throated and grasslands (Jahn 1997) in Kompong Pipit Anthus cervinus and Yellow Wagtail Thom support internationally important Motacilla flava. As the flood rises in the populations of Greater Spotted Eagle early wet season, deepwater rice fields are Aquila clanga and a few Imperial Eagles important feeding habitats for Javan Pond Aquila heliaca (both Vulnerable) between Heron Ardeola speciosa, Watercock December and April, during the boreal Gallicrex cinerea and Purple Swamphen winter (P.J.A.D. unpublished data). Many Porphyrio porphyrio, Cinnamon Ixobrychus other raptors exploit this rich prey base cinnamomeus and Yellow Bitterns during the dry season, including large Ixobrychus sinensis (P.J.A.D. unpublished numbers of Eastern Marsh Harriers Circus data). spilonotus, and smaller numbers of Pied Harriers Circus melanoleucos, Black Kites Aerial insectivores occur in dense flocks Milvus migrans, Brahminy Kites Haliastur over grassland and scrub mosaic habitats, indus, which collectively must represent including Red-rumped Swallow Hirundo one of the most important raptor assem- daurica and Sand Martin Riparia riparia blages in South-east Asia. (P.J.A.D. unpublished data).

The largest-known wintering population of Bird communities of the swamp Manchurian Reed-warbler Acrocephalus forest and short-tree shrublands tangorum (Vulnerable) uses a variety of The inundated forest around the lake itself tall grass and scrub habitats, and Asian is rather species-poor in comparison with Golden Weaver Ploceus hypoxanthus other forest types (evergreen, semi-

24 evergeen, deciduous and deciduous dipte- observation platform, and at least 27 are rocarp) in lowland Cambodia. Nonetheless, visible from more than one platform. The it supports some important populations of potential margin of error in estimating resident birds. Most notable is the Grey- numbers is therefore considerable in a headed Fish Eagle Ichthyophaga ichthyae- year when large numbers of birds chose to tus (Near Threatened), which is wide- nest in trees that are not visible from spread and may occur at higher densities observation platforms: there exists poten- around the Tonle Sap than in any other tial to underestimate numbers by up to part of its wide Asian range. Estimates 35%. Counts made elsewhere (along the suggest that at least 15 pairs may breed lakeshore and at post-breeding dispersal in the Prek Toal Core Area, and the popu- sites) provide useful comparatives to the lation around the lake as a whole may colony counts, and a means of assessing exceed 100 pairs (Goes 2001, Goes and numbers for species that cannot be count- Hong 2002). It is a dry season breeder in ed at colonies, e.g. ibises. Boeng Chhma and Prek Toal Core Areas (Goes 2001), when adults are able to take The monitoring programme is undergoing advantage of the abundant fish build up continual improvements, to provide a sim- behind fishing lot traps. One other notable ple and sustainable method to generate element to the resident swamp forest bird annual population estimates. Nonetheless, fauna is that it supports one of very few the current platform-based monitoring sys- inland populations of Mangrove Whistler tem has already proved invaluable in a) Pachycephala grisola, perhaps a remnant enabling numerical assessments of the associated with elements the floodplain conservation significance of the Prek Toal flora that show affinities with coastal colonies, b) providing an extremely effec- mangrove habitats. tive tool for managing the Prek Toal colonies, including protection, enforce- Short-tree shrublands provide important ment, tourism and colony demarcation, c) roosting, loafing and cover habitats for evaluating the success of conservation species feeding in the more productive activities, and d) detecting threats else- adjacent grassland and wetlands. where in the landscape. Populations of many species for which the TSBR is internationally important use this For each of the species of global signifi- habitat, including all stork species, cor- cance, the following summary accounts morants and darters, herons and egrets, give minimum population estimates for most birds of prey and Manchurian Reed Prek Toal Core Area within a global con- Warbler. No bird species occurs exclusive- text, highlighting aspects of the species ly in this habitat, however. ecology that are important to their conser- vation. Globally important bird population statistics and ecological notes Greater Adjutant Leptoptilos dubius Estimating numbers of breeding large IUCN Endangered waterbirds at the colonies in the Prek Toal The Prek Toal Core Area population num- Core Area is complex and potentially bered a minimum of 51 breeding pairs in prone to large variances, largely due to the 2003-4 (WCS unpublished data), making it logistical difficulty of viewing the colonies. the second largest colony in the world, Only conspicuous tree-nesters can be and representing just over 10% of the counted at the colonies; this immediately global population (BirdLife International rules out ibises, which nest out of sight, 2004a). This number may indeed be high- low down in scrub. Storks, pelicans, er, because colonies are not always visible Darters and cormorants are tree-nesters, from the counting platforms (Goes 2005). and can be counted from the network of Greater Adjutant was also reported to 7-10 observation platforms. These species breed in the Boeung Chhma/Moat Khla and may occupy 690-880 trees each year, of Stung Sen Core Areas, but follow-up field which almost 250 are not visible from any surveys only found empty, collected nests 25 (Goes and Hong 2002). Up to 82 birds the Northern Plains breeding populations have been counted along the Prek Toal (see Greater Adjutant). Core Area lakeshore (Goes 2005), and post-breeding flocks of at least 85 have Spot-billed Pelican Pelecanus philippen- been counted on feeding grounds in inun- sis IUCN Vulnerable dated grasslands in Stung Sen-Santuk- Prek Toal Core Area supports the largest Baray IBA in Kompong Thom province in single colony of this pelican in the world, June-July (e.g. Goes 2001). estimated to number 900-1,100 pairs in 2005 (Goes 2005, WCS unpublished This stork, and its congener the Lesser data), representing approximately 20% of Adjutant, also breeds in Cambodia’s the global population (BirdLife International Northern Plains; twenty-one Greater 2004a). Although pelicans disperse widely Adjutant nests were counted in two across Cambodia and beyond post-breed- Northern Plains colonies in 2004-5 (Tan et ing, the species is better adapted to al. 2005). Adjutants in the Northern Plains remain in deeply flooded habitats than any breed during the wet season and/or early other large waterbird species, and as such dry season (between August and a proportionately larger number may remain February), a little earlier, but in some in the flooed habitats of the TSBR during cases overlapping with, the breeding peri- the wet seaso n. od for adjutants in Prek Toal (December- May). This implies that they comprise at Milky Stork Mycteria leucura IUCN least partially separate breeding popula- Vulnerable tions. Further research is required to better Up to ten pairs are estimated to breed understand seasonal movements and any with the Painted Stork colonies in the Prek linkages between these populations, which Toal Core Area. This number is rather disperse widely across Cambodia during insignificant in a global context, but this is non-breeding periods. the only known inland breeding population in mainland South-east Asia; the species Lesser Adjutant Leptoptilos javanicus breeds primarily in mangroves elsewhere IUCN Vulnerable (BirdLife International 2001). The A minimum number of 209 nesting pairs Cambodian population may soon be the was counted in the Prek Toal Core Area in only population in South-east Asia, with 2004-5, an increase from 149 pairs in only 10 birds remaining in Malaysia 2003-4 (WCS unpublished data). This is (Malayan Nature Society in litt. to C. the largest known breeding population in Shepherd, September 2005). South-east Asia, representing approxi- mately 10% of the global population Painted Stork Mycteria leucocephala (BirdLife International 2004a). Other IUCN Near Threatened (much smaller) colonies were reported in The Prek Toal Core Area supports a breed- Dei Roneat IBA in 2000-2001, and ing population in the region of 1,000 pairs Boeung Chhma/Moat Khla and Stung Sen (Goes and Hong 2002), equivalent to 20% Core Areas in 2001 (Goes and Hong of the global population. Accurate popula- 2002). The highest count (168) of the tion estimates for this species are ham- species anywhere in Cambodia in recent pered by the large proportion of birds that decades, perhaps a post-breeding aggrega- nest out of view of platforms each year. tion, was made in Veal Srangai IBA in April 2002 (Goes and Davidson 2002), when only 40 pairs were estimated to Darter Anhinga melanogaster IUCN breed in the Prek Toal Core Area (Goes Near Threatened and Hong 2002). Understanding the con- This species has increased dramatically servation requirements of the Cambodian since monitoring began in 2001, when population of this species will require 280 nests were counted; in 2004-5, that research into the seasonal movements and total had risen to 1,870, a factor of eight any linkages between the Tonle Sap and increase over four years, or a doubling of

26 the population annually. If this exponential lake. However the western extremity of increase continues (a similar pattern has these same ‘flooded forests’ which abut been shown by cormorants also), it will agricultural lands on the western flood- present management problems through plain, and where White-shouldered Ibis potential occupancy of nesting trees might be most likely found, has never required by other species that breed later been well surveyed, most survey work in the season, and with protection. having been focused on the ‘eastern’ lake- side. White Shouldered Ibis Pseudibis davi- soni IUCN Critical Black-headed Ibis Threskiornis Historically the species was described as a melanocephalus IUCN Near Threatened common bird in Cambodia in 1924 Prek Toal Core Area supports the largest (Delacour and Jabouille 1925), or less breeding colony of the species in South- numerous than Woolly-necked Stork, but east Asia (e.g. Goes 2001), but numbers nevertheless very abundant in the region cannot be reliably counted there because of Siem Reap and Angkor (Delacour they nest out of sight in thick shrubland. 1924). Other historical records and Post-breeding counts from Ang Trapeang accounts, in the Tonle Sap region include; Thmor Sarus Crane Reserve in Banteay Kompong Thom, common in this province Meanchey province peaked at 1,600 in during the 1920’s, (although the only spe- 2003 (Goes 2005). If only half of these cific record given is of five resting in a were breeding adults, 400 pairs are equiv- marsh near Kompong Thom, April 1924) alent to 8-15% of the global population (Delacour and Jabouille 1925); one seen (Goes and Hong 2002). at Angkor Wat roosting on a temple in 1924 (Delacour and Jabouille 1925); large Bengal Florican Houbaropsis bengalen- numbers seen between Kompong Thom sis IUCN Endangered and Siem Reap in December 1927 and Counts of displaying males at three sites January 1928 (Delacour 1929), at Siem monitored from 2002-4 gave an estimate Reap many observed in the 1920’s around of 64–122 at three sites covering a com- Boeng Tonle Sap (Delacour 1928); two bined area of 170km2 (Davidson et al. in collected at Siem Reap in August 1938 prep.). Examination of MRC/GTZ (1999) (Eames and Ericson 1996). land-cover data and 2002 Landsat This Critically Endangered species is now Thematic Mapper (TM) raw satellite very rare with populations persisting only imagery suggest that additional grassland in Indochina and on Borneo. The most areas within and around the Tonle Sap important populations in Indochina now floodplain may cover more than 2,000 km2 are found in Cambodia, with birds concen- (Davidson 2004), almost ten times the trated in four main areas. area surveyed. It is therefore possible that In the Tonle Sap region in contrast to the the total number of Bengal Floricans in species historical abundance the only and around the Tonle Sap floodplain will recent record come from two relatively be found to equal or exceed the total small areas of the southeastern floodplain number (530; BirdLife International 2004a) in Khompong Thom Thom Province, where thought to exist in the only other popula- up to 20 birds have been sighted on a tion, in the Indian subcontinent (Davidson number of occasions between 1999 and et al. in prep.). Breeding male floricans dis- 2004. Despite extensive survey work for play in a variety of grassland landscapes, Bengal Floricans in similar areas of the which are maintained by various combina- eastern and northern floodlplain there have tions of cultivation and fallowing, burning been no other survey records. Also the and livestock grazing land-uses. At a land- species has never been observed in the scape scale, territorial males aggregate in northwest in the extensive areas of scrub, larger areas of short grassland for display- forest, watercourses and pools around the ing, but tolerate a broad range of structur- colonies of large waterbirds in the Prek al habitat composition and land-use within Toal area of the inner floodplain of the their individual territories (Davidson 2004). 27 Much still needs to be learnt about the produced 6-10 birds. Smaller numbers are florican: it appears to have an exploded recorded at other outer floodplain sites lek mating system, in which female breed- (e.g. in Stung-Chikreng-Kompong Svay ing requirements may not be well reflected IBA; Seng et al. 2003). Greater Spotted by male habitat associations; annual pro- Eagles, together with occasional Imperial ductivity is not known and urgently Eagles Aquila heliaca (also IUCN requires detailed study. Floricans breeding Vulnerable), congregate in the cultivated within the floodplain make short-distance parts of the floodplain to exploit the high migrations across a 15-30km wide agricul- densities of rodents. Given the extent of tural belt to open woodland mixed with suitable habitat, the population using the patchy grasslands. The florican’s non- Tonle Sap floodplain’s agro-ecosystems is breeding (wet-season) distribution, habitat potentially larger than the population win- preferences and threats are poorly under- tering in Thailand, estimated at 60-100 stood, and are a priority for further birds (Mallalieu in prep.). research (Davidson 2004). Manchurian Reed Warbler Sarus Crane Grus antigone IUCN Acrocephalus tangorum IUCN Vulnerable Vulnerable The outer floodplain grasslands in Stung Since the discovery of a wintering popula- Sen-Santuk-Baray IBA regularly host flocks tion of Manchurian Reed Warblers (present of up to 30 Sarus Cranes (e.g. Goes and from at least December to early May) in Davidson 2002) migrating between their the Tonle Sap floodplain in 2001 breeding grounds in the northern and (Davidson 2001), the species has been north-eastern plains of Cambodia (e.g. proven locally common in a variety of Davidson et al. 2001) and non-breeding habitats in the middle and outer floodplain, congregations at Ang Trapeang Thmor including tall, dry grassland and dry shrub- Sarus Crane Reserve just north-west of land edge and freshly burnt tall grass and the Tonle Sap floodplain in Banteay scrub habitats, as well as sedge beds, Meanchey province (Goes 2004, Hong and Phragmites stands and lotus swamps Goes 2001). Since 2002, a steadily grow- (Goes and Davidson 2003, P.J.A.D. ing non-breeding population of Sarus unpublished data). There is so much habi- Cranes have been spending the dry season tat that superficially seems suitable for it (December to April) in the Prolay grassland in the TSBR, that if it occurs across much of Stung District, Kompong Thom of the floodplain at the densities we know province; by February 2005 this popula- it to occur in at one or two localities, it tion numbered 30+ (Davidson 2005), the will very likely prove to be too numerous only known non-breeding congregation of to be considered IUCN Vulnerable under this bird in the TSBR. small population criteria. Plans to survey representative habitats in different parts of Greater Spotted Eagle Aquila clanga the floodplain are being made for early IUCN Vulnerable 2006. Only one other site, in Thailand, is Historical evidence indicates that currently know to regularly support signifi- Cambodia once supported a regionally sig- cant numbers (tens of birds) in winter nificant wintering population of Greater (Round 1994), where a close association Spotted Eagle (e.g. Thomas and Poole with Phragmites beds (now a scarce habi- 2003). Up to six are recorded each winter tat in Thailand) has been demonstrated (January-April) in the 120km2 of grass- (Round and Rumsey in prep.). land-scrub-agricultural landscape in the Kruos Kraom area of Kompong Thom Yellow-breasted Bunting Emberiza province (e.g. Davidson 2001, Goes and aureola IUCN Near Threatened Davidson 2001, Goes et al. 2004), and a This small passerine, known in Khmer as single day-count at the site in March 2004 Chap Prey Veng (“the sparrow of Prey

28 Veng”) on account of its former abun- dance, has undergone widespread declines REPTILES at wintering sites across much of its range (BirdLife International 2004a). Large num- The Tonle Sap is perhaps the single most bers still occur in the TSBR: flocks num- important wetland for reptile conservation bering 5,000 birds have been noted as in Southeast Asia (Goes 2005). The enor- recently as 2000 (Goes 2000, P.J.A.D. mous extent of wetland habitat formerly unpublished data). The traditional floating supported an abundant reptile fauna (e.g. rice agro-ecosystems of the Tonle Sap Zhou 2002), which has been subject to floodplain are crucial wintering habitat for sustained high levels of exploitation for the species now that much of this habitat over a decade (Bailleux 2003). in its former strongholds in the Mekong Nonetheless, the TSBR is still believed to Delta (e.g. Prey Veng; Thomas and Poole support internationally significant popula- 2003) and similar habitats in Thailand tions of at least eight globally threatened (Round 2002) have been converted to reptile species, including the Critically intensive rice cultivation. Endangered Siamese Crocodile Crocodylus siamensis, at least six species of freshwa- Locally extinct large waterbirds ter turtle, including the Endangered Yellow-headed Temple Turtle Hieremys One large waterbird species has been annandalii, and Burmese Python Python extirpated from the Tonle Sap (and with it molurus (Near Threatened). Relatively little South-east Asia as a whole), the Greater herpetofaunal survey work has been con- Flamingo Phoenicopterus ruber, which was ducted, and many reptiles are now diffi- last recorded in 1935 (Delacour and cult to observe in the field because their Jabouille 1931, Thomas and Poole 2003). populations have been reduced to very One more large waterbird, the Great White low densities, thus recent status assess- Pelican Pelecanus onocrotalus was former- ments are generally based more on sur- ly numerous (Thomas and Poole 2003), veys of markets than animals observed in but is now on the brink of extirpation, the wild. The following accounts focus on with just two recent records, one reported the key species. from Boeng Chhma in January 1996 (Edwards 1999), and one photographed near Prek Toal on 25 March 2001 (Goes Globally threatened species and Falise 2001). Black-faced Spoonbill Platalea minor (IUCN Endangered) was Siamese Crocodile Crocodylus siamensis described by Delacour (1929) as “seen in The Siamese Crocodile was once abun- some numbers near Kompong Thom”. dant and widespread in the TSBR (e.g. There have been no recent records, and in Nao and Touch 1994), but wild popula- the absence of specimen evidence, the tions have plummeted in recent decades species’s occurrence is best treated as due largely to chronic over-harvesting of unconfirmed due to potential confusion eggs, young and adults to stock crocodile with Eurasian Spoonbill P. leucorodia, farms (Anon. 2004, Platt et al. 2004). which was recorded for the first time in Since 2000, wild crocodiles have contin- Cambodia at Prek Toal in March 2004 ued to be widely reported in the TSBR, (Goes et al. 2004). and at least 105 wild crocodiles were col- lected in the Tonle Sap region between 1998 and 2002 (Anon. 2004).

Widespread, dedicated crocodile interview, field sign and spotlight surveys were con- ducted in the TSBR from June 2000 to September 2001 (Platt et al. 2004). However, these spotlight surveys along almost 170km of waterways in the Prek Toal, Boeng Chhma-Moat Khla and Stung

29 Sen Core Areas did not yield any crocodile mensis x C. rhombifer in farms in Siem sightings, and no crocodile signs were Reap (Platt et al. 2004). The existence of encountered during daylight surveys (Long a large, unregulated crocodile farming 2002, Platt et al. 2004). industry in Cambodia is still the primary Interview reports indicate that very small threat to wild crocodiles in the TSBR, pro- numbers of Siamese Crocodiles persist in viding strong incentives for villagers to several areas of TSBR: the three core harvest the last remaining wild individuals. areas, Dei Roneat, Fishing Lot no.6 and inundated forest in Battambang province Habitat loss and degradation appears to adjacent to the Prek Toal Core Area (Platt have contributed little to their decline: sig- et al. 2004). Reported collection of hatch- nificant areas of suitable but unoccupied lings from Boeng Chhma-Moat Khla and crocodile habitat remains (Platt et al. Prek Toal Core Areas, inundated forest 2004). The TSBR probably retains the sin- adjacent to the Prek Toal Core Area, and gle most extensive contiguous tract of the Dei Roneath wetlands indicates that suitable habitat for Siamese Crocodiles in reproduction was occurring in these areas Cambodia. If adequate in situ protection of (Platt et al. 2004). However, densities are wild crocodile populations can be achieved probably so low that their viability may be through enlisting the cooperation of fish- questionable (Platt et al. 2004), and the ing lot owners and fishermen, conserva- total population may not exceed a few tion management prospects for the Tonle pairs (Long 2002). Sap Siamese Crocodile population would significantly improve, and reintroduction In 2005, two wild Siamese Crocodiles into secure areas of the TSBR could be (one female and one male) were caught in warranted, alongside rigorous monitoring Fishing Lot no. 1 by fishermen from near- to evaluate recovery efforts and detect by Kbal Taol village, and taken to Pursat future population trends (Platt et al.2004). Provincial Department of Fisheries, for future tag and release back into the same Esturine Crocodile Crocodylus porosus Fishing Lot (Heng Sovannara and Sun Historical Occurrence Visal, WCS, verbally, September 2005). There is strong anecdotal evidence that One adult Siamese Crocodile was discov- Estuarine Crocodile C. porosus—which is ered dead following a fire in inundated for- not globally threatened—formerly occurred est-scrub habitat at Moat Khla in March in the Tonle Sap, based on verbal reports 2005, and a wild crocodile sighting was and bas-reliefs of very large, aggressive reported from an area north of Boeng (human-attacking) crocodiles (Platt in Chhma Core Area by the Kompong Kleang prep.). The measurements of a crocodile fishing community in June-July 2005 mandible collected in swamp forest along (Heng Sovannara, WCS, verbally the northern shore of the lake provide the September 2005). first compelling physical evidence that Estuarine Crocodile formerly occurred sym- Platt et al. (2004) found no evidence to patrically C. siamensis in the Tonle Sap suggest that crocodilians other than C. ecosystem, as it does with other con- siamensis currently inhabit the TSBR. geners elsewhere (Platt in prep.). Recent However, there is the possibility that some interview survey data indicate that C. of the “wild” crocodiles present around porosus was extirpated from the lake 30 the Tonle Sap now may be animals that to 50 years ago (Platt et al. 2004). Its have escaped or been released from farms large size and aggressive behaviour would (Long 2002), and there are reports from have made it especially vulnerable to 2003-4 that captive crocodiles (of heavy persecution (Platt in prep.). unknown genetic stock) have escaped from Prek Toal (Sun Visal, WCS, verbally Freshwater turtles September 2005). This is a major concern Turtle populations in lowland wetland because of the presence of Cuban habitats throughout Cambodia have plum- Crocodile C. rhombifer and hybrid C. sia-

30 meted as a result of over-harvesting, Asian Box Turtle Cuora amboinensis chiefly for both legal and illegal export to IUCN Vulnerable China and Vietnam, for use in traditional Considered to be the second most numer- medicine and as food (e.g. Walston 2005), ous turtle in the TSBR after Malayemys and also for domestic consumption (Touch subtrijuga, but it is now considered et al. 2000). The TSBR is no exception: uncommon in the TSBR, which is consid- turtles here have sustained heavy harvest- ered to support a small population, and ing pressure since the early 1980s (Keng becoming harder to find in the wild (Long 2000, 2003). Most recent turtle records 2003). Recently recorded from Prek Toal derive from markets and captive individu- Core Area and a captive individual was als, few of which have verifiable locality photographed in Sary village near data. Eight species are reported or known Kompong Thom town (Stuart and Platt to occur in the TSBR, all of which are 2004). This turtle occurs in lowlands globally threatened. More detailed informa- throughout Cambodia, and its global range tion on the individual threatened species is extends from north-east India to the given below. Philippines and Indonesia (Stuart et al. 2001). It is believed to still be wide- Inundated forest-scrub with numerous spread in the lowlands of Cambodia, pools is considered the most important which probably support the largest popula- habitat for turtles during the dry season tion of the species in Indochina (Touch (Long 2003), particularly in areas with Saeng Tana et al. 2000, Stuart et al. high numbers of fruiting trees (Sun Visal, 2001). WCS, verbally September 2005). Most turtle species are thought to breed in the Black Marsh Turtle Siebenrockiella dry season, when human disturbance and crassicollis IUCN Vulnerable the risk of fire are highest; indeed, use of This species tends to occur more com- fire is reported as a method for hunting monly in floodplain ponds away from the turtles in forest habitats in the floodplain Great Lake itself, where it is considered during the dry season (Balzer et al. 2002). rare, but is not favoured for its meat, Some species lay eggs in submerged sub- which is considered by many to be either strates within ponds that later dry-out, foul-smelling or even inedible (Holloway et others spend the dry season partially or al. 2000). This turtle ranges from wholly submerged in mud at the bottom of Myanmar to Indonesia; the Cambodian ponds that dry out. Both are susceptible population is potentially large and may be to collection by professional hunters with the most important population in the dogs. Turtle nests have a variety of natu- region (Touch Saeng Tana et al. 2000). ral predators, including the macaques (locally numerous), rats (abundant) and Yellow-headed Temple Turtle Hieremys some birds (e.g. coucals and corvids, both annandalii IUCN Endangered locally common). Considered the third most common species in the TSBR within Kompong River Terrapin Batagur baska Critically Chhnang province, based on interview sur- Endangered veys there in 2000 (Holloway et al. The discovery of two complete shells of 2000), and known from the Prek Toal area this species, found buried in lake sedi- (Stuart and Platt 2004). Cambodia is ments in 1985, confirmed the historical probably the most important country in occurrence of the “Royal Terrapin” Indochina (including Thailand) for the con- (Cambodia’s national reptile) in the Tonle servation of this species (Touch et al. Sap Lake (Platt et al. 2003). Elderly resi- 2000), and the TSBR may be the most dents reported that small numbers were important area for its conservation in present in the early 1900s. Surveys in Cambodia (Holloway et al. 2000). It is 2000-2001 found no evidence of the know to occur in Prek Toal Core Area species continued existence and it is con- (Stuart and Platt 2004), has been record- sidered almost certainly extirpated from ed in markets at Chong Kneas, Kompong the TSBR (Platt et al. 2003). 31 Thom town (Stuart and Platt 2004), and reported from the TSBR, but there are no was reported by hunters in Kompong Leng confirmed specimens or photographs. It Commune, Chornouk Village and Kompong was reported during interview surveys in Chhnang town (all Kompong Chhnang 1998 (Doroshenko et al. 1998), and there Province). continue to be unconfirmed reports (D. Emmett, Conservation International and Giant Asian Pond Turtle Heosemys Sun Visal, WCS, in litt. 2005), although it grandis IUCN Vulnerable was not recognised during interviews in Reported from the TSBR within Kompong Kompong Chhnang province in 2000 Chhnang province (under the name (Holloway et al. 2000). Despite very few Orange-headed Temple Turtle), based on confirmed recent records, Cambodia is interview surveys there in 2000, and is suspected to support a regionally impor- thought to be scarce (Holloway et al. tant population, perhaps the most impor- 2000). This species half-buries itself in tant in South-East Asia (Touch et al. muddy substrates in ponds that subse- 2000). quently dry out during the dry season. Elongated Tortoise Indotestudo elonga- Malayan Snail-eating Turtle Malayemys ta (IUCN Endangered) subtrijuga IUCN Vulnerable This species has been recorded in markets Known colloquially as the “ricefield turtle”, in Kompong Chhnang (e.g. Holloway et al. this species is reportedly the most numer- 2000), but its occurrence within the TSBR ous turtle around the Tonle Sap Lake has not been confirmed with either speci- (Holloway et al. 2000, Keng 2003). It is men or photographic evidence. favoured for its meat and is also used in traditional medicine (Balzer et al. 2002). It Pythons occurs in floating ricefields in Kompong Thom province between August and Burmese Python Python molurus IUCN December; outside this period it is consid- Near Threatened ered by local people to inhabit inundated The Stung Sen and Boeung Chhma Core forest habitats (Balzer et al. 2002). It Areas are reportedly important areas for occurs in flooded ricefields as well as nat- both Burmese and Reticulated Pythons ural wetland habitats, and is declining Python reticulatus (Long 2003). Pythons across its range, which stretches from are hunted heavily in both areas (Long Thailand and Indochina to Indonesia 2003), for their skins, meat and blood to (Stuart et al. 2001). serve both domestic and international mar- kets. Some fishermen in the Prek Toal Asiatic Softshell Turtle Amyda carti- Core Area use specific net types to try laginea IUCN Vulnerable and catch pythons (Sun Visal, WCS, ver- Strongly favoured for its meat, it is vari- bally September 2005) while others catch ously regarded as common, in the Tonle them by hand. Immature individuals are Sap Lake, although has undergone a seri- occasionally caught in gillnets set by fish- ous decline (Keng 2003), and quite rare, ers targeting fish or water snakes. in Kompong Chhnang (Holloway et al. Female pythons make their nests within 2000). This turtle is also widespread in dense scrub, on the ground, during the dry Asia, occurring from Myanmar to season, typically laying 20-30 eggs. In Indonesia, and inhabits various wetland May 2005, three dead pythons were dis- habitats from estuaries to montane covered curled around their eggs in the streams (Touch et al. 2000, Stuart et al. Prek Toal Core Area, following forest fires 2001). (Sun Visal, WCS, verbally September 2005). Asian Giant Softshell Turtle Pelochelys Although Burmese and Reticulated cantorii IUCN Endangered Pythons clearly occur sympatrically in the This enigmatic species has often been TSBR, it is not clear what their relative

32 statuses are, or whether any ecological The enormous scale of this snake harvest segregation exists. Reticulated Python is is a very recent phenomenon. Since the not IUCN Red-Listed (IUCN 2004). late-1990s, snakes have become one of the main protein sources for captive-bred Selected additional reptile species crocodiles in the burgeoning number of accounts crocodile farms around the lake. In addi- tion to the species also targeted for their Watersnakes skins, a proportion of the catch of several species is also used for human consump- The abundance of reptiles within the TSBR tion (Stuart et al. 2000, Brooks et al. is perhaps best illustrated by what is the 2005). The watersnake harvest has two largest exploitation of any single snake peaks, between July and August (wet sea- assemblage in the world (Stuart et al. son), and again in November-December 2000): an estimated 3.8 million snakes (early dry season) (S. Brooks verbally were harvested between June 2004 and September 2005). The first and largest January 2005 (Brooks et al. 2005). This peak of the harvest coincides with the harvest principally targets species in the breeding of E. enhydris (Brooks et al. sub-family Homalopsinae (Homalopsine 2005) and the second peak occurs just watersnakes). The rainbow Watersnake E. prior to the breeding seasons of many of enhydris is the most abundant species the other species, as the water starts to caught, accounting for 72-ca.80% of the recede (S. Brookes verbally September number of individuals in the overall catch 2005). It is not known what effect this (Stuart et al. 2000, Brooks et al. 2005). has on recruitment. The Tonle Sap Watersnake E. longicauda is the second-most abundant species, and The seasonal distribution and habitat use accounts for almost 50% of the total of watersnakes in the Tonle Sap is still caught in the southern TSBR (Brooks et al. very poorly understood. The highest catch 2005). This poorly known species is success is achieved by hunting in the shal- endemic to the Tonle Sap Lake and River, low edge of the lake, which can move up and is Cambodia’s only known endemic to several kilometres back and forth each reptile. As such, it is of special conserva- year (Brooks et al. 2005). Although fac- tion interest, and is a focus of the tors affecting catchability cannot be sepa- Cambodian Snake Trade Project, (Brooks rated from density effects, this observa- et al. 2005). Other species targeted (in tion is corroborated by research else- addition to pythons – see above) include where, which has shown that some of Puff-faced Watersnake Homalopsis bucca- these species are closely associated to ta and Bocourt’s Watersnake Enhydris shallow-water edge habitats and mud-root bocourti, both of which have been collect- tangle (Voris and Karns 1996, Murphy et ed over several decades for their skins, al. 1999, both cited in Brooks et al. which are exported to Thailand, Vietnam 2005). Water snakes are not caught in and China (Stuart et al. 2000, Long 2003, open water at the height of the dry sea- Brookes et al. 2005). The Bocourt's water son, when water has receded from the snake is also traded alive for its meat, forest (in March-April). This indicates that supplying restaurants locally and regional- the snakes may remain in the forest; both ly. Long (2003) considers Bocourt’s E. enhydris and H. buccata have been Watersnake to be locally rare as a result found aestivating in the dry floodplain of of over-exploitation. Other species that the lake during this period (Brookes et al. occur in the harvest include Cylindrophios 2005), inferring a close year-round associ- ruffus, Xenochrophis piscator, Erpeton ation with forest habitats. tentaculatum, Enhydris plumbea, Enhydris subtaeniata (misidentified as Enhydris Additionally during the dry season other jagorii in Stuart et al. 2000), Boiga occel- aquatic snakes are caught as a fisheries lata, Boiga cyanea, Naja spp. and Ptyas by-catch in the open water of the lake. korros. Species so far recorded are the granulated

33 file snake, Acrochordus granulatus and a sea snake, Enhydrina spp, yet to be fully FISH identified. These species do not occur in Overview of the Lower Mekong the large scale harvest occurring in the inundated forest and are therefore likely to Basin and Tonle Sap fish fauna be associated with the deep water habitat. Fish are the largest vertebrate group in the Tonle Sap ecosystem, both in terms of number of species as well as biomass (Lamberts 2001). The Mekong Basin as a AMPHIBIANS whole supports an exceptional freshwater fish diversity, numbering between 1,200 This is the least-studied of the vertebrate and 1,700 species (Coates et al. 2003), groups; almost no information exists on and in terms of overall biodiversity the amphibians in the TSBR. Two species Mekong River is among the top three occur in the deepwater rice agro-ecosys- rivers in the world, after the Amazon and tem in Kompong Thom province, the frog the Zaire (Dudgeon 2000). About 500 Hoplobatrachus rugulosus and the toad species have been described for the Bufo melanostictus (Balzer et al. 2002). Mekong system in Cambodia, which Each is reportedly abundant, particularly includes the Tonle Sap ecosystem between June and September. The frog is (Rainboth 1996), but the precise number a local delicacy and is commonly seen in of species that occur in the Tonle Sap is Kompong Thom markets. The toad is used simply not known. Fisheries data tends to as an anthelmintic for cattle, is also eaten, underestimate the number of species, due and is sometimes exported to China to uncertainty over identifications and the (Balzer et al. 2002). McDonald et al. use of local names that often cover multi- (1997) report seeing a medium-sixed tree ple biological species. The lack of a practi- frog inhabiting the thatched roof of a cal, comprehensive fish species identifica- floating home in Moat Khla; this was the tion guide for use in the field by local data only amphibian observed in the wild during collectors contributes to this uncertainty one month of dry season field surveys in (Lamberts 2001). The Tonle Sap ecosys- the seasonally inundated swamp forests tem is expected to lack localised and short-tree shrublands. endemism (Rainboth 1996).

It is likely that the amphibian fauna of the In terms of fish biomass, the Tonle Sap is TSBR exhibits close affinities with that in widely believed to be one of the most pro- the adjacent lowlands. A collection of 18 ductive (and most heavily harvested) amphibian species was recently made in freshwater ecosystems in the world. the vicinity of the Angkor Wat temple Annual inland fish production in the lake complex; a number of these would be between 1994 and 1999 was estimated expected to occur in the TSBR (B. Stuart at 179,500-246,000 tons, which in terms in litt. 2005). of yield per hectare per year is equal to or greater than that in Amazonian, Bangladeshi, Indonesian and Thai flood- plain ecosystems (Lieng and Van Zalinge 2001, also presented in Baran 2004).

Fish migration and life cycle patterns The Tonle Sap fish fauna is characterised by extensive and diverse migration pat- terns, both within the Tonle Sap ecosys- tem itself, and between the Tonle Sap and the Mekong. A chiefly passive annual migration of large numbers of eggs, fry, juvenile and adult fish into the Tonle Sap

34 Lake begins as the flow in the Tonle Sap White fish are mainly associated with the River reverses with the onset of the wet main channels and streams, and exhibit season. This migration, of fish at different strong lateral and longitudinal migrations, stages of development, is for many including into floodplains. They move into species very important, as it determines the Tonle Sap as the flood rises in the the initial stock levels and therefore the early wet season (from May), remaining potential final production (Lamberts 2001). there to feed and reproduce, until the Migration out of the Tonle Sap ecosystem waters begin to recede (from November), is also essential for many species (simply when they return to the turbid (“white”) for survival), and minimum levels of fish waters of the Tonle Sap and Mekong migrating into and out of the ecosystem Rivers, where they spend the majority of are another essential parameter determin- each annual cycle (e.g. Nao 1997, Hortle ing the lake’s productivity (Lamberts et al. 2004). The white fish includes many 2001). Fish migrations are generally classi- cyprinids (Cirrhinus microlepis, Hampala fied as either lateral (from main macrolepidota, Barbodes altus, water–floodplain) or longitudinal (from Leptobarbus hoeveni, Osteochilus lake–channel–Mekong River). melanopleura, Morulius chrysophekadion), various Pangasius sp., Siluridae (e.g. Most descriptions of the fish fauna of the Wallago attu, Micronema apogon) and Lower Mekong Basin generally differenti- Notopteridae (e.g. Notopterus chitala and ate two classes, black fish and white fish, N. notopterus) (Baran 2004). based on life cycles, water quality require- ments and migratory behaviour/distribution Opportunists are small, fast-growing patterns, which in turn are very closely species, able to make maximum use of the linked to the seasonal fluctuations in flood period for prolific reproduction hydrology (e.g. Lamberts 2001, Baran and/or growth. This group (which are also 2004, Hortle et al. 2004). Two further classified as white fish by some authors) categories can usefully be added to this comprises mainly cyprinids (e.g. simplistic overview, opportunists and Henicorhynchus siamensis, Thynnichthys mainstream associates (Mekong thynnoides, Dangila spilopleura), which are Secretariat 1992 cited in Lamberts 2001). harvested in large quantities for the manu- facture of prahoc, fish sauce and feed for Black fish are chiefly residents in the caged fish. “Trey Riel” Henicorhynchus Tonle Sap, inhabiting relatively clear-water spp. are the most common and account swamps and plains year round and make for the largest catch by weight of any limited lateral migrations. The waters they genera in Cambodia (van Zalinge et al. inhabit are tea-coloured by chemicals dis- 2000). solved from floodplain vegetation, the decomposition of which increases acidity Mainstream associates are only a mar- and depletes oxygen, stresses to which ginal group in the catches of the Tonle black fish are adapted (Hortle et al. 2004). Sap ecosystem, primarily occurring in Most species are specifically adapted to main channels of larger rivers. The group such conditions, and can breathe air, are includes clupeids, sciaenids and soleids. able to move overland in search of fresh waterbodies, and some can survive out of Complex variations exist in the migration water for long periods (Hortle et al. 2004). strategies adopted by white and black Black fish are mostly carnivorous and fish. Most respond to hydrological detritus feeders, and include the changes and use hydrological events as Channidae (Snakeheads), Clariidae, cues for the timing of their migration (e.g. Bagridae (Mystus spp.) and Anabantidae Poulsen et al. 2002). Many longer-dis- (Baran 2004). tance migratory fish species move into Tonle Sap at the beginning of the annual flood (May-June) and return to the Tonle Sap and Mekong River systems at the end 35 of the flood (October-November), produc- • Sharp declines in the catch of larger ing two peak migration periods. The and more valuable fish, and Tonle Sap Lake forms part of the Lower • Decreasing mean catch per capita and Mekong Migration System, and is a key unit effort site for both lateral (from floodplain to river channel) and longitudinal (up and Each of these features is characteristic of down the Mekong) migrating species, proven declines in freshwater fisheries including several globally and regionally elsewhere (Welcomme 1995). threatened fishes. Many fish species that Standardised long-term and large-scale are resident in the TSBR make short-dis- monitoring data are required for fish popu- tance movements each year in response to lations in the TSBR to statistically prove the flood-pulse, to utilise the different these observed declines, however. Several floodplain habitats. fish species that occur in the Tonle Sap are globally threatened. Several others are Rises in water levels in the early wet sea- of conservation concern. son trigger most migratory species to move down the Mekong and into the Fish diversity, distribution and habi- Tonle Sap, spending the flood season tat associations feeding in the fertile floodplain habitats, One hundred and forty-three fish species particularly the inundated swamp forest were identified from five sources (Lim et and short-tree shrublands, which provide al. 1999 van Zalinge et al. 1999, key breeding, nesting, spawning, feeding Lamberts 2001, Neou et al. 2001, Balzer and nursery areas for many fish species et al. 2002) as having been recorded in fish (and their predators, e.g. waterbirds), the Tonle Sap Great Lake and its flood- and are consequently essential for fish- plain, the Tonle Sap River and the transi- eries productivity (Lieng and van Zalinge tional zone between the two (the principal 2001, Poulsen et al. 2002). Shrub habi- channel, old channel and ox-bow lakes tats on the Tonle Sap floodplain are key around the mouth of the lake) since 1995 for these life cycle stages in many fish (Annex 2). A further 14 potentially differ- species. Throughout Cambodia a four- ent taxa were not conclusively identified month fishing ban is instituted to coincide to species level. with the main fish breeding period, in order to promote the sustainable develop- Lim et al. (1999) compared 1995-7 fish- ment and use of fishery resources. In the eries catch data with previous studies st TSBR, this ban falls between June 1 and (1936-1976), and found that only 54% of st October 31 (Baran 2004). species that had been caught historically (based on Chevey 1936, Bardach 1959, Fish population trends Fily and d’Aubenton 1966, Rainboth As early as 1959, local fish catch declines 1996) were still present. This indicates a around the Tonle Sap were being reported potentially substantial loss in fish species by fishermen (Bardach 1959). It is widely richness within the TSBR, but differences perceived that total catches on The Tonle in fishing techniques employed and the Sap are now close to a maximum (e.g. areas being compared (e.g. previous stud- van Zalinge and Touch Seang Tana 1996, ies appeared to include data from the Baran et al. 2001, Baran 2004). The abun- Mekong River) were not accounted for in dant observational evidence for this the analysis. includes: Sampled catches from four locations in • An overall decrease in average fish the Tonle Sap Lake, one in the Tonle Sap size, River and one in the intermediate zone, • An increase in the proportion of small were dominated by fish species belonging fish in catches, to three orders, the Cypriniformes (chiefly • A decrease in the catch of medium carps and minnows), Siluriformes (catfish- sized fish, es) and Perciformes (perch-like fishes) (Lim

36 et al. 1999). Cyprinidae was the dominant of some Cyprinidae (in the genera family, represented by 40 species. The Cyclocheilichthy, Leptobarbus, Hampala Tonle Sap River and intermediate zone and Labeo) than the Tonle Sap River supported more species than the lake (Chevey 1936 in Lim et al. 1999). itself, although this may simply reflect the timing of fishing activity from which the Globally Threatened Fish Species data was taken (November-April, peaking Mekong Giant Catfish Pangasianodon in December-February) falling outside the gigas (IUCN Critically Endangered; CITES main period that longer-distance migratory Appendix I) fishes are using the Tonle Sap lake and The Mekong Giant Catfish Pangasianodon floodplain. gigas is one of the largest and rarest freshwater fish in the world. It is endemic The Tonle Sap River fish fauna is dominat- to the Mekong River, with a historical ed by the genera Thynnichthys, range encompassing the Tonle Sap Lake Puntioplites, Dangila (Cyprinidae) and and River, and the Mekong River from Pristolepis (Nandidae); the transitional southern China south to Vietnam. It zone is characterised by the genera measures up to three metres in length and Cyclocheilichthys, Cirrhinus, Leptobarbus, can weigh in excess of 300 kg. It is most Osteocheilus and Morulius (Cyprinidae); easily differentiated from other species by and the lake itself is inhabited by its size, lack of teeth and barbels (in indi- Barbodes, Hampala (Cyprinidae), viduals over 50cm), and eight or nine Micronema (Siluridae), Pangasius pelvic fin rays. It has a broader head and (Pangasiidae), Trichogaster (Belontidae) mouth than its close relative P. hypoph- and Channa (Channidae) (Lim et al. 1999). thalmus (Roberts and Vidthayanon 1991). Species diversity is generally higher in the Fisheries in the Upper Mekong Basin floodplain and inundated forest during the began targeting the species as early as flood, but higher in the braided, deltaic 1890, when Pavie reported harvests of mouth of the Tonle Sap (transitional zone) 6,000 fish from Luang Prabang (Roberts during the migration period (Lim et al. and Vidthayanon 1991). By 1935 fisher- 1999). Ricefields in the middle-outer flood- men were targeting the species through- plain support at least 60 fish species out the Mekong Basin (Giles 1935), and it (Balzer et al. 2002). Fish species inhabit- has since disappeared from Luang Prabang ing floodplain and riparian forest mainly and north-east Thailand fisheries (Smith spawn between May and August (Lim et 1945, Phukasawan 1968, Mengumpun al. 1999), except some Cypriniformes and 2000), and has been declining in fisheries Clupeiformes (includes shads and sar- in far northern Thailand since at least dines), the Pleuronectiformes (flatfishes) 1990 (Mengumpun 2000). In 2003, its and some Tetraodontiformes (triggerfishes global threat status was elevated to and boxfishes). Local people report the Critically Endangered (IUCN 2004), on eggs of four species from ricefield habitats account of an estimated rate of decline— in Kompong Thom: two catfish, Clarias based on annual catch data (used as a macrocephalus and C. batrachus, Channa proxy for population size)—of more than striata (Chevron Snakehead) and Anabas 80% in two decades in the Mekong River testudineus (Climbing Perch) (Balzer et al. Basin area prior to the late 1990s (Hogan 2002). The areas and dates of spawning 1998). It has been listed on CITES for some families, including Engraulidae, Appendix I since 1975. Belonidae, Mastacembelidae, Soleidae and Cygnoglossidae, are still unknown (Lim et Knowledge of the species ecology is al. 1999). The transitional zone is also scant, and very little data exists on its considered important fish spawning and natural migratory or spawning behaviour. nursery habitat (Lim et al. 1999). Growth The Tonle Sap Lake and Tonle Sap River studies showed that the floodplain and appear to provide important habitat for the lake habitats sustained more rapid growth species, especially Fishing Lot #2, which 37 may be an important nursery area, sustain- Jullien’s Golden Carp Probarbus jullieni ing a significant population of young (rear- (IUCN Endangered; CITES Appendix I) ing) fish, while the Tonle Sap River is a This distinctive fish, also known as the migratory corridor for mature fish moving Seven-line Barb, can weigh over 70kg, to the Mekong River (Hogan et al. 2001, although 5-20kg individuals are more com- Zeb Hogan in litt. 2005). mon now (Roberts and Baird 1995). It is distributed in several major South-east The population status of the Mekong Giant Asian rivers, chiefly inhabiting mainstream Catfish is unclear. Catches have been channels, where it occurs in deep, slow monitored from the Tonle Sap River dai reaches, with sand or gravel substrates, fishery since 1999, since when between and abundant mollusc populations four and 11 fish have been caught annual- (Roberts and Warren 1994, Rainboth ly (Hogan et al. 2001). As water levels 1996). In the Mekong basin, it performs begin to drop in October and November, spawning and trophic (feeding) migrations, Mekong Giant Catfish move out of the and has been recorded in the Tonle Sap Tonle Sap Lake and into the main Mekong Lake (Poulsen et al. 2004). Known River channel. They inhabit the deep upstream spawning migrations take place pools of the Mekong River during the dry between October and February from season, and probably spawn in the Kompong Cham in central Cambodia to Mekong River either in the Stung Treng Chiang Khong in northern Thailand, where area or in the upper Mekong near Chiang it spawns between late December and Khong (north Thailand). early February. Trophic migrations occur throughout the Mekong, take place mainly Fishing-induced mortality represents the at the onset of the flood season (May- most immediate threat to the Mekong June), and are mainly undertaken by juve- Giant Catfish. A fundamental problem is niles and subadults, which take several that fishermen continue to catch and sell years to mature (Sokheng et al. 1999, the fish despite its conservation status Poulsen et al. 2004). (Sretthachuea 1995), although in Cambodia an ongoing purchase and Jullien’s Golden Carp is undergoing a seri- release initiative was launched in 2000 ous long-term decline and the most obvi- (Hogan et al. 2001). This problem is fur- ous (but not necessarily only) cause is ther complicated by the difficultly of moni- over-fishing with gillnets during the repro- toring commercial and subsistence fish- ductive migrations and spawning periods eries; catches of Mekong Giant Catfish (Roberts and Warren 1994), especially the often go unreported. Other potential November-January spawning migration threats to the species include waterway (Roberts and Baird 1995). modification, clearance of inundated swamp forest and water pollution. Laotian Shad Tenualosa thibaudeaui (IUCN Endangered) Leaping Barb Chela caeruleostigmata This rather small fish species (most large, (IUCN Critically Endangered) spawning fish weigh 400-500g) is endem- Not recorded from the Tonle Sap Lake, ic to the Mekong basin, and its main area but it could occur given its Cambodian of occurrence appears to be from the range, the fact that it appears to make Tonle Sap north. It is migratory, moving migratory movements back into large downstream into Cambodia from the mid- rivers at the end of the flood cycle in dle Mekong in Thailand/Laos in July. It March or April, and its preference for the may follow the turbid floodwaters all the surface waters of large rivers and inundat- way to the Tonle Sap, probably moving ed swamp forests (Fishbase 2005). It is into the Great Lake as it fills with water probably used to make prahoc and is from the Mekong. As water levels in the occasionally seen in the aquarium trade. Great Lake fall, it migrates back down the Tonle Sap to the Mekong (Rainboth 1996). In the mid-1900s it was abundant, one of

38 the principal species caught in the Tonle Warren 1994). Further investigation is Sap fishery (Bardach 1959), but the last needed to find out whether it shares recorded large scale migrations were in spawning grounds, general distribution and 1984. This drastic decline is probably due migration periods with P. jullieni (Roberts to several factors, including dam construc- and Warren 1994). tion and over-fishing in southern Laos (Roberts 1993). Giant Pangasius Pangasius sanitwongsei (IUCN Data Deficient) Tricolor (or Bala) Sharkminnow This large Pangasid is endemic to the Balantiocheilos melanopterus IUCN Lower Mekong and Chao Phraya river Endangered basins, and although listed as common by This rather small Cyprinid inhabits the Rainboth (1996), it is becoming increas- Mekong and Chao Phraya basins, the ingly rare throughout its range (Poulsen et Malay Peninsula, Sumatra and Borneo, but al. 2004). Little information is available on is becoming rare or extinct in many river its ecology and distribution (www.fish- basins, and is commercially important in base.org), but it spawns just prior to the both aquaculture and the aquarium trade rainy season (Rainboth 1996). It has been (Fishbase 2005). It formerly occurred in reported during local knowledge surveys the TSBR (e.g. Rainboth 1996), but is at the southern end of the Tonle Sap Lake now rare and was not taken in fish 1995- (Poulsen et al. 2004), but it is unclear 97 fish catches analysed by Lim et al. whether there are specimens from the (1999); it is depicted on the Department Tonle Sap. of Fisheries Endangered Fishes of Cambodia poster. Additional fish species of conserva- tion concern Asian Bonytongue/Asian Arowana Giant Barb Catlocarpio siamensis Scleropages formosus IUCN Endangered; Catlocarpio siamensis is endemic to the CITES Appendix I middle and lower Mekong River (from This familiar fish is threatened throughout Thailand-Laos to Vietnam), and is consid- its range (Indonesia, Malaysia, Thailand, ered one of the world’s largest cyprinid Cambodia and Vietnam) by demand for fish, measuring up to 3m and weighing up the aquarium trade and to a lesser extent to 200kg, although fish weighing more its meat, which commands a moderate than 50 kg are rarely caught today price (Fishbase 2005). It inhabits tannin (Rainboth 1996, Mattson et al. 2002). stained blackwater streams (Rainboth Adults prefer large, deep pools in the 1996). It is unclear whether it occurs in Mekong for at least part of the year, but the TSBR. juveniles are mostly seen in swamps and small tributaries (Sokheng et al. 1999). It Thicklip Barb Probarbus labeamajor IUCN is a migratory species, but its movements Data Deficient appear to be poorly understood (Mattson At least one other Probarbus species, et al. 2002). Giant Barb is caught inciden- Thicklip Barb Probarbus labeamajor, and tally in the Tonle Sap dai fishery, especial- perhaps the Thinlip Barb P. labeaminor ly dai #2 where the river is deepest and (also IUCN Data Deficient), both of which narrowest and the current is swiftest, are very similar in appearance to P. making escape difficult (Hogan et al. jullieni, has been recorded in the Tonle 2001b). Its numbers have declined drasti- Sap, based on Local Ecological Knowledge cally: in Cambodia, the total catch of (Poulsen et al. 2004). Little is known Giant Barb declined from 200 tonnes in about either due to confusion with P. jul- 1964, to 50 individual fishes in 1980, and lieni, but each apparently migrate together just 10 fishes in 2000 (Mattson et al. with P. jullieni, and the population of P. 2002). Individuals rarely survive to reach labeamajor has been decimated by an reproductive maturity, and its catch needs intensive gillnet fishery which targets the to be strictly regulated by size (Rainboth species in southern Laos (Roberts and 1996). It is an urgent candidate for an 39 IUCN Red List evaluation and immediate Albulichthys albuloides conservation attention (Hogan et al. This relatively small Cyprinid occurs in the 2001b). Mekong, Chao Phraya and Mae Khlong basins and Borneo (Fishbase 2005), and is Puntioplites bulu (no English name) depicted on the Department of Fisheries Endangered Fishes of Cambodia poster. This fish was formerly common, but has Adults are common in the Tonle Sap Lake, recently become very rare (Rainboth where it is of commercial value when sold 1996); it is depicted on the Department of fresh (Fishbase 2005). Fisheries Endangered Fishes of Cambodia poster. It occurs from Indonesia to penin- Elephant-ear Gourami Oxonedus exodon sular Thailand and Cambodia, inhabiting This medium sized (<60 cm) Mekong midwater to bottom depths in large low- Basin endemic inhabits pools of large land rivers and lakes, and moves into inun- rivers and inundated forests during the dated forest habitats when water levels rainy season. It reproduces in non-flowing rise (Fishbase 2005). These habits suggest waters near the river shore during lowest it may occur in the TSBR. water levels (Rainboth 1996) and under- takes lateral migrations into floodplain Sabretoothed Thryssa Lycothrissa croco- areas during the flood season, returning to dilus the Mekong River or other permanent A relatively small (<30 cm) anchovy, water bodies during the dry season, trig- which is distributed from the Lower gered by changes in water levels (Sokheng Mekong Basin and parts of Thailand to et al. 1999). It is unclear whether it western Indonesia and Borneo (Poulsen et occurs in the TSBR, but it is depicted on al. 2004). It is depicted on the the Department of Fisheries Endangered Department of Fisheries Endangered Fishes Fishes of Cambodia poster. of Cambodia poster, but is common from the Mekong Delta north into the TSBR, Botia genus into which it performs migrations in response to wet season floods, and Several species in this genus were record- spawns in the TSBR’s floodplain habitats ed in the Tonle Sap during the first half of (Poulsen et al. 2004). the 20th Century (Botia hymenophysa and B. modesta; Bardach 1959), but there Four-barred Tigerfish Datnioides quadri- have been no recent records (E. Baran fasciatus verbally 2005). Each of these species is This small (<30cm) but strikingly marked therefore of conservation concern. Perciforme is widely distributed from India to Indonesia and New Guinea, which Key areas of conservation impor- occurs in brackish waters and inland lakes. tance for fish It is unclear whether it occurs in the There are three well-known key areas of TSBR, but it is depicted on the conservation importance for fish, each of Department of Fisheries Endangered Fishes which seasonally supports significant con- of Cambodia poster. centrations of fish:

Wallago leeri 1) Prek Toal Core Area (Fishing Lot #2) – the most productive fishery This catfish occurs from Thailand to on the Tonle Sap; Indonesia, inhabiting large rivers and 2) Boeng Chhma Core Area (Fishing entering inundated forest (Roberts 1993, Lots #5, 6 & 7) – supports the Rainboth 1996). It is unclear whether it highest diversity of fish habitats; occurs in the TSBR, but it is depicted on 3) Rivermouths (e.g. Stung Sen river the Department of Fisheries Endangered mouth and the Pi Stuon Fish Fishes of Cambodia poster, and is of com- Sanctuary) – these areas concen mercial importance (Fishbase 2005). trate nutrients, which in turn support rich feeding aggregations of fish. 40 Other important habitats must include per- fish sanctuaries to the existing eight are manent wetlands in the floodplain (e.g. in also planned under Community Fisheries Veal Srangai IBA). These are a high priori- agreements (Cheng Penh verbally 2005). ty for research because there is currently Whatever the approach to revising and no information on them. expanding fish conservation areas, it will be imperative to have a clear understand- Although it is often stated that the inun- ing of the biophysical features that are dated forest is a rich feeding and spawn- important for the selection and designa- ing ground for many species during the tion of new sanctuary areas and the exci- flood season (e.g. Lieng and van Zalinge sion of any inappropriate areas. 2001), no studies have been conducted This understanding does not currently on the importance or ecological role of the exist. floodplain habitats to overall fish species richness and diversity, or indeed to any Conservation of the many migratory fish individual species (E. Baran verbally populations utilising the Tonle Sap is 2005). This is a key area for research. dependent on factors operating elsewhere in the Mekong Basin. Many species only Research into fish abundance and species use the Tonle Sap seasonally (most if not composition in the eight Fish Sancturies all long-distance migratory forms only began in April 2005 (Cheng Phen verbally occur during the wet season). The 2005). Only two of the eight fish sanctu- Mekong mainstream in the Lower Mekong aries are clearly of elevated importance to Basin, anywhere from the Mekong Delta fish in comparison to other (superficially up to the Chinese border is where many very similar) parts of the Tonle Sap. One, species spend the dry season. Deepwater Pi Stuon, at the mouth of the Stung Sen pools in the Mekong are key dry season River, contains a deepwater area (average refuge habitats for some (Poulsen et al. dry season depth is 4-5m), with a rocky 2002), e.g. the Mekong Giant Catfish. bottom (sediments are apparently flushed Migratory fish departing the lake before through by the river current), and incorpo- the flood has receded very far often return rates a narrow belt of inundated forest to the Mekong via a route other than the along the lakeshore. It supports large fish Tonle Sap River. These fish are heavily aggregations. The other, Kompong Phluk, exploited at various locations, including is apparently a slightly deeper water area under the NR-6 bridges in Kompong Cham than most of the lake. The other six fish (E. Baran verbally 2005). sanctuaries are located in the open water column, and none appear to support par- ticular concentrations of fish. The experi- mental protection of two fish sanctuaries (Pi Stuon and Ba Lat) using concrete tubes has two functions, the primary purpose being protection from fishing of the lower water column, the other being creation of shelter for brood stock and spawning habi- tat for some species (Cheng Phen verbally 2005).

The most suitable locations for fish sanc- tuaries in the Tonle Sap Lake, River and adjacent floodplains will be determined, new large scale fish sanctuaries estab- lished by the Department of Fisheries as recommended, and existing fish sanctuar- ies situated in unsuitable locations will be decommissioned (Anon. 2003). Additional

41 The invertebrate fauna of the Tonle Sap is Studies of insects resumed in September little studied and consequently poorly 2005 in and around the Prek Toal Core known, although a preliminary collection Area, with funding and technical assis- made on 27-28 May 2003 in Prek Toal tance from the Royal Belgian Institute of using sweep nets and light traps (Constant Natural Sciences, in collaboration with the and Grootaert 2003), yielded 218 species, Sam Veasna Centre for Conservation in not including Diptera (true flies, gnats, Siem Reap. mosquitoes and crane flies), Lepidoptera (butterflies and moths) and Odonata (drag- Odonata (dragonflies) onflies). These 218 species comprised five Eight dragonfly species were identified dur- Blattoptera (cockroaches), 81 Coleoptera ing a visit to Prek Toal on 7 February (beetles and weevils), 26 Heteroptera (true 2003 (P.K. Batchelor in litt. to Sam bugs, a suborder of Hemiptera), 29 Veasna Centre for Wildlife Conservation, Homoptera (a group including cicadas, 2003): Pseudagrion rubriceps; plant hoppers, treehoppers, spittlebugs, Brachythemis contaminata; Crocoothemis aphids, mealybugs, whiteflies and scale servilia; Orthetrum safina; Rhyothemis insects), 46 Hymenoptera (ants, bees, phyllis; Rhyothemis variegata; Tholymis sawflies, and wasps) and 31 Orthoptera tillarga and Trithemis pallidinervis. (crickets, grasshoppers, katydids, mantids, and walking sticks). Apis (honeybees) Three of the four species of honeybee At least two species of Diptera, of the known from Cambodia (Apis dorsata, A. family Dolichopodidae, collected in Prek cerana, A. florae and A. andreniformis) Toal during 2003 were previously unde- were identified during interviews with Prek scribed (Constant and Grootaert 2003). Toal villagers in 2003 to occur, or to have Constant and Grootaert (2003) predict occurred, in the Tonle Sap floodplain that a low invertebrate species richness is (Petersen 2003). Apis dorsata and Apis expected from the Prek Toal area, com- florea reportedly still occur in the area. prising chiefly aquatic and semi-aquatic One person reported that Apis cerana species. Nothing is known of how these existed in the area prior to 1995, but for- insects spend the flood season, whether it est fires and wood cutting activities since is as larvae, pupae, or adults in the emer- then had destroyed most suitable nesting gent trees, or whether perhaps the flood- sites, large cavities (minimum volume 15- plain is colonised annually from outside 20 liters) in trees (Petersen 2003). Apis the inundation zone. andreniformis was not recognized by the Insects must form a key component of the respondents and may not exist in the ecosystem food chain’s lower trophic lev- TSBR. els, and may be a particularly important resource during drought years when fish A. dorsata appears to be a year-round resi- becomes scarce (MRC 2003). Two distinc- dent of swamp forest and short-tree tive aquatic insects that are common to shrubland habitats. It nests in the open in abundant in the ricefield ecosystem of large, prominent colonies that are easy to Kompong Thom, a giant water bug collect. All respondents reported that over- Lethocerus sp. and a water beetle proba- harvesting, forest fires and cutting for tim- bly from either the Dytiscidae or the ber and firewood had contributed to a Hydrophilidae (Balzer et al. 2002), form steep decline of A. dorsata populations important food items in the local human since 1995 (Petersen 2003). A. florea diet. 42 prefers low, dense shrub for nesting, and Sap channel and the floodplains”, and makes small nests that yield low returns made up 85% of the zoobenthos by per unit harvesting effort; they do not weight (Nguyen and Nguyen 1991 cited in appear to be at risk in the TSBR (Petersen Lamberts 2001). The remainder of 57 2003). identified zoobenthic species included 15 insects, eight oligochaete worms and five Crustacea (crabs and shrimps) crustaceans (Nguyen and Nguyen 1991 At least one crab, and one shrimp species cited in Lamberts 2001). One species of occurs in ricefield habitats in Kompong ampullarid snail, probably in the genus Thom (Balzer et al. 2002). Five morpho- Pila, was found in the ricefield ecosystem logically distinct forms of crab are distin- in Kompong Thom (Balzer et al. 2002). guishable in these ricefield agro-ecosys- Other, unidentified snails and snail shells tems (all but one of which is abundant), were collected from the rivers and the indicating as many as five species may lakes nearby. occur; van Amerongen (1999 cited in Balzer et al. 2002) suggests that all rice- ZOOPLANKTON field crabs in the Tonle Sap belong to the The zooplankton of the Tonle Sap ecosys- genus Somanniathelphusa. tem was partly described from work con- Somanniathelphusa crabs are generally ducted in the late 1980s (Nguyen and considered rice pests (Balzer et al. 2002), Nguyen 1991 cited in Lamberts 2001). Of and as such are collected from June to the 46 species identified, half are rotifers, December, but only eaten in times of food about one third are cladocerans and the scarcity. remaining seven species are copepods. As The shrimp species, identified as with phytoplankton, the number of species Macrobrachium lanchesteri (Mogensen in in the flooded areas is higher than in the Balzer et al. 2002), is abundant between river, the composition of the zooplankton September and December, when it is har- communities varies throughout the year, vested for food. McDonald et al. (1997) and zooplankton density (in the Tonle Sap mention that local informants report two channel) increases by a factor greater than species of shrimp, which are commercially 100 during the dry season. In the flood exploited from the deeper waters of the season, zooplankton density in the flooded lake itself. areas is almost one hundred times higher than in the river (Nguyen and Nguyen Mollusca (snails and mussels) 1991 cited in Lamberts 2001). The Mekong Basin, including in the Tonle Sap, exhibits an extremely high diversity of molluscs (snails and mussels) (Rainboth 1996). For example, the stenothyrid and pomatiopsid snails (gastropods) include over 120 species, at least 111 of which are endemic to the river (Davis 1979 cited in MRC 2003), the greatest known biodi- versity of snails in a freshwater system in the world (MRC 2003). Twenty-nine species of mollusc (comprising over 50% of all species present) were identified in the zoobenthos of “the Mekong, the Tonle

43 With approximately 470,000 people living • All major migratory routes used by within the inundation zone and over 1 mil- fish to depart the Tonle Sap Lake and lion in all parts of the floodplain including floodplain (the Tonle Sap River, and, adjacent urban centres, coupled with the before the flood has receded very far, highest incidence of poverty in Cambodia flooded ‘overland’ routes, e.g. under (Keskinen 2003), pressure on natural the NR-6 bridges in Kompong Cham resources is high. Access to natural (E. Baran verbally 2005) are heavily resources is very unequal, land tenure is over-fished. Lamberts (2001) notes insufficient and many areas are effectively concern within the Department of common land, which is driving increasing Fisheries over the collection of large competition for resources, and with it a quantities of fry in the Mekong, the cycle of overexploitation and degradation. possible impact of heavy and noisy boat traffic along migration paths, and Over-exploitation of animals the use of illegal fishing methods (e.g. • Populations of some species have brush parks) along migration corridors. indisputably declined to the verge of extirpation from the TSBR as a result • The recent watersnake harvest, proba- of chronic over-harvesting (e.g. bly the largest exploitation of a snake Siamese Crocodile, Platt et al. 2004). assemblage anywhere in the world, Similar declines are suspected for may well be unsustainable, and other reptiles, especially turtle species declines in several species (particularly (e.g. Holloway et al. 2000). Hunting of Enhydris bocourti) are reported by Irrawaddy Dolphin has probably driven local fishermen (Stuart et al. 2000, the species to extinction in the TSBR Brooks et al. 2005). (Baird and Beasley 2005). • Unsustainable harvesting of eggs and • Over-fishing through inappropriate chicks from breeding colonies was the practices (e.g. use of mosquito net- primary threat to large waterbirds until ting, Troeng 2000) is a major concern very recently (Parr et al. 1996, Ear- for some fish, particularly the larger Dupuy et al. 1997, Goes and Hong and more valuable species, e.g. 2002). This threat has been minimised Mekong Giant Catfish, Giant, Thicklip within the Prek Toal Core Area since and Thinlip Barbs and Jullien’s Golden permanent ranger presence at popula- (or Seven-line) Barb. Suspending fish- tion monitoring platforms through the ing operations at dai #2 in the Tonle breeding season (Goes 2005), but Sap River would decrease the chance remains a problem in colonies else- of capture of the large endangered where that are not currently monitored fishes (Hogan et al. 2001). or afforded protection, e.g. Beong Identifying the source area(s) where Chhma-Moat Khla and Dei Roneath over harvesting occurs is not always (Seng et al. 2003). straightfor ward, however, and for some or many species, it may be • Bengal Florican was heavily hunted occurring in areas outside the TSBR prior to conservation in parts of (e.g. Laotian Shad; Roberts 1993). Kompong Thom province (Goes et al. 2001), and is probably still threatened by hunting away from conservation areas (Seng et al. 2003).

44 Habitat Loss and Degradation cambodiana is the preferred tree species for construction (Gum 1998, Rundel Inundated Forest-scrub 2000). Significant areas of inundated forest-scrub, a critical fish habitat (e.g. Lieng and van Land-use conversion Zalinge 2001), have been destroyed through clearance and conversion to rice cultivation, Grasslands and agro-ecosystems burning, harvesting for fuelwood and These multiple-use landscapes have poorly construction materials and collection for developed regulatory and protection mech- firing brick kilns (Giesen 1998, Troeung 2001, anisms, hence are vulnerable to rapid Baran 2004). changes in agricultural practice, either through intensification or abandonment. • In the 1930’s, an estimated 1 million Episodic cultivation of deepwater rice has ha of natural vegetation around the created a mosaic landscape of great biodi- Tonle Sap declined to approximately versity importance in parts of the flood- 564,000 ha in the 1980’s plain, and has probably been instrumental (Woodsworth 1995). in the creation and maintenance of grass- lands in the floodplain (Davidson 2004, • By the late 1990’s, approximately Evans et al. 2005). Abandonment of deep- 361,000 ha of natural forest and water rice cultivation and reversion to tall 157,000 ha of degraded forest were scrub due to low economic returns from estimated extant (Giesen 1998). deepwater rice harvests represents a sub- stantial concern. However, agricultural • In 2002, the floodplain supported intensification poses a more urgent threat. approximately 22,000 ha of forest, Intensification is a major cause of biodiver- 448,000 ha of shrub and almost sity loss in Thailand and Vietnam (Tran 730,000 ha of grassland (Keskinen Triet et al. 2000; Round 2002; Buckton & and Huon 2002, cited in Baran 2004). Safford 2004), particularly conversion to irrigated rice production with associated It is very likely that each of these estimates high levels of chemical inputs and the loss was based on a different dataset and different of dry season stubble and fallow habitats. interpretation processes, but the overall Rapid expansion of dry-season irrigated impression is clear: the area of forested rice now presents a serious and immediate habitat in the TSBR has declined over the threat to landscapes of the outer Tonle past five decades. Forest cover at a local Sap floodplain. In 2004–5, at least 15 scale has been monitored more consistently: dams were constructed to irrigate over 60 close to 30% of the inundated forest in km2 for intensive rice production within Battambang province was cleared between and surrounding two sites in Kompong 1965 and 1992, including 10,688 ha in Thom covering approximately 200km2 Fishing Lot no.2 (the Prek Toal Core Area) (Evans et al. 2005). This is expected to (Troeung 2001). Firewood collectors are greatly reduce habitat suitability for grass- relatively unselective, but Hymenocardia land dependent species due to reduced wallichii, Lumnitzera aff. racemosa, structural diversity, high pesticide inputs Terminalia cambodiana, Barringtonia acu- and greatly reduced opportunities for non- tangula, Coccocera anisopodum and Vitex crop species to survive, as well as reduce holodenon are preferred, and Diospyros open access resource availability (e.g. 45 grazing, fishing) to local communities Urban and domestic (Evans et al. 2005). The potential impact of untreated urban and domestic solid and liquid waste, Only 5% of the grasslands inside the which is discharged directly into TSBR country’s Important Bird Areas are repre- waterbodies (especially from floating vil- sented within Cambodia’s protected-areas lages), is large, particularly if fish migra- network (Seng Kim Hout et al. 2003). tion routes become polluted (Lamberts Development of Cambodia’s agricultural 2001). Impacts on water quality are most sector is a key objective of the govern- severe during the dry season when shal- ment’s poverty reduction strategy, of low waters quickly become stagnant and which irrigation projects form an important temperatures can be high. component. Urgent priority should there- fore be given to mitigating agricultural Petro-chemical and Noise intensification, through research into agro- Oils spills pose a threat to waterways and ecosystem biodiversity, the incorporation associated habitats around ports (e.g. of grasslands and agro-ecosystems within Chhong Kneas) and waste oil, sludge and biosphere core areas and/or the integration bilge are often discarded into the water in of conservation priorities into land-use these locations. There is concern about management strategies for multiple-use the impact of persistent noise pollution zones supporting grasslands and agro- from motorised boat traffic along the main ecosystems (Davidson et al. in prep.). Tonle Sap channel on migration of fish (e.g. Lamberts 2001). There have also Pollution been fears that oil exploration may begin on the Tonle Sap (Goes 2002); it is not Agro-chemical clear whether these remain. Increasing use of agro-chemicals, especially pesticides, has been highlighted as a major Mekong Basin-wide threats concern in the TSBR and Cambodia as a The predicted effects of damming of the whole (Yang et al. 2001, EJF 2002, upper Mekong River in China include CEDAC 2004). Approximately one third reductions in sediment deposition, soil fer- (ca.80) of 241 pesticides on the market in tility and biological productivity within the 2000 were classified by the World Health Tonle Sap floodplain, and alterations to Organisation as hazardous or very haz- the chronology of the wet-season flood ardous (Yang et al. 2001). These include regime (Kummu et al. 2005, in press a & endosulfan, monocrotophos, DDT, brodifa- b). Any combination of these could have coum, cypermethrin, diazonin, carbofuran serious long-term consequences, and alter and phosphamidon, all of which are the current seasonal cycle of processes known to have caused mass wildlife mor- that maintains the ecosystem’s exception- talities in other countries (EJF 2002). It is al productivity and its key populations of suggested that the use of these agro- animals and plants. chemicals will have a relatively limited effect on overall fisheries productivity due to the size and hydrology of the ecosys- tem (Lamberts 2001), but impacts to bio- civersity and human populations at the local level can be considerable (EJF 2002, CEDAC 2004). 46 Introduction of exotic (non-native) species spp., which specialize in feeding on can cause irreversible alterations to the already moribund plants. Confirmed evi- ecology of aquatic and terrestrial environ- dence of the occurrence of Pomacea gigas ments, through competition, predation or in the TSBR could not be traced, but given transmission of diseases, and is a key how widespread it is in Thailand, there is threat theme that requires addressing in a high possibility in could enter the Tonle the TSBR. Sap ecosystem if it hasn’t already. It is a favoured prey of Asian Openbill, the popu- Fish lation of which has increased rapidly in About 17 exotic fish species have estab- Thailand’s central plains potentially as a lished wild populations in the Lower result of the abundance of P. gigas (Round Mekong Basin (Welcomme and 2002). Toxic agro-chemicals are used to Vidthayanon 2003), most of which origi- eradicate the snail, which can then accu- nated from aquaculture. No specific infor- mulate in species higher up the food mation could be traced on their effects on chain. The case of the apple snail is an the fish fauna of the Tonle Sap. example of the disastrous consequences Cambodia’s DoF recently prohibited aqua- that can follow from an inappropriate culture of one exotic species, the red-bel- introduction (Welcomme and Vidayathon lied pacu Piaractus brachypomus, a mem- 2003). ber of the Piranha family from South America (Hortle et al. 2004). Plants A variety of exotic plant species have Invertebrates become established in the TSBR. Two The inappropriate introduction of European major invasives present the greatest honeybees (Apis mellifera) poses a serious known problems: Water Hyacinth threat through disease introduction, chemi- Eichornia crassipes and Giant Mimosa cal pesticide dependence (mites and foul- Mimosa pigra. brood control) and disruption of ecosys- tem processes. Introduction has occurred Water hyacinth Eichornia crassipes, a near Siem Reap but is being contained (D. native of South America, reported present Jump in litt. October 2005). Further intro- since the 1980s at least (Hellsten et al. ductions of A mellifera as part of bee- 2003), appears to be increasing greatly in keeping schemes should be vigorously dis- abundance along the shore of Tonle Sap, couraged by donors and other agencies as it has in many tropical river systems in (Petersen 2003). Southeast Asia (McDonald et al. 1997). Water hyacinth is most apparent along Pomacea gigas (Apple Snail), which feeds waterways with slow moving currents and on the growing basal stems of aquatic in pools and ditches. While this species plants, has had a major impact on aquatic has some economic value as a pig fodder habitats and agro-ecosystems in Thailand, and in fish baits, and has some local uses, degrading natural vegetation and severely e.g. edible flowers, fertilizer and as rope reducing rice yields (Round 2002, (McDonald and Veasna 1996), its negative Welcomme and Vidayathon 2003). qualities greatly outweigh any economic Affected habitats become denuded of their value. Water hyacinth has a remarkably fish populations, and the introduced snails high rate of growth, and quickly forms compete with native species such as Pila dense mats of vegetation that deoxy-

47 genate water. It competes strongly with native aquatic species and impacts on fish production through lowered water quality (McDonald et al. 1997).

Several alien woody species occur in the floodplain, some of which are greatly expanding their cover and dominance. The Giant Mimosa Mimosa pigra, also a native of South America, reported present since at least 1990 (Hellsten et al. 2003), cur- rently presents the most serious problems. Giant Mimosa is already a serious pest in other parts of Asia and northern Australia (e.g. Jantunen 2003). This aggressive plant rapidly invades fallow fields and heavily disturbed shrubland and swamp forest habitats after clearance or burning. Once established, it forms dense impene- trable thickets of spiny growth that choke out other native species, provides very lit- tle value as wildlife habitat, and is detri- mental to fisheries (Goes 2005). The largest patches occur in abandoned fields in the south-western part of the TSBR, especially in Pursat province. There are also substantial thickets in the delta area and around Siem Reap (Hellsten et al. 2003). Two other alien Mimosa species, M. invisa and M. pudica, are also estab- lished around Tonle Sap but have been less aggressive (Rundel 2000 citing McDonald et al. 1997).

Two grass species have also widely invad- ed wetland margins of Tonle Sap. Brachiaria mutica and Echinochloa stagni- na were introduced at some time in the past from Africa (Rundel 2000) as high quality species for grazing, and have become dominant species in many areas of the lake. Their ecological effects in the TSBR are undocumented.

48 The TSBR is one of the most productive Long-tailed Macaque freshwater ecosystems in the world. It (widespread and locally common, probably supports unique floral and faunal assem- one of the larger populations in South-east blages characterised by a high relative Asia) abundance and adaptations to seasonally Germain’s Silver Leaf Monkey exploit specific components of an ecosys- (poorly known but certainly important tem that experiences extreme stresses globally) during the course of its annual cycle of flooding and drought-like conditions. It is Smooth and Hairy-nosed Otters one of the most important wetlands for (both poorly known but potentially impor- biodiversity conservation in Southeast tant in South-east Asia) Asia, supporting numerous narrowly or regionally endemic plant species, and an exceptional number of internationally Fishing Cat important populations of globally threat- (very poorly known but potentially impor- ened, Near Threatened and Data Deficient tant in South-east Asia) animal species. The key conservation research priorities Biodiversity within the TSBR is not exten- for mammals are: sively documented and is rather poorly understood. A great deal remains to be ·Further surveys to assess the status, learned about the status, distribution and distribution and conservation signifi- ecology of most individual species, ecolog- cance of the Germain’s Silver Leaf ical relationships within and between indi- Monkey population vidual species and whole assemblages, ·Further surveys to assess the status, and patterns of seasonal migrations, distribution and conservation signifi- changes in abundance and habitat use. cance of Smooth and Hairy-nosed The following breakdown by taxonomic Otter populations group attempts to highlight both a) the ·Further surveys to assess the status, key known facts about the TSBR fauna distribution and conservation signifi- and flora, and b) species or taxonomic cance of the Fishing Cat population groups that are potential conservation pri- ·Any significant populations of these orities but for which there is insufficient species warrants follow-up conserva- information. tion measures

MAMMALS BIRDS

Relatively few mammal species occur in Birds have been more extensively studied the TSBR for a tropical lowland ecosys- than any other group, 210 species have tem. These include six IUCN Red Listed been recorded, and globally and/or Asia- species, each of which inhabits the sea- regionally significant populations of over sonally inundated swamp forests and 25 species occur, 17 of which are IUCN short-tree shrublands. Populations that are Red-Listed. These most significant habi- potentially of global or regional conserva- tats, species and populations are: tion significance are:

49 • Globally significant colonies of Greater Kompong Chhnang provinces and Lesser Adjutants, Spot-billed (Davidson 2004), coupled with contin- Pelican, Painted and Milky Storks, uing detailed studies of florican ecolo- Black-headed Ibis, Darter and Indian gy and behaviour to better understand Cormorant that breed in seasonally its conservation requirements; inundated swamp forest in the Prek • Conduct more widespread wet-season Toal Core Area; surveys in seasonally inundated • Probably one of the largest known swamp forest to assess status and breeding populations of Grey-headed distribution and conservation signifi- cance of breeding Masked Finfoot Fish Eagle; populations; • A unique bird community in perhaps • Conduct wide-ranging dry season sur- the largest remnant tract of seasonally veys of seasonally inundated swamp inundated grasslands and agro-ecosys- forest to assess status and distribution tems in South-east Asia, including dur- and conservation significance of ing the dry season, probably the breeding Grey-headed Fish Eagle popu- world’s largest breeding population of lations; Bengal Florican, the largest known • Conduct mist-netting surveys in grass- wintering population of Manchurian land, short-tree shrubland and wetland Reed Warbler, very small numbers of habitats to assess distribution, habitat White-shouldered Ibis and Asia-region- use and population size of wintering ally important populations of Greater Manchurian Reed Warbler in dry sea- Spotted and Imperial Eagles, and in son; the wet season post-breeding aggrega- • Survey Aquila eagle roosts in outer tions of Greater and Lesser Adjutants floodplain grassland and short-tree and Black-headed Ibis. shrubland habitats during the dry sea- son to assess the size of the wintering The key conservation research priori- population. ties for birds are:

• Further develop monitoring systems of large waterbird populations to improve understanding of precise numbers, dis- persal and interchange with other pop- ulations in Cambodia to facilitate con- servation planning; • Further surveys of potentially suitable seasonally inundated swamp forest habitat for breeding large waterbird colonies; • An assessment of grassland extent and quality within the TSBR, and the status and distribution of Bengal Florican within remaining grasslands, in western Siem Reap, Banteay Meanchey, Battambang, Pursat, south- ern Kompong Thom and north-eastern

50 At least eight, perhaps ten globally threat- • Conduct amphibian surveys to estab- ened and Near Threatened reptile species lish baseline data on species diversity, occur in the TSBR, but all species are distribution and abundance poorly known and difficult to survey in the wild. FISH • Siamese Crocodiles probably still Fish catch data spans a longer period than occur in several localities but in very for any other taxonomic group, but despite small numbers, at least some popula- the enormous economic importanc e of the tions may not be viable Tonle Sap fishery to Cambodia as a whole, • The largest known harvest of a snake fish ecology in the TSBR is remarkably assemblage in the world (chiefly of six poorly understood. Many fish species that species of watersnake) occurs in the occur in the TSBR are endemic to the seasonally inundated forests and Mekong Basin itself, or the Mekong Basin short-tree shrublands during the wet and one or two other South-east Asian season and into the early dry season catchments. • An exceptional freshwater turtle assemblage has been reduced to small • At least five globally threatened fish but nonetheless globally or regionally occur in the Tonle Sap, including significant populations of at least six Mekong Giant Catfish, for which globally threatened species, the most Fishing Lot #2 may seasonally be of signficant being Yellow-headed great global conservation importance Temple Turtle and perhaps Asian • Several other species are of conserva- Giant Softshell Turtle, with key popu- tion concern based on apparent sharp lations of Black Marsh Turtle, Asian declines Box Turtle, Malayan Snail-eating Turtle • Most fish species migrate in response and Asiatic Softshell Turtle to the highly seasonal fluctuations in • Large populations of Burmese and hyrdrology, but these movements are Reticulated Pythons have been report- poorly understood beyond some gen- ed from Boeng Chhma Core Area and eral basic principles adjacent Moat Khla, and Stung Sen • Seasonally inundated swamp forest Core Area and short-tree shrubland habitats pro- • The amphibian fauna is undocumented vide key nesting, spawning and rear- ing habitats for many species, but the Key conservation research priorities for relative importance of specific habitats herpetofauna are: to specific species and basic under- standing of ecological requirements • Investigate potential to re-introduce and precise movements for almost all wild Siamese Crocodiles to secure species is lacking sites • General trends apparent from fish • Monitor watersnake harvests and con- catch data are that there have been duct field studies to assess sustain- sharp declines in the catch of large, ability and conservation requirements rare fish species, the proportion of • Manage key sites (e.g. Prek Toal and small fish caught is increasing as aver- Boeng Chhma Core Areas) to minimise age fish size decreases, and mean or remove exploitation and monitor catch per capita and unit effort is recovery of crocodile, turtle and snake decreasing populations 51 • Only two of the eight designated Fish and perhaps genera to science. Clearly, Sanctuaries appear to support signifi- further collections ar required in all major cant concentrations of fish habitat types to establish a baseline of data. Too little is currently known to Key conservation research priorities for assess the conservation status of individ- fish are: ual species or taxonomic groups.

• Conduct detailed ecological studies of the globally threatened species and FLORA other species of conservation concern Knowledge of the TSBR flora is based on to assess their status and establish historical studies and one month-long sur- sustainable fishing practices to allow vey in 1997. From these, we know that: populations to recover, including a dedicated study of the Mekong Giant • The TSBR supports the largest remant Catfish population in Fishing Lot #2 tract of a unique seasonally inundated (Prek Toal Core Area) freshwater swamp forest formation, • Conduct detailed studies of the sea- which although relatively species poor sonal use by fish communities of per- is highly distinctive and rich in both manent wetlands in the middle and regional and narrow endemics. outer floodplain, on which there is cur- • This swamp or gallery forest has an rently no information, to assess their understorey of lianas, made up of a conservation importance to “black group of species that form their own fish” distinctive formation – short-tree • Develop and implement a standardised shrublands – in the absence of the monitoring programme to annually larger species in the middle floodplain. assess the status of fish populations • Exceptional examples of these occur in the main fisheries in the TSBR in Prek Toal, Boeng Chhma (and adja- • Pursue international collaboration, data cent Moat Khla) and Stung Sen Core sharing and fisheries management Areas development for key populations of • The TSBR supports what may be the long distance migratory “white fish” largest remaining area of seasonally utilising the Tonle Sap seasonally, but inundated grasslands (of diverse struc- which are affected by factors operat- ture and community composition) in ing elsewhere in the Mekong Basin, South-east Asia particularly the Mekong mainstream between the Mekong Delta and the Key conservation research priorities for Chinese border, where many species flora are: spend the dry season. • Further surveys to assess distribution and conservation status of regional INVERTEBRATES and narrow endemic species, and The invertebrate fauna is only now begin- identify important examples of each ning to be studied. The first collection, for conservation action made over two days in late May 2003 at • Detailed study of grassland species to Prek Toal, has so far yielded 218 identi- assess grassland community unique- fied species (excluding one large order and ness and their regional conservation two smaller orders), including new species significance

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62 ANNEX 1. PLANT SPECIES RECORDED IN THE TSBR Dominant or locally common species in each broad vegetation formation are indicated with an * Family Scientific Name Seasonally inundated Aquatic Grasslands Outer Source swamp forest, tropical herbaceous and agro- floodplai (s) evergreen forest and communities ecosystems n pools short-tree shrublands and ricefields 1 Alangiaceae Alangium ridleyi King * 1,?4 Alismataceae Ott elia alismoides Pers. 1 Sag ittaria sag ittifolia Lour 1 Amaranthaceae Ac hyran thes aqua tica R.Br. *1 Alternan thera sessilis (L.) DC . * 1,2 Cen trostac hys aqua tica Wall. 1 Annonaceae Gagnep. * 1,2 Popow ia diospyrifolia Pierre * 1,2 Uncaria homomalla * 4 Uvar ia pierrie Fine t & G agnep. * 1,4 Apocynaceae Paramer ia glan dulifera Ben th. * 1,4 Theve tia nere ifolia Juss. ex Steu dl 1 Araceae Pistia stratiotes L. * * 1,2 Asclepiadaceae Stenocau lon kleinii W. & A rn. * 1,2 unidentified sp. 1 Asteraceae Ec lipta alba (L.) Hassk. * 1,2 Grangea ma deraspa tana (L.) Poir. * * 1,2 Balsaminaceae Impa tiens sp. * 1,2 Barraginiceae Cordia sp. *4 Bombaceae Ceiba pen tan dra Gaer tn. 1 Boraginaceae Colden ia procum bens L. * 1,2 Heliotrop ium indicum L. * * 1,2 Caesalpiniaceae Crudia chrysantha * 1,4 Cynome tra c f. dongna iensis *4 Cynome tra c f. inaequ ifolia *4 Cynometra ramiflora * 4 Cynome tra sp. 1 dasyrrac his *1 Sindora coc hinc hinensis Lam. 1 Tamar indus indicus L. 1 Campanulaceae Sphenoc lea zeylan ica Gaer tn. 1 Capparidaceae Cappar is m icran tha DC. 1 Crataeva nurva la Ham. * 1,4 Crataeva roxburg hii R.Br. *1 Maerua decan dra Gagnep. *1 Celastraceae Lop hope talum wightianum Arn. 1 Lop hope talum fimbriatum *4 Combretaceae Calycop teris floribunda Lam. * 1,4 Com bretum dec iduum Coll. & Hemsl. *2 Com bretum trifoliatum Ven t. * 1,2,4 Com bretum qua drangu lare Kurz. *1 Lumn itzera aff. racemosa Willd. * 1,2 Quisqua lis indica L. *1 2 Quisqua lis densiflora *4 Term ina lia cam bodiana Gagnep. * 1,2,4 Commelinaceae Comme lina salic ifolia Roxb. *1 Connaraceae Connarus semidecan drus Jac k 1 Convolvulaceae Ipomoea aqua tica Forsk. * * 1,4 Ipomoea c hrysioides Ker.-Gaw l. * * 1,4 Ipomoea obscura Ker.-Gaw l. *1 Merrem ia hederaceae (Burm.f.) Hall.f. 1 Cucurbitaceae Tric hosan thes sp. * 1,2 63 Family Scientific Name Seasonally inundated Aquatic Grasslands Outer Source swamp forest, tropical herbaceous and agro- floodplai (s) evergreen forest and communities ecosystems n pools short-tree shrublands and ricefields

Cyperaceae Ac tinosc irpus grossus (L.f.) Goe tgh. & D.A. Simpson * * 1,2,3 Cyperus digitatus Roxb. 3 Cyperus iria L. 3 Cyperus aff. pilosus Vahl * * 1,2 Fimbristylis dic hotoma (L.) Vahl 3 Fimbristylis dura (Zoll. ex Maritz.) Merr. 3 Fimbristylis um bellaris (Lam. ) Vahl 3 Rhync hospora submarg ina ta Kuk. 3 Rhync hospora sp. * 1,2,3 Sc leria sp. 1 Dilleniaceae Tetracera sarmen tosa Vahl *1 Droceraceae Drocera burmann ii Vahl 1 Drocera indica L. 1 Ebenaceae Diospyros bejau dii Lecom te *4 Diospyros cam bodiana H. Lec. *1 Diospyros sylva tica Roxb. 1,4 Diospyros sp. *4 Elaeocarpaceae Elaeocarpus griffithii Mast. * 1,4 Elaeocarpus lacunosus Wall. *1 Elaeocarpus ma dope talus Pierre * 1,4 Elaeocarpus sp. *1 Euphorbiaceae Antid esma ghaesem billa Gaer tn. * 1,4 Breyn ia rhamno ides Arg. * 1,4 Bridelia cam bodiana Gagnep. * * 1,2 Coccoceras an isopo dum Gagnep. * 1,4 Croton cau datus Gagnep. * 1,4 Croton krabas Gagnep. * * 1,2,4 Croton me kongens is Gagnep. * 1,2 Homono ia ripar ia Lour. * 1,4 Hymenocar dia wallic hii Tul. * 1,2,4 Jatrop ha curcas L. 1 Mallotus coc hic hinensis Lour. * 1,4 Melanolepis vitifolia Gagnep. 1 Phyllan thus julien ii Beille *1 Phyllan thus reticu latus Poir. *1 Phyllan thus taxo diifolius Beille * 1,4 Phyllan thus sp. * * 1,2,4 Flacourtiaceae Homa lium brevidens Gagnep. * 1,4 Homalium dasyanthum 1 Homa lium griffithianum *4 Hydnocarpus an thelmintica Pierre * 1,4 Hydnocarpus saigonens is Pierre * 1,4 Guttiferae Calop hyllum sp. 1 Cratoxylon prun ifera Dyer. 1 Garc inia loure iri Pierre * 1,4 Hippocrataceae Salac ia verrucosa Wight 1 Hydrocharitaceae Elodea cana dense Mic hx. *2 Hydroc haris dubia (Bl.) Blac k. 1 Lauraceae Cryp tocarya oblong ifolia Bl. * 1,4 Lecythidaceae Barring ton ia acu tangu la (L) Gaer tn. * * 1,2,4 Barringtonia micrantha * 4 Careya arborea * 4

64 Family Scientific Name Seasonally inundated Aquatic Grasslands Outer Source swamp forest, tropical herbaceous and agro- floodplai (s) evergreen forest and communities ecosystems n pools short-tree shrublands and ricefields

Careya sp. 1 Lemnaceae Lemna minor L. * 1,2 Lentibulariaceae Utricu laria aurea Lour. * * 1,2 Lythraceae Lagers troem ia sp. * 1,4 Ammann ia bacc ifera Pollini 1 Malphigiaceae Hiptage triacantha * 1,4 Marsiliaceae Marsilea qua drifolia L. 1 Marantaceae Schummanianthus dichotomus * 1 Melicaceae ? Dysoxylum procerum 1 Menyanthaceae Nymp hiodes indica (Thwa ites) Kuntze 1 Mimosaceae Acacia spiralis * 4 Acac ia thailan dica Nielson * * 1,2 Albi zia lebekkoides (A.P. DC.) Ben th. * 1,4 Entero lobium saman (Jacq ) Prain. 1 4 Mimosa invisa 5 Mimosa pigra L. 1 4 Mimosa pu dica 5 Nep tunia oleracea Lour. 1 Moraceae Capparis microcantha * 4 Cudran ia cam bodiana Gagnep. * 1,4 Ficus heterop hylla L.f. * 1,4 Ficus sp. *1 Myrtaceae Syzygium c inereum Wall. *1 Syzygium sterrophyllum * 1 Nelumbonaceae Nelum bo nuc ifera (L) Gaer tn. * 1,2 Nymphaeaceae Nymp haea nouc halii Burm.f. 1 Onagraceae Jussiaea linifolia Vahl *2 Ludwigia adscen dens (L.) Hara. * * 1,2 Ludwigia hyssopifolia (G. Don ) Exell * * 1,2 Palmeae Calamus godefroyi * 4 Calamus pa lustris Griff. *1 Calamus salic ifolius Becc. *1 Papilionaceae Aeschynomene indica L. * * 1,2,3 Butea frondosa Roxb.(monosperma (Lam ) Taub er t) 1,4 Dalberg ia entadoides Pierre * 1,2,4 Dalberg ia pinna ta Pierre * 1,2 Den drolobium lanceo latum (Dunn ) Sc hindler 1 Derris lao tica Gagnep. * * 1,2 Sesban ia paludosa Prain (Sesban ia javan ic * * 1,4 Poaceae Arundine lla setosa Trin. 3 Bam busa arun dinacea Ait. * 1,4 Brac hiaria mutica Stap f. *1 Ch amaerap his m inuta Mez. 1 Ch rysopogon festuco ides (J. Presl) Veldk. 3 Cym bopogon cam bog iensis (Balansa ) E..G & A Camus 3 Desmostac hya bipinna ta (L.) Stap f. 3 Ec hinoc hloa stagn ina (Retz.) Beauv. * 1,2 Eragros tis a trovirens (Desf.) Trin. 3 Eragros tis brown ii (Kunth) N ees. 3 Impera ta cylindrica (L.) P. Beauv. 3 Leersia hexan dra Sw. 1,3 Lep toc hloa sinensis Nees. 1

65 Family Scientific Name Seasonally inundated Aquatic Grasslands Outer Source swamp forest, tropical herbaceous and agro- floodplai (s) evergreen forest and communities ecosystems n pools short-tree shrublands and ricefields

Oryza rufipogon Griff. 3 Oryza sativa L. * 1,2 Pan icum luzonense J. Presl. 3 Pan icum repens L. 3 Paspa lum scrobicu latum L. 1,3 Phragm ites karka Trin. * * 1,3 Pseudovossia cam bogens is (Bal.) Camus. * 1,2 Sacc harum spon taneum L. * 1,4 Sacc iolep is interrup ta A. Ch ase 1 Sacc iolep is mysuro ides A. Ch ase 1 Sc lerostac hya fusca (Roxb.) A. Camus. 1,3 Setaria pallida-fusca Stap f. * 1,2 Sorg hum nitid um (Vahl) Pers. 3 Theme da arun dinacea (Roxb.) Ridl. 3 Vossia cusp idata Griff. 1 Podostemonaceae Dalzellia carinata 1 Polygonaceae Polygonum barbatum L. *1 Polygonum tomen tosum Willd. **2 Pontederiaceae Eic hhornia crassipes (Martius) Solms * * 1,2 Monoc horia hastata Solms 1 Monoc horia vag ina lis Presl. 1 Potamogetonacea Potamoge ton sp. 1 Rhamnaceae Zizyphus jujube 1 Zizyphus sp. 1 Rubiaceae (Naucl a Antocep halus indicus A. Ric h. 1 Garden ia cam bodiana Pitard. *1 Morinda persicae folia Ham. * * 1,2,4 Mitragyna c f. diversifolia *4 Ran dia long ifolia Ben th. * 1,2 Ran dia sp. *1 Step hegyne c f. diversifolia *4 Step hegyne parv iflora Korth. * 1,4 Uncar ia sp. 1 Rutaceae Ferona luc ida Tiejs.& Binn. 1 Salicaceae Salix tetrasperma Roxb. 1 Salviniaceae Salvinia cucu llata Roxb. * * 1,2 Sapindaceae Cardiospermum halicaca bum L. * 1,2 Sc hleic hera oleosa (Lour.) Oken 1 Sapotaceae Mimusops eleng i L. * 1,4 Scrophulariaceae Bacopa sp. * 1,2 Limnop hila geo ffray i Bona ti. 1 Limnop hila repens Ben th. 1 Simaroubaceae Saman dura harman dii Perre. *1 Sterculiaceae Bue ttneria pilosa Roxb. * 1,4 Meloc hia corc horifolia L. * 1,2 Trapaceae Trapa bicorn is Osbec k * 1,2 Trapa na tans L. *5 Tiliaceae Grew ia sinua ta Wall. * 1,4 Brown low ia paludosa Kost. * * 1,2 5 Sicrea go defroyana * 1,4 Verbenaceae Gme lina asiatica L. * * 1,2 Vitex holoa denon D.Dop * * 1,2,4

66 Family Scientific Name Seasonally inundated Aquatic Grasslands Outer Source swamp forest, tropical herbaceous and agro- floodplai (s) evergreen forest and communities ecosystems n pools short-tree shrublands and ricefields

Vitex negun do L. 1 Vitaceae Cissus hexangu laris Thore l * 1,2 Xanthophyllaceae Xan thop hyllum glaucum Wall. * 1,4 Xyridaceae Xyris comp lana ta R. Br. 1 Xyris indica L. 1 Xyris intersita 1 Xyris pauc iflora Willd. 1 Pentapetes pheonicea * 4 Unknown family Drypetes thorelii * 4 Styxis harmandiana * 4 Stryc hnos sp. *4 Desmo dium sp. *4 Glossocarya siamens is Craib. *4

Sources: 1 = Mc Dona ld et al. (1997) 2 = Lamberts (2001) 3 = Davidson (2004) 4 = Rollet (1972) 5 = Rundel (2000)

1 Rollet (1972) mentions Alangium cf. costatum, a synonym for A. ridleyi 2 ?Rollet (1972) may have mistaken for Q. indica 3 ?Rollet (1972) may have mistaken for A. thailandica; Khmer names very similar 4 Listed by Rundel (2000) citing McDonald et al. (1997), but not traced incopy of McDonald et al. (1997) viewed. 5 Rollet (1972) gives genus as Schoutenia

67 ANNEX 2. FISH SPECIES RECORDED IN THE TSBR

Family Species Global Spawning habitat2 Tonle Sap Tonle Inundated Ricefield Refs threat River and Sap forest2 agro- status Delta2 Lake2 ecosyste ms4

Anabantidae Anabas testudineus floodplain * * 1,2,4 Anguillidae Anguilla bicolor *4 Ariidae Arius spp./Hemipimelodus 1 Ariidae Arius stormi Mekong * 2 Ariidae Arius truncatus 3 Ariidae Hemipimelodus borneensis 3 Bagridae Hemibagrus spilopterus *4 Bagridae Heterobagrus bocourti 3 Bagridae Mystus albolineatus floodplain * * 2,4,5 Bagridae Mystus atrifasciatus floodplain * * 2 Bagridae Mystus filamentus floodplain * * 1,2,3 Bagridae Mystus multiradiatus floodplain * * 2,3 Bagridae Mystus mysticetus floodplain * * * 2,3,4 Bagridae Mystus nemurus floodplain * * 1,2 Bagridae Mystus singaringan floodplain * * 2 Bagridae Mystus wicki floodplain * * 2 Bagridae Mystus wolffi floodplain * * 2 Bagridae Mystus wyckioides floodplain * * 2,5 Bagridae Pseudomystus siamensis *4 Belonidae Xenentodon cancila unknown ** *1?,2,3,4 Belonidae Xenentodon canciloides unknown ** 2 Belontiidae Trichogaster pectoralis floodplain * * 1,2,3,4 Belontiidae Trichogaster sp. *4 Belontiidae Trichogaster trichopterus * 3,4 Belontiidae Trichograster microlepis floodplain * * 1,2,3,?4 Belontiidae Trichopsis schalleri *4 Belontiidae Trichopsis vittata *4 Chandidae Parambassis apogoniodes 1 Chandidae Parambassis notatus / siamensis *4 Chandidae Parambassis wolffi floodplain * * * 1,2,4 Chandidae Pseudambasis notatus 3 Channidae Channa lucius floodplain * * 1,2,4 Channidae Channa micropeltes floodplain * * 1,2,4 Channidae Channa orientalis 1 Channidae Channa striata floodplain * * 1,2,4 Clariidae Clarias batrachus floodplain * * * 1?,2,4 Clariidae Clarias macrocephalus floodplain * * * 1?,2,4 Clupeidae Clupeichthys aesarnensis Tonle Sap River * 2,?4 Clupeidae Clupeichthys goniognathus Tonle Sap River * 1,2,?4 Clupeidae Clupeoides borneensis Tonle Sap River * 2 Clupeidae Tenualosa thibaudeaui EN Tonle Sap River * * 1,2 Clupeidae Tenualosa toli Tonle Sap River * 2 Cobitidae Acantopsis sp. *4 Cobitidae Botia helodes floodplain * * * ?1,2,4 Cobitidae Botia lecontei floodplain * * ?1,2, Cobitidae Botia modesta floodplain * * * ?1,2,4 Cynoglossidae Cynoglossus cynoglossus estuaries? * 2 Cynoglossidae Cynoglossus feldmanti estuaries? * 2 Cyprinidae Albulichthys albuloides inundated land * * 1,2 Cyprinidae Amblyrhynchichthys truncatus inundated land * * 2 Cyprinidae Barbodes altus rivers and floodplain * * * 1,2,4 Cyprinidae Barbodes gonionotus rivers and floodplain * * * 1,2,3,4 Cyprinidae Balantiocheilos melanopterus EN 7 Cyprinidae Catlocarpio siamensis ?floodplain * * 1,2 68 Family Species Global Spawning habitat2 Tonle Sap Tonle Inundated Ricefield Refs threat River and Sap forest2 agro- status Delta2 Lake2 ecosyste ms4 Cyprinidae Cirrhinus microlepis floodplain * * * 1,3,4 Cyprinidae Cirrhinus molitorella floodplain * * 2 Cyprinidae Cosmochilus harmandi inundated land * * 1,2 Cyprinidae Crossocheilus reticulatus 3 Cyprinidae Cyclocheilichthys apogon 1,3,4? Cyprinidae Cyclocheilichthys armatus inundated land * * 1,2,4? Cyprinidae Cyclocheilichthys enoplos inundated land * * 1,2,3 Cyprinidae Cyclocheilichthys lagleri inundated land * * 2 Cyprinidae Cyprinus carpio unknown * 1,2 Cyprinidae Dangila cf. cuvieri floodplain * * 1?,2 Cyprinidae Dangila kuhli floodplain * * 1?,2 Cyprinidae Dangila lineata floodplain * * 1?,2,3 Cyprinidae Dangila spilopleura floodplain * * 1,2,3 Cyprinidae Danio aequipinnatus 1 Cyprinidae Esomus longimanus 3 Cyprinidae Esomus metallicus * 3,4 Cyprinidae Hampala dispar inundated land * * 1,2,3 Cyprinidae Hampala macrolepidota inundated land * * * 2,3,4 Cyprinidae Henicorhynchus caudimaculatus rivers and floodplain * * ?1,2 Cyprinidae Henicorhynchus cryptopogon rivers and floodplain * * ?1,2 Cyprinidae Henicorhynchus siamensis rivers and floodplain * * * ?1,2,3,4 Cyprinidae Hypsibarbus lageri 3 Cyprinidae Labeo erythropterus 3 Cyprinidae Labiobarbus siamensis *4 Cyprinidae Leptobarbus hoevenii floodplain * * 1,2,3,4 Cyprinidae Lobocheilos melanotaenia floodplain * * 2 Cyprinidae Luciosoma bleekeri * 3,4 Cyprinidae Luciosoma setigerum floodplain * * 1,2 Cyprinidae Morulius chrysophekadion rivers * * 1,2,3 Cyprinidae Opsarus koratensis rivers? * 2 Cyprinidae Opsarus pulchellus rivers? * 2 Cyprinidae Osteochilus hasselti floodplain * * * 1,2,3,4 Cyprinidae Osteochilus melanopleurus floodplain * * * 1,2,3,4 Cyprinidae Osteochilus microcephalus 3 Cyprinidae Osteochilus schlegeli floodplain * * 1,2 Cyprinidae Parachela maculicanda floodplain * * 2 Cyprinidae Parachela siamensis * 1,3,4 Cyprinidae Paralaubuca typus unknown * * * 1,2,3,4 Cyprinidae Probarbus jullieni EN rivers and floodplain * * 1,2 Cyprinidae Probarbus labeamajor DD 8 Cyprinidae Puntioplites bulu inundated land * * 1,2 Cyprinidae Puntioplites proctozysion inundated land * * * 1,2,4 Cyprinidae Puntius brevis inundated land * * * 1,2,3,4 Cyprinidae Puntius masyai inundated land * * 2 Cyprinidae Rasbora aurotaenia rivers and ponds * 1,2,3 Cyprinidae Rasbora borapetensis *4 Cyprinidae Rasbora caudimaculata 3 Cyprinidae Rasbora daniconius rivers and ponds * * 2,4 Cyprinidae Rasbora hobelmani rivers and ponds * 2 Cyprinidae Rasbora pausisquamis rivers and ponds * 2 Cyprinidae Rasbora paviei rivers and ponds * 2 Cyprinidae Rasbora tornieri rivers and ponds * * 2,4 Cyprinidae Rasbora trilineata *4 Cyprinidae Sikukia gudgeri 1

69 Family Species Global Spawning habitat2 Tonle Sap Tonle Inundated Ricefield Refs threat River and Sap forest2 agro- status Delta2 Lake2 ecosyste ms4 Cyprinidae Systomus orphoides rivers and floodplain * * 1,2,3 Cyprinidae Systomus partipentazona *4 Cyprinidae Thynnichthys thynnoides floodplain * * * 1,2,3,4 Datnioididae Datnioides microlepis 1 Eleotridae Oxyeleotris marmorata floodplain * * 1,2,4 Engraulidae Coilia lindmani Tonle Sap River * * ?1,2,3 Engraulidae Coilia macrognathos Tonle Sap River * * ?1,2 Engraulidae Lycothrissa crocodilus unknown * * 2,3 Engraulidae Macrochirichthys macrochirus unknown * * 1,2 Engraulidae Setipinna melanochir unknown * 1,2 Gobiidae Glossogobius aureus floodplain * * 2 Gyrinocheilidae Gyrinocheilus aymonieri 5 Hemiramphidae Hyporhamphus limbatus unknown * * 1,2,3 Mastacembelidae Macrognathus maculatus unknown * 2 Mastacembelidae Macrognathus siamensis unknown * * 1,2,?3,4 Mastacembelidae Macrognathus taeniagaster unknown * * 2,4,?3 Mastacembelidae Mastacembelus favus unknown * * * 1,2,4 Nandidae Nandus nandus *4 Nandidae Pristolepis fasciata floodplain * * * 1,2,3,4 Notopteridae Chitala blanci inundated forest * * * 2 Notopteridae Chitala ornata inundated forest * * * * 1,2,4 Notopteridae Notopterus notopterus inundated forest * * 1,2,4 Pangasiidae Pangasionodon gigas CR * * 6 Pangasiidae Pangasionodon hypophthalmus 1,3 Pangasiidae Pangasius bocourti 5 Pangasiidae Pangasius conchophilus *4 Pangasiidae Pangasius djambal 5 Pangasiidae Pangasius krempfi 5 Pangasiidae Pangasius larnaudii floodplain * * 1,2,3,5 Pangasiidae Pangasius siamensis floodplain * * 1,2 Polynemidae Polynemus borneensis unknown * 2 Polynemidae Polynemus longipectoralis 3 Polynemidae Polynemus mutifilis 1 Sciacnidae Boesemania microlepis unknown * 1,2 Siluridae Belodontichthys dinema floodplain * * 1,2,3 Siluridae Kryptopterus cheveyi 3 Siluridae Kryptopterus cryptopterus river * * 1,2,3 Siluridae Kryptopterus schilbeides 3 Siluridae Micronema apogon floodplain * * 1?,2,3 Siluridae Micronema bleekeri floodplain * * 1?,2,5 Siluridae Micronema micronema floodplain * * * 1?2,4,5 Siluridae Ompok bimaculatus floodplain? * * * 2,4,5 Siluridae Ompok hypophthalmus floodplain? * * * 1,2,4 Siluridae Ompok sp. cf. eugeniatus floodplain? * * 2 Siluridae Wallago attu floodplain? * * * 1,2,4 Siluridae Wallago leeri floodplain? * * 2 Sisoridae Bagarius/Glypthorax spp. 1 Soleidae Achiroides leucorhynchos 1 Soleidae Euryglossa harmandi large rivers? * 2 Soleidae Euryglossa orientalis estuaries? * 2 Soleidae Euryglossa panoides estuaries? * 2 Synbranchidae Monopterus albus * 4,5 Syngnathidae Doryichthys boaja *4 Tetraodontidae Chelonodon fluviatilis estuaries * 2

70 Family Species Global Spawning habitat2 Tonle Sap Tonle Inundated Ricefield Refs River and Sap forest2 agro- Delta2 Lake2 ecosyste ms4 Tetraodontidae Chelonodon nigroviridis estuaries * * 2 Tetraodontidae Monotreta cambodgiensis * 1,3,4 Toxotidae Toxotes chatareus floodplain * 1,2,3 Toxotidae Toxotes microlepis floodplain 2

References: 1 = van Zalinge et al. 1999 2 = Lim et al. 1999 3 = Lamberts 2001 4 = Balzer et al. 2003 5 = Neou et al. 2003 6 = Hogan et al. 2001 7 = Rainboth 1996 8 = Poulsen et al. 2004

71 ANNEX 3. BIRD SPECIECS RRECORDED IN THE TSBR

Open water Seasonally Herbaceous Global (lake, inundated Grasslands English Name (sensu Robson TS wetlands Scientific Name Threat lakeshore and swamp forest and agro- 2000) Status (including Status major and short-tree ecosystems sedge beds) tributaries) shrublands Rain Quail Coturnix coromandelica * Blue-breasted Quail Coturnix chinensis * Lesser Whistling-duck Dendrocygna javanica * ** Comb Duck Sarkidiornis melanotos *** Cotton Pygmy-goose Nettapus coromandelianus ** Spot-billed Duck Anas poecilorhyncha *** Small Buttonquail Turnix sylvatica * Barred Buttonquail Turnix suscitator * Fulvous-breasted Woodpecker Dendrocopos macei * Rufous Woodpecker Celeus brachyurus * Laced Woodpecker Picus vittatus * Common Flameback Dinopium javanense * Indian Roller Coracias benghalensis * Common Kingfisher Alcedo atthis *** Stork-billed Kingfisher Halcyon capensis ** White-throated Kingfisher Halcyon smyrnensis ** * Black-capped Kingfisher Halcyon pileata ** * Collared Kingfisher Todiramphus chloris ** Pied Kingfisher Ceryle rudis ** Blue-tailed Bee-eater Merops philippinus **** Chestnut-winged Cuckoo Clamator coromandus * Large Hawk Cuckoo Hierococcyx sparverioides * Indian Cuckoo Cuculus micropterus * Oriental Cuckoo Cuculus saturatus * Plaintive Cuckoo Cacomantis merulinus * Drongo Cuckoo Surniculus lugubris * Green-billed Malkoha Phaenicophaeus tristis * Greater Coucal Centropus sinensis *** Lesser Coucal Centropus bengalensis *** Red-breasted Parakeet Psittacula alexandri * Silver-backed Needletail Hirundapus cochinchinensis ** Brown-backed Needletail Hirundapus giganteus ** Asian Palm Swift Cypsiurus balasiensis * Fork-tailed Swift Apus pacificus * Barn Owl Tyto alba * Spotted Wood Owl Strix seloputo * Spotted Owlet Athene brama * Large-tailed Nightjar Caprimulgus macrurus * Rock Pigeon Columba livia * Spotted Dove Streptopelia chinensis * Red Collared Dove Streptopelia tranquebarica * Peaceful Dove Geopelia striata * Bengal Florican Houbaropsis bengalensis EN * Sarus Crane Grus antigone VU * Masked Finfoot Heliopais personata VU * Slaty-breasted Rail Gallirallus striatus ** White-breasted Waterhen Amaurornis phoenicurus ** Baillon's Crake Porzana pusilla * Ruddy-breasted Crake Porzana fusca * White-browed Crake Porzana cinerea ** Watercock Gallicrex cinerea ** Purple Swamphen Porphyrio porphyrio *** Common Moorhen Gallinula chloropus ** Pintail Snipe Gallinago stenura ** 72 Open water Seasonally Herbaceous Global (lake, inundated Grasslands English Name (sensu Robson wetlands Scientific Name Threat lakeshore and swamp forest and agro- 2000) (including Status major and short-tree ecosystems sedge beds) tributaries) shrublands Black-tailed Godwit Limosa limosa ** Whimbrel Numenius phaeopus ** Spotted Redshank Tringa erythropus ** Common Redshank Tringa totanus ** Marsh Sandpiper Tringa stagnatilis ** Common Greenshank Tringa nebularia ** Green Sandpiper Tringa ochropus * Wood Sandpiper Tringa glareola * Common Sandpiper Actitis hypoleucos * Red-necked Stint Calidris ruficollis * Temminck's Stint Calidris temminckii ** Long-toed Stint Calidris subminuta * Curlew Sandpiper Calidris ferruginea * Broad-billed Sandpiper Limicola falcinellus * Ruff Philomachus pugnax * Red-necked Phalarope Phalaropus lobatus * Greater Painted-snipe Rostratula benghalensis * Pheasant-tailed Jacana Hydrophasianus chirurgus *** Bronze-winged Jacana Metopidius indicus *** Eurasian Thick-knee Burhinus oedicnemus * Black-winged Stilt Himantopus himantopus ** Pacific Golden Plover Pluvialis fulva * Grey Plover Pluvialis squatarola * Little Ringed Plover Charadrius dubius * Oriental Plover Charadrius veredus * Grey-headed Lapwing Vanellus cinereus ** Red-wattled Lapwing Vanellus indicus * Oriental Pratincole Glareola maldivarum * Brown-headed Gull Larus brunnicephalus * Black-headed Gull Larus ridibundus * Caspian Tern Sterna caspia * Whiskered Tern Chlidonias hybridus *** White-winged Tern Chlidonias leucopterus * Osprey Pandion haliaetus **** Black Kite Milvus migrans * Brahminy Kite Haliastur indus **** White-bellied Sea Eagle Haliaeetus leucogaster ** Grey-headed Fish Eagle Ichthyophaga ichthyaetus NT * * Short-toed Snake Eagle Circaetus gallicus * Pied Harrier Circus melanoleucos * Shikra Accipiter badius * Japanese Sparrowhawk Accipiter gularis * Greater Spotted Eagle Aquila clanga VU * Imperial Eagle Aquila heliaca VU * Common Kestrel Falco tinnunculus * Peregrine Falcon Falco peregrinus *** Little Grebe Tachybaptus ruficollis * Darter Anhinga melanogaster NT * * * Little Cormorant Phalacrocorax niger ** * Indian Cormorant Phalacrocorax fuscicollis ** * Great Cormorant Phalacrocorax carbo ** * Little Egret Egretta garzetta **** Grey Heron Ardea cinerea **** Purple Heron Ardea purpurea **** Great Egret Casmerodius albus ****

73 Open water Seasonally Herbaceous Global (lake, inundated Grasslands English Name (sensu Robson wetlands Scientific Name Threat lakeshore and swamp forest and agro- 2000) (including Status major and short-tree ecosystems sedge beds) tributaries) shrublands Intermediate Egret Mesophoyx intermedia **** Cattle Egret Bubulcus ibis *** Chinese Pond Heron Ardeola bacchus **** Javan Pond Heron Ardeola speciosa *** Little Heron Butorides striatus * Black-crowned Night Heron Nycticorax nycticorax * Yellow Bittern Ixobrychus sinensis *** Cinnamon Bittern Ixobrychus cinnamomeus *** Black Bittern Dupetor flavicollis *** Greater Flamingo Phoenicopterus ruber Glossy Ibis Plegadis falcinellus ** Black-headed Ibis Threskiornis melanocephalus NT * * White-shouldered Ibis Pseudibis davisoni CR * * Eurasian Spoonbill Platalea leucorodia * Great White Pelican Pelecanus onocrotalus * Spot-billed Pelican Pelecanus philippensis VU * * * Milky Stork Mycteria cinerea VU * * * Painted Stork Mycteria leucocephala NT * * * Asian Openbill Anastomus oscitans *** Woolly-necked Stork Ciconia episcopus ** Black-necked Stork Ephippiorhynchus asiaticus NT * * Lesser Adjutant Leptoptilos javanicus VU * * * Greater Adjutant Leptoptilos dubius EN * * * Blue-winged Pitta Pitta moluccensis * Black-and-red Broadbill Cymbirhynchus macrorhynchos * Brown Shrike Lanius cristatus * Mangrove Whistler Pachycephala grisola * Racket-tailed Treepie Crypsirina temia ** Large-billed Crow Corvus macrorhynchos ** Ashy Minivet Pericrocotus divaricatus * Pied Fantail Rhipidura javanica * Black Drongo Dicrurus macrocercus * Ashy Drongo Dicrurus leucophaeus * Black-naped Monarch Hypothymis azurea * Asian Paradise-flycatcher Terpsiphone paradisi * Common Iora Aegithina tiphia * Blue Rock Thrush Monticola solitarius ** Asian Brown Flycatcher Muscicapa dauurica ** Tickell's Blue Flycatcher Cyornis tickelliae * Siberian Rubythroat Luscinia calliope * Bluethroat Luscinia svecica * Oriental Magpie Robin Copsychus saularis * Common Stonechat Saxicola torquata * Pied Bushchat Saxicola caprata * White-shouldered Starling Sturnus sinensis ** Asian Pied Starling Sturnus contra * Common Myna Acridotheres tristis ** White-vented Myna Acridotheres grandis ** Sand Martin Riparia riparia ** Barn Swallow Hirundo rustica ** Red-rumped Swallow Hirundo daurica ** Asian House Martin Delichon dasypus ** Yellow-vented Bulbul Pycnonotus goiavier * Streak-eared Bulbul Pycnonotus blanfordi * Zitting Cisticola Cisticola juncidis *

74 Open water Seasonally Herbaceous Global (lake, inundated Grasslands English Name (sensu Robson wetlands Scientific Name Threat lakeshore and swamp forest and agro- 2000) (including Status major and short-tree ecosystems sedge beds) tributaries) shrublands Bright-headed Cisticola Cisticola exilis * Yellow-bellied Prinia Prinia flaviventris * Plain Prinia Prinia inornata ** [Spotted Bush Warbler] * Bradypterus thoracicus Lanceolated Warbler Locustella lanceolata * Rusty-rumped Warbler Locustella certhiola ** Black-browed Reed Warbler Acrocephalus bistrigiceps *** Manchurian Reed Warbler Acrocephalus tangorum VU * * * Blunt-winged Warbler Acrocephalus concinens * Oriental Reed Warbler Acrocephalus orientalis ** Thick-billed Warbler Acrocephalus aedon * Dusky Warbler Phylloscopus fuscatus ** Radde's Warbler Phylloscopus schwarzi * Yellow-browed Warbler Phylloscopus inornatus * Striated Grassbird Megalurus palustris * White-crested Laughingthrush Garrulax leucolophus * Striped Tit Babbler Macronous gularis * Australasian Bushlark Mirafra javanica * Oriental Skylark Alauda gulgula * Scarlet-backed Flowerpecker Dicaeum cruentatum * Ruby-cheeked Sunbird Anthreptes singalensis * Purple-throated Sunbird Nectarinia sperata * Olive-backed Sunbird Nectarinia jugularis * House Sparrow Passer domesticus * Plain-backed Sparrow Passer flaveolus * Eurasian Tree Sparrow Passer montanus * White Wagtail Motacilla alba ** Yellow Wagtail Motacilla flava *** Richard's Pipit Anthus richardi * Paddyfield Pipit Anthus rufulus * Red-throated Pipit Anthus cervinus * Streaked Weaver Ploceus manyar * Baya Weaver Ploceus philippinus * Asian Golden Weaver Ploceus hypoxanthus NT * Red Avadavat Amandava amandava * Scaly-breasted Munia Lonchura punctulata * Yellow-breasted Bunting NT * Emberiza aureola

This species list has been compiled from records published in Recent Sightings in Cambodia Bird News nos.1-13, Goes 2001 and other articles in Cambodia Bird News 8, and Goes and Hong 2002.

75 ANNEX 4. MAMMAL SPECIES RECORDEDIN THE TSBR

English Name Scientific Name Global Seasonally Grasslands and agro- Threat inundated ecosystems Status swamp forest and short-tree shrublands

Loris sp. Nycticebus sp. ? * Long-tailed Macaque Macaca fascicularis NT * Germain’s Silver Leaf Monkey Trachypithecus villosus germaini DD * Hairy-nosed Otter Lutra sumatrana DD * Smooth Otter Lutrogale perspicillata VU * Asiatic Jackal Canis aureus * Fishing Cat Prionailurus viverrinus VU * Leopard Cat Prionailurus bengalensis ? Small Asian Mongoose Herpestes javanicus * Giant Flying Squirrel Petuarista sp. * Variable Squirrel Callosciurus finlaysonii * Lyle’s or Large Flying Fox Pteropus lylei/vampyrus * Bandicoot Rat Bandicota indica * Lesser Bandicoot Rat Bandicota savilei * Field Rat Rattus rattus * Ricefield Rat Rattus argentiventer * Lesser Ricefield Rat Rattus losea * Field Mouse Mus cervicolor *

76