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The Avifauna of the Mo Singto Forest Dynamics Plot, Khao Yai National Park, Thailand

By Philip D. Round, Andrew J. Pierce, Wangworn Sankamethawee & George A. Gale
 
Abstract
The 169 species of birds recorded on the 30 ha Mo Singto Forest Dynamics Plot, Khao Yai National Park, Thailand, are a subset of the 329 species recorded in the headquarters area of the park. Most of the Mo Singto-recorded species are typical of evergreen forest interior but the transient occurrence of a small number of other species, inhabitants of forest edge or more open habitats, is documented. Almost one third of species found on the plot were moderate to long-distance migrant, non-breeding, visitors. The largest foraging guilds were foliage-gleaning insectivores and sallying insectivores, together accounting for over one-third of all species. Though obligate frugivores were poorly represented, most insectivorous birds incorporated fruits in their diet.

The nearly 30-year history of avian recording at Mo Singto, with intensive community studies having been conducted in the past decade, and the location of the study plot near the submontane-montane transition, make it an ideal site for continuation of detailed monitoring, particularly that related to the impact of climate change.
INTRODUCTION
Observations of birds at Mo Singto, in Khao Yai National Park, have been made for as long as observations have been conducted on gibbons (Brockelman & Gittins, 1984; Marshall et al., 1972; Marshall & Sugardjito, 1986), and for almost the same duration as on birds in the wider park area. Systematic compilation of a Khao Yai avifauna began soon after the park’s establishment (Dickinson, 1963, 1967; Dickinson & Tubbbb, 1964, 1966; McClure, 1974). Both gibbon researchers and birdwatchers collected information on birds on the Mo Singto Forest Dynamics Plot (hereafter MFDP) from at least the early 1980s. A comprehensive inventory of the birds of Khao Yai and adjacent areas in what is known as the Dong Phaya Yen Forest Complex, was given in Lynam et al. (2006) with updates in Pobprasert et al. (2008).

The establishment of a marked grid in 1998 provided a new opportunity and incentive to conduct detailed studies of the bird community involving individual colour-marking, territory mapping of birds, evaluating census methods, and studies of breeding biology and social behaviour (Dhanasarnpaiboon & Round, 2004; Gale et al., 2009; Nimnuan et al., 2004; Pierce, 2004, 2005, 2009; Pierce & Pobprasert, 2007; Pierce and Round, 2006; Pierce et al., 2004a, 2004b, 2007; Pobprasert & Gale, 2010; Pobprasert & Pierce, 2010; Pobprasert et al., 2008; Praditsup et al., 2007; Round & Gale, 2008; Round et al., 2006; Sankamethavee et al., 2009, 2010; Savini & Sukumal, 2009; Sukumal & Savini, 2009a, 2009b; Sukumal et al., 2010; Steward, 2010; Tokue, 2007.)

Detailed inventories and long-term data sets for tropical forest sites are rare. Round (1984) assessed avifaunal changes at another Thai site, Doi Suthep-Pui, over a 50-year period. Turnover, encompassing both apparent losses due to hunting and habitat degradation, and many additions (“new” records), was documented over this period. Assessment of faunal change is complex: some turnover may be stochastic in nature; some species may be extirpated through human activity; other “genuinely new” species may colonize due to changes in the environment or appear for short periods in response to ephemeral disturbance. Increased observer effort through time at any site will inevitably accumulate new records. Increasingly, climate change is recognized as a major anthropogenic disturbance that not only impacts habitats and plant distribution, but also causes distributional and behavioural changes in animals (e.g. Crick & Sparks, 1999; Hickling et al., 2006; Schaefer et al., 2006).

Because of MFDP’s location in the heart of the 2,169 km2 Khao Yai National Park its avifauna is exempt from hunting and most forms of direct human disturbance (though even there, some collection of high-value forest products, such as Aquilaria wood, is still carried out; Brockelman et al., 2011). The location of MFDP may prove to be an especially fortuitous choice for studying climate change-related impacts on the bird community since the plot is situated close to the lowland-montane ecotone (for Thailand and tropical SE Asia usually placed at 900–1000 m elevation; Neal 1967; Whitmore, 1975). Round & Gale (2008) demonstrated major changes in the relative abundance of two pheasant species (one lowland, one montane) in and around MFDP over a two-decade period which they hypothesized was related to warming temperatures.

The purpose of this paper, therefore, is to provide an updated inventory of bird species recorded on the Mo Singto Forest Dynamics Plot and to discuss this in the larger context of the park of which it is a small part. This will provide a baseline for assessing future turnover in the bird community, and will also be useful as a reference for ecological studies of interactions such as seed dispersal and predation.
STUDY AREA & METHODS
Details of MFDP are set out in Brockelman et al. (2011). In brief, it is an area of 30 ha, located in closed-canopy, moist, evergreen forest habitat at elevations of 725 to 815 m. It lies within 500 m of the forest edge and park headquarters. The range of habitats and species diversity of plants and animals in the immediate surroundings was likely increased by the large expanse of forest clearings, formerly settled and cultivated areas, which became Imperata grassland and scrub when the park was established in 1962. Though grasslands are still burnt during management, clearings are gradually reverting to scrub and secondary forest. No clearings exist within the plot itself.

The species listed (Table 1) are all those recorded within the confines of the present-day plot. This includes those recorded opportunistically by birdwatchers and biologists from 1980 onwards; and those noted during systematic censuses of MFDP during 2002–2006 and subsequently (Pierce & Round, 2006; Round et al., 2006; Pobprasert et al., 2008; Gale et al., 2009). It also includes species seen in the airspace over the plot. Observations were carried out year-round and the records also include species detected by mist-netting and capture in baited spring-traps. Nocturnal birds were detected both on and in the immediate vicinity of the plot by listening for calls at night while walking forest trails (though much less search effort was concentrated at night than during the day). All sight photographic and aural records (both those of the authors and those submitted by outside observers) were authenticated by peer-review.

The plot constitutes a small part of a plateau in the north-west quadrant of the park. This relatively homogeneous habitat and elevational zone, extending to the rim of the northern park scarp, and southwards to the summit of the Khao Khieo–Khao Rom scarp (1,350 m), and eastwards to the Haew Suwat waterfall, covers an area of roughly 60 km2. This corresponds to the “headquarters area” (hereafter Hq area) of the park. It is traversed by roads and trails and therefore frequently visited. At higher elevations, the forest grades into a montane forest type. At lower elevations, around the northern rim, it grades into a drier, semi-evergreen or mixed deciduous facies.

Besides providing a comprehensive listing of bird species known for MFDP, we have also listed all other species known for Hq area with reference to the baseline inventory for the park in Lynam et al. (2006) and Pobprasert et al., 2008 (Table 2). We have omitted species listed by other authors where it cannot be established whether the records were from the Hq area as defined above. We discuss those that might have the potential to either colonise, disperse or stray into MFDP due to habitat change induced by ecological succession, human-induced or natural disturbance, or during random turnover. Taxonomic order and nomenclature are based on Robson (2008).
RESULTS
Species Found on MFDP
The 169 species recorded on MFDP (both residents and migrants or winter visitors) are a subset of the total of 329 species which have been recorded in the Hq area of the park as a whole (Table 1, Table 2). Of the MFDP-recorded species, 112 are thought to be resident (either on MFDP itself or in the immediately adjacent Hq area); one is a wet-season breeding visitor; 47 species are considered non-breeding (winter) visitors, seven are spring and autumn passage migrants. At least two species recorded on MFDP are represented in the park by both resident and wintering populations (Table 1).

Of these, 128 are more or less regular on MFDP, while another 41 are considered rare (< 10 sightings). In general, MFDP was very intensively surveyed and most rare species were considered to be genuinely rare, either because they were birds that tended to occur at low density and had large territories (e.g. most raptors) or were those normally characteristic of more open habitats. At least one aerial feeder (e.g. Asian House-martin) may have been under-recorded due to the difficulty of observing them from within closed canopy forest.
Table 1. List of bird species recorded on the Mo Singto Forest Dynamics Plot, Khao Yai National Park.
Seasonal status: resident/presumed resident (R); non-breeding visitor (N); breeding visitor (B); passage migrant (P); [ ] rarely recorded on plot (< ten records).
Habitats: grassland or scrub (G); forest or woodland (F); aerial feeders (A)
Guilds: Following Johns (1986), terrestrial insectivore–frugivore (TIF), bark-gleaning insectivore (BGI), arboreal insectivore–frugivore (AIF), arboreal faunivore/frugivore (FF), foliage-gleaning insectivore (FGI), terrestrial insectivore–faunivore (TIV), piscivore (P), diurnal or nocturnal raptor (R), sallying insectivore (SaI), sweeping insectivore (SwI), terrestrial insectivore (TI), insectivore–nectarivore (IN).
Common Name Scientific Name Seasonal
Status
Habitat Guild
Scaly-breasted Partridge
Red Junglefowl
Silver Pheasant
Siamese Fireback
Malaysian Night-heron
Chinese Pond-heron
Jerdon’s Baza
Black Baza
Oriental Honey-buzzard
Crested Serpent-eagle
Crested Goshawk
Shikra
Japanese Sparrowhawk
Besra
Mountain Hawk-eagle
Eurasian Woodcock
Spotted Dove
Barred Cuckoo-dove
Emerald Dove
Thick-billed Green-pigeon
Wedge-tailed Green-pigeon
Mountain Imperial-pigeon
Vernal Hanging-parrot
Chestnut-winged Cuckoo
Large Hawk-cuckoo
Hodgson’s Hawk-cuckoo
Indian Cuckoo
Banded Bay Cuckoo
Asian Emerald Cuckoo
Violet Cuckoo
Drongo Cuckoo
Greater Coucal
Coral-billed Ground- cuckoo
Green-billed Malkoha
Oriental Bay Owl
Mountain Scops-owl
Collared Scops-owl
Oriental Scops-owl
Spot-bellied Eagle-owl
Brown Wood-owl
Collared Owlet
Asian Barred Owlet
Brown Boobook
Great Eared Nightjar
Brown-backed Needletail
Asian Palm-swift
Orange-breasted Trogon
Red-headed Trogon
Oriental Dollarbird
Banded Kingfisher
Black-backed Kingfisher
Blue-eared Kingfisher
Blue-bearded Bee-eater
Chestnut headed Bee-eater
Northern Brown Hornbill
Oriental Pied Hornbill
Great Hornbill
Wreathed Hornbill
Green-eared Barbet
Moustached Barbet
Blue-eared Barbet
Rufous Woodpecker
Greater Yellownape
Laced Woodpecker
Greater Flameback
Black-and-buff Woodpecker
Heart-spotted Woodpecker
Great Slaty Woodpecker
Long-tailed Broadbill
Dusky Broadbill
Silver-breasted Broadbill
Banded Broadbill
Hooded Pitta
Blue Pitta
Eared Pitta
White-bellied Erpornis
Black-winged Cuckooshrike
Rosy Minivet
Swinhoe’s Minivet
Scarlet Minivet
Black-naped Oriole
Silver Oriole
Large Woodshrike
Bar-winged Flycatcher-shrike
Common Iora
Great Iora
Ashy Drongo
Bronzed Drongo
Lesser Racket-tailed Drongo
Greater Racket-tailed Drongo
Hair-crested Drongo
Black-naped Monarch
Asian Paradise-flycatcher
Japanese Paradise- flycatcher
Eastern Jungle Crow
Common Green Magpie
Racket-tailed Treepie
Tiger Shrike
Grey-backed Shrike
Black-throated Sunbird
Ruby-cheeked Sunbird
Little Spiderhunter
Thick-billed Flowerpecker
Yellow-vented Flowerpecker
Plain Flowerpecker
Fire-breasted Flowerpecker
Blue-winged Leafbird
Asian Fairy-bluebird
White-rumped Munia
Pin-tailed Parrotfinch
Grey Wagtail
Common Rosefinch
Velvet-fronted Nuthatch
Common Hill-myna
Orange-headed Thrush
White’s Thrush
Siberian Thrush
Grey-sided Thrush
Eyebrowed Thrush
Siberian Blue Robin
White-throated Rock-thrush
Slaty-backed Forktail
White-crowned Forktail
Blue Whistling-thrush
Hainan Blue Flycatcher
Hill Blue Flycatcher
Verditer Flycatcher
Rufous-bellied Niltava
Mugimaki Flycatcher
Slaty-backed Flycatcher
Dark-sided Flycatcher
Asian Brown Flycatcher
White-rumped Shama
Sultan Tit
Grey-headed Flycatcher
Black-headed Bulbul
Black-crested Bulbul
Stripe-throated Bulbul
Grey-eyed Bulbul
Puff-throated Bulbul
Ashy Bulbul
Asian House-martin
Barn Swallow
Red-rumped Swallow
Asian Stubtail
Martens’s Warbler
Plain-tailed Warbler
Eastern Crowned Warbler
Sulphur-breasted Warbler
Claudia’s Warbler *
Arctic Warbler
Pale-legged Leaf-warbler
Two-barred Warbler
Yellow-browed Warbler
Radde’s Warbler
Chestnut-flanked White-eye
Everett’s White-eye
Large Scimitar-babbler
White-browed Scimitar-babbler
Pin-striped Tit-babbler
Puff-throated Babbler
Scaly-crowned Babbler
Abbott’s Babbler
Black-throated Laughingthrush
White-crested Laughingthrush
Lesser Necklaced Laughingthrush
Dark-necked Tailorbird
Common Tailorbird
Arborophila chloropus
Gallus gallus
Lophura nycthemera
Lophura diardi
Gorsachius melanolophus
Ardeola bacchus
Aviceda jerdoni
Aviceda leuphotes
Pernis ptilorhynchus
Spilornis cheela
Accipiter trivirgatus
Accipiter badius
Accipiter gularis
Accipiter virgatus
Nisaetus nipalensis
Scolopax rusticola
Streptopelia chinensis
Macropygia unchall
Chalcophaps indica
Treron curvirostra
Treron sphenurus
Ducula badia
Loriculus vernalis
Clamator coromandus
Hierococcyx sparverioides
Hierococcyx nisicolor
Cuculus micropterus
Cacomantis sonneratii
Chrysococcyx maculatus
Chrysococcyx xanthorhynchus
Surniculus lugubris
Centropus sinensis
Carpococcyx renauldi
Phaenicophaeus tristis
Phodilus badius
Otus spilocephalus
Otus lettia
Otus sunia
Bubo nipalensis
Strix leptogrammica
Glaucidium brodiei
Glaucidium cuculoides
Ninox scutulata
Eurostopodus macrotis
Hirundapus giganteus
Cypsiurus balasiensis
Harpactes oreskios
Harpactes erythrocephalus
Eurystomus orientalis
Lacedo pulchella
Ceyx erithacus
Alcedo meninting
Nyctyornis athertoni
Merops leschenaulti
Anorrhinus austeni
Anthracoceros albirostris
Buceros bicornis
Aceros undulatus
Megalaima faiostricta
Megalaima incognita
Megalaima australis
Micropternus brachyurus
Chrysophlegma flavinucha
Picus vittatus
Chrysocolaptes lucidus
Meiglyptes jugularis
Hemicircus canente
Mulleripicus pulverulentus
Psarisomus dalhousiae
Corydon sumatranus
Serilophus lunatus
Eurylaimus javanicus
Pitta sordida
Pitta cyanea
Anthocincla phayrei
Erpornis zantholeuca
Coracina melaschistos
Pericrocotus roseus
Pericrocotus cantonensis
Pericrocotus speciosus
Oriolus chinensis
Oriolus mellianus
Tephrodornis gularis
Hemipus picatus
Aegithina tiphia
Aegithina lafresnayei
Dicrurus leucophaeus
Dicrurus aeneus
Dicrurus remifer
Dicrurus paradiseus
Dicrurus hottentottus
Hypothymis azurea
Terpsiphone paradisi
Terpsiphone atrocaudata
Corvus levaillanti
Cissa chinensis
Crypsirina temia
Lanius tigrinus
Lanius tephronotus
Aethopyga saturata
Chalcoparia singalensis
Arachnothera longirostra
Dicaeum agile
Dicaeum chrysorrheum
Dicaeum minullum
Dicaeum ignipectus
Chloropsis cochinchinensis
Irena puella
Lonchura striata
Erythrura prasina
Motacilla cinerea
Carpodacus erythrinus
Sitta frontalis
Gracula religiosa
Zoothera citrina
Zoothera aurea
Zoothera sibirica
Turdus feae
Turdus obscurus
Luscinia cyane
Monticola gularis
Enicurus schistaceus
Enicurus leschenaulti
Myophonus caeruleus
Cyornis hainanus
Cyornis banyumas
Eumyias thalassinus
Niltava sundara
Ficedula mugimaki
Ficedula hodgsonii
Muscicapa sibirica
Muscicapa dauurica
Copsychus malabaricus
Melanochlora sultanea
Culicicapa ceylonensis
Pycnonotus atriceps
Pycnonotus flaviventris
Pycnonotus finlaysoni
Iole propinqua
Alophoixus pallidus
Hemixos flavala
Delichon dasypus
Hirundo rustica
Hirundo daurica
Urosphena squameiceps
Seicercus omeiensis
Seicercus soror
Phylloscopus coronatus
Phylloscopus ricketti
Phylloscopus claudiae
Phylloscopus borealis
Phylloscopus tenellipes
Phylloscopus plumbeitarsus
Phylloscopus inornatus
Phylloscopus schwarzi
Zosterops erythropleurus
Zosterops everetti
Pomatorhinus hypoleucos
Pomatorhinus schisticeps
Macronous gularis
Pellorneum ruficeps
Malacopteron cinereum
Malacocincla abbotti
Dryonastes chinensis
Garrulax leucolophus
Garrulax monileger
Orthotomus atrogularis
Orthotomus sutorius
R
R
R
R
[R]
[N]
[R]
N
F TIF
F TIF
F TIF
F TIF
F P
F P
F P
F R
R,N F R
R F R
R F R
R F R
P F R
[R] F R
[R] F R
N F TI
[R] G TF
R F AF
R F TF
R F AF
[R] F AF
R F AF
R F AF
[N] F FGI
N F FGI
[R] F FGI
[N] F FGI
R F FGI
N F FGI
[R] F FGI
R F FGI
[R] G TIV
R F TIV
R F FGI
R F R
R F R
R F R
[N] F R
[R] F R
[R] F R
R F R
R F R
R F R
R F SwI
R A SwI
R A SwI
R F FGI
R F FGI
[R] F SaI
R F TI
[P] F P
[R] F P
R F SaI
R A SaI
R F FF
R F FF
R F FF
R F FF
R F AIF
R F AIF
R F AIF
R F BGI
R F BGI
R F BGI
R F BGI
R F BGI
R F BGI
[R] F BGI
R F FGI
[R] F FGI
R F FGI
B F FGI
B F TIV
R F TIV 
R F TIV 
R F FGI 
N F FGI
N F FGI
N F FGI
R F FGI
N F FGI
[N] F FGI
R F FGI
R F FGI
[R] F FGI
R F FGI
N F SaI
R F SaI
N F SaI
R F SaI
N F IN
R F SaI
R,N F SaI
[P] F SaI
[R] F TIV
R F FGI
[R] F AIF
[P] F TIV
[N] F TIV
R F IN
R F IN
R F IN
R F AF
R F AF
[R] F AF
R F AF
R F FGI
R F FGI
R [G] AF
N F AF
N F TI
[N] G AF
[R] F BGI
R F AF
N F TIF
[N] F TIF
[P] F TIF
[N] F TIF
N F TIF
N F TI
N F TI
[R] F TI
R F TI
N F TI
R F SaI
R F SaI
N F SaI
[N] F SaI
[N] F SaI
[N] F SaI
N F SaI
N F SaI
R F TI
R F FGI
N F SaI
R F AIF
R F AIF
R F AIF
R F AIF
R F AIF
R F AIF
[N] A SwI
N A SwI
N A SwI
N F TI
N F FGI
N F FGI
P F FGI
N F FGI
N F FGI
P F FGI
N F FGI
N F FGI
N F FGI
N F FGI
N F AIF
R F AIF
R F TI
R F FGI
R F FGI
R F TI
[R] F FGI
R F FGI
R F TI
R F TI
R F TI
R F FGI
[R] F FGI
* Two taxa, Claudia’s Warbler Phylloscopus claudiae and Hartert’s Warbler P. goodsoni constitute a monophyletic group with Blyth’s Leaf-warbler P. reguloides, with which they were formerly treated as conspecific (Olsson et al., 2005). The Khao Yai-wintering taxon is tentatively listed as P. claudiae but the possibility that it might instead be P. goodsoni, recently recorded in N Laos by Fuchs et al. (2007) has not been excluded with certainty.
Table 2. List of additional bird species recorded in headquarters area of Khao Yai National Park.
Common Name Scientific Name

Forest/
Woodland

Grassland/Scrub Water-bodies
Blue-breasted Quail
Lesser Whistling-duck
Little Grebe
Asian Openbill
Black Stork
Yellow Bittern
Von Schrenck’s Bittern
Cinnamon Bittern
Black Bittern
Little Heron
Eastern Cattle Egret
Little Egret
Spot-billed Pelican
Little Cormorant
Oriental Darter
Common Kestrel
Oriental Hobby
Osprey
Black-shouldered Kite
Black-eared Kite
Himalayan Griffon
Short-toed Snake-eagle
Eastern Marsh-harrier
Pied Harrier
Chinese Sparrowhawk
Grey-faced Buzzard
Common Buzzard
Eastern Imperial Eagle
Booted Eagle
Black Eagle
Rufous-bellied Eagle
Changeable Hawk-eagle
Slaty-legged Crake
White-breasted Waterhen
Ruddy-breasted Crake
Common Moorthen
Masked Finfoot
Yellow-legged Buttonquail
Barred Buttonquail
Black-winged Stilt
Pacific Golden Plover
Grey-headed Lapwing
Red-wattled Lapwing
Pintail Snipe
Whimbrel
Common Sandpiper
Green Sandpiper
Wood Sandpiper
Spotted Redshank
Oriental Pratincole
White-winged Tern
Rock Pigeon
Pale-capped Pigeon
Oriental Turtle-dove
Orange-breasted Green- pigeon
Ashy-headed Green-pigeon
Pin-tailed Green-pigeon
White-bellied Green- pigeon
Red-breasted Parakeet
Moustached Hawk-cuckoo
Himalayan Cuckoo
Plaintive Cuckoo
Asian Koel
Lesser Coucal
Buffy Fish-owl
Blyth’s Frogmouth
Grey Nightjar
Large-tailed Nightjar
Himalayan Swiftlet
White-throated Needletail
Silver-backed Needletail
Fork-tailed Swift
House Swift
Indian Roller
Stork-billed Kingfisher
Ruddy Kingfisher
White-throated Kingfisher
Black-capped Kingfisher
Common Kingfisher
Blue-tailed Bee-eater
Blue-throated Bee-eater
Common Hoopoe
Lineated Barbet
Coppersmith Barbet
Eurasian Wryneck
Grey-capped Pygmy Woodpecker
Lesser Yellownape
Grey-headed Woodpecker
Blue-winged Pitta
White-browed Shrike- babbler
Chestnut-fronted Shrike- babbler
Slender-billed Oriole
Maroon Oriole
Ashy Woodswallow
Black Drongo
Crow-billed Drongo
Brown Shrike
Burmese Shrike
Olive-backed Sunbird
Crimson Sunbird
Golden-fronted Leafbird
Plain-backed Sparrow
Red-throated Pipit
Olive-backed Pipit
Blyth’s Pipit
Richard's Pipit
Paddyfield Pipit
Forest Wagtail
White Wagtail
Eastern Yellow Wagtail
Yellow-billed Grosbeak
Chestnut Bunting
Yellow-breasted Bunting
White-vented Myna
Common Myna
Chestnut-tailed Starling
White-shouldered Starling
Purple-backed Starling
Chinese Blackbird
Japanese Thrush
Dusky Thrush
Siberian Rubythroat
Rufous-tailed Robin
Blue Rock-thrush
Grey Bushchat
Eastern Stonechat
Pied Bushchat
Red-flanked Bluetail
Chinese Blue Flycatcher
Vivid Niltava
Yellow-rumped Flycatcher
Green-backed Flycatcher
Taiga Flycatcher
Ferruginous Flycatcher
Blue-and-white Flycatcher
Oriental Magpie-robin
Yellow-vented Bulbul
Red-whiskered Bulbul
Himalayan Black Bulbul
Asian House-martin
Common Sand-martin
Yellow-bellied Warbler
Sunda Bush-warbler
White-tailed Leaf-warbler
Chinese Leaf-warbler
Dusky Warbler
Buff-throated Warbler
Yellow-eyed Babbler
Japanese White-eye
Chestnut-capped Babbler
Black-browed Reed- warbler
Blunt-winged Warbler
Oriental Reed-warbler
Thick-billed Warbler
Lanceolated Warbler
Baikal Bush-warbler
Bright-capped Cisticola
Rufescent Prinia
Yellow-bellied Prinia
Plain Prinia
Coturnix chinensis
Dendrocygna javanica
Tachybaptus ruficollis
Anastomus oscitans
Ciconia nigra
Ixobrychus sinensis
Ixobrychus eurhythmus
Ixobrychus cinnamomeus
Dupetor flavicollis
Butorides striata
Bubulcus coromandus
Egretta garzetta
Pelecanus philippensis
Phalacrocorax niger
Anhinga melanogaster
Falco tinnunculus
Falco severus
Pandion haliaetus
Elanus caeruleus
Milvus lineatus
Gyps himalayensis
Circetus gallicus
Circus spilonotus
Circus melanoleucos
Accipiter soloensis
Butastur indicus
Buteo buteo
Aquila heliaca
Aquila pennata
Ictinaetus malayensis
Lophotriorchis kienerii
Nisaetus limnaeetus
Rallina eurizonoides
Amaurornis phoenicurus
Porzana fusca
Gallinula chloropus
Heliopais personatus
Turnix tanki
Turnix suscitator
Himantopus himantopus
Pluvialis fulva
Vanellus cinereus
Vanellus indicus
Gallinago stenura
Numenius phaeopus
Actitis hypoleucos
Tringa ochropus
Tringa glareola
Tringa erythropus
Glareola maldivarum
Chlidonias leucopterus
Columba livia
Columba punicea
Streptopelia chinensis
Treron bicinctus
Treron phayrei
Treron apicauda
Treron sieboldii
Psittacula alexandri
Hierococcyx vagans
Cuculus saturatus
Cacomantis merulinus
Eudynamys scolopaceus
Centropus bengalensis
Ketupa ketupu
Batrachostomus affinis
Caprimulgus jotaka
Caprimulgus macrurus
Aerodramus brevirostris
Hirundapus caudacutus
Hirundapus cochinchinensis
Apus pacificus
Apus affinis
Coracias benghalensis
Pelargopsis capensis
Halcyon coromanda
Halcyon smyrnensis
Halcyon pileata
Alcedo atthis
Merops philippinus
Merops viridis
Upupa epops
Megalaima lineata
Megalaima haemacephala
Jynx torquilla
Dendrocopos canicapillus
Picus chlorolophus
Picus canus
Pitta moluccensis
Pteruthius flaviscapis
Pteruthius aenobarbus
Oriolus tenuirostris
Oriolus traillii
Artamus fuscus
Dicrurus macrocercus
Dicrurus annectans
Lanius cristatus
Lanius collurioides
Cinnyris jugularis
Aethopyga siparaja
Chloropsis aurifrons
Passer flaveolus
Anthus cervinus
Anthus hodgsoni
Anthus godlewskii
Anthus richardi
Anthus rufulus
Dendronanthus indicus
Motacilla alba
Motacilla tschutschensis
Eophona migratoria
Emberiza rutila
Emberiza aureola
Acridotheres grandis
Acridotheres tristis
Sturnus malabaricus
Sturnus sinensis
Sturnus sturninus
Turdus mandarinus
Turdus cardis
Turdus eunomus
Luscinia calliope
Luscinia sibilans
Monticola solitarius
Saxicola ferreus
Saxicola maurus
Saxicola caprata
Tarsiger cyanurus
Cyornis glaucicomans
Niltava vivida
Ficedula zanthopygia
Ficedula elisae
Ficedula albicilla
Muscicapa ferruginea
Cyanoptila cyanomelana
Copsychus saularis
Pycnonotus goiavier
Pycnonotus jocosus
Hypsipetes leucocephalus
Delichon dasypus
Riparia riparia
Abroscopus superciliaris
Cetia vulcania
Phylloscopus ogilviegranti
Phylloscopus yunnanensis
Phylloscopus fuscatus
Phylloscopus subaffinis
Chrysomma sinense
Zosterops japonicus
Timalia pileata
Acrocephalus bistrigiceps
Acrocephalus concinens
Acrocephalus orientalis
Acrocephalus aedon
Locustella lanceolata
Bradypterus davidi
Cisticola exilis
Prinia rufescens
Prinia flaviventris
Prinia inornata
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HABITAT USE
Most species that occur on MFDP are inhabitants of the edge or interior of evergreen forest. Some are shared with either deciduous forest or secondary growth. Many have a broad altitudinal range, but there are no exclusively montane species. Some rarer MFDP species are larger birds that naturally occur at low density (Table 1). Great Slaty Woodpecker Mulleripicus pulverulentus, for example, is scarce in the Hq area of the park and is mostly found in association with lowland, old-growth forests (Lammertink et al., 2009). Mountain Hawk-eagle Nisaetus nipalensis and Malaysian Night-heron Gorsachius melanolophus may be similarly scarce. The former is assumed to require large territories, while the latter is secretive in addition to its apparent scarcity and seems primarily to inhabit lowlands (Lekagul & Round, 1991; Thewlis et al., 1998). Scaly-crowned Babbler Malacopteron cinereum may be representative of lowland species that approach their upper altitudinal limit on the plot. It appears to be patchy and thinly distributed in the Hq area of Khao Yai.

Some others have a special association with a particular habitat (e.g. Pin-tailed Parrotfinch Erythrura prasina is a semi-nomadic specialist in seeding bamboo (Lekagul & Round, 1991; Wells, 2007). The scarcity of some others is harder to explain. Velvet-fronted Nuthatch Sitta frontalis occurs across a wide range of elevations, from lowlands to the montane zone. Its extreme scarcity on the plot (and, incidentally elsewhere in the Hq area) may be due to a preference for slightly more open forest than is found around the interior of Khao Yai. The species evidently bred on or near the plot in 2005, when there were four sightings during March–August, one of which involved two adults and two fledglings. None was recorded in subsequent years, however. At least two rare, resident, plot-recorded species, Blue-eared Kingfisher Alcedo meninting and Slaty-backed Forktail Enicurus schistaceus, are riparian species associated mostly with larger streams that move into MFDP (presumably from the Lamtakhong River) in mid- to late wet season as the few small streamlets that traverse the plot become swollen.

Five resident species recorded infrequently on MFDP but typical of more open habitats include Greater Coucal Centropus sinensis, Spotted Dove Streptopelia chinensis, Racket-tailed Treepie Crypsirina temia, Common Tailorbird Orthotomus sutorius and White-rumped Munia Lonchura striata; Table 1). These are either characteristic of, or regularly enter, open, deciduous woodland and bamboo (Lekagul & Round, 1991) and all but Racket-tailed Treepie are regularly present in edge habitats around the headquarters. There were several records of Racket-tailed Treepie, a bird of lowland deciduous habitats and scrub, often close to standing water (Lekagul & Round, 1991; Robson, 2002) in both the headquarters area of the park and on MFDP during 2004–2005 but the species evidently did not establish itself and has since seemingly since vanished from the park headquarters area. A sixth species, Plain Flowerpecker Dicaeum minullum, may also be placed with this group, in that it is regularly present in disturbed, more open woodland habitat around park headquarters. It probably tends to be associated with taller, better quality forest than the other members of this group, occurring elsewhere across a wide altitudinal range to an elevation of 1700 m (Lekagul & Round, 1991).

Most other species that occur infrequently on MFDP are migrants or non-breeding visitors found in a range of habitats. Chinese Pond-heron Ardeola bacchus is a waterbird that occasionally enters closed forest along streams.

SEASONAL STATUS
Assigning seasonal status to birds is not always straightforward. Many species are polytypic, with both discrete resident and migrant subspecies or populations occurring in the country, and it is not always clear which taxon or population occurs in the park Hq area or on MFDP. Only 114 of 169 MFDP-recorded species (67.5%) are considered residents and for only approximately half of these are there confirmed breeding records on MFDP. Most of the remainder are non-breeding winter visitors. A number of species that are widespread in forest habitats elsewhere in the country (Large Hawk-cuckoo Hierococcyx sparverioides, Black-winged Cuckooshrike Coracina melaschistos, Ashy Drongo Dicrurus leucophaeus, Lesser Racket-tailed Drongo D. remifer, Hair-crested Drongo D. hottentottus, Verditer Flycatcher Eumyias thalassinus and Grey-headed Flycatcher Culicicapa ceylonensis) are only known on the plot and in the Hq area of Khao Yai as non-breeding visitors. Hair-crested Drongos breed in lower elevation, deciduous forest around the park boundaries, but the great abundance of this species in evergreen forest, both on the plot and elsewhere around the park Hq during the winter months, lends support to the idea that these are more likely migrant D. h. brevirostris from countries to the north, rather than resident or short-distance migrant, up-slope dispersant D. h. hottentottus. The absence of resident Ashy Drongo from the plot is more puzzling since residents breed both in lower elevation deciduous forest around the eastern park margins, and in hill slope and montane evergreen forests elsewhere (Lekagul & Round, 1991; Lynam et al., 2006). Lesser Racket-tailed Drongo is mainly montane and possibly resident in Khao Yai on the small areas of >1000 m habitat that lie 8–10 km S of the plot. (Lynam et al., 2006). But both the Lesser Racket-tailed Drongos and Ashy Drongos that occur on MFDP during winter months are similarly more likely to be migrants from further afield than local dispersants.

The absence of resident populations of some of these species may be due to a variety of factors. The relative homogeneity of the closed evergreen forest habitat; the elevation of the plot, above the upper elevational limits of some species (itself presumably a consequence of preference for more open, deciduous or forest mosaic habitats) and historical (biogeographical) factors may all contribute. The Khao Yai Hq area is at too great an elevation to support the full complement of lowland (deciduous or forest mosaic) species, yet lacks any compensating diversity of either montane or other evergreen forest species (Lynam et al., 2006). The same authors argued that the relatively depauperate nature of the resident evergreen forest avifauna might stem from Khao Yai constituting a relatively isolated block of evergreen forest which during Pleistocene interpluvial periods, would have been smaller than at present, surrounded by deciduous habitats, with even fewer connections to other evergreen isolates elsewhere in Thailand: to the north, south and east.

In a study of mixed species foraging flocks on MFDP, Nimnuan et al. (2004) found 58 species (residents and migrants) that regularly participated. Although residents contributed 74% of individuals found in flocks, the sightings were biased towards resident species as the survey covered only the months May–October, when most migrants were absent. Even so, the most numerous birds counted (136 of 956 individuals, 14 % of all sightings) were one or two species of (migrant) leaf-warbler. If migrant species are excluded from the analysis, the relatively depauperate nature of the MFDP avifauna is immediately apparent. Just 11 species of resident bird contributed 67% of sightings of birds in flocks. The two predominant arboreal species, White-bellied Erpornis Erpornis zantholeuca (69 sightings) and Black-naped Monarch Hypothymis azurea (53 sightings) together contributed 17% of all individuals observed. The single most abundant understorey-middle storey species on MFDP is the Puff-throated Bulbul Alophoixus pallidus (3.4 individuals/ha, Gale et al., 2009) which is chiefly associated in territorial groups and contributed few (1.5%) mixed flock sightings.

FORAGING GUILDS
The largest guild among MFDP-recorded species was that of foliage-gleaning insectivores (46 species, 35.5%) followed by terrestrial insectivores and insectivore-faunivores (23 species in total); sallying insectivores (19 species), both diurnal and nocturnal raptorial birds (18 species), arboreal omnivores (16 species, mostly hornbills, barbets and bulbuls); arboreal frugivores (including granivores, not listed separately; 14 species). If granivores are excluded from the arboreal frugivore assemblage, this leaves only four pigeons, four flowerpeckers and two mynas. Though eight species of bark-gleaning insectivores (mostly woodpeckers) are listed for the plot, only four (Greater Flameback Chrysocolaptes lucidus, Laced Woodpecker Picus vittatus, Greater Yellownape Chrysophlegma flavinucha and Heart-spotted Woodpecker Hemicircus canente), listed in declining order of abundance (Round et al., 2006), are frequent.

Assigning species to guilds is necessarily imprecise. The seeds of small fruits were detected in the faeces of many ostensibly insectivorous species that were handled, including Abbott’s Babbler Malacocincla abbotti, scimitar-babblers Pomatorhinus spp. and all three laughingthrushes (Dryonastes chinensis and Garrulax spp.). Both Laced Woodpecker and Greater Flameback were seen eating fruit. Among the commoner resident insectivores, only Hill Blue Flycatcher Cyornis banyumas has not yet been observed to take fruit. Other true flycatchers (Verditer Flycatcher Eumayias thalassinus, Taiga Flycatcher Ficedula albicilla) and “flycatcher guild” sallying insectivores such as Black-naped Monarch are likely to take fruits on occasion. While most neotropical trogons take a considerable proportion of fruit in their diet (Pizo, 2007; Remsen et al., 1993; Riehl & Adelson, 2008), the Asian species are primarily foliage-gleaning insectivores (Ali & Ripley,?,? 1983? ;Johnsgard,?,? 2000; Steward,? 2010; Wells, 1999). Nonetheless both Red-headed Trogon Harpactes erythrocephalus and Orange-breasted Trogon H. oreskios at Khao Yai have now been observed to take a small proportion of fruits.

In comparison with Sundaic lowland forest bird communities such as those at (e.g.) Pasoh and at Kuala Lompat, Malaysia (Francis & Wells, 2003; Wong, 1986), Khao Yai appears to have more generalist omnivores and fewer specialist insectivores or frugivores.

DISCUSSION
A further 160 species (loosely divided into 60 forest or woodland inhabiting species, 70 grassland or scrub-associated species and 30 waterbirds and riparian species) have been recorded in the Hq area without being either seen or heard on MFDP (Table 2: Lynam et al., 2006; Pobprasert et al., 2008). The occurrence of so many species elsewhere in the Hq area relative to the number on MFDP can be attributed to the much larger area, and greater range of habitats, around the park Hq as a whole compared with the closed forest habitat of the plot. These additional habitats encompass grassland (both tall grass and mown grass or lawn), scrub, forest edge, roadside verge, parkland, large streams with associated riparian habitats, reservoirs, human habitations, power lines and other installations which provide nesting of foraging niches for a few species.

The 60 forest-inhabiting species comprise 23 resident/presumed residents and 37 migrants/non-breeding visitors, all of which may have the potential to occur on MFDP. At least two of the resident species, Lesser Yellownape Picus chlorolophus and Grey-capped Pygmy Woodpecker Dendrocopos canicapillus (the latter reported as “absent from the Hq area” by Lynam et al., 2006, but presence since noted in at least one season: author), seem to share the habitat preference of Velvet-fronted Nuthatch in that they occur across a wide range of elevations, often in more open forest. However they also occur in tall, mature, closed-canopy, lowland forest. The scarce lowland resident Moustached Hawk-cuckoo Cuculus vagans, two passage migrants (Ruddy Kingfisher Halcyon coromanda and Chinese Sparrowhawk Accipiter soloensis) and the scarce winter visitor Rufous-tailed Robin Luscinia sibilans have all been seen on the Mo Singto Nature Trail or elsewhere in the vicinity of the plot margins, and are therefore almost certain to occur on MFDP occasionally. A small number of birds (e.g. White-tailed Leaf-warbler Phylloscopus ogilviegranti, White-browed Shrike-babbler Pteruthius flaviscapis and Chestnut-fronted Shrike-babbler P. aenobarbus) are montane residents, and could conceivably disperse downslope to the elevation of the plot although the park populations of these species are relatively small, due to the limited areas of montane habitat and therefore the chances of detecting them may be slight.

The occurrence of some others, mainly lowland deciduous woodland, species (e.g. Red-breasted Parakeet Psittacula alexandri, Lineated Barbet Megalaima lineata and Golden-fronted Leafbird Chloropsis aurifrons) that occur chiefly around the margins of the headquarters plateau, at the deciduous-evergreen ecotone, is unlikely at present but could be plausible, especially if warming temperatures cause significant drying of the moist forest habitat.

The apparent absence from MDFP of some forest birds may simply be due to the difficulty of observing them in the canopy of enclosed forest—most records of (e.g.) migrant flycatchers are from forest edge, from the roads or other open areas.

The 70 species of grassland-scrub inhabiting birds recorded elsewhere in the Khao Yai Hq area may be divided into 39 non-breeding winter visitors and 31 residents. Their absence from MFDP is noteworthy given the relative proximity of such open habitats. Many of these open-country species (e.g. Red-wattled Lapwing Vanellus indicus) have colonized the headquarters area of the park within the past 20 years or so. Common Myna Acridotheres tristis, first recorded c. 2000, and White-vented Myna A. grandis, found as recently as 2006 (PDR own data) are even more recent arrivals. Natural habitat perturbation (e.g. major tree-falls caused by whirlwinds, affecting areas up to 0.3 ha) has occurred just off-plot at Mo Singto, and such infrequent natural events could in future create transient conditions on MFDP that might permit the temporary occurrence of smaller open country birds. Additionally most migrant species, including inhabitants of open country, have the potential to stray, and occur in unusual situations, and could therefore occur on the plot from time to time. Most waterbirds, on the other hand, have no likely prospects of being found on the plot due to the lack of significant water bodies (with the possible exception of riparian species such as kingfishers that may move up flood-swollen smaller tributaries in the wet season).
DISCUSSION
While the avifauna of the Mo Singto Plot is largely composed of birds typical of the interior of evergreen forest, that around the larger headquarters area of Khao Yai National Park contains a wider array. In addition to evergreen forest species (including a very few montane species) are a few species more typical of deciduous forest habitats and bamboo; those that inhabit grassland and scrub, and waterbirds. Even so, some of these species occur at low densities or are distributed patchily around the headquarters area, or are rare or occasional transient visitors.

What Future Changes Might We Expect?
Some turnover of species is a natural process which might be expected even without any human disturbance. Since Khao Yai was established as a park in 1962 the most obvious trends (inferred or observed) have been decreased direct persecution of birds and increased use by tourists. There has been a gradual increase in infrastructure (roads, buildings and other facilities) but impact on forest appears to have been slight. So far as known, no species has been lost since avifaunal recording started.

Habitat succession
Grasslands around the park Hq are derived from former cultivated fields and have been variously maintained by either burning or mowing as a means to maintain open grassy areas for ungulates (especially Sambar Rusa unicolor). While the respective merits of the two grassland management regimes (burning and mowing) have not been fully assessed it is nonetheless undeniable that many areas of former grassland around the park headquarters are gradually reverting to scrub and secondary forest, and this will undoubtedly cause some grassland-inhabiting species recorded in the park Hq area to decline or even disappear. Blue-breasted Quail Coturnix chinensis (already rare in the park), buttonquails Turnix spp., Red-whiskered Bulbul Pycnonotus jocosus, Bright-capped Cisticola Cisticola exilis, prinias Prinia spp., and grassland-inhabiting migrants such as Blunt-winged Warbler Acrocephalus concinens are representatives of this group. None of these species has been recorded on MFDP. The loss of grassland will impact on the overall species richness of the park Hq area without adding any further forest-living species.

Human persecution
Species that were formerly hunted (e.g. pheasants Lophura spp. and Red Junglefowl Gallus gallus; pigeons, especially Mountain Imperial-pigeon Ducula badia) would probably have increased in the first decade or more after the park’s establishment and are now more or less common. Assuming that populations of these have probably reached carrying capacity, further population increase would seem unlikely (though some may have become more easily detectable due to behavioural habituation).

Illegal forest product collecting continues (Brockelman et al., 2011) and has the potential to pose a possible threat should any bird species be targeted by poachers. The communally roosting Red-whiskered Bulbul is known to be widely targeted elsewhere by commercial bird trappers supplying the cagebird trade, and numbers in the park Hq area, seemingly already diminishing, possibly due to habitat change, could be potentially at risk from poachers. Hornbill chicks, widely targeted elsewhere (and certainly at risk in peripheral regions of Khao Yai), are not known to have been poached in the immediate Hq area. Evidence has emerged of a trade in owl carcasses from Malaysia, (apparently destined for the restaurants in China; Shepherd & Shepherd, 2009) while owls are much persecuted in Laos, and frequently seen in captivity (Duckworth et al., 1999). They could therefore also be at elevated risk in peripheral areas of Khao Yai and other Thai protected areas.

Increased recreational use
The main direct human uses are tourism-related. In general there has been a great increase in vehicular traffic, and also foot traffic on park trails that approach within 100–200 m of the plot. There may be occasional cases where nests are abandoned due to inadvertent disturbance, or birds killed by vehicular collision, but because both roads and trails affect relatively limited areas, the impact is likely to be small. Road construction may well have contributed to the colonisation of the park Hq by open country species, however, since roads provide a narrow ribbon of edge habitat connecting the park margins with the park interior.

Canalization and concreting of small sections of riverbank, and the expansion of tourist facilities along waterways, could impact some riparian species, including bank-nesters such as kingfishers and forktails in the heavily used and developed Hq area.

A possible indirect effect of increased recreational use on the avifauna could be increased usage of forest around park Hq by the generalist omnivore Northern Pig-tailed Macaque Macaca leonina. Formerly either shy or scarce, macaques have learnt to utilize garbage and (illegal) hand-outs from tourists, and, now habituated, routinely beg for food along roadsides. They have become much more detectable in the past two decades, suggesting the possibility that their total population has increased. Macaques are the most frequently recorded predator of bird nests on MFDP (Pierce & Pobprasert, in prep.). If presently observed levels of nest predation by macaques are higher than formerly this might have the potential to induce changes in the productivity of the resident avifauna.

Climate change
Round & Gale (2008) postulated that changes in the relative abundance of two pheasant species in the park headquarters area were due to rising temperatures that favour species characteristic of lowland, semi-evergreen forest at the expense of those that favour moister, montane or submontane conditions. Formerly only Silver Pheasant Lophura nycthemera was found on MFDP but in the past 15–20 years sightings of this species have been overtaken by lowland-inhabiting Siamese Fireback L. diardi. Other species that favour moist, sub-montane and montane, evergreen forest that are at present relatively frequent on MFDP, but that could decline, include Red-headed Trogon, the broadbills Serilophus lunatus and Psarisomus dalhousiae, and possibly Common Green Magpie Cissa chinensis, Hill Blue Flycatcher, and Fire-breasted Flowerpecker Dicaeum ignipectus. Corresponding lower elevation species that might increase in abundance include Orange-breasted Trogon, Banded Broadbill Eurylaimus javanicus, Racket-tailed Treepie, Hainan Blue Flycatcher Cyornis hainanus and Plain Flowerpecker. As already indicated, other species at present more typical of deciduous woodland and edge habitats might colonise the plot if rising temperatures induce significant change in the vegetation towards a semi-evergreen or deciduous facies.

Migrant species are at elevated risk from climate change (Coppppack & Both, 2002; Butler, 2003). Migrants have evolved migration strategies in response to a narrow range of environmental conditions on their breeding and wintering areas, each of which is affected by climate-change to a differing extent. While some tropical or sub-tropical wintering species may have adjusted their arrival times on their Palearctic breeding grounds in synchrony with earlier warm temperatures, other species have failed to do so. In some, this has led to loss of synchrony between an earlier-peaking food supply and production of young (which has not advanced as correspondingly early) reducing breeding success and survival of young (Both et al., 2006). Changing weather patterns have likewise impacted the wintering areas of migrant birds. The winter survival of many western Palearctic breeding, Afrotropical wintering, migrants has been adversely affected by reduced rainfall and increased desertification in north equatorial Africa (Peach et al., 1991; Baillie & Peach, 1992; Sanderson et al., 2006).

There is much less information on the Eastern Palaearctic-Indo-Malayan migration system. However, the preponderance of migrant birds in Khao Yai, both Palaearctic and Sino-Himalayan species, offers significant opportunity for detailed studies of their ecology and survival.
FUTURE RESEARCH
This paper is largely descriptive. Continued monitoring of avifaunal change on the Mo Singto Plot and elsewhere in the park headquarters is a priority for the future. Besides maintaining the inventory and documenting turnover of species, this should include monitoring of annual population levels of both resident and migrant bird species, through continued use of territory mapping combined with distance sampling or other detectability-based estimates. Preliminary data on density and biomass (Round et al., 2006) is available for a wider range of species than treated in Gale et al. (2009) but needs revising and updating in the light of the latter’s caveats on interpretation. Monitoring of bird population levels combined with detailed studies that yield information on (e.g.) productivity and habitat use by birds, together with detailed monitoring of forest dynamics may help understand the implications of climate-related environmental change for biodiversity conservation.
ACKNOWLEDGEMENTS
We thank Warren Brockelman, Will Duckworth and James Steward for commenting on drafts of this paper. We are grateful to the Department of National Parks, Wildlife and Plants Conservation for facilitating our work in Khao Yai. We thank the Biodiversity Research and Training Programme (BRT) for supporting fieldwork during 2003–2006. Philip Round is also supported by The Wetland Trust.

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Kindly submitted by:

Philip D. Round, Department of Biology, Faculty of Science, Mahidol University, Rama 6 Road, Bangkok 10400, Thailand. E-mail: frpdr@mahidol.ac.th

Andrew J. Pierce, Conservation Ecology Program, School of Bioresources & Technology, King Mongkut’s University of Technology Thonburi, 83 Moo. 8 Thakham, Bangkhuntien, Bangkok 10150, Thailand. E-mail: george.and@kmutt.ac.t

George A. Gale, Conservation Ecology Program, School of Bioresources & Technology, King Mongkut’s University of Technology Thonburi, 83 Moo. 8 Thakham, Bangkhuntien, Bangkok 10150, Thailand. E-mail: george.and@kmutt.ac.t

Wangworn Sankamethawee, present address: Walai Rukhavej Botanical Research Institute, Mahasarakham University, Mahasarakham 44150, Thailand
   
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