io s m b THE NATURAL HISTORY MUSEUM 2 1 AUG 2012 PURCHASED TRING LIBRARY ' *mZm vi$§r mm 1.5P" ylM ;/ «y Forktail 28 (2012) f OBC Council Stephen Browne (Co-Chairman, Conservation Committee) David Buckingham (Co-Chairman, Conservation Committee) Mike Edgecombe (Promotions) John Gregory (Treasurer) Tim Loseby (Art and Photographic Editor) Steve Rowland Tony Sawbridge Graeme Spinks Brian Sykes (Chairman) Margaret Sykes (Membership Secretary / Secretary) Jo Thomas (Promotions) Richard Thomas (Internet) Publications Committee Stuart Butchart, Nigel Collar, Tim Loseby, Rene Pop, Nigel Redman, Simon Roddis, Brian Sykes, Richard Thomas Conservation Committee Nick Brickie, Stephen Browne, Dave Buckingham, Francis Buner, Mike Crosby, John Fellowes, Jim Wardill, Simon Wotton The Oriental Bird Club has been established for ornithologists throughout the world, both amateur and professional, who share a common interest in the region's birds and wish to assist in their conservation. 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Cover picture: Camiguin Flawk Owl, new species, Camiguin Sur, Philippines, 8 June 201 1 by Rob Hutchinson ISSN 0950-1746 © Oriental Bird Club 2012 FORKTAIL Number 28, 2012 Acting Editor N. J. Collar Associate Editors Jeremy Bird, David Buckingham, Stuart Butchart, Will Duckworth, Eben Goodale, John Pilgrim and Jack Tordoff THE NATURAL HISTORY MUSEUM 2 I AUG 2012 PURCHASED TRING LIBRARY CONTENTS P. C. RASMUSSEN, D. N. S. ALLEN, N. J. COLLAR, B. DeMEULEMEESTER, R. 0. HUTCHINSON, P. G. C. JAKOSALEM, R. S. KENNEDY, F. R. LAMBERT & L. M. PAGUNTALAN Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex . 1 M. MONIRUL H. KHAN & NASIM AZIZ Bird species diversity in five protected areas of Bangladesh . 21 MARY ROSE C. POSA & DAVID ALEXANDER MARQUES Peat swamp forest birds of the Tuanan research station, Central Kalimantan, Indonesia, with notes on habitat specialists . 29 MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE An annotated checklist of birds and conservation issues in Salkhala Game Reserve, an isolated Important Bird Area in Azad Kashmir, Pakistan . 38 SIMON P. MAHOOD & JONATHAN C. EAMES A review of the status of Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis . 44 TILLTOPFER Seasonal changes in plumage coloration of Orange Bullfinches Pyrrhula aurantiaca . 49 D. KHAMCHA & G. A. GALE The use of tree-fall gaps by a forest interior avian frugivore in a tropical evergreen forest . 53 SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M. THOMPSON Status and distribution of the endangered Baer's Pochard Aythya baeri in Bangladesh . 57 ZHIXIN ZHOU, YUE SUN, LU DONG, CANWEI XIA, HUW LLOYD & YANYUN ZHANG Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area . 62 DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU Sex bias in a wintering population of Dunlin Calidris alpina in central Taiwan . 67 JEREMY P. BIRD, BERRY MULLIGAN, ROURS VANN, PHILIP D, ROUND & JAMES J. GILROY Habitat associations of the Manchurian Reed Warbler Acrocephalus tangorum wintering on the Tonle Sap floodplain and an evaluation of its conservation status . 71 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE The avifauna of Alor and Pantar, Lesser Sundas, Indonesia . 77 H. L. WRIGHT, N. J. COLLAR, I. R. LAKE, BOU VORSAK & P. M. DOLMAN Foraging ecology of sympatric White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigantea in northern Cambodia . 93 LE MANH HUNG, MARK B. ROBBINS, NATHAN H. RICE & ERICK A. GARCIA-TREJO Survey of the avifauna at Muong Nhe Nature Reserve, Dien Bien province, Vietnam . 101 T. E. MARTIN, D. J. KELLY, N. T. KEOGH, D. HERIYADI, H. A. SINGER & G. A. BLACKBURN The avifauna of Lambusango Forest Reserve, Buton Island, south-east Sulawesi, with additional sightings from southern Buton . 107 PHILIP D. ROUND, JOHN M. HOBDAY, RUNGSRIT KANJANAVANIT & JAMES S. STEWARD A nesting pair of Gecinulus woodpeckers in a likely zone of intergradation between Pale-headed Woodpecker G. grantia and Bamboo Woodpecker G. viridis . 1 1 3 PAUL J. LEADER & GEOFF J. CAREY Zappey's Flycatcher Cyanoptila cumatilis, a forgotten Chinese breeding endemic . 121 ALAIN HENNACHE, SIMON P. MAHOOD, JONATHAN C. EAMES & ETTORE RANDI Lophura hatinhensis is an invalid taxon . 129 Forktail 28 (2012) Short Notes SIMON P. MAHOOD, DAVID P. EDWARDS & FELICITY A. EDWARDS Bar-winged Wren Babbler Spelaeornis troglodytoides: a first record for Vietnam, with speculation for 1 7 further avifaunal additions . 1 36 ZHU LEI, ZHANG JUN, QIU JING, WEI QIAN, DONG LEI & SUN YUE-HUA Long-tailed Duck Clangula hyemalis and Red-breasted Goose Branta ruficollis : two new birds for Sichuan, with a review of their distribution in China . 138 DELIP K. DAS Western Hoolock Hoolock hoolock preying on chicks of Greater Racket-tailed Drongo Dicrurus paradiseus in Lawachara National Park, Bangladesh . 142 FRANK E. RHEINDT & JAMES A. EATON Notes on the life-history and taxonomy of Muscicapa dauurica umbrosa, an overlooked Bornean canopy bird . 144 YANG LIU & SERGEY PYZHJANOV Apparent inter- and intraspecific brood-parasitism in a nest of Tufted Duck Aythya fuligula . 147 DAVID J. KELLY & NICOLA M. MARPLES Annual survival rate and mean life-span of Lemon-bellied White-eyes Zosterops chloris flavissimus on Kaledupa island, Wakatobi, south-east Sulawesi, Indonesia . 148 CHANG-YONG CHOI & HYUN-YOUNG NAM Migrating dragonflies: famine relief for resident Peregrine Falcons Falco peregrinus on islands . ) . 1 49 DONG-WON KIM, CHANG-WAN KANG, HWA-JUNG KIM, YOUNG-SOO KWON & JIN-YOUNG PARK Breeding of the Japanese Murrelet Synthliboramphus wumizusume in South Korea . 151 NORIMASA SUGITA, TOSHITAKA N. SUZUKI, CRAIG A. BARNETT & KEISUKE UEDA An intraspecific adult killing in female Japanese Great Tits Parus major minor . 1 53 VLADIMIR DINETS Nesting Fork-tailed Swifts Apus pacificus in north-eastern Vietnam . 155 ADAM C. STEIN & GALINA NOSACHENKO Notes on the 2009 autumn crane migration in Muraviovka zakaznik, Amur oblast, Russian Federation . 1 56 JAMES W. BURNHAM & ERIC M. WOOD Woolly-necked Stork Ciconia episcopus at Napahai wetland, Yunnan, China . 1 58 TIM ROBINSON First records of Javan Munia Lonchura leucogastroides in Peninsular Malaysia . 1 59 GERARD E. RYAN Brahminy Kites Haliastur indus fishing with Irrawaddy dolphins Orcaella brevirostris in the Mekong River . 161 MOHAMMAD IRHAM, E. MEIJAARD & S. (BAS) van BALEN New information on the distribution of White-fronted Microhierax latifrons and Black-thighed Falconets M. fringillarius in Kalimantan, Indonesia . 162 WEN-LOUNG LIN, SI-MIN LIN & HUI-YUN TSENG Breeding ecology of the Northern Boobook Ninox japonica totogo in central Taiwan . 164 SIMON P. MAHOOD & JAMES A. EATON The vocalisations of Red-collared Woodpecker Picus rabieri . 1 67 JAMES A. FITZSIMONS, ERIK MEIJAARD, IWAN HUNOWU, DEWI PRAWIRADILAGA, JANELLE L. THOMAS & JOHNY S.TASIRIN Diet of the Speckled Boobook Ninox punctulata in north Sulawesi, Indonesia . 169 TOSHITAKA N. SUZUKI Mobbing to death of a Japanese Long-eared Bat Plecotus sacrimontis by two species of tit . 171 Guidelines for contributors inside back cover Forktail 28 (2012) Editorial notes The re-styling of Forktail last year was designed in large part to allow for a larger number of contributions to the journal, and the upshot was that the issue for 201 1 appeared rather slim by comparison with those of the previous five years. In fact with 1 1 full papers and 21 short notes it was an average size issue, the mean number of full papers and short notes for the years 2000-2010 being 1 1.7 and 16.8 respectively. This year, somewhat by contrast, the journal carries 18 papers and 19 short notes, and the space gains of last year's makeover have proved invaluable. The trend seems to be universal: all ornithological journals are growing in size in response to the increasing quantity of material that is competing for a place in them. In a club as small and as prudently managed as OBC, however, the budgetary constraints on publications are significant and I express my gratitude to OBC Council for its approval of the considerable costs that this relatively large and colour-rich issue has incurred. The other costs that the journal incurs are on the time of its acting and associate editors, all of whose positions are strictly honorary. Assuming that contributions to Forktail will continue to increase, the burden of editorship will need further redistribution, not only in terms of more or new human shoulders but also in relation to the annual editorial time-frame. Having only recently established 31 March as the deadline for the receipt of material for publication in the same year, we now propose to bring this deadline forward to 14 February so as to allow an extra six weeks for the various necessary processes of moving from manuscript to publication — a six- month period, given the publication date of mid-August which Forktail now rigidly observes. I once more thank Jez Bird, Dave Buckingham, Stuart Butchart, Will Duckworth, Eben Goodale, John Pilgrim and Jack Tordoff for their help through the year, Brian and Margaret Sykes for their unswerving support, and Peter Creed for his super-fast, efficient and stylish layout of the end product. N. J. Collar Forktail 28 (2012) Plate. Species in the Philippine Hawk Owl Ninox philippensis complex: (1) Mindoro Hawk Owl N. mindorensis (a, dark individual: b, pale individual); (2) Mindanao Hawk Owl N. spilocephala; (3) Luzon Hawk Owl N. philippensis', (4) Cebu Hawk Owl, new species; (5) Romblon Hawk Owl Ninox spilonota, new subspecies from Tablas; (6) Romblon Hawk Owl, nominotypical from Sibuyan; (7) Camiguin Hawk Owl, new species; (8) Sulu Hawk Owl Ninox reyi. Original painting by John Gale. FORKTAIL 28 (2012): 1-20 Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex P. C. RASMUSSEN, D. N. S. ALLEN, N. J. COLLAR, B. DeMEULEMEESTER, R. 0. HUTCHINSON, P. G. C. JAKOSALEM, R. S. KENNEDY, F. R. LAMBERT & L. M. PAGUNTALAN We show, based on morphology and especially vocalisations, that the Philippine Hawk Owl Ninox philippensis requires treatment as seven allopatric species and at least one additional subspecies. Morphological distinctions between three groups of taxa are striking, and although taxa within one major group are relatively similar in plumage they vary rather consistently in size and proportions. It has not been possible until now to resolve the species limits in this complex due mainly to the lack of sound recordings of key taxa, a problem now rectified. Vocalisations differ significantly between all seven species, the limits of which are incongruent with all previous taxonomies. Taxa from Mindoro (mindorensis), Mindanao [spilocephala), Camiguin Sur (named herein), and the Sulu Islands ( reyi ) exhibit especially great vocal differences from all other taxa along with smaller but consistent differences in plumage and morphometries. Although specimens have been in museum collections for many years, two of these species and one subspecies have heretofore remained undescribed, and we formally name these taxa for science. The recommended species-level treatment and English names of the N. philippensis complex are: Luzon Hawk Owl N. philippensis; Mindanao Hawk Owl N. spilocephala; Mindoro Hawk Owl N. mindorensis; Sulu Hawk Owl Ninox reyi; Romblon Hawk Owl N. spilonota; Camiguin Hawk Owl new species; and Cebu Hawk Owl new species. INTRODUCTION During the heyday of ornithological discovery in the Philippines (approximately 1850-1910), as many as seven species were recognised in the islands’ endemic hawk owl complex (e.g. McGregor 1909-1910). Since 1945, however, it has been treated as a single polytypic species, Philippine Hawk Owl Ninox philippensis (Delacour & Mayr 1945), recently considered to contain eight subspecies (Dickinson 2003). These subspecies group into three ’distinctive plumage types: one with all-streaked underparts and plain crown ( philippensis , proximo., ticaoensis and centralis of Luzon and many other islands); one with mottled or barred breast, streaked lower underparts, and spotted crown (. spilocephala of Mindanao); and one with barred to nearly plain underparts (the ‘unstreaked’ group: mindorensis of Mindoro, spilonota of several small islands, and reyi of the Sulu Islands) (Collar & Rasmussen 1998; for main islands and distribution of taxa see Figure 1). This last group of three described races is highly disjunct, and the distribution of the race spilonota as currently defined in particular is biogeographically peculiar because the Cebu population is surrounded by members of th e philippensis group, and the Camiguin Sur population is from a small island off northern Mindanao and well away from other taxa in spilonota. Within the unstreaked group there are marked differences in size and proportions, as well as more subtle distinctions in plumage. However, it has not been possible to resolve relationships between these racial groups based on morphology owing to intra-island plumage variation and, for some taxa, small sample size. Sound recordings until recently were available only for taxa from a few islands, and most were incomplete and of poor quality. Given the complexity of the vocal repertoire in this group, the small sample of recordings previously available precluded further analysis of species limits. The sample was, however, adequate to establish that Mindoro mindorensis differs profoundly in vocalisations from Luzon nominotypical philippensis, prompting the separation of Mindoro Hawk Owl N. mindorensis (Konig etal. 1999). However, this cursory treatment left other unstreaked forms united with N. philippensis, although in the absence of acoustic data it seemed plausible that they could separate out as a single species for which the name with priority was Ninox reyi (Collar & Rasmussen 1 998). Within this species, however, there were clearly multiple undescribed taxa based on morphology, although the populations on Cebu and Tablas were feared possibly extinct (Collar & Rasmussen 1998, Collar etal. 1999). There the matter unsatisfactorily rested, in the absence of adequate or sometimes any vocal evidence from all the taxa, including the four insular populations comprising the form spilonota. However, recent fieldwork has resulted in nearly complete sampling with extensive, high-quality recordings of the vocal repertoire of the key island populations of Ninox philippensis sensu lato. Taxa that differ in plumage also differ in vocalisations, so much so that their treatment as conspecific in a group with innate vocalisations such as owls is untenable. Surprisingly, however, some unstreaked taxa that resemble each other closely are also divergent in vocalisations, and cannot be maintained as taxa below the species level. We propose here that seven vocally well-defined allopatric species are involved in the Philippine Hawk Owl complex, of which two are new species (described herein) with distinctly divergent vocalisations. Because individuals from four islands now known to pertain to four different taxa were included in the description of N. spilonota, we designate a lectotype and hence type locality for this ambiguous name. We also describe an additional island taxon here that shows only moderately distinctive vocalisations and morphology, and which we consider better treated at the subspecies level. METHODS Acoustic analyses We analysed sound recordings of all taxa known or suspected to be critical to an analysis of species-level taxonomy of the Philippine Hawk Owl {sensu lato). The majority of recordings we used were made by ROH during trips specifically targeting islands that hold morphologically 'distinctive Ninox taxa for which we previously lacked or had poor representation of vocalisations. Several other recordings were made by co-authors and others, and most of these are available in full on AVoCet (avocet.zoology.msu.edu, AY). (To access individual numbered recordings on AVoCet, use e.g. http:/ /avocet.zoology.msu.edu/recordings/14561.) A few recordings were assembled from other sound archives (Macaulay Library, http://macaulaylibrary.org/, ML; National Sound Archive, London, http://www.bl.uk/nsa, NSA; xeno-canto, http:// www.xeno-canto.org/, XC). 2 P. C. RASMUSSEN etal. Forktail 28 (2012) Figure 1. Map of the known distribution of taxa ofthe Philippine Hawk Owl Ninox philippensis ( sensu lato). Taxa and groups are Philippine Hawk Owl Ninox philippensis ( sensu stricto ) nominotypical philippensis group, of Luzon, Samar, Leyte, and small surrounding smaller islands; centralis, of Panay, Negros, Bohol, Siquijor (unlabelled small island between southern Negros and Bohol) and surrounding smaller islands; proximo, of Masbate; and ticaoensis, of Ticao; Mindanao Hawk Owl N. spilocephala: Mindanao and smaller surrounding islands except Camiguin Sur; Sulu Hawk Owl N. reyi: larger islands of Sulu and Tawi Tawi provinces; Camiguin Hawk Owl new species; Mindoro Hawk Owl N. mindorensis: Mindoro; Romblon Hawk Owl N. spilonota: Tablas and Sibuyan islands, Romblon Province; Cebu Hawk Owl new species. By island (listed alphabetically), the number of recordings used is listed below, with recordist (initials used for co-authors) and (where applicable) abbreviation for sound archive where recording is held (for recordings by FRL first uploaded to AVoCet but also on xeno-canto, only the AV number is provided here): Biliran, 1 (RSK: ML#38695); Bohol, 2, (F. Verbelen [FV]: AV#8971 -8972); Camiguin Sur, 19 (ROH: 13554-13557, AV#13559, AV#13567, AV# 13575, AV# 13577, AV# 13593, AV# 13598, AV# 13602, AV#13605, AV#13609, AV#136l4-136l 5, AV#13618, AV#13622; LMP: AV#13552-13553); Cebu, 13 (DNSA: XC#79316, AV#1 1320-1 1322; LMP: AV#10469- 10470, AV#10805; PGCJ: AV#10804, AV#10806; BD: AV#12609- 12612); Leyte, 2 (RSK: ML#38671, ML#38674); Luzon, 34 (PCR: AV#2 168-2 174; FV: AV#8970; FRL: XC#30725-30728; ROH: AV# 1 2420, AV# 12450-1 2454, AV# 13551, AV# 1 3648- 1 35 54; D. Edwards: XC#35238; P. Noakes: XC#40819, 40821, XC#40823, XC#40825-40826, XC#40828; G. Wagner: XC#23 1 16); Masbate, 1 (LMP: AV#14563); Mindanao, 23 (FRL: AV#8056-8057, AV#8088, AV#81 1 1-8113; ROH: AV# 12455- 12467; S. Harrap: NSA Wildlife ref. #132605-132606; B. F. King: NSA Wildlife ref. #54931); Mindoro, 6 (ROH: AV#11507, AV#13655; P. Morris: NSA#65216 W1CDR0000309 BD24, NSA#W1CDR0000307 BD1, NSA#W1CDR0000309 BD21, NSA#W1CDR0000309 BD24); Negros, 5 (FRL: AV#10664, AV#10699-10701, AV# 10800- 10801); Sibuyan, 8 (ROH: AV#13637, AV#1364l- 13647); Siquijor, 2 (DNSA: AV#14564- 14565); Tablas, 15 (DNSA: AV#10803,AV#1 1323-1 1324; BD: AV#11325; ROH: AV#1 1508-1 1515, AV# 12606- 12608); Tawi Tawi, 17 (DNSA: AV#10802; ROH AV#14566-14581). Recordings are highly variable in length, quality and documentation, but we have extensive, Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 3 good to excellent material from all the above islands except Biliran, Leyte and Masbate; material for Siquijor is rather inadequate. Sounds were studied, measured and graphed in Raven Pro 1.3 (Raven 2012). Measurements taken (where possible) for each recording were maximum number of notes/strophe, maximum and minimum note length per recording; minimum inter-note spacing; maximum and minimum fundamental frequencies (one each per recording), maximum frequency difference within a single note and between strophes; and maximum note bandwidth at a single point. Principal components analyses (PCAs) were done in SYSTAT 13 (SYSTAT 2012) using these measurements except for maximum note bandwidth (excluded because hisses were difficult to measure precisely). After an initial run that showed that all recordings from islands populated by nominotypical philippensis or centralis grouped together, these groups were combined in the analysis of all groups. Also, because inclusion of atypical song sequences resulted in much greater variance in early analyses, only songs considered to be typical were included in further analyses. Thus, typical songs of all taxon groups (with philippensis and centralis combined) were used in the main PCA. However, because of the great vocal differences of Camiguin Sur, reyi , mindorensis and spilocephala from each other and all other (‘core’) groups (those with non-extreme song characteristics), the core groups ( philippensis plus centralis , Tablas spilonota, Sibuyan spilonota, and Cebu) were not well resolved. Therefore, a subsequent PCA was undertaken using just the core groups, and for this recordings from the different islands were graphed separately. For taxa (all but Camiguin Sur, reyi, mindorensis and spilocephala ) that typically give lengthy series starting with single notes and building after a few minutes to a multi-note climax, only one of each of the above measurements was taken. It should be noted that many recordings and analyses of this type of strophe are likely to be incomplete, as the recordist may have begun recording only after hearing the bird, or the initial notes may be very soft and cannot be picked up by a recording. For taxa that typically give a series of short strophes each of which climaxes individually, each of the above measurements was taken for each good-quality strophe. It must also be borne in mind that many recordings made by a single recordist on a single night, or even possibly over multiple nights at the same locality, are likely to be of the same individual owls, leading to some potential pseudoreplication in our analyses (a problem we could not avoid but which is unlikely to bear on the outcome). Moreover, many recordings will have been made after playback, and this has not typically been documented by the recordist. Many recordings are duetted between pair members, while others appear to be counter-singing between birds in neighbouring territories; it is not easy on present knowledge to distinguish these. However, the ability to do so is not critical to our analyses, as the differences between taxa here considered species are so marked. We chose the above measurements because they are little affected by such problems. However, given the great variation among taxa in vocalisations, few if any song characteristics are shared among all taxa, so choosing appropriate measurements was challenging. In the vocal transcriptions presented in Results, notation follows Rasmussen & Anderton (2005). Lower case signifies relatively low volume compared to SMALL CAPITALS then to FULL CAPITALS, which is much louder than lower case. The forward slash / signifies a rise in frequency and the backslash \ a frequency drop between elements. Run-together syllables signify no pause, an apostrophe (’) extremely short breaks (e.g. in a trill), a hyphen (- ) indicates a very short pause, a comma a mid-length break, and an open underscore _ a still longer pause. Ellipsis (...) is used to indicate the continuation of a vocalisation as previously transcribed, not to indicate pauses or fading out. Sonagrams (spectrograms) prepared in Raven Pro 3.1 are presented for each taxon along with the corresponding waveform (oscillogram, in Raven units, which are unique to Raven software and hence not indicated on the Y axis), which shows power versus time, and hence allows visualisation of rhythm better than the sonagram alone. The corresponding spectrum plots power versus frequency, hence allowing visualisation of power peaks. The area of the sonagram highlighted in grey is that on which the spectrum is constructed. Mensural analyses We studied specimens of Ninox philippensis (. sensu lato ) held (in alphabetical order of acronym) in the American Museum of Natural History, New York (AMNH); Academy of Natural Sciences, Philadelphia (ANSP); Natural History Museum, Tring, UK (BMNH); Carnegie Museum of Natural History, Pittsburgh (CM); Cincinnati Museum of Natural History, Cincinnati (CMNH); Delaware Museum of Natural History, Greenville (DMNH); Field Museum of Natural History, Chicago (FMNH); Institut Royal des Sciences Naturelles, Brussels (IRSNB); Museum of Comparative Zoology, Boston (MCZ); Bell Museum of Natural History, Minneapolis (MMNH); Museum National d’Histoire Naturelle, Paris (MNHN); Naturalis, Leiden (NCB); Philippine National Museum, Manila (PNM); Royal Ontario Museum, Toronto (ROM); Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt (SFN); Staatliches Museum fur Tierkunde, Dresden (SMTD); Staatliches Naturhistorisches Museum, Braunschweig (SNHM); University of Michigan Museum of Zoology, Ann Arbor (UMMZ); National Museum of Natural History, Washington, DC (USNM); Peabody Museum, Yale University, New Haven (YPM); and Museum fiir Naturkunde, Berlin (ZMB). All specimens of this complex available at the above museums during our visits were studied, and most of them photographed and measured. In total, 177 specimens of the following established taxa were included in mensural analyses: 40 philippensis (5 Catanduanes, 5 Leyte, 2 Lubang, 26 Luzon, 1 Marinduque, 1 Polillo);4 proxima (Masbate); 1 ticaoensis (Ticao); 36 centralis (2 Bohol, 27 Negros, 7 Siquijor); 16 spilonota (1 Tablas, 12 Sibuyan, 1 Cebu, 2 Camiguin Sur); 43 spilocephala (10 Basilan, 31 Mindanao, 2 Siargao); 25 mindorensis (Mindoro); and 12 reyi (1 Siasi, 9 Tawi Tawi [8 main island, 1 Bongao], 1 Sibutu, 1 Sulu). Not all measurements were available for all specimens (see below), hence numbers in certain analyses are smaller. For specimens from most of the above collections, PCR measured a wide array of characters, and then after preliminary analyses chose the following as most useful: culmen from cere; upper mandible height at cere; auriculars maximum length (including filamentous extensions); tail length (measured by inserting ruler between two central rectrices); tarsus length; unfeathered (bristled) portion of tarsus; length of middle claw; wing length (flattened), and maximum width of dark and light bands on central portion of one central rectrix. Broken, heavily worn or incompletely grown feathers were not measured. NJC also measured culmen from skull for specimens from some museums. Univariate statistics and PCAs were run in SYSTAT 12. Sexes were combined for analyses, because most specimens of key taxa were not labelled as to sex. Although it would be possible to examine sexual size dimorphism in the better-represented taxa, and it may be significant, we leave that to future studies and do not consider the interpretation of our results to hinge on the matter. Colour and pattern analyses Plumage characters were documented at major collections with holdings of multiple key taxa. Photographs were taken of nearly all specimens examined and were used for later comparisons, but only as a general guide and mnemonic. In addition to traditional assessment of species limits under the Biological Species Concept, we apply the system proposed by Tobias 4 P. C. RASMUSSEN etal. Forktail 28 (2012) et al. (2010) to measure the degree of phenotypic differentiation between taxa. In this system an exceptional difference (a radically different coloration, pattern or vocalisation) scores 4, a major character (pronounced difference in body part colour or pattern, measurement or vocalisation) 3, a medium character (clear difference reflected, e.g., by a distinct hue rather than different colour) 2, and a minor character (weak difference, e.g. a change in shade) 1 ; a threshold score of 7 is required for taxa to be considered separate species, but only three plumage characters, two vocal characters, two biometric characters (assessed for effect size using Cohen’s d where 0.2-2 is treated as minor, 2-5 medium, 5-10 major and >10 exceptional) and one behavioural or ecological character may be counted (Tobias etal. 2010). However, we observe that, in the case of nightbirds where vocalisations are crucial for species recognition, this system may not give enough weight to single key vocalisations and may give too much weight to plumage characteristics, which in owls often show relatively high degrees of individual variability. Because the original diagnoses of heretofore named taxa placed in spilonota and reyi were invariably based on inadequate material, often involving single specimens and without reference to the most similar taxa, and because spilonota proves to contain four different taxa, we provide new diagnoses based (where possible) on larger samples. We are constrained by the paucity of specimens from Cebu, Tablas and Camiguin Sur, but good photographs now exist of these and other taxa. Photographs often show features no longer existing or never apparent in specimens, so we rely strongly on them to supplement specimen material in these diagnoses. We have few data on immatures, so these are not included in the diagnoses. In general immatures are less well-marked and fluffier than adults. Owing to the generally small sample sizes and poor labelling of key taxa we were unable to analyse whether sexual dichromatism exists in these owls. However, we suspect that it may, as in some other Ninox owls. In photographs that show what is almost certainly a singing pair for each ‘unstreaked’ taxon, one individual has noticeable whitish streaking on the lower underparts, while the other is plain or barred below. Whether this is a simple sexual difference is, however, unclear, given that the white-streaked birds are in a minority among specimens. Further fieldwork is needed to clarify this issue. RESULTS Acoustic analyses The following are vocal comparisons by taxon or island (see Methods for conventions used). For simplicity, we focus on a few individual recordings and then summarise variations. Overall song patterns and quality are summarised in Table 1, and univariate statistics for measurements of songs in Table 2. N. p. philippensis (Luzon) (AV#2168, AV#2171, Luzon). — The non-duetted song of N. p. philippensis (Table 1, Figure 2a) begins quietly, with single, short, rather sharp cuk notes (each note comprising a short upslurred then rapidly downslurred element, upper frequency limits c.0.9 kHz, frequency range c.0.6 kHz, note duration c.0. 11s) spaced c.2-2.5 s apart (at first slightly more than 2 s, then gradually accelerating slightly to just under 2 s apart). The series becomes louder, and after several single notes the bird then begins to add in soft, lower-pitched (0.66 kHz) preliminary notes, single or doubled, as in boo, /CUK and bu-bu,/CUK. There are several iterations of this, in which the first element becomes progressively louder, but the main subsequent element is still greatly accented, more strident and yapping, and much broader-band, and the couplets or triplets are separated by pauses of just over 1 s. This then changes into a four-element repeating motif in which the first note is mellow and low as before, and the subsequent three notes are broader-band, louder and sharper, the last slightly softer and tailing off in pitch. In the four- element motif, the second and third notes are farther apart than the third and fourth, hence the rhythm seems halting; in addition the third note is primarily upturned, while the second and third are primarily downturned, and the third note has a slightly lower maximum frequency. The four-note strophes are c.2 s in duration. The series becomes slightly higher-pitched overall towards the end, just exceeding 1 kHz, and the pauses between motifs are c.2 s long. The series lasts 75-95 s and ends suddenly, or becomes intermittent, with longer pauses (up to 8 s) between motifs, e.g.: cuk_cuk_cuK_cuK_cuK_boo,/CUK_boo,/CUK_bu-bu,/CUK_bu- bu,/CUK_boo,/CUK,CRIK-COok_boo,/CUK,CRIK-COok ... boo,/ C UK, CRIK- CO ok... Shorter versions and song fragments (SOM la. 12453) may be given, as may long series of single note types, and excited duets with multiple birds joining in that may appear to have more than four notes per motif (SOM lb. 13551). N. p. centralis (Bohol, Negsros) (AV#8971, Bohol; AV#10699-10700, Negros). — Song on both islands is similar to that of N. p. philippensis. A Bohol recording (SOM lc.8972) of a duet is similar to duets of N. p. philippensis. AV# 10699, from Negros, has much longer (c.5 s) pauses between initial notes than similar song strophes studied for nominotypical philippensis, while those of AV# 10700 (SOM ld-e. 10700) are c.2 s, as in nominotypical philippensis , and the strophes last 120-168 s. N. p. centralis (SSquijor) Song (SOM 1 f . 1 12a) is similar to that of nominotypical philippensis, except that it is hoarser and the later strophes contain more (sometimes several more) notes, especially in duets, which have many short hoarse notes (SOM lg.99). N. p. spilocephala (Mindanao) The long song of spilocephala (Table 1, Figure 2d, continued as SOM 1 j) differs dramatically from all others in consisting of very slow, long, mellow, dove-like notes, singly or in couplets. The strophe starts simple and becomes somewhat more complex, but slows down considerably. The song (e.g. AV# 12462) starts with single mellow, slightly slurred hUUUu notes that eventually become longer, with the addition of an initial very soft low segment to the note (now oo/hUU\Ju ), then adding a short final element after a short pause (now oo/hUUUu, Huh). The fundamental frequency is 0.37-0.53 kHz, with only the main ( hUUUu ) note reaching the highest frequencies. All notes are slightly slurred and convex (especially the main note) and have the same quality. Early notes in a strophe are 0.4 s and the last couplets are 1.3 kHz. The notes have one strong harmonic. Pauses between early notes are c.2. 5 s and between later couplets c.4 s. The entire strophe may last over 190 s. A transcription is: hUU\Ju_hUUUu_ ... h UUUuJj UUUu_oo/hUUUu_oo/hUUUu_ ... oo/hUUUu_oo/ hUUUu_oo/ hUUUu, huu_oo/ hUUUu, huu oo/hUUUu oo/ hUUUu, huu ... oo/hUUUu, huu oo/hUUUu, huu oo / hUUUu, huu ... Duets (e.g. AV# 14263, Figure 2e) may be much shorter, e.g. 11s, with the primary singer starting with just a few single notes that quickly lead into oo/hUUUu, huu couplets like those described above. The duetting bird sporadically adds in slightly but noticeably higher couplets in which the first, stressed note is mostly upslurred and the second, slightly softer note is mostly downslurred, transliterated as IVAUUUU, JVUUUu. The lowest frequency of this couplet is 0.36 and the highest c.0.73. The couplet length is 1.2 s. The single noticeable harmonic produced by this second individual reaches c. 1 .3 kHz. These same WA UUUU, IVUUUu couplets may be given with another distant bird giving this same type instead of the oo/hUUUu, Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 5 Table 1 . Qualitative summary characteristics of songs of each taxon and/or island population of Ninox philippensis ( sensu loto) studied. Song characteristic Gives long Increasing Maximum note number/ Note pace accelerating (+), even (0), or decreasing (-) Pause length even (0) or decreasing (-) between 1st note of multi¬ note strophe lowest (-), highest (+), intermediate (i) or 1st note of multi¬ note strophe softest (-), loudest (+),or All notes of multi-note strophe nearly the same length (0) or different (in order from shortest taxon steady series note number? strophe within strophes strophes no difference (0) no difference (0) to longest) philippensis Yes Yes 4 + 0 to — - - 3, 2, 4,1 spiiocephaia Yes Yes 2 + - + + 2,1 reyi Yes Yes c.20 0 Irregular 0 or — 0 or — 0 CamiguinSur Occasionally Occasionally 22 - Irregular - - Shortest at start, longest at end mindorensis Yes Occasionally 1 main note, several grace notes + 0 0 - 1-4,5 Cebu Yes Yes 5 (strophes run-together) + 0 - - 1,4, 3, 2 (frequent; also other patterns) labhs spilonota Yes Yes 4-5 (strophes well separated) + - + - 1,3,2 (usually) Sibuyan spilonota Yes Yes 5 (strophes well separated) + - i - 2, 1-4, 3 Song characteristic Taxon Most notes short (|), long ( — ), or intermediate (0) Most notes near-vertical (|) vs horizontal (-) or slurred (~) Harsh notes present Harmonics prominent Quality of main notes Degree of variability (from 0 to 5, 5 highest) philippensis 0 1 As response to playback, not in songs No Emphatic but mellow 3 spiiocephaia — Rarely No Mellow, clear 2 reyi 1 1 No No Hollow, clucking, percussive 2 CamiguinSur 1 1 to ~ Yes, at start of each strophe Yes Rapid hooting barks, like mid-sized dog 2 mindorensis — ~ (slightly) Yes No Thin whistles, peeps, metallic screeches 3 Cebu 0 ~ (greatly) Yes No Shrill, slightly croaking 5 1ab\as spilonota 0 ~to | Yes In some note types Sweet whistles to hoarse plaintive barks 3 Sibuyan spilonota 0 ~ Slight Yes Robust, musical, slightly hoarse, mournful 4 Table 2. Univariate summary statistics [mean±SD (range, n)] of measurements of song characteristics of each taxon and/or island population of Ninox philippensis (sensu lato). Song characteristics Max. interstrophe Taxon/island Max. notes/strophe frequency difference (Hz) Max. frequency in song series (Hz) Min. frequency in song series (Hz) Max. note length (s) Min. note 1 (s) Min. internote interval (s) Delta frequency within note (Hz) philippensis 4.2±0.8 303.4±66.3 111 6.6±90.2 585.7±90.7 0.3±0.1 0.1 ±0.1 0.6±2.1 479.0±148.6 (including centralis, proximo, ticaoensis ) (2-7,34) (177.0-420,34) (864-1364,55) (386-787,54) (0.1-0.7,52) (0.0-0.3,52) (0.1-15.0,52) (107-1068,52) spiiocephaia 2±0 82.1 ±31 .8 693.4±92.2 402.1 ±45.5 0.5±0.1 0.2±0.04 0.3±0.1 271.0±69.7 (2,11) (55.0-166.0,11) (549.0-854.0,22) (325.0-508.0,22) (0.3-0.6,22) (0.1-0.3,22) (0.1-0.4,20) (178-417,22) reyi 18.8±8.1 284.6±85.2 1048.8±90.7 427.8±64.6 0.1±0.1 0.01 ±0.0 0.08±0.04 529.0±165.4 (6-40,15) (155-420,15) (834-1179,18) (331-528,18) (0.0-0.3,18) (0.01-0.02,18) (0.01-0.14,18) (376-1006,18) mindorensis 2.4±1 .5 365.7±68.7 2364.6+182.7 1451 .0±208.6 0.8±0.2 0.2±0.1 0.6±0.5 648.8±183.5 (1-5,11) (221-443,8) (2070-2716,12) (1282-1840,12) (0.5-1.0,12) (0.1-0.6,12) (0.2-1.6,12) (351-909,12) Camiguin Sur 22.8±5.6 122.9±50.2 631 .8±36.8 327.6±53.0 0.2±0.4 0.1±0.1 0.3±0.9 284.0±78.3 (8-39,34) (39-221,34) (568-699,47) (241-434,47) (0.1-0.3,44) (0.0-0.3,43) (0.1-5.4,34) (96.4-445.1,43) Sibuyan 3.8±0.4 110.6±28.9 1559.6±238.4 783.4±95.6 0.3±0.1 0.1±0.1 0.8±1.7 639.2+236.5 (3-4,7) (66-133,7) (1280-1911,9) (584-935,9) (0.2-0. 5, 9) (0. 1-0.3, 8) (0.2— 5.0, 8) (239-935,8) Tablas 3.6±0.5 201.4±80.6 1 784.1 ±192.2 923.6±190.5 0.4±0.1 0.2±0.1 2.3±3.2 681.2±229.6 (3-4,11) (111-332,11) (1484-2213,18) (569-1545,18) (0.2-0.6,18) (0.1-0.4, 18) (0.1-8.5,18) (306-1121,18) Cebu 4±0.8 181.5±83.9 1466.7±96.7 829.2±73.0 0.4±0.2 0.1±0.0 0.2±0.1 505.6±1 30.5 (3-5,10) (99-376, 10) (1220-1545,12) (711-996,12) (0.2-0.9,12) (0.03-0.1,12) (0.05-0.3,12) (254-734,12) 6 P.C. RASMUSSEN etol. Forktail 28 (2012) Figure 2. Typical songs of each major taxon in the Philippine Hawk Owl Ninox philippensis ( sensu lato ) complex. For each song, the first row is the waveform, the second is the sonagram, and the third is the spectrum: (a): Philippine Hawk Owl N. p. philippensis, probably non-duetted long song of single bird singing in different directions, hence the variations in amplitude late in the song series; AV#21 71, PCR, Luzon; (b): Camiguin Hawk Owl Ninox new species 1 , duetted series of short strophes; AV#1 3554, ROH, Camiguin Sur; (c): Camiguin Hawk Owl Ninox new species 1 , long song by single bird; AV#1 3622 ROH, Camiguin Sur; (d): Mindanao Hawk Owl N. spilocephala, long song by single bird (series continued in SOM 1 j); NSA Wildlife ref. #132605, S. Harrap, Mindanao; (e): Mindanao Hawk Owl N. spilocephala, short duetted songs; AV#12461, ROH, Mindanao; (f): Sulu Hawk Owl N. reyi, series of short probably non-duetted strophes; AV#1 4577, ROH, Tawi Tawi; (g): Mindoro Hawk Owl, first half of long duet (continued in SOM 1 1); AV#1 1 507, ROH, Mindoro; (h): long song of single Romblon Hawk Owl Ninox spilonota new subspecies; AV#10803, DNS A, Tablas; (i-j): long song of single Romblon Hawk Owl N. spilonota on Sibuyan, AV#1 3637, ROH, Sibuyan; (k): long song of single Cebu Hawk Owl Ninox new species 2, AV#1 261 2, BD, Cebu; (I): duetted song of Cebu Hawk Owl Ninox new species 2; AV#1 0805, LMP, Cebu. a) Luzon 10 IS time (s) 20 2S 30 1 ' » * t V V * «> _ 05 10 kHz XS 20 25 L L 1 i III UI ' f r r f If ft* , 35 40 45 time (s) so SS 60 I 1 ^ I, i i i.i l I' 1 100 “ JO - - " V - 05 1.0 . 15 kHz 20 25 n m fit • UI UI rrr . Ml 70 7S time (s) BS 90 5 ' r . !U (II .111 . IK IK , m IK )0 05 kHz 1-5 20 25 1 . . . „z 95 100 105 110 115 120 time (s) .HI . in . Ui . m OS It) , , 1.S 2.0 25 kHz h . . — 4U- 120 125 >30 IBS KO time (s) i 1 i .ui .in .in .in b) c) S 10 IS Camiguin Sur time(s) 20 IS 30 4j^nnini»wi « * - v - Camiguin Sur 10 IS 20 25 •IS SO SS 60 BS 40 4S SO 65 70 75 80 time (s) 85 '' ttiinni | kHz d) e) f) 05 1.0 15 20 25 Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 7 Huu couplets, or the latter type may grade into the former type (e.g. AV#8088). Otherversions include (AV#805 6) aseries oflong dove- like notes ending in a stressed barking upslur, a version in which the first note of each couplet is more accented and staccato (AV#81 1 1), or single short gruff notes. None of the vocalisations of spilocephala closely resembles those of any other taxon in the group, although there is some resemblance to the much shorter, sharper, two-note portion of- the long song of philippensis. N. p. mindorensis (Mindoro) The song (Table 1, Figure 2g, continued in SOM ll) is distinctly high-pitched, and composed ol long series of thin, slightly descending squealed whistles fiiiiew (each c.0.8 s long) separated by pauses of c. 1 s, then gradually changing into descending squealed whistles that become hoarse raspy downslurred screeches, e.g. jiiikshreiv , the whistle-screeches often preceded by several very short high-sounding tittering toots, 0.1 -0.3 s. The screeches and toots 45 . 50 55 60 T tt V't H 1 ♦ ■» ♦ ♦ 4 ¥-■+*- — - fihvvrvmHir fcrr r ' ' TTtTT' 1 11T i) — *- — - + Sibuyan 10 15 time (s) 20 25 3 * 05 W 15 Z0 2 kHz -4 - 4 - 4 - 4 - 4- Sibuyan 1 1JS 140 h 1 1 'i k) 10 is ill ! " HI 'nil ' ih m U i -i > ■> k 4 4 4 ^ 1 4 4 4 f f H i ID)) i) nn i) m in m ) o n 4 >4 »4 4 4 >4 ‘4 >4 Mf lM If f 4 65 70 75 80 as time (s) j it M i \) i> h \ \) |) m (n i\ 35 40 45 time Is) i | 1 ' lli' 1 ill! ' 1 4 III ) 50 55 t kHz 15 2.0 2. i» >4 M it t 8 P.C. RASMUSSEN etal. Forktail 28 (2012) are duetted, both birds often chiming in simultaneously giving both note types, which may overlap closely. Whistled notes may rise slightly at the start before falling again, and others may be entirely screeched. Duetted song may also consist of much longer series almost entirely of long whistled notes, some of them strongly frequency-modulated (SOM lm-n). The song oi mindorensis bears no resemblance to those of philippensis , spilocephala , reyi or Camiguin Sur birds. It somewhat resembles songs of Tablas, Cebu and Sibuyan birds (as noted by Allen [2006], except that no recordings were then available for Sibuyan) in being composed of thin whistles and hoarser rasps, but it is very different in frequency and structure, especially in its much longer-drawn whistles and screeches, the near-constant frequencies of the long whistled notes, presence of high tittering toots, much harsher screeches, and lack of rhythmic repetition of lower notes. N. p. spilonota (Tafoias and Sibuyan) The song of Tablas birds (Table 1, Figure 2h. 10803) consists of long slow series of single short, steeply falling wistful whiny whistles \FYew separated by pauses of several seconds, and later in a strophe often changing into a hoarse version \FYuRsh, then into two-note and finally three- to (occasionally) four-note versions, e.g. (tut-)TUT- TIY\lEw-hut near the end. Hoarser note-types have a few widely spaced harmonics (SOM lo). The maximum fundamental frequency is 2 kHz, and that only at the start of strongly downslurred notes, while most of the notes are 1-1.7 kHz. Song in Sibuyan (Table 1, Figs. 2i— j. 13637) is similar to that in Tablas, but sounds slightly more croaking and is even slower, with many introductory notes nearly level, barely downslurred, and with strong harmonics. The later notes in a strophe are typically clearly four-noted, and each note is more similar to the others except the longer, stressed, downslurred third note, e.g. TOOT, TOOT, TIYUEW, TOOT. N. p. spilonota (Cebu) In Cebu, song (Table 1 , Figure 2k) is highly variable in rhythm and note type, although in quality notes are of two main types, gruff staccato chucks and plaintive short downslurs, and (less often) metallic treefrog-like upslurred abrupt bwick! notes (several of which may be given in quick succession), low clear abrupt duit! bell-tones (sometimes quickly doubled), and hoarse white-noise screeches lasting c.0. 6 s (Figure 21, SOM lq-r). Overall, compared to Tablas and Sibuyan birds, to which it is most similar, the song in Cebu is much faster (but relatively low-pitched, with notes mostly below 1.5 kHz and even down to 0.8 kHz) and lacks the longpauses between notes within a strophe so characteristic of these other taxa. Non-duetted strophes are long series of short IKYeut notes (separated by pauses of only c.0.9 s), running into djuk, / KYEUr_djuk , IKYeut... series and even four-noted, more complex versions. Duets are medium-length strophes starting with several low soft djuk and gwick notes, running into djuk, \KYEUr-givuck, djuk, YKYEUr-gwuck , etc., petering out after c.30 s and then starting again after a pause of a few seconds. Despite similarities to Tablas and Sibuyan birds the numerous notable and consistent differences exhibited by Cebu birds are very striking. N. p. spilonota (Camiguin Sur) (AV# 13554, 13556-13557). — Duetted strophes (the common type available to us; Table 1, Figure 2b) differ greatly from those of all other known taxa. They begin with a few sporadic, very soft, very low (0.28 kHz) mid-length gruff notes, then turn into a low, very hoarse growl (e.g. 0.9 s) that quickly leads into rapid laughing mellow barks (0.36-0.69 kHz), recalling the chorused yapping of distant mid-sized dogs. The duetted notes typically overlap only partially, so the strophe sounds faster than is each individual’s contribution. and the rhythm is non-uniform and jerky. Each note (excepting the growl) is strongly convex (highest part in the middle), and note shape is quite uniform; individual note length is 0.10-0.15 s, and pauses between note-pairs in the first two-thirds ol the main strophe can be as long as 0.1 s, then to 0.2 s between this part (signified by CUR notes below) and the accented ending ( BOO notes). Strophes (including introductory grace notes) last 5-1 1.5 s and are separated irom each other by 7-18 s. Strophes slow down slightly near the end, with the final few notes slightly longer, louder and higher- pitched. There are often a few stray loud notes just following the well-defined strophe (SOM lh. 13557). Harmonics are prominent throughout the entire strophe following the growl, which has very broad bandwidth. The highest detectable harmonics on available recordings are at c.2.6 kHz, and there are up to five bands of harmonics above the fundamental frequency, although only the first harmonic (peaking at J.25 kHz) is very prominent. Growls do not occur elsewhere within the strophe but the introductory growls are duetted, with the second individual joining in just after the first, or they may be shorter growls separated by short pauses, e.g.: (huk__huk__)burrrrrrrr’CUR,CUR ’r,CUR ’ur, CUR, CUR ’ur, CUR ’ur, CUR dr, CUR, CUR ’r,CUR r, CUR r, CUR’ ur, CUR’ CUR-/ BOO-BOOR-BU’U _ (boo) Lone birds (Figure 2c, SOM 1 i) also give shorter, low-intensity series of c.6 boo notes, the series lasting c.2 s, and the fourth note highest and stressed (e.g. AV# 13602), or long series starting with single notes, then two, the first stressed, then three, first still stressed, then grading into strophes of up to seven low hoots followed by up to six higher stressed hoots, the first of the higher notes being most stressed, but all evenly spaced and of approximately the same length (e.g. AV# 13622). N. p. reyi (Tawi laws) (AV# 10802). — The song of this taxon is extraordinarily different in tone from those of all other known taxa (Table 1, Figure 2f), being a hollow wooden knocking, usually in duetted short strophes of a few to several seconds, separated from other such strophes by short pauses (1-2 s). Each strophe starts with several quiet low clucks at the rate of 1/0.1 s, then gets louder and higher-pitched, although each note has broad bandwidth; the strophe then slows down, often dropping in pitch and becoming more emphatic and higher again. The duet has a stuttering rhythm overall and there are no harmonics. Each note has broad bandwidth, the earlier ones c.45 kHz and the louder, higher ones c.80 kHz, and note length is c.0.0 1 s for earlier notes and 0.02 s for most others, some perhaps as long as 0.03 s, e.g. kt-kt-kt-kt-kt-kt-kt-/KRT-KRT-KRT-KRT- KR T\ -KLAK-KLAK-KLAK-KR T-KR T-KR T-KR T-KR T-KR 7V TOK'.-TOK! Strophes may be much shorter, e.g. kt-kt-kt-KR T-/T OK!- T OK!, and successive strophes in a series can vary considerably in length (SOM lk). They can also be more uniform, but still with slightly higher pitch and emphasis near the end, or they can be mostly uniform kt- series until a sudden speeding up and more clattery pattern at the end. This clattery effect occurs when clacking notes are introduced by a very short low grace note that increases the hollow knocking quality. The latter notes are evidently responsible for the taxon’s local name of ‘lukluk’ (Allen 1998). This form also gives a simple song of hollow triplets continued for longperiods, e.g. tluk’tluk’tluk_tluk’tluk’tluk ... tluk’tluk’tluk. The motifs are c.0.7 s in duration and separated by usually 3-5 s (commonly 4.5 s), and consist of three principal elements, each preceded by a very short, lower, introductory element. The frequency range of the principal elements is about 0. 5-0.9 kHz. This song type somewhat resembles in quality the typical song of Cinnabar Hawk Owl Ninox ios of Sulawesi, although that species’s song is distinctly two-noted (King 2005, Hutchinson et al. 2006), and no songs comparable to the other types of N. reyi are known. Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 9 Very occasionally a longer, very hoarse growl or hoarse hoot is interspersed with the clacking notes. Principal components analysis The PCA using all groups (with philippensis and centralis combined; Table 3, Figure 3a) shows that Tawi Tawi reyi, Camiguin Sur spilonota , Mindanao spilocephala and Mindoro mindorensis are highly differentiated from each other and from ‘core’ groups (those with intermediate characteristics) on the vocal characteristics measured. Factor 1 contrasts mainly frequency and note length with number of notes, so factor scores for individuals of the high-pitched, long-noted mindorensis are uniformly higher on this axis than other groups, while those of the short-noted, low-pitched Camiguin Sur spilonota and (to a lesser extent) reyi are lowest on this axis. Factor 2 is more difficult to interpret, but it contrasts mainly frequency differences within and between notes with length of shortest notes and length of shortest internote pauses; on this axis reyi and Camiguin Sur spilonota separate out clearly from each other, and spilocephala separates out cleanly from the philippensis group. The PCA using only core groups (with all island forms of philippensis and centralis graphed separately, and including Tablas spilonota , Sibuyan spilonota and Cebu spilonota ; Table 3, Figure 3b) shows that all philippensis/ centralis forms group together, with the two Siquijor recordings somewhat marginal, while Cebu, Tablas and Sibuyan are all fairly similar to each other on vocal measurements, Tablas birds being somewhat intermediate between those of Cebu and Sibuyan. Factor 1 is mainly a contrast between frequency and note length versus number of notes/strophe and degree of inter¬ strophe frequency change, while Factor 2 mainly contrasts number Table 3. S ummary results of principal components analyses of measurements of vocal characteristics for all groups and core groups only ( philippensis and centralis, Tablas spilonota, Sibuyan spilonota, and Cebu). Component loadings All groups Coregroups Factor 1 Factor 2 Factor 1 Factor 2 Factor3 Max. no. notes/strophe -0.736 0.189 -0.468 0.461 -0.075 Max. note length/series 0.772 -0.337 0.556 0.461 0.165 Min. note length/series 0.562 -0.629 0.169 -0.507 0.697 Min. inter-note spacing 0.470 -0.537 0.324 -0.598 0.168 Max. fundamental frequency/series 0.883 0.339 0.895 0.249 -0.020 Min. fundamental frequency/series 0.898 0.127 0.759 -0.058 -0.464 Max. intranote frequency difference 0.590 0.587 0.519 0.469 0.607 Max. inter-strophe frequency difference 0.416 0.589 -0.650 0.282 0.376 Variance explained 3.781 1.657 2.733 1.405 1.273 Percent of total variance explained 47.263 20.711 34.157 17.557 15.911 of notes, maximum note length, and frequency change within notes versus minimum note length and minimum pause length. Summary of vocalisations To summarise the main points of the vocal evidence reviewed above and in Tables 1 and 2, the Camiguin Sur population and reyi from Tawi Tawi give many more notes per strophe than do other forms. Tawi Tawi reyi is strikingly divergent from all others in giving almost exclusively very short percussive toneless notes in extremely rapid, rhythmic strophes. Camiguin Sur birds and, to a lesser extent, Mindanao spilocephala are much lower-pitched than other taxa, while Mindoro mindorensis is much higher-pitched than others. The song of Mindanao spilocephala is unique in several ways, including the stressed first note in the couplets, low number of notes per strophe, and consistently mellow tone. Songs of nominotypical philippensis and centralis are similar to each other; they are in the middle of the pitch range for the entire group, and differ from other taxa in the halting, regular rhythm of later notes, strongly convex note shape, without whistles and normally without growls, and mellow barking quality. Recordings from Siquijor appear to show rather more distinctive vocalisations, but further and higher-quality material is required before taxonomic conclusions can be drawn. While Cebu birds are similar in frequency range and number of notes to Tablas and Sibuyan spilonota, their songs have several peculiarities including the rapid, continuous series, varied unmatched note types, and often erratic delivery. Tablas and Sibuyan spilonota , while by no means identical vocally, do share most characteristics, and on present data appear to represent distinctive races. The other taxa summarised here all show an extraordinary degree of vocal differentiation in a group of birds for which vocal communication is innate and of paramount importance in species recognition. Mensural analyses Our sample of Ninox philippensis sensu lato, with all island populations kept separate (Table 4), shows that, while we do not have adequate sample sizes of most taxa for significance testing, mensural differences between philippensis, centralis, proxima and ticaoensis as curirently recognised are not striking. Flowever, in a PCA of nominotypical philippensis, centralis , proxima and ticaoensis (Figure 4, Table 6), centralis from Siquijor separates out from almost all centralis from Negros and Bohol on Factor 1, a size axis, while the sole ticaoensis specimen included falls at the extreme small end of this axis. Masbate proxima is medium-sized on this axis (Figure 4, Table 6). Measurements (Table 4) also show that all island populations of spilocephala are quite similar in size and proportions. We therefore combine these poorly if at all differentiated forms for subsequent analyses. Table 5, in which only clearly differentiated taxa (named or unnamed) are included, shows that there is considerable size variation, with philippensis, spilocephala and mindorensis all being small, while reyi and the four spilonota populations from Camiguin Key Q) spilocephala O Camiguin 9 'eyi □ philippensis □ Cebu | Sibuyan Tablas ^ mindorensis Factor 1 Figure 3. Results of PCAs of measurements of vocal characteristics for (a) all groups (philippensis and centralis combined) and (b) core groups (philippensis and centralis from different islands graphed separately; Tablas spilonota, Sibuyan spilonota, and Cebu). 10 P. C. RASMUSSEN etal. Forktail 28 (2012) Sur, Tablas, Sibuyan and Cebu are all large in most measures. PCAs (Figure 5, Table 6) consistently show this same pattern. On Factor 1 , which is a strong size axis, taxa are either small or large, although a few individuals in both groups are intermediate in size. On Factor 2, the highest-loading variables are the two tail-banding measurements (Figure 5a, Table 6). On this axis ^philippensis, with its broadly banded tail and to a lesser extent the Cebu bird, score high, in contrast to narrow tail-banded taxa including spilocephala, mindorensis and the Camiguin Sur birds. Although spilocephala is small, it has the longest (and densest) ear-covert extensions of all taxa (readily visible in photographs of live birds; Figure 6b), with philippensis not too different; all the larger taxa have relatively shorter and less prominent ear-covert extensions, which are rarely apparent in photographs. In fact, auricular length is much the highest-loading variable on Factor 3 of the PCA (Figure 5b, Table 6), on which spilocephala loads highest, with considerable overlap with philippensis and a few Sibuyan birds. Figure 4. Results of PCAs of skin specimen measurements of Ninox philippensis philippensis, N. p. centralis, N. p. proximo and N. p. ticaoensis, with all islands shown with different symbols but previous subspecies groupings within dashed polygons. Summary statistics in Table 6. Table 4. Summary statistics for univariate measurements [mean±SD (range, n); (in mm)] of island populations of Ninox philippensis sensu lato, using previously recognised racial divisions. Not included here are mindorensis and taxa formerly united in spilonota. Ht = height; L = length. Variable Island Culmen from skull Culmen from cere Upper Mandible Ht Auricular extension L Tail L Tarsus L Tarsal feathering Mid-daw l Wing L Dark tail-bands Light tail-bands philippensis Luzon 12.7±0.7 (11.8-14.4,25) 7.4+0.4 (6.8-8.7,26) 27.913.6 (18.1-33.6,26) 82.714.5 (72.4-92.2,24) 31.111.8 (26.9-36.0,25) 15.812.2 (12.8-20.8,25) 11.410.6 (10.0-13.1,26) 173.515.7 (164-190,26) 8.811.1 (6.7-10.7,26) 3.910.8 (2.4— 5.7, 26) Polillo 13.4(1) 7.9(1) 28.6(1) 83.2(1) 30.5(1) 16.9(1) 13.0(1) 8.4(1) 3.7(1) Catanduanes 13.1±0.4 (12.4-13.6,5) 7.3(1) - 79.716.8 (74.6-89.6,4) 29.711.8 (27.8-32.4,5) - 11.810.8 (10.8-12.9,5) 17214.2 (169.0-175.0,2) 9.210.9 (8.1-10.3,4) 3.111.4 (2.1-5.1,4) Marinduque 13.2(1) 7.8(1) 22.9(1) 84.0(1) 32.3(1) 21.5(1) 12.0(1) 174.0(1) 9.8(1) 2.6(1) Lubang 12.7±0.7 (12.2-13.2,2) 7.110.1 (7.1-7.2,2) 26.314.4 (23.2-29.5,2) 84.316.8 (79.5-89.2,2) 29.014.2 (26.0-32.0,2) 14.211.2 (13.4-15.1,2) 11.111.3 (10.1-12.0, 2) 169.519.2 (163.0-176,2) 8.910.6 18.5-9.4, 2) 4.510.8 (3.9-5. 1,2) Leyte 12.9±0.8 (12.4-14.3,5) 7.410.4 (6.8-7. 9, 5) 30.112.9 (27.9-35.0,5) 75.612.9 (71.4-79.5,5) 30.312.7 (26.8-33.9,5) 16.612.3 (13.8-20.3,5) 11.410.3 (11.0-11.7,5) 167.219.2 (154-175,4) 9.011.0 (7.8-9.8, 5) 3.310.4 (3.0— 4.0, 5) centralis Negros 12.910.7 (11.6-14.6, 25) 7.610.3 (7.1-8.8, 25) 25.313.3 (19.3-32.7,20) 78.713.0 (73.8-86.4,24) 28.411.7 (25.6-31.1,25) 14.312.4 (9.6-18.8,17) 11.710.6 167.716.1 (10.3-13.1,25) (158.0-184.0,21) 8.311.1 (6-10.6,24) 3.710.7 (2.5-5.2,24) Bohol 13.310.1 (13.3-13.4, 2) 7.410.3 (7.2-77,2) 25.412.1 (23.9-26.9,2) 80.810.8 (80.2-81.4,2) 30.010.4 (29.7-30.3,2) 15.710.5 (15.4-16.1,2) 10.310.5 (10.0-10.7,2) 187.515.0 (184-191,2) 8.611.0 (7.9-9.3, 2) 3.510.2 (3.4— 3.7, 2) Siquijor 14.010.7 (13.6-15.3,5) 8.110.5 (7.5-8.8, 6) 24.514.8 (18.4-91.0,6) 86.114.0 (80.9-91.0,6) 30.812.4 (28.2-34.4,5) 14.611.7 (13.1-17.5,6) 11.710.2 (11.6-12.1,5) 17617.3 (166-186,7) 8.511.5 (6.5-10.8,6) 3.410.6 (2.8-4.5, 6) proximo Masbate 13.110.5 (12.7-13.6,4) 7.510.3 (7. 1-7.9, 4) 25.412.9 (21.5-28.5,4) 80.311.1 (78.7-81.3,4) 28.911.9 (26.3-30.6,4) 14.710.9 (14.0-15.9,4) 12.010.7 (11.2-12.9,4) 173.314.9 (170.0-179.0,3) 9.011.1 (8.2-10.6,4) 3.410.3 (3.0-3.8.4) ticaoensis Ticao 12.0(1) 7.7(1) 24.5(1) 81.0(1) 29.3(1) 13.4(1) 10.1(1) 173.5(1) 10.2(1) 4.5(1) spilocephala Siargao 15.2+0.3 (15.0-15.4,2) 8.610.3 (8.4— 8.8, 2) 33.312.2 (31.8-34.9,2) 81.911.4 (80.9-82.9, 2) 31.210.3 (31.0-31.4,2) 15.710.3 (15.5-16.0,2) 12.410.7 (11.0-12.9,2) 178111.3 (170-186,2) 7.610.7 (7. 1-8.1, 2) 3.410.5 (3. 0-3.9, 2) Mindanao 13.710.7 (12.1-15.0, 29) 7.810.4 (7.0-8.5,30) 29.314.0 (23.5-40.2,30) 74.614.4 (68.3-82.7,24) 29.511.8 (25.5-33.2) 16.812.2 (13.1-21.0,25) 11.510.6 17014.3 (10.3-12.7,29) (162.0-181,26) 7.210.9 (5.9-9.6,28) 2.610.5 (1.2-3.7,28) Basilan 14.210.7 (12.7-15.2,9) 8.110.7 (7.3-9.4,10) 29.113.8 (23.9-37.6,10) 77.516.8 (67.5-87.0,10) 29.611.1 (27.8-31.5,10) 14.511.5 (11.9-16.4,9) 11.811.0 (10.0-13.3,9) 170.619.6 (159-188,10) 7.611.4 (5.7-10.0, 9) 2.710.5 (2.0-3.6.9) reyi Tawi-Tawi 24.3±0.9 (23.0-26.0,7) 13.7+1.2 (10.7-15.2,10) 8.110.3 (7.7-8.5,10) 21.015.2 (15.7-31.3,10) 98.514.1 35.712.6 (92.9-107.5,10) (31.9-39.4,10) 17.712.4 (14.4-21.1,8) 12.510.7 186.718.4 (11.5-13.5,10) (172-195,10) 6.711.2 (5.2-9.2,10) 3.210.8 (2.4-4.3,10) Sibutu 25.0(1) 14.1(1) 7.8(1) 88.7(1) 34.7(1) 12.7(1) 173.0(1) 5.3(1) 2.7(1) Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 11 Table 5. Summary statistics for univariate measurements [mean±SD (range, n); (in mm)] of major taxon groups of Ninox philippensis (sensu lato). Variable Island Culmen from skull Culmen from cere Upper Mandible Ht Auricular extension L Tail L Tarsus L Unfeathered tarsus Mid claw L Wing L Dark tail-bands Light tail-bands philippensis 13.0±0.7 (11.6-15.3,75) 7.5±0.4 (6.8-8.8,73) 26.6±3.7 (18.1-35.0,67) 80.9±4.6 (71.4-92.2,73) 29.8±2.2 (25.6-36.0,75) 15.3±2.3 (9.6-21.5,63) 11.6±0.7 (10.0-13.1,76) 171.9±7.2 (154.0-191.0,68) 8.7+1. 1 (6-10.8,75) 3.6±0.8 (2.1-5.7,75) mindorensis 21.2±0.6 (20.0-22.0,12) 12.6±0.6 (11.7-14.3,24) 7.2±0.4 (6.4-8.3,25) 23.5±3.7 (15.5-30.3,25) 82.6±4.1 (74.1-91.7,25) 29.3±2.4 (24.2-36.1,25) 13.0±1.9 (8.4-16.0,24) 11.2±0.8 (9.6-12.5,25) 164.4±5.7 (154-176,25) 5.2±0.8 (4.0-7.1,24) 2.9±0.6 (2.0-3.9,24) Tablas 15.5(1) 8.5(1) 24.5(1) 96.1(1) 30.2(1) 15.8(1) 12.6(1) 188.0(1) 8.6(1) 3.2(1) Sibuyan 26.0±1.1 (25.0-28.0,8) 16.4±0.8 (15.2-17.6,11) 9.1±0.4 (8.8-9.9,11) 22.6±3.5 (18.3-30.5,11) 101.9±4.6 33.0±1.1 (91.6-107.0,12) (31.2-34.4,11) 1 5.5±1 .3 (13.0-17.5,11) 13.7±0.7 (12.4-14.7,12) 194.4±5.4 (185-202,12) 7.5±0.6 (6.4-8.6,11) 3.4±0.5 (2.5-4.0,11) Cebu 22.0(1) 13.1(1) 8.4(1) 23.8(1) 98.5(1) 38.4(1) 195(1) 5.8(1) Camiguin 16.2±0.1 (16.2-16.3,2) 9.4±0.1 (9.3— 9.4, 2) 25.1±2.0 (23.7-26.6,2) 97.7±10.9 (90.0-105.4,2) 35.2±3.7 (32.6-37.8, 2) 19.8±1.5 (18.8-20.9, 2) 13.5±1.1 (12.7-14.3,2) 184±4.2 (181-187,2) 6.5±0.5 (6.1-6.8,2) 2.4±0.8 (19-3.0,2) spilocephala 13.9±0.8 (12.1-15.4,40) 7.9±0.5 (7.0-9.4,42) 29.4±3.9 (23.5-40.2,42) 75.8±5.3 (67.5-98.0,36) 29.6±1.6 (25.5-33.2,40) 16.2±2.2 (11.9-21.0,36) 11.6±0.8 (10.0-13.3,40) 170.3±6.5 (159.0-188.0,38) 7.3±1.0 (5.7-10.0,39) 2.7±0.5 (1.2-3.8,39) reyi 24.2±1.0 13.7±1.1 8.1±0.3 21.0±5.2 97.5±4.7 35.6±2.5 17.7±2.4 12.5±0.6 186.7±8.2 6.6±1.2 3.2±0.8 (23.0-26.0,12) (10.7-15.2,11) (7.7-8.5,11) (15.7-31.3,10) (88.7-107.6(12) (31.9-39.4(11) (14.4-21.1,8) (11.5-13.5,11) (172-195,15) (5.2-9.2,11) (2.4-4.3,11) Key O spilocephala O Camiguin • reyi □ philippensis □ Cebu ■ Sibuyan ■^Tablas — If mindorensis Figure 5. Results of PCAs of skin specimen measurements of Ninox philippensis ( sensu lato), with all major taxa shown with different symbols and those with more than two specimens grouped within dashed polygons. Summary statistics in Table 6. Unfortunately, sample sizes of all three new taxa formally described in this analysis are very small, and therefore we can only guess at the ranges of their measurements. However, the sample size for Sibuyan is moderate, and while the single Tablas specimen included falls just within the range of measurements for Sibuyan birds (on the small side for all but one measure: Table 4), it consistently falls outside all Sibuyan birds, on the small side of Factor 1 (a size axis), in the PCA (Figure 5). This could either indicate minimal or moderate difference in size between these two morphologically and acoustically fairly similar taxa. There is great plumage variation in photographs of Cebu birds, and we assume they also vary in size as with other taxa. The only Cebu specimen known is as large as the smallest Sibuyan birds but has a proportionately long tarsus, the measurement of which has been independently verified (Table 4, Figure 5). Camiguin Surbirds may vary less in size and plumage than other taxa, based on the two specimens studied and photographs of at least three further individuals. Colour and pattern analyses Most taxa in the. Ninox philippensis ( sensu lato) complex are at least moderately variable in plumage, particularly in the case of most of the small island forms formerly united within spilonota or reyi. Given the small sample sizes available for most of these taxa, delineating diagnostic plumage characteristics is problematic. Plumage or structural characters that are unique to a single taxon are scarce. It may not even be possible to attribute every individual Table 6. Summary results of principal components analyses of measurements of morphological characteristics for all groups and philippensis ( sensu stricio) island taxa only (philippensis, centralis, proximo, ticaoensis). Ht = height; I = length; w = width. All groups philippensis only Component loadings Factor 1 Factor 2 Factor 3 Factor 1 Factor 2 Factor 3 Factor4 Culmen from cere 0.75 -0.26 0.31 0.68 -0.41 0.13 0.05 Upper mandible ht 0.79 -0.17 0.34 0.62 -0.40 0.24 -0.33 Auricular 1 -0.25 -0.06 0.76 0.25 0.64 -0.42 0.18 Taill 0.83 0.08 -0.28 0.61 0.41 0.24 0.06 Tarsus 1 0.73 0.09 -0.09 0.55 0.45 0.29 0.05 Unfeathered tarsus 1 — — — 0.42 0.33 -0.15 -0.53 Mid-daw 1 — — — 0.62 -0.02 -0.29 0.43 Wing 1 0.84 0.24 0.01 0.14 0.02 0.75 0.03 Dark tail-band w -0.03 0.70 0.55 -0.43 0.45 0.26 -0.46 Light tail-band w 0.01 0.88 -0.20 -0.37 0.21 0.49 0.54 Variance explained 3.17 1.44 0.85 2.49 1.47 1.37 1.13 by components Percent of total 39.60 17.99 15.30 24.87 14.72 13.69 11.26 variance explained 12 P. C. RASMUSSEN etal. Forktail 28 (2012) Figure 6. Photographs of (a) Luzon Hawk Owl N. philippensis, Dolores, Quezon, Luzon, 1 6 January 201 2 (R. J. Quisumbing); (b) Mindanao Hawk Owl N. spilocephala, PICOP, Mindanao, February 201 1 (R. O. Hutchinson); (c) Mindoro Hawk Owl N. mindorensis, Sablayan, Mindoro, December 2008 (James Eaton); (d) Romblon Hawk Owl N. spilonota, Tablas, 4 March 201 2 (Marc Thibault); (e) Camiguin Hawk Owl (new species), Camiguin Sur, 8 June 201 1 (R. O. Hutchinson); (f) Cebu Hawk Owl (new species), Cebu, 3 January 2012 (Christian Artuso); (g) Sulu Hawk Owl N. reyi, Tawi- Tawi, January 2012 (R. O. Hutchinson). Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 13 to taxon without recourse to measurements or knowledge of island of origin. However, on a combination of characters, each of the taxa we recognise below can be readily identified. The main plumage and other external features that differ between taxa in the Ninox philippensis ( sensu lato ) complex are summarised in Table 7. While plumage differences between N. philippensis ( sensu stricto ) and all other taxa are straightforward and consistent, those between all other taxa are less obvious and are only diagnostic in combination, although the conformation of the streaks of the lower underparts of spilocephalus is highly distinct except from a very few individuals of reyi. The profound differences in vocalisations demonstrated above are entirely inconsistent with treatment as a single species, but these differences are largely congruent with additional evidence from mensural analyses and plumage and soft-part colours. Under the Biological Species Concept, we consider that Ninox philippensis ( sensu lato ) represents seven species, ofwhich two formerly treated under the race spilonota are undescribed species and a third is an undescribed subspecies. Under the Tobias criteria (Tobias et al. 2010), all unstreaked forms differ from the taxa in N. philippensis sensu stricto by their exceptional vocalisations (score 4) and unstreaked versus streaked plumage (score 3), plus various other characters not necessary to enumerate here; total at least 7. Diagnoses N. p. philippensis Our evidence indicates clearly that the streak-breasted taxa form a highly consistent group very weakly differentiated in plumage, morphometries and voice, and without attempting to score these characters, all of which appear to be minor, we propose that the taxa philippensis, proximo., centralis and ticaoensis be considered a single species, Luzon Hawk Owl Ninox philippensis. Indeed, given the variability in plumage, size and voice we feel that the case for subspecific separation in this widespread northern and central Philippine species is not solid. The Masbate rac cproxima (diagnosed on a sample size of two as ‘similar to philippensis but larger and with a relatively shorter tail’, but also darker above, with less pronounced pale tail-bars, reduced pale spots on wing-coverts and wings, and coarser, darker stripes below: Mayr 1945) is particularly doubtful, and we subsume it here in the nominotypical. The small dark Ticao race ticaoensis (diagnosed in a comparison with proxima on a sample size of three as having ‘the upperparts darker, the ventral streaking sharper and darker, and the light barring on the tail narrower’: duPont 1972), is at best marginally distinct. Birds from Siquijor, bearing the name centralis (diagnosed as ‘much larger than proxima and with a long tail’, with no rufous or tawny tones in the ‘dark earth brown’ of the upperparts, much reduced pale spotting on the scapulars and upperwing-coverts, ill-defined streaking below with an ochraceous wash to the white feathers: Mayr 1945), are certainly larger than and, on our sample, vocally distinguishable from other island populations currently placed with centralis , so we restrict centralis to the Siquijor population (for which the name was established) and place populations from other islands except Ticao in the nominotypical, but further study is needed. Diagnosis (from large series of specimens and many photos of live birds; see Plate and Figure 6a). — Size small, with compact, sleek plumage. Head has prominent if small and/or buff-tinged whitish supercilium. Auriculars are uniform plain dark brown, concolorous with or slightly paler than crown, with fairly long, prominent filamentous extensions; facial disk not well defined. Above, uniform brown above from crown to rump, lacking barring or speckling. Scapular patch with some largely white to pale buff feathers with dark brown edgings, or large white/buff patch on mid-distal portion of dark brown feather; barring or mottling lacking. Larger outer wing-coverts have large white spots, usually one but sometimes two per feather; barring is lacking. Tail banding is variable in intensity but most have fairly weakly marked pale bands that are much narrower than the dark bands. Below, throat narrowly white, with fine dark streaks; underparts prominently streaked dark brown or rufescent-brown, either crisply or indistinctly, on a white or rufous- tinged background. The streaks are not edged with darker colour or intruded by spots or bars. The streaks become more distinct, narrower, and longer on the central lower underparts, where they typically cover the central half to third of the feather, the remainder being white or whitish, and the undertail-coverts are white without markings or with a few faint and/or narrow dark markings. Bare parts: irides bright yellow, often appearing slightly paler, more lemon-yellow on outer ring; narrow orbital ring pale tan, not conspicuous; cere and bill pale olive-yellow, the culmen ridge and tip slightly yellower; tarsi and toes bright yellow, claws almost entirely black, the bases paler. Vocalisations are long, mid-pitched series of strongly convex barking notes starting with single notes and ending with about four halting notes, the first note softest, without whistles and usually without growls. Table 7. Main qualitative plumage and soft-part character differences between taxa in the Ninox philippensis ( sensu lato ) complex. Morphological characteristic Taxon Crown pattern White supercilia Filamentous ear- covert extensions Throat-patch Dark-streaked underparts Barring below Iris colour Bill colour Claw colour philippensis unmarked well-marked long not apparent yes no yellow, paler outer ring mostly dark mindorensis finely barred small but prominent shorter, less apparent intermediate; apparent but not large or bright white no yes, narrow and/or broken yellow, mustard inner, lemon outer pale dull yellow, sides greener mostly dark Tablas spilonota barred shorter, less apparent not apparent no obsolete yellow mostly dark Sibuyansp/'/o/wfo speckle-barred not present shorter, less apparent not apparent no yes, prominent to obsolete yellow mostly mustard yellow mostly dark Cebu speckle-barred small but prominent shorter, less apparent well-marked, bright white no at most, broken spotty barring yellow pale olive, culmen and tip yellower mostly dark Camiguin Sur coarsely barred not present shorter, less apparent well-marked, bright white no yes, strong overall blue-grey to whitish mostly mustard- yellow, sides greener mostly pale spilocephala spotted well-marked very long, profuse not apparent yes no yellow mostly dark reyi prominently barred very small shorter, less apparent well-marked, bright white occasionally yes, strong at least on breast mustard-yellow, may show pale greenish- yellow outer ring dull greenish yellow mostly dark 14 P. C. RASMUSSEN etal. Forktail 28 (2012) A/, p. spiloeephala Tweeddale (1879) established this taxon simply on the basis of the 'bright rufous’ to ‘pale tawny rufous’ spotting on the forehead and crown, a feature absent in the otherwise similar form from Luzon with which he compared his extensive (6 males, 1 3 females) material from Mindanao, noting that this material was variable in coloration above and pattern below. He also observed, without proposing these points as possessing diagnostic force, that male spiloeephala were longer-winged than male philippensis , but that spiloeephala by his measurements was shorter-legged than R. B. Sharpes of philippensis. However, Mayr (1945) pointed out that spiloeephala is ‘the principal connecting link’ between the extremes represented by philippensis on the one side and mindorensis on the other, and in this he was clearly referring to the fact that spiloeephala possesses underparts which conflate the streaking of philippensis and the barring of mindorensis. Specimen material confirms the apparent intermediacy of plumage for spiloeephala , which shows brown and white streaking in varying degree on the belly, as in philippensis and its Visayan representatives. However, this streaking is only vague on the breast, which is interrupted by a disorganised and inconsistent pattern of bars and mottling in many specimens. In addition, the streaking of the lower underparts differs from philippensis in being dark-edged and irregular (see Diagnosis below). The spotting and stippling on the crown (which extends onto the mantle) is similar between spiloeephala and the bar-breasted forms. Thus the form spiloeephala differs significantly from all the others in plumage, morphometries and voice, and we propose that this also be considered a species under the name Mindanao Hawk Owl. Its mam song and other vocalisations are highly distinct from that of any other form (Tobias score 3). Moreover, in its combination of barred breast and streaked belly, it displays what we judge to be a major character difference from any other taxon (score 3). It also has exceptionally long, full auricular extensions (from data in Table 5 effect size vs philippensis — 0.74; score 1). Diagnosis (from large series of specimens and photos of at least three live birds; see Plate and Figure 6b). — Size small, with especially short tail. Plumage compact and sleek. Head has a very small, indistinct or absent white patch between eye and supercilia. Auriculars appear nearly uniform dark brown but they are in fact vaguely speckled and/or barred; filamentous auricular extensions normally very long, full and profuse, more so than for any other taxon. Above, crown, nape and upper mantle of adults is strongly and usually evenly spotted with buff, the spots becoming weak on the upper mantle and disappearing. In an immature, the crown is unspotted brown but there is a collar of weak pale buff spots on the upper mantle. Mid- to lower mantle to rump is uniform dark brown. Uppertail-coverts may have a little vague fine pale barring. Scapular patch similar to those in philippensis group. Larger wing-coverts have both large white spots and narrow buff barring. Tail is typically more distinctly banded, with narrower dark bands, than for philippensis. Below, throat is narrowly white with medium-broad, distinct blackish-brown shaft- streaks, becoming broader on upper breast. Sides of neck and breast are weakly barred and spotted dark brown and buff, and central breast is mostly streaked. Lower underparts have mostly longitudinal dark markings but these are typically somewhat rounded, with dark chevron tips and irregular dark markings edging the paler brown internal markings. Undertail-coverts white, often with tiny dark spots. Bare parts: irides pale yellow; narrow bare orbital ring tan-brown, not conspicuous; cere and bill pale olive, more yellow on culmen ridge near tip; feet bright yellow, claws nearly all black, paler at bases. Vocalisations are low-pitched, mellow, slow, and two-noted, with first note of each couplet stressed. N. p. mindorensis Ogilvie Grant (1896a) characterised this form as ‘nearly allied to N. spiloeephala... in having the top of the head and nape barred [sic] with buff’ but ‘the whole of the underparts... tawny buff, transversely barred with brown, while in... N. spiloeephala , though the breast is generally like that of the present species, the belly and flanks are always white, with longitudinal reddish-brown shaft- stripes’. He distinguished mindorensis from spilonota (which the describers had listed as including the Mindoro population) simply by its ‘much smaller size’. Our examination of specimens confirms this basic diagnosis. In addition, as noted above, the barring on the tail is denser in mindorensis than in spiloeephala. Konig et al. (1999) mistakenly suggested that mindorensis is larger and that the number of its tail- bars is fewer than the Philippine Hawk Owl sensu lato. Dutson et al. (1992) and Brooks et al. (1995) were the first in the modern era to realise that the form mindorensis is so different from Luzon birds that it appears to represent a separate species. After hearing tapes by P. Morris made in 1999, Konig et al. (1999) concurred and established it as Mindoro Hawk Owl N. mindorensis. However, they made this move without knowing the songs of other unstreaked forms, which they retained with N. philippensis ( sensu lato). Only now is it confirmed that mindorensis possesses a highly distinctive voice, with few similarities to those of the other unstreaked forms. Compared to any other unstreaked form, mindorensis is distinctly smaller (see Table 5: wing and tail shorter, with effect size vs reyi -2.79 and -3.39 respectively; Tobias score 2), with a weaker pale throat than Camiguin Sur and Cebu birds and reyi but more prominent than in spilonota (score 1), especially narrow underparts barring (score 1), high-pitched song (score 2), and very short toots in climax song (score 2). There are other unique vocal characteristics as well (not scored), such as the unusually long notes. Diagnosis (from large series of specimens and photos of numerous live individuals; see Plate and Figure 6c). — Size small, with small bill, tarsus mostly feathered. Head moderately patterned, with small short white supercilia; facial disk rather pale brown, small, with moderately distinct rim, and usually rather short filamentous auricular extensions. Above, crown and nape finely barred dark brown and buff to whitish, but bars broken resulting in overall appearance of speckling. Speckling grades out on upper mantle, and rest of central mantle uniform, with vague fine barring at edge of mantle and on rump. Scapular patches are extensively white with irregular dark margins and dark spots. Wing-coverts are extensively and finely barred, with dark bands predominating, and with white spots/bands on outer webs of larger coverts that have tiny dark spots and scrawls within the white portions. Primaries have prominent broad dark brown bands and narrower buff bands on outer webs, and very vaguely banded inner webs. Tertials virtually unbanded and with very fine, weak pale mottling, appearing uniform brown. Tail banding is obscure and both dark and light bands appear narrow. Below, throat-patch pale buff, not very clear-cut but more so than for Sibuyan and Tablas birds. Underparts highly variable but typically the most finely barred of any taxon in the N philippensis [sensu lato ) complex. Underparts vary in tone from pale buff to saturated, and may have fairly prominent white streaking below. Bare parts: irides have mustard-yellow inner ring, brighter lemon-yellow outer ring. Bill and cere pale dull yellow, tinged greenish on sides. Tarsi and toes dull olive -yellow, claws black, paler near bases. Vocalisations are high-pitched long whistles often ending with a screech, and often starting with tittering high toots. N. p. spilonota from Tablas and Sibuyan Bourns & Worcester (1894) gave the name spilonota to unstreaked birds from ‘Cebu, Sibuyan, Tablas, Mindoro’, but their description made no comparison with any other Ninox taxon, so the diagnostic features of spilonota are not immediately obvious. The description Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 15 refers to an overall colour of ‘fulvous brown’ above, with light rufous speckling from head to mantle and wing-coverts, a tail ‘nearly black with nine narrow transverse bands of light rufous brown’, underparts ‘rufous brown... many feathers of breast and abdomen with fulvous brown spots’. McGregor (1905) noted that the scapulars, normally white in many Ninox , are reduced to ‘bars of pale fulvous’ in the specimen to which he had access. We now verify that this character holds in the far more extensive material we have examined from Sibuyan. On Sibuyan and Tablas, spilonota is diagnosed by its plain head without white supercilia (score 2), lack of notably paler throat (score 3), lack of white spots in scapulars (score 2), and distinctive song pattern and quality (score 3). Diagnosis (from several Sibuyan specimens and two photos of a live bird, and one Tablas specimen and two photos of a live bird; see Plate and Figure 6d). — Size large, with large bill. Plumage rather lax, and general appearance very plain. Head plain brown, lacking pale supercilia or any strongly contrasting pattern, and with indistinct facial disk. Above, finely speckled buff on dark brown crown and nape, mantle unmarked, scapulars and tertials entirely and distinctly barred dark brown and buff, without large white scapular spots; uppertail rather narrowly but distinctly barred dark, with narrower buff bars. Below, no prominent pale throat -patch. Underparts typically have ochraceous background, either lightly and indistinctly marked with large darker brown broken bars and spots, sometimes looking almost plain ochraceous-brown, or completely barred below, with fairly even, dark brown bars on ochraceous background; rarely has fairly prominent white streaking on lower underparts. Bare parts: irides bright yellow, not differing between inner and outer rings; narrow bare orbital skin dull tan- brown, not standing out from surrounding feathers; bill and cere pale olive -yellow, yellowest along culmen and at tip. Vocalisations are short falling whistles, often hoarse, starting singly then changing into a three-note (Tablas) or four-note (Sibuyan) version. Populations of both islands are covered by this diagnosis; however, Tablas birds differ from those on Sibuyan and require a name. Ninox spilonota fisheri subsp. n. Romblon Hawk Owl Diagnosis. — Smaller than N. s. spilonota. Vocalisations differ from those of N. s. spilonota as indicated above. Holotype. — USNM 314875, female, Badajos [now San Agustin] , Tablas, 1 8 September 1 892, collected by D. C. Worcester and F. S. Bourns; Ex. Menage Collection no. 373. Culmen from cere 15.5 mm, wing 188 mm, tarsus 30.2 mm, tail 96.1 mm. Description of holotype. — Medium-large, weakly marked round- headed owl, largely barred above and nearly plain ochraceous below. Head. — S mall, with short, inconspicuous partially concealed whitish supercilium extending to about half-way over eye; auriculars appear rather dark brown with inconspicuous buff spotting, their filamentous extensions fairly weak and short; forehead and crown dark brown moderately narrowly and distinctly barred and spotted. Forehead to hindcrown and sides of neck profusely covered with small, closely spaced buff spots and short buff bars. Throat pale buff overall (appearing Munseli 2.5Y 8/4), each larger throat feather with a narrow darker brown shaft-streak and often with narrow whitish outer webs and/or bases. Rictal bristles largely pale buff, with black rachis that extend well beyond the barbs, some exceeding the bill tip in length. Upperparts. — Barring/spotting on crown becomes less regular and less conspicuous on sides of neck and nape and grades into a rather vague pattern on upper mantle of broad dark brown and narrower buff (Munseli 10YR 7/6) banding. Scapulars lack any white but have bold, broad but irregular rich buff (Munseli 10YR 8/6) and dark brown banding, the dark brown banding with narrow blackish edgings. Lower mantle and rump are essentially unstreaked warm dark brown (Munseli 10YR 3/4). There is a single all-white abnormal-appearing, badly worn feather in the upper mantle. Wings . — Small external coverts of carpal area nearly uniform warm dark brown, with very vague lighter buff speckling. Median wing- coverts fairly distinctly barred with narrow buff bars and broad dark brown; greater coverts yet more distinctly and broadly banded. Tertials very broadly and evenly banded rather dark brown with narrow, widely spaced, moderately distinct dull buff bars. Primaries have broad dark and narrower dull bull banding with narrower, irregular dull buff uniform dark brown inner webs. Underparts. — Pale buff throat becomes darker, richer buff on lower throat and grades into rich ochre underparts (Munseli 7.5YR 6/8). Medium brown shaft-streaks on lower throat grade into rather indistinct, broken but fairly broad medium brown bars on upper breast, which break up and fade out on lower breast. Except for flank feathers overlying thighs, which are indistinctly banded medium brown, lower underparts are nearly solid rich ochre 6/8, with a few tiny darker spots and vague mottling. Lower centre of belly has some feathers with a few broad whitish tips, and undertail-coverts are buffy-white with narrow buff tips. Tibial and tarsal feathering is ochre-buff mottled whitish, and extends more than half-way to distal end of tarsus. Tail. — Uppertail-coverts are dark brown with a few narrow ill-marked dull buff bars. Uppertail has moderately prominent broad dark brown and rather narrow pale dull buff bands. Counting from base of feather (with uppertail-coverts moved aside), there are 9 clearly discernible pale bands and 10 dark ones. Bare parts. — In the dried specimen cere and bill are fairly dark horn, with a paler, more orange -yellow culmen ridge. Distal end of tarsus and toes have sparse, stiff, pale bristles. Toes and claws on dried specimen are medium horn, tips of claws slightly darker. Remarks. — Appears to be quite closely related to the nominotypical form on Sibuyan, but differences at least in size and vocalisations dictate recognition as a new subspecies. Etymology. — We name this form in honour of the late Tim Fisher, dedicated Philippine ornithologist and co-author of Kennedy ei-u/.(2000). N. p. spilonota from Camiguin Sur Birds on Camiguin Sur are unique in their boldly, broadly barred underparts (Tobias score 2), the combination of no white supercilium with an extensive white throat (score 2), grey to whitish eyes (score 3), and mostly pale claws (no score available; would be 1 ). Vocally they are highly distinctive as well, with their low-pitched, rapid, multi-noted hooting duets (score 3 or 4). This taxon has always been treated within the race spilonota , but clearly under both the traditional Biological Species Concept and the scoring system employed here it requires a name and recognition at the level of species. Ninox leventisi sp. n. Camiguin Hawk Owl Diagnosis (from FMNH 284397 and 399384 and photographs oi at least three living birds; see Plate, Figure 6e, and journal front cover). — Size large, with long, deep bill. Plumage appears full and lax compared to other members of the N. philippensis complex, except that ear-coverts have only short filamentous extensions visible only at very close range. Face drab warm brown, the auriculars barred but appearing plain at a distance; lacks pale supercilium but may have small white patch above bill extending just over eye; face shows no contrast with rest of head, and facial disk border is not well marked. Above, crown very distinctly and regularly barred buff and dark brown, more narrowly and obscurely barred on upper mantle, the barring becoming vague and irregular on lower mantle to rump. Wing-coverts are entirely boldly barred dark brown, buff and white. Scapular patch includes feathers with 16 P. C. RASMUSSEN etal. Forktail 28 (2012) large unbarred white areas on outer web, just tip and base being barred. Primaries are dark brown, the outer webs mainly dark buff edged darker brown, with narrow pale buff bars. Tertials are mottled and incompletely but prominently banded. Uppertail has rather narrow dark and pale bars. Below, large bright white throat- patch with a few narrow dark streaks extends to top of breast, with some largely white feathers in uppermost central breast; white throat-patch normally mostly hidden, but shown to be extensively white in song display. Sides of neck and entire underparts heavily and distinctly banded dark brown and buff. Underparts from breast to abdomen have some irregular white or light buff banding; banding below is heavier, broader and more distinct, with generally darker, warmer brown tone on underparts than other taxa. Undertail-coverts white with extensive dark barring and spotting. Undertail irregularly banded with broad dark and narrow buffy bands, banding sometimes obsolete. Bare parts: irides grey to whitish or very pale yellow-green ( versus yellow in all other taxa); narrow but distinct bare eye-ring mustard-yellow; bill and cere mustard-yellow, more olive-green at base of bill; legs and feet mustard-yellow, tarsi less extensively feathered than in most other taxa in complex, mostly thinly bristle-covered; claws mostly pale, only the tips fairly dark (claws mostly dark in other taxa, judging from photos of live birds). Vocalisations are very low-pitched, typically short strophes repeated after brief pauses, with many rapid, irregular, barking notes per strophe. Holotype. — FMNH 284397, female, Catarman Mountain, Catarman, Camiguin province, Camiguin Sur Island, 17June 1968, D. S. Rabor/W. S. Anguila; edge of second-growth, approx. 1,5002 (2,000-4,0002 on label; Balete et al. [2006] state 1, 5002, c. 450 m). ‘Irish [sic] dark white’. Culmen from cere 16.2 mm, wing 187 mm, tarsus 37.8 mm, tail 90 mm. Paratype. — FMNH 399384, female, Mt Timpong, Matugnao, Mahinog, 13 June 1969, D. S. Rabor. Specimens of other species taken on 13 June 1969 on Mt. Timpong were from 3,1502 ore. 950 m (Balete et al. 2006). Culmen from cere 16.3 mm, wing 181 mm, tarsus 32.6 mm, tail 105.4 mm. Description of holotype. — Crown dark brown, distinctly and regularly barred with buff from forehead to nape and sides of neck, the barring becoming less distinct on upper mantle and obsolete on lower mantle. Scapulars moderately distinctly barred, outer scapulars largely white with dark brown tips and bases. Wing- coverts heavily barred with buff, the greater coverts barred mostly white. Tertials moderately barred buff on dark brown. Uppertail has broad dark and narrow dull buff bands. No pale supercilium, and facial disk dark brown, weakly barred, and weakly defined. Throat extensively bright white, the white extending well onto the upper central breast. Rest of underparts heavily and distinctly but irregularly banded dark brown on buff, with some short broad white bars on central and lower underparts. Tarsal feathering dull brown with narrow darker bars. Remarks. — This is evidently the first known owl with grey or whitish eyes. It is vocally highly distinct, normally giving a rapid, low-pitched duet, the two birds facing and leaning towards each other while very close together, with white throat puffed out and pulsating, wings drooped. Etymology. — We name this new species for Anastasios P. Leventis, whose generous long-term commitment to BirdLife International has been crucial in the stable development of the organisation, and whose particular support for NJC has allowed him to work extensively on Philippine birds and conservation issues over the past decade. N. p. spilonota from Cebu The Cebu population of N. p. spilonota is diagnosed from Tablas and Sibuyan birds, to which it is in vocal, morphological and geographical terms most closely related, by its combination of fairly prominent white supercilium and barred wing-coverts (Tobias score 1), barred back (score 1), prominent (when singing) white throat strongly edged dark (score 3), relatively small bill (score at least 1), long tarsus (score at least 1), and fast-paced song (score 2) with multiple unmatched note types(score 2). Because n=l in the morphometric sample (Table 4) we cannot generate an effect size but the differences in bill and leg length are striking. We consider that, under both the traditional Biological Species Concept and the scoring system utilised here, this heretofore unnamed taxon is a new species. Ninox rumseyi sp. n. Cebu Hawk Ow! Diagnosis (based on holotype, BMNH 1955. 6. N30. 4747, and photographs of at least nine living birds; see Plate and Figure 6f). — Size medium-large, with long tarsi (based on type specimen), but relatively small-billed for its size class. Prominently marked round- headed owl, mostly barred above, weakly marked or uniform below, with white or near-white vent. Plumage compact and sleek, as with most other taxa in complex; ear-coverts have only short filamentous extensions. Head shows prominent short white or whitish supercilia; plain dark ear-coverts contrast with prominent well-marked white throat-patch bordered below and on sides by conspicuous dark markings (apparent in life, especially when singing); crown strongly marked with rows of pale, almost contiguous speckles between dark bars. Above, including wing- coverts and tertials, strongly and narrowly barred dark brown and bull, the barring varying from strong to obscure on lower mantle. Scapular spots are mostly white, with narrow dark brown edgings and warm buff between the white and brown areas. Below, weakly and irregularly speckled or marked with short broken bars on pinkish-buff to fulvous background, usually heaviest on breast; sometimes with broad white streaking on lower underparts and/or flanks, or virtually unmarked pinkish-buff to rich fulvous below, with or without indistinct white streaking on lower underparts. Undertail-coverts white (rarely pale buff), usually with very narrow dark fringes to longest feathers. Central rectrices have relatively broad, distinct dark bands. Bare parts: irides pale to bright lemon-yellow, the inner and outer rings not differing; narrow eye-ring indistinct and dull brown, not standing out from adjacent feathers; bill and cere dull pale olive, tip and culmenridge yellower; legs and feet bright yellow, tarsi about half-feathered, otherwise thinly bristle-covered; claws mostly black, only the proximal portions paler. Vocalisations are mid-pitched, irregular strophes delivered rapidly and irregularly with multiple note types. Holotype. — BMNH 1955.6.N.20.4747, female, Cebu, 17 March 1888, collected F. S. Bourns and D. C. Worcester. Ex. Norwich Castle Museum. Culmen from cere 13.1 mm, wing 195 mm, tarsus 38.4 mm, tail 98.5 mm. Description of holotype. — Crown very dark brown, heavily speckled with buff in rows that resemble bars; weak short pale supercilium; dark brown auriculars. Rear crown and nape heavily speckled with rows of pale buff spots, becoming more barred on upper mantle, then weaker and more mottled on central mantle, which is nearly uniformly dark brown; inner scapulars are more distinctly barred dark brown and buff, and outer scapulars are largely white with very dark brown edgings, golden-buff between the edgings and white inner portions; wing-coverts lightly barred, more prominently so on greater coverts and tertials; uppertail broadly barred very dark brown and narrowly barred dull buff. Throat white, but mostly hidden due to preparation style. Breast pale ochraceous, moderately barred and mottled dark brown, the bars breaking up into small dull brown speckles on lower breast, flanks, and even smaller, fewer speckles on central lower underparts. Tarsal feathering and undertail-coverts buffy white, with a few irregular dull brown speckles. Forktail 28 (2012) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 17 Remarks. — The holotype is the sole known specimen in existence. Photographs show this species to be highly variable in underparts colour and pattern, less so in other characters. In plumage closest to the much smaller N. mindorensis, from which it most obviously differs in the more widely spaced bars below (where these exist). Vocally fairly distinctive, although recognisably closer to N. spilonota and N. mindorensis than to other taxa in song characteristics and quality. Thought possibly extinct (Brooks et al. 1995b) until its rediscovery in 1998 (Pagantalan & Jakosalem 2008). Etymology. — We name this new species for Stephen J. Rumsey, dedicated conservationist and ornithologist who has given great support of BirdLife International but who has also helped promote research and conservation on the island of Cebu. N. p. reyi This was the third form of resident Ninox to be described from the Philippines; but Tweeddale’s (‘1878’ = 1879 fide Dickinson et al. 1991) description of spilocephala apparently was too recent for Oustalet (1880) to be aware of it. Thus Oustalet (1880) only had philippensis for comparison, finding that reyi is immediately distinguished by its larger size, longer wings (reaching beyond the tail-tip), and russet (‘ roux ’) plumage with transverse brown bars on head and shoulders. Mayr (1945), having no specimens to hand and evidently misreading or ignoring Oustalet as well as Sharpe’s (1897) Latin account of Ninox everetti (a synonym of reyi), misattributed Sulu birds to the spilocephala group, and duPont & Rabor (1973) mysteriously failed to remarkon this, despite having eight reyi to hand by 1972 (material in DMNH). The form reyi is distinguished from all other taxa by its very pronounced, even barring on head and breast (Tobias score 2), combination of tiny white supercilia with large bright white throat (score 2), and its extraordinarily distinct song (4) as well as other unscored song pattern characters. Diagnosis (from numerous specimens and photos of three live individuals; see Plate and Figure 6g). — Size large, with large, broad bill. Head shows tiny white patch in supercilium, concentrically dark-barred ear-coverts, and relatively well-defined facial disk. Above, usually distinctly, regularly and rather narrowly barred dark brown on whitish or buffy background; barring becomes muted on upper mantle, and mantle vaguely barred, mottled and vermiculated. Outer scapulars largely or at least partly white, with intricate dark patterning intruding to vane on some; wing-coverts prominently banded, the banding on tertials more muted. Uppertail with prominent dark bands of medium width. Below, throat -patch bright white and very well defined, visible even in perched birds looking down. Breast distinctly and evenly dark- barred, but lower underparts highly variable, often being broadly white-streaked or even irregularly dark-mottled on white background; equally the lower underparts can be fulvous, either fairly plain or regularly dark-barred. Some individuals appear almost black-and-white barred overall, lacking warm tones; others (especially young birds) resemble spilocephala in underparts pattern, but the dark-streaking pattern on the lower underparts is not as distinct or regular as in spilocephala. Bare parts: irides bright yellow, often paler on outer ring; bill dull greenish-yellow; feet and toes mustard-yellow, claws black except for paler bases. Vocalisations are very short rhythmic strophes of rapid toneless clucks, accented at the end. Lectotype designation The description of Ninox spilonotus (spelling thus) Bourns & Worcester 1894 was explicitly based on specimens from Cebu, Sibuyan, Tablas and Mindoro. The description does not indicate how many specimens were available, but the only specimens of the last three of these taxa at the NMNH (formerly USNM) are now considered the syntypes. We consider that Bourns & Worcester’s (1894) spilonotus was based on four diagnosable taxa, and hence designation of a lectotype is necessary. The first specimen listed, from Cebu, is nowBMNH 1 955. 6. N. 20.4747, and is not currently considered a syntype (it does not bear a type label, and is not listed in Warren & Harrison 1973). However, Recommendation 74D in the current code (ICZN 2000; http://www.nhm.ac.uk/hosted- sites/iczn/code/index.jsp?nfv=true&article=74) states that the lectotype should be ‘...preferably of the institution containing the largest number of syntypes of the nominal species-group taxon...’. On this basis we exclude the BMNH Cebu specimen from further consideration as the lectotype, because NMNH has three syntypes. Subsequent to the description of spilonota, Ninox mindorensis Ogilvie Grant 1896 was described as a new species, based on a BMNH specimen (97.6.14.51) collected by J. Whitehead. This excludes the NMNH Mindoro specimen (USNM 314876) from further consideration as the lectotype of spilonotus. Therefore, we are left to choose between the two NMNH specimens, USNM 314877 from Sibuyan and USNM 314875 from Tablas. Neither specimen was figured at the time, and both agree approximately equally well with the description. We therefore arbitrarily fix USNM 314877 from Sibuyan, which precedes Tablas in the ‘Habitat’ list on the original description, as the lectotype of Ninox spilonotus. This furthermore implies that BMNH 1955.6.N. 20.4747, USNM 314876, and USNM 314875 are paralectotypes of the name Ninox spilonotus. Note, however, that the first and last of these specimens are herein designated as types of a new species and a new subspecies, respectively. DISCUSSION Earlier treatments of taxa in the complex Treatments of the Philippine Ninox philippensis complex have inevitably varied over time, and separate into two periods of activity, 1940-1945 and 1990-2000. First, Peters (1940) grouped the taxa into three species, ( 1 ) monotypic N. philippensis from Luzon, Marinduque, Masbate, Ticao, Guimaras, Negros, Leyte and Siquijor; (2) monotypic N. spilonota from Mindoro, Tablas, Sibuyan and Cebu; and (3) polytypic N. spilocephala, consisting of N. s. mindorensis from Mindoro, nominotypical spilocephala from Mindanao and Basilan, N. s. reyi from Sulu and Bongao (Sulu Islands), and N. s. everetti from Siasi (Sulu Islands). Shortly afterwards, Delacour & Mayr (1945) united these six forms (plus two ,proxima and centralis, added by Mayr) as races of one species, Ninox philippensis, falling into three groups based on simple shared plumage characters (some of which were mistaken): (1) philippensis group (‘upperparts plain, underparts boldly striped’) — iV. p. philippensis (Luzon, possibly also Marinduque, Samar and Leyte), N. p. proxima (Ticao and Masbate) and N. p. centralis (Siquijor; apparently also Panay, Guimaras, Negros); (2) spilocephala group (‘head and neck spotted or barred, underparts striped or variegated’) — N. p. spilocephala (Mindanao, Basilan), N. p. reyi (Sulu, Tawi Tawi, Bongao) and N. p. everetti (Siasi); and (3) mindorensis group (‘head and neck spotted or barred; underparts entirely vermiculated or barred’) — N. p. mindorensis (Mindoro) and N. p. spilonota (Tablas, Sibuyan, Cebu). Sibley & Monroe (1990) accepted these groupings but rationalised them geographically north-south, making the mindorensis group (which, owing to chronological precedent, they called the spilonota group) second and spilocephala third. However, Dickinson et al. (1991) recast the arrangement (adding ticaoensis, described in 1972) rather more radically to give: N. p. philippensis (Luzon, Polillo, Catanduanes, Marinduque, Samar, Buad, Biliran and Leyte); N. p. spilocephala (Dinagat, Siargao, Mindanao and Basilan); N. p. reyi (Sulu, Siasi, Tawi Tawi and the adjacent Sanga 18 P.C. RASMUSSEN etal. Forktail 28 (2012) Sanga and Bongao, and Sibutu); N. p. centralis (Semirara, Carabao, Boracay, Panay, Guimaras, Negros, Siquijor and Bohol); N. p. spilonota (Tablas, Sibuyan, Cebu and Camiguin Sur); N. p.proxima (Masbate); N. p. ticaoensis (Ticao); and N. p. mindorensis (Mindoro). Kennedy et al. (2000) retained this treatment in spite of the queries by Collar & Rasmussen (1998) and the split of mindorensis by Konig et al. (1999). On the basis of our data (and reiterating our doubt about the validity of ticaoensis ) we propose the following arrangement (ranges derived from Dickinson et al. 1991, Dickinson 2003): Luzon Hawk Owl Ninox philippensis N. p. philippensis Biliran, Bohol, Boracay, Buad, Carabao, Catanduanes, Guimaras, Leyte, Lubang, Luzon, Marinduque, Masbate, Negros, Panay, Polillo, Samar, Semirara N p. ticaoensis Ticao N. p. centralis Siquijor Mindanao Hawk Owl Ninox spilocephala Mindanao, Dinagat, Siargao, Basilan Mindoro Hawk Owl Ninox mindorensis Mindoro Romblon Hawk Owl Ninox spilonota N. s. spilonota Sibuyan N s. fisher i Tablas Cebu Hawk Owl Ninox rumseyi Cebu Camiguin Hawk Owl Ninox leventisi Camiguin Sur Sulu Hawk Owl Ninox reyi Sulu, Siasi, Tawi Tawi and the adjacent Sanga Sanga and Bongao, Sibutu Figure 7 maps the newly defined species limits of the N. philippensis ( sensu lato ) complex. We use ‘Luzon’ rather than ‘Philippine’ as the common name for N. philippensis ( sensu stricto ) to avoid confusion with N. philippensis ( sensu lato). For N. spilonota ( sensu stricto) we use ‘Romblon’ because, although no member of this taxon is definitely known to occur on Romblon Island proper (there are unconfirmed reports), both Sibuyan and Tablas are part of Romblon province, so this seems the most appropriate geographical name for this species. Relationships How can we explain the curious circumstance in which rather similar-plumaged unstreaked Ninox populations are scattered on six well-separated islands or island groups within the Philippine archipelago while streak-breasted (or part-streak-breasted) populations occupy the largest islands ? One possibility, consistent with the speculation in Collar & Rasmussen (1998) that all unstreaked forms are monophyletic, is that they represent the remnants of an earlier stock of Ninox which occupied the archipelago before being displaced from almost all islands which had a Pleistocene land-bridge by a later invasion of streak-breasted birds. Militating against this notion is the fact that the streak¬ breasted birds are generally smaller than the unstreaked forms. Even so, Ninox spilonota of Tablas and Sibuyan, N. rumseyi of Cebu and N. mindorensis of Mindoro seem to form a monophyletic group. Their distribution mirrors that of some Philippine Hypsipetes bulbuls, of which the forms on Mindoro, Tablas, Romblon and Cebu are more closely related to each other than to the bulbuls of Negros, Panay, Greater Luzon and Greater Mindanao (Oliveros & Moyle 2010). Vocally spilonota, rumseyi and mindorensis all share certain characteristics including descending thin, squealed whistles and croaky hisses, and all are at least relatively high-pitched. Ninox mindorensis often particularly resembles the much larger rumseyi Figure 7. Map of species of the Ninox philippensis sensu lato species complex as recognised herein. of Cebu in plumage pattern, although it is typically more narrowly and clearly barred below, and has the throat-patch duller and less conspicuous. Ninox rumseyi, which is geographically surrounded by Ninox philippensis, can only be diagnosed morphologically from mindorensis and spilonota on a combination of characters, although its song is moderately autapomorphic. Both forms of Ninox spilonota lack both strong head pattern and markedly paler throat- patch, but one specimen ofiV. r. spilonota from Sibuyan (FMNH 358295) shows a white-streaked underparts pattern similar to many N. reyi specimens, and thus recalls a weakly marked N. spilocephala. The relationships of Mindanao’s spilocephala remain unclear. Its different voice and certain other features (e.g. well-developed auriculars) are inconsistent with its being the link (Mayr 1945) between the plain-headed, streak-breasted philippensis and the unstreaked taxa. On the other hand its Sulu replacement reyi, although completely different from any other Philippine taxon in voice, shows plumage characters in some individuals (dark teardrop¬ shaped streaking on lower underparts) reminiscent of spilocephala, hinting at a shared ancestry. Given reyi' s geographical isolation from all taxa except spilocephala, this inference is more parsimonious than the notion that reyi is more closely related to a more widespread unstreaked form. (The possibility that reyi is more closely related to Ninox ios of Sulawesi, based on the similarity noted above in the quality if not the pattern of their songs, is not supported by their very dissimilar morphologies; see Rasmussen 1999.) The derivation of Ninox leventisi of Camiguin Sur, despite the fact that this island lies just off Mindanao, is no less problematic. Although at least superficially similar in plumage to other unstreaked taxa, leventisi is the most highly autapomorphic of all taxa (although vocally N. reyi is even more so). In fact, leventisi Forktail 28 (201 2) Vocal divergence and new species in the Philippine Hawk Owl Ninox philippensis complex 19 and spilocephala are strikingly dissimilar to each other, in size, plumage texture and fullness, development of filamentous auricular extensions, pattern of lower underparts, and soft-part colours. The only character that links leventisi with spilocephala to the exclusion of other taxa is the low pitch of their songs, albeit these are very different in other ways. Indeed, the long solo song of leventisi (SOM) has closer resemblances to that of philippensis than to any other taxon, and it may well be more closely related to philippensis than to the geographically closer spilocephala or the morphologically closer spilonota. Resolution will have to await molecular analysis. For the moment, we note a high level of concordance between the species limits proposed here and zones of turnover proposed by Peterson (2006). A separate issue is that of generic limits of Ninox, which at least on morphology is probably a non-monophyletic group (Rasmussen 1999). A recent molecular phylogeny (Wink et al. 2009) shows a monop hyletic Ninox but taxon sampling therein was not dense. Although we think that all the species formerly united under Ninox philippensis are indeed fairly closely related, this remains to be established, most likely by molecular analyses. The conservation status of these various forms needs full evaluation elsewhere. However, it should be noted at once that five species — mindorensis , spilonota, rumseyi, leventisi and reyi — are likely to be at risk. On Mindoro, the Sablayan Penal Colony holds the largest remaining area of lowland forest on Mindoro, and is critically important for populations of lowland bird species endemic to Mindoro (Brooks etal. 1995a, Mallari cZTz/. 2001). The Mindoro Hawk Owl is also known from forest in Mts Iglit-Baco National Park (Gonzalez & Dans 1998), Naujan Lake National Park (Ticsay & Ledesma 1998), and Mt Halcon between 1,150-1,250 m (ROH). Satellite images taken in the late 1980s suggested Tablas then held two forest parcels together covering under 0.5 km2 (Goodman & Ingle 1993) and recent visits confirm that the amount of habitat there is tiny, although the species occurs in tall secondary as well as primary forest on the island (Allen 2006, DNSA, ROH). Meanwhile, although forest covered over half of Sibuyan’s 233 km2 land surface at the start of the 1 990s, logging was proceeding apace and, in the absence of intervention, all lowland areas were predicted to be cleared within a few years (Goodman & Ingle 1993); illegal logging inside Mt Guiting-guiting Natural Park continues (M. Wallbank verbally 2009 to DNSA). On Cebu, where it was rediscovered in 1998 after a gap of 110 years, the total population of Cebu Hawk Owls may be just 192 pairs (and possibly many fewer) scattered among 1 1 forest patches ( Jakosalem et al. in press). On Camiguin Sur forest covers just the centre of the small island, and numbers of the species cannot be high (Heaney & Tabaranza 2006). Finally, of the six islands in the Sulu archipelago from which the owl has been recorded, the current status of forest on Siasi appears on satellite photos to show that almost the entire island’s forest has been replaced by orchards; Sulu (Jolo) has apparently also been largely converted except around the volcanic cones; Sanga- Sanga’s last forest patch was cleared in 1992-1993; and Bongao retains only a small patch of forest on its sacred mountain (DNSA, ROH). Sibutu may have significant areas of secondary forest, although that seen by DNSA in 1995 was of low stature. The main island of Tawi Tawi still has much secondary and some primary forest (Dutson etal. 1996, DNSA, ROH). However, on Tawi Tawi N. reyi is also found in forest edge, mature mangroves and large trees in the vicinity of villages. ACKNOWLEDGEMENTS Work on early drafts of this paper in the mid-1990s benefited greatly from information supplied by T. M. Brooks, G. C. L. Dutson, S. J. Harrap, B. F. King, P. A.J. Morris, R. Ranft, R. J. Timmins and M. P. Walters. More recently we have had the help of f. A. Eaton, C. Espanola, the late T. Fisher, P. C. Gonzales, N. Icarangal, Jr., H. C. Miranda, Jr., M. Lagerqvist, F. E. Rheindt, I. Sarenas, P. Simpson, M. Thibault, C. Tipp and P. Verbelen. DNSA's work on Tawi Tawi was greatly facilitated by Governor Hadji Sadikul Sahali and his family, and by Eddie Ali Dean. Some of DNSA’s studies on Tablas and Tawi Tawi were sponsored by ZGAP (Zoologische Gesellschaft fur Arten- und Populationsschutz). For permission to examine specimens in their care and other assistance (institutions given in full in main text) we thank staff including P. R. Sweet, T. Trombone and M. Shanley (AMNH), L. Joseph (ANSP), R. P. Prys- Jones and M. P. Adams (BMNH), the late B. C. Livezey (CM), G. K. Hess and J. Woods (DMNH), D. E. Willard (FMNH), G. Lenglet (IRSNB), the late R. A. Paynter, A. Pirie and J. Trimble (MCZ), R. Zink (MMNH), C. Voisin (MNHN), R. W. R. J. Dekker (NCB), L. Alvarez (PNM), B. Millen (ROM), G. Mayr (SFN), the late S. Eck (SMTD), J. Hinshaw (UMMZ), C. Milensky (USNM), N. Rice (YPM) and S. Frahnert (ZMB). For permission to use their photographs in Figure 6 we thank C. Artuso, J. Eaton, R. J. Quisumbing and M. Thibault. We also thank the National Geographic Society Committee on Research and Exploration for financial support to PCR (NGS CRE 8919-11). Two referees, one of them Lawrence Heaney, the other anonymous, made very helpful comments to improve the manuscript. The colour plate is an original work by John Gale commissioned for this paper. REFERENCES Allen, D. (1998) On the birds of Tawi Tawi Province in the Philippines. Bull. Tsurumi Univ. 35: 73-154. Allen, D. (2006) New records and other observations of birds on the island of Tablas, Romblon province, Philippines. Forktail 22: 77-84. Balete, D. S., Tabaranza Jr., B. R. & Heaney, L. R. (2006) An annotated checklist of the birds of Camiguin Island, Philippines. Fieldiana Zool. No. 106: 58-72. Bonaparte, C. (1855) [Les principals especes nouvelles... dans son recent voyage en ecosse et en Angleterre.] C. R. Acad. Sci. Paris 41 : 651-661 . Bourns, F. S. & Worcester, D. C. (1894) Preliminary notes on the birds and mammals collected by the Menage Scientific Expedition to the Philippine Islands. Occas. Pap. Minnesota Acad. Nat. Sci. 1: 1-64. Brooks, T., Dutson, G., Gabutero, L. & Timmins, R. (1995a) Siburan - key area for birds on Mindoro. Oriental Bird Club Bull. 21 : 28-31 . Brooks,! M„ Magsalay, P., Dutson, G. & Allen, R. (1 995b) Forest loss, extinction and last hope for birds of Cebu. Oriental Bird Club Bull. 21 : 24-27. Collar, N. J. & Rasmussen, P. C. (1998) Species limits in the Ninox philippensis complex. Ostrich 69: 398 (Proc. 22 Internatn. Orn. Congr.). Collar, N. J., Mai lari, N. A. D. & Tabaranza, B. R. (1999) Threatened birds of the Philippines. Manila: Bookmark, Inc. & Haribon. Delacour, J. & Mayr, E. (1945) Notes on the taxonomy of the birds of the Philippines. Zoologica 30: 105-1 17. Dickinson, E. C., ed. (2003) The Howard & Moore complete checklist of the birds of the world. Third edition. London: Christopher Helm. Dickinson, E. C., Kennedy, R. S. & Parkes, K. C. (1991) The birds of the Philippines: an annotated check-list. Tring, U.K.: British Ornithologists' Union (Check-list no. 12). duPont, J. E. (1972) Notes on Philippine birds (no. 2): birds of Ticao. Nemouria 6. duPont, J. E. & Rabor, D. S. (1973) South Sulu Archipelago birds: an expedition report'! Nemouria 9. Dutson, G.C. L., Evans,! D., Brooks,! M„ Asane, D. C., Timmins, R. J. &Toledo, A. (1992) Conservation status of birds on Mindoro, Philippines. Bird Conserv. Internatn. 2: 303-325. Dutson, G., Allen, D. & Brooks, ! (1996) Tawi-Tawi - extreme Philippine birding. Oriental Bird Club Bull. 24: 32-35. Gonzales, J. C.T.& Dans, A.T. L. (1998) Birds and mammals of the fragmented forests along the Anahawin River, Mt. Iglit-Baco National Park, Mindoro Island, Philippines. Sylvatrop 8: 43-6! Goodman, S. M. & Ingle, N. R. (1993) Sibuyan Island in the Philippines - threatened and in need of conservation. Oryx 27: 1 74-1 80. 20 P. C. RASMUSSEN etal. Forktail 28 (2012) * Heaney, L. R. &Tabaranza, B. R. (2006) Mammal and landbird studies on Camiguin Island, Philippines: background and conservation priorities. Field iana Zool. 106: 1-13. Hutchinson, R„ Eaton, J. & Benstead, P. (2006) Observations of Cinnabar Hawk Owl Ninox ios in Gunung Ambang Nature Reserve, North Sulawesi, Indonesia, with a description of a secondary vocalisation. Forktail 22: 120-121. ICZN (2000) International Code of Zoological Nomenclature. 4th ed. International Trust for Zoological Nomenclature, London. Jakosalem, P. G. C., Collar, N. J. & Gill, J. A. (in press) Habitat selection and conservation status of the endemic Ninox hawk-owl on Cebu, Philippines. Bird Conserv. Internatn. Kennedy, R. S„ Gonzales, P. C., Dickinson, E. C., Miranda, Jr., H. C.& Fisher, T. H. (2000) A guide to the birds of the Philippines. Oxford: Oxford University Press. King, B. F. (2005) The song of Cinnabar Hawk Owl Ninox ios in North Sulawesi, Indonesia. Forktail 21: 173-174. Konig, C., Weick, F. & Becking, J.-H. (1999) Owls: a guide to the owls of the world. Robertsbridge, East Sussex, U.K.: Pica Press. Mallari, N. A. D„ Tabaranza, B. R. & Crosby, M. J. (2001) Key conservation sites in the Philippines: a Haribon Foundation & BirdLife International directory of Important Bird Areas. Makati City, Philippines: Bookmark, Inc. Mayr, E. (1945) Original descriptions of N. p. proximo (p.l 08) and N. p. centralis (p.l 1 0) in Delacour & Mayr (1945) above. McGregor, R. C. (1905) Birds from the islands of Romblon, Sibuyan, and Cresta de Gallo. Bull. Bureau Gov. Lab. Manila 25: 5-23. McGregor, R. C. (1909-1910) A manual of Philippine birds. Manila: Bureau of Printing. Ogilvie Grant, W. R. (1896a) On the birds of the Philippine Islands. Part VII. The highlands of Mindoro. Ibis (7)2: 457-477. Oliveros, C. H. & Moyle, R. G. (2010) Origin and diversification of Philippine bulbuls. Mol. Phyl. Evol. 54: 822-832. Oustalet, E. (1880) Description de deux oiseaux nouveaux des lies Souloo. Bull. Hebdo. Assoc. Sci. France 2: 205-206. Paguntalan, L. M. J. & Jakosalem, P. G. (2008) Significant records of birds in forests on Cebu island, central Philippines. Forktail 24: 48-56. Peters, J. L. (1940) Check-list of birds of the world, 4. Cambridge, Mass.: Harvard University Press. Peterson, A.T. (2006) Taxonomy is important in conservation: a preliminary reassessment of Philippine species-level taxonomy. Bird Conserv. Internatn. 16: 155-173. Rasmussen, P. C. (1999) A new species of hawk owl Ninox from Indonesia. Wilson Bull. 111:457-464. Rasmussen, P. C. & Anderton J. C. (2005) Birds of South Asia: the Ripley guide. Washington D.C. and Barcelona: Smithsonian Institution and Lynx Edicions. Raven (2012) Raven interactive sound analysis software. Available athttp:/ /www.birds.cornell.edu/brp/raven/RavenVersions.html#Ravenl4 Sharpe, [R.] B. (1897) [Two species of owls.] Bull. Brit. Orn. Club 6: 47. Sibley, C. G. & Monroe, B. L. (1990) Distribution and taxonomy of birds of the world. New Haven: Yale University Press. SYSTAT (2012) Statistical and graphical software. Available at: http:// www.systat.com/SystatProducts.aspx Ticsay, M. V. & Ledesma, M. M. (1998) Faunal inventory of Naujan Lake National Park and its adjacent watershed, Oriental Mindoro, Philippines. Sylvatrop 8: 62-78. Tobias, J. A., Seddon, N„ Spottiswoode, C. N., Pilgrim, J. D„ Fishpool, L. D. C. & Collar, N. J. (2010) Quantitative criteria for species delimitation. Ibis 152: 724-746. Tweeddale, Marquis of (1878 [1879]) Contributions to the ornithology of the Philippines. No XL On the collection made by Mr A. H. Everett at Zamboanga, in the island of Mindanao. Proc. Zool. Soc. London 1878: 936-954. Wink, M., El-Sayed, A.-A„ Sauer-Gurth, H. & Gonzalez, J. (2009) Molecular phylogeny of owls (Strigiformes) inferred from DNA sequences of the mitochondrial Cytochrome b and the nuclear RAG-1 gene. Ardea 97: 581-591. P. C. RASMUSSEN, Dept, of Zoology and Michigan State University Museum, Michigan State University, East Lansing, Ml 48824, USA; and Bird Croup, Department of Zoology, Natural History Museum, Akeman St, Tring, Herts HP23 6AP, UK. Email rasmus39@gmail.com D. N. S. ALLEN, 97 Sussex Way, London N7 6RU, UK N. J. COLLAR, BirdLife International, Wellbrook Court, Girton Road, Cambridge CB3 0NA, UK; and Bird Group, Department of Zoology, Natural History Museum, Akeman St, Tring, Herts HP23 6AP, UK B. DeMEULEMEESTER, Koningin Astridlaan 154, 9000 Gent, Belgium. Email bram_dm@yahoo.co.uk R. O. HUTCHINSON, 26 Sutton Avedue, Chellaston, Derby DE73 6RJ, UK. Email robhutchinson@birdtourasia.com P. G. C. JAKOSALEM, Philippines Biodiversity Conservation Foundation, Inc. c/o Negros Forest Ecological Foundation, Inc. South Capitol Road, Bacolod City, 6100 Philippines R. S. KENNEDY, Frederick and Amey Geier Collections and Research Center, Museum of Natural History & Science, Cincinnati Museum Center, 1301 Western Ave, Cincinnati, OH 45203; and Ornithology Department, Museum of Comparative Zoology, Harvard University, 26 Oxford St, Cambridge, MA 02138; current address 18 Riverview Rd, Durham NH 03824, USA F. R. LAMBERT, El 802 A Perdana Condo, Jalan PJU 8/1, Damansara Perdana, Petaling Jaya, 47820 Selangor, Malaysia L. M. PAGUNTALAN, Philippines Biodiversity Conservation Foundation, Inc. c/o Negros Forest Ecological Foundation, Inc. South Capitol Road, Bacolod City, Philippines Supplementary online material To be found under Forktail 28 page on www.orientalbirdclub.org. SOM 1. Additional songs of taxa in the Philippine Hawk Owl Ninox pbilippensis ( sensu Into ) complex. SOM la.12453: short isolated duetted segment of song of Philippine Hawk Owl N. p. pbilippensis, AV# 12453, ROH, Luzon; SOM lb. 1355 1 : long song by a single bird, joined near the end with another in a presumed duet; Philippine Hawk Owl N. p. pbilippensis , AV# 1 355 1 . ROH, Luzon. SOM 1 c.8972: series of short duetted strophes by Philippine Hawk Owl N. p. centralis , F. Verbelen, Bohol; SOM ld-e. 10700; long series by Philippine Hawk Owl N. p. centralis , FRL, Negros; SOM If. 1 12a: long series by Philippine Hawk Owl/V. p. centralis , DNSA, Siquijor; SOM lg.99: three short duetted strophes by Philippine Hawk Owl N. p. centralis , DNSA, Siquijor, with many hoarse notes. SOM lh. 13557: duetted series of short strophes with stray notes between main strophes; Camiguin Hawk Owl Ninox sp. 1. SOM li: single Camiguin Hawk Owl Ninox sp. 1 singing, giving several more notes/strophe than in Figure 2c. SOM lj: continuation of long song of Mindanao Hawk Owl N. spilocepbala , starting where Figure 2d ends. SOM lk: duet by Sulu Hawk Owl N. reyi , series of short variable strophes. SOM 11: continuation of duet of Mindoro Hawk Owl N. mindorensis, starting where Figure 2gends; SOM lm-n: whistled duet of Mindoro Hawk Owl N. mindorensis , with many highly frequency-modulated notes. SOM lo: Variant song and note-types of Romblon Hawk Owl Ninox spilonota newssp. on Tablas; SOM lp.l 1510: duetted song type of Romblon Hawk Owl Ninox spilonota on Sibuyan. SOM lq-r: various alternate song types of Cebu Hawk Owl Ninox new sp. 2. FORKTAIL 28 (2012): 21-28 Bird species diversity in five protected areas of Bangladesh M. MONIRUL H. KHAN & NASIM AZIZ Birds of five protected areas (Lawachara, Satchari, Rema-Kalenga, Chunati and Teknaf ) in north-east and south-east Bangladesh were studied during 2005-2008 by strip transect sampling and opportunistic surveys. Of 239 species of birds recorded, 189 were residents, 39 winter visitors, 6 summer visitors and 5 vagrants; 40 (17%) were 'Very Common', 66 (28%) 'Common', 48 (20%) 'Fairly Common' and 85 (35%) 'Uncommon'. Population densities of eight species, selected as indicators of forest condition, were estimated; two understorey species, Red Junglefowl Gallus gallus and Puff-throated Babbler Pellorneum ruficeps, increased in the study period, perhaps owing to understorey regeneration increasing the carrying capacity and nesting sites, plus (for the junglefowl) reduced hunting pressure. However, illegal logging of timber trees continues in some areas, probably causing the decline of Oriental Pied Hornbill Anthracoceros albirostris. Densities of the other five indicators (Red-headed Trogon Harpactes erythrocephalus, Greater Racket-tailed Drongo Dicrurus paradiseus, White-rumped Shama Copsychus malabaricus, Hill Myna Gracula religiosa and White-crested Laughingthrush Garrulax leucolophus) remained more or less unchanged. INTRODUCTION Bangladesh has a network of protected areas under the jurisdiction of the Forest Department, but very little information is available on the birds of these areas. Moreover, there had been no attempt to use birds as indicators of ecological change in the protected areas, although birds are one of the best indicators of such change (Morrison 1986, Temple & Wiens 1989, Browder et al. 2002). Figure 1 . Forested areas of Bangladesh showing the locations of five NSP (Nishorgo Support Project) sites where the survey was conducted. Panchagarh Thakurgaon ® Nilphamari Dinajpur 0 Ranggi ® Kurigran r © Gaibandha © \ Joypurhat © Chapai Nawabganj © Nator Naogaon Bogra 0 Sherpur © . , Netrakona Jamalpur © Sunamgani © T Sylket W 4“ Forest Types Mixed Evergreen Forest ■ Deciduous Forest 1 Mangrove Forest Others © District Headquarter .. . River or Sea District Boundary 100 100 Kilometer Scale 1:3,80,000 91 Different species of bird occur in different vertical strata of an area and are adapted to specific types of plant and animal food. Therefore, avian population density and species diversity reflect the changes in their habitat conditions, and indicate the health of different vertical strata of the forest. Determining the extent of change in ecological systems is critical to the long-term conservation of biotic diversity (Canterbury et al. 2000). In its 147,570 km2 Bangladesh harbours nearly 700 species of bird, which is 50% of the total of the Indian subcontinent and 7% of the world’s total (Harvey 1990, Khan 2008). However, the natural forests and other wilderness areas of the country are under great pressure from legal and illegal overexploitation, together with improper management. Forest cover has been reduced by more than 50% since the 1970s (IUCN-Bangladesh 2000). Bangladesh has less than 0.02 ha of forest land per person — one of the lowest forest- to-population ratios in the world (IUCN-Bangladesh 2000). The urgent need to conserve the remaining tracts and their inhabitants, O O Table 1. Strip transect locations in five NSP sites where the bird survey was conducted. Name of project site Transect location Geographic locations of transect ends Transect length (km) Lawachara National Park Eastern 24°1 9.9'N 91 °47.6'E; 24°20.2'N 91 °47.5'E 0.50 (area: 1,250 ha; year of Central 24°19.7'N 91°47.2'E; 24°19.8'N 91°47.5'E 0.61 establishment: 1996) Central 24°1 9.8'N 91°47.2'E; 24°20.2'N 91 °47.2'E 0.50 Central 24°19.5'N 91047.2'E;24°19.7'N 91°47.6'E 0.70 Western 24°19.2'N 91°47.TE; 24°19.4'N 91°46.8'E 0.52 Western 24°1 8.8'N 91 °46.4'E; 24°1 9.1 'N 91°46.9'E 0.89 Satchari National Park Central 24°07.5'N 91°26.7'E; 24°06.6'N 91°27.2'E 1.94 (area: 243 ha; year of Central 24°07.6'N 91°27.0'E; 24°07.3'N 91°27.2'E 0.56 establishment: 2006) Northern 24°07.4'N 91°26.7'E; 24°07.5'N 9r27.0'E 0.50 Rema-Kalenga Northern 24°10.7'N 91°37.6'E; 24°09.6'N 91°38.0'E 2.02 Wildlife Sanctuary Central 24°09.6'N 91 °38.0'E; 24°09.8'N 91 °37.5'E 0.78 (area: 1,795 ha; year of Northern 24°10.2'N 91°37.5'E; 24°10.3'N 91°37.9'E 0.80 establishment: 1996) Southern 24°06.9'N 91 °37.5'E; 24°06.4'N 91 °37.8'E 1.11 Chunati Wildlife Eastern 21°55.4'N 92°03.5'E; 21°55.3'N 92°02.7'E 1.41 Sanctuary Central 21°55.3'N 92°02.7'E; 21°55.5'N 92°02.3'E 0.76 (area: 7, 764 ha; year of Eastern 21°55.7'N 92°02.5'E; 21°56.1'N 92°03.5'E 1.91 establishment: 1986) Northern 21°57.3'N 92°04.1'E; 21°57.2'N 92°03.7'E 0.65 Northern 21°57.2'N 92°03.7'E;21°57.4'N 92°04.0'E 0.65 Teknaf Game Reserve Northern 21°05.8'N92°09.8'E;21°05.2,N92°10.2'E 1.25 (area: 11,615 ha; year of Northern 21°05.2'N 92°10.2'E; 21°05.4'N 92°09.5'E 1.27 establishment: 1983) Northern 21 °06.3'N 9201 1 ,7'E; 21°05.5'N 92°1 0.8'E 0.74 Central 21°05.2'N 92°1 1 ,9'E; 21°03.9'N 92°1 1 ,6'E 2.49 Central 21 °03.9'N 92°1 1 ,6'E; 21 °04.5'N 92°1 1 ,9'E 1.21 22 M. MONIRUL H. KHAN & NASIM AZIZ Forktail 28 (2012) while simultaneously developing non-consumptive uses, gave rise to the Forest Department’s five-year Nishorgo Support Project (NSP), which was implemented by the International Resources Group (IRG) of Washington DC, together with its local partners. The work reported here was supported by NSP, which used the results as a tool to understand the impact of newly introduced co-management on live protected areas. In our survey, the temporal changes of population densities of eight indicator species of bird were studied so that the ecological health of five protected areas could better be monitored. All species of bird were recorded, together with their relative abundance. The study was conducted on a participatory basis, involving birdwatchers, local communities and Forest Department officials in order that everybody would become aware of what is happening to the birds in these sites. There are 28 protected areas in Bangladesh with a total area of 8,718.87 km:, covering only 5.9% of the total area of Bangladesh. All the five protected areas (Lawachara, Satchari, Rema-Kalenga, Chunati and Teknaf; Fig. 1, Table 1) where the bird survey was conducted are mainly covered by mixed evergreen forests, but the areas also have some bushy and bamboo-covered areas. METHODS Survey methods were selected on the basis ol simplicity and effectiveness so that local stakeholders, including people with little or no education, could participate. Strip transect sampling (Buckland et al. 2001) and opportunistic survey methods were selected. The survey was conducted in the breeding season (February- August), so that there would be more bird activity. The baseline survey was conducted in 2005, and was repeated in 2006, 2007 and 2008, in the same season and following the same transects, so that the results could be compared across different years. Strip transect samplingwas found most suitable to estimate the population densities of eight indicator species of bird (see below for choice rationale). In this method the observers slowly walked (c.1.5 km/hr) in relatively straight lines through the study areas and counted the indicator birds from both sides. The observation- range on each side (half-width of the strip) was 20 m, which was found suitable in terms of visibility. Thus the strip width was 40 m. The initial location of the bird was recorded. The observation-range was measured by an infrared rangefinder (Bushnell Yardage Pro). Any bird sighted beyond the pre-decided observation-range or coming from behind the observer was not counted. The survey was conducted in early mornings and late afternoons when the birds were most active. This method assumes that all birds in the strip are recorded. Transects were located mainly in the rich parts of the protected areas where the bird habitats were least disturbed. Depending on reserve size, the total lengths of transects in each site varied from 3.00 to 6.96 km (Table 1), but transects were periodically repeated 12-15 times per year. The eight indicator species of bird, suggested at a meeting of experts organised in IRG, Dhaka, were: Red Junglefowl Gallus gallus , Oriental Pied Hornbill Anthracoceros albirostris, Red-headed Trogon Harpactes erythrocephalus , Greater Racket-tailed Drongo Dicrurus paradiseus , White-rumped Shama Copsy clous malabaricus , Hill Myna Gracula religiosa , White-crested Laughingthrush Garrulax leucolophus and Puff-throated Babbler Pellorneum ruficeps. They were selected because: (1) they are primarily forest birds; (2) they live in different vertical strata of the forest (junglefowl, laughingthrush and babbler in lower stratum; trogon, drongo and shama in middle stratum; and hornbill and myna in upper stratum); (3) they are noisy (and thus less likely to be missed during counts); and (4) they are breeding residents. It was assumed that the improvement or degradation of the forest condition would have a direct impact on the feeding and breeding of these species, which in turn would show changes in population densities (Morrison 1986, Temple & Wiens 1989, Canterbury et al. 2000, Browder et al. 2002, Lammertink et al. 2009). Any important or interesting observation or information relating to diet, foraging guild and threats was recorded opportunistically at any time in the field. The birds were identified from Ali & Ripley (1987), Grimmett et al. (1998) and Rasmussen & Anderton (2005). Relative abundance was assessed by classifying sighting frequencies into four: ‘Very Common’ — recorded on 76- 100% observation-days, ‘Common’ — recorded on 51-75% observation-days, ‘Fairly Common’ — recorded on 26-50% observation-days, and ‘Uncommon’ — recorded on 25% or fewer observation-days. RESULTS During the avian breeding season (February- August) of four consecutive years (2005-2008), 239 species of bird were recorded in the five protected areas, of which 189 were residents, 39 winter visitors, 6 summer visitors and 5 vagrants (see Appendix). The relative abundance shows that 40 (17%) species were ‘Very Common’, 66 (28%) ‘Common’, 48 (20%) ‘Fairly Common’ and 85 (35%) ‘Uncommon’. Based on principal diet 131 (55%) species were insectivorous, 32 (13%) carnivorous, 29 (12%) omnivorous, 23 (10%) granivorous and herbivorous, 17 (7%) frugivorous, and 7 (3%) nectarivorous. Based on principal foraging guild 100 (42%) species were middle-canopy foragers, 78 (33%) ground foragers, 36 (15%) upper-canopy foragers, and 25 (10%) undergrowth and bush foragers. The site lists show that the total number of species and the total number of primarily forest species for Lawachara were, respectively, 167 and 90 (54%), Satchari 1 53 and 84 (55%), Rema-Kalenga 206 and 91 (44%), Chunati 162 and 53 (33%), and Teknaf 188 and 73 (39%). The percentages indicate how the ratio of total species to total forest species varied across the five sites. Annual mean density estimates from 2005 to 2008 revealed that two understorey indicator species (Red Junglefowl and Puff- throated Babbler) increased at every site and at all sites combined (Fig. 2). Oriental Pied Hornbill declined and the other five indicator species remained more or less unchanged (Fig. 2). Three species of global conservation concern were recorded: White-rumped Vulture Gyps bengalensis (Critically Endangered; nesting colony found in Rema-Kalenga), Lesser Adjutant Leptoptilos javanicus (Vulnerable; one adult in Rema-Kalenga, 10 March 2008) and White-cheeked Partridge Arborophila atrogularis (Near Threatened; parent and hatchling in Lawachara, 15 March 2008). Eleven nationally threatened species (according to IUCN- Bangladesh 2000; since the list is out of date the White-rumped Vulture is not included) were recorded: Kalij Pheasant Lophura leucomelanos, Grey Peacock Pheasant Polyplectron bicalcaratum , Oriental Pied Hornbill, Red-headed Trogon, Dollarbird Eurystomus orientalis , Spot-bellied Eagle Owl Bubo nipalensis. Figure 2. Comparison of the population density of eight indicator bird species across five NSP sites during 2005-2008. ■ In 2005 □ In 2006 □ In 2007 □ In 2008 Forktail 28 (2012) Bird species diversity in five protected areas of Bangladesh 23 Brown Fish Owl Ketupa zeylonensis. Tawny Fish Owl Ketupa flavipes, Malayan Night Heron Gorsachius melanolophus , Lesser Adjutant and Streaked Spiderhunter Aracbnothera magna. DISCUSSION The total bird species (239) recorded in five NSP sites represents over 30% of the birds recorded in Bangladesh (Khan 2008, Siddiqui et al. 2008), and almost 18% recorded in the Indian Subcontinent (Grewal et al. 2002). The species were recorded mainly during transects while counting indicator birds, so some may have been missed either because they occurred away from the transects or because of observer inattention. The cryptic nature of some birds might also have contributed to the crudeness of the relative abundance. An increase in density of two indicator species (Red Junglefowl and Puff-throated Babbler) would suggest that the forest understorey is regenerating and increasing the carrying capacity and nesting sites for these two species. Community patrolling and awareness, and other programmes conducted by NSP has played a key role in reducing understorey firewood exploitation and hunting pressure. However, White-crested Laughingthrush did not respond like the junglefowl and babbler, possibly having different limiting factors; it is a rare species in Bangladesh and was found only in Chunati, which is more open compared to four other NSP sites. Illegal removal of timber trees and forest fruits still persists, and probably caused the decline of the Oriental Pied Hornbill over the four years. This bird lives in the upper canopy and suffers severely if large trees are removed from the forest. Habitat loss remains the main threat to birds in all five NSP sites. Illegal felling of trees and bamboo, and conversion of natural forests to monoculture plantations and agricultural, fields, were witnessed during the survey. Hunting and trapping of birds, together with the taking of nestlings as cagebirds (particularly mynas, parakeets and hornbills), is another threat. Large-scale illegal harvest of forest fruits, particularly Artocarpifs chaplasha and Bixa sp., is a growing threat for frugivorous birds. Moreover, the number of visitors in Lawachara and Satchari is becoming excessive. These threats should be reduced in order to maintain a healthy status of birds in the study’s five protected areas. ACKNOWLEDGEMENTS The authors are grateful to Nishorgo Support Project, Bangladesh Forest Department, together with United States Agency for International Development, International Resources Group (IRG) and its partners in Bangladesh (CODEC, RDRS, NACOM and IUCN) for providing necessary support to this work. Sincere thanks to Philip DeCosse and Paul Thompson, who provided strong support and advice during the survey. Also thanks to all volunteers (particularly Dhrubo Kundu, Shafiqur Rahman, Ahsanul Haque, Mustafezur Rahman, Samiul Mohsanin, Shehab Rayhan and Jewel Ahmed) and local participants. REFERENCES Ali, S. & Ripley, S. D. (1987) Compact handbook of the birds of India and Pakistan together with those of Bangladesh, Nepal, Bhutan and Sri Lanka. 1 0 vols. Delhi, India: Oxford University Press. Browder, S. F. Johnson, D. H. & Ball, I. J. (2002) Assemblages of breeding birds as indicators of grassland condition. Ecol. Indicators 2: 257-270. Buckland, S.T., Anderson, D. R., Burnham, K. P., Laake, J. L., Borchers, D. L. & Thomas, L. (2001) Introduction to distance sampling: estimating abundance of biological populations. Oxford, UK: Oxford University Press. Canterbury, G. E„ Martin, T. E„ Petit, D. R„ Petit, L. J. & Bradford, D. F. (2000) Bird communities and habitat as ecological indicators of forest condition in regional monitoring. Conserv. Biol. 14(2): 544-558. Grewal, B., Harvey, B. & Pfister, 0. (2002) A photographic guide to the birds of India including Nepal, Sri Lanka, The Maldives, Pakistan, Bangladesh and Bhutan. London, UK: Christopher Flelm. Grimmett, R„ Inskipp, C.& Inskipp, T. (1 998) Birds of the Indian subcontinent. Delhi, India: Oxford University Press. Harvey, W. G. (1990) Birds in Bangladesh. Dhaka, Bangladesh: University Press Limited. lUCN-Bangladesh (2000) Red book of threatened birds of Bangladesh. Dhaka, Bangladesh: IUCN -The World Conservation Union. Khan, M. M. H. (2008) Protected areas of Bangladesh: a guide to wildlife. Dhaka, Bangladesh: Nishorgo Program, Bangladesh Forest Department. Lammertink, M., Prawiradilaga, D. M., Setiorini, U., Naing, T. Z., Duckworth, J. W„ & Menken, S. B. J. (2009) Global population decline of the Great Slaty Woodpecker (Mulleripicus pulverulentus). Biol. Conserv. 142: 1 66— 179. Morrison, M. L. (1986) Bird populations as indicators of environmental changes. Current Ornith. 3:429-451. Rasmussen, P. C. & Anderton, J. C. (2005) Birds of South Asia: the Ripley guide. Vols. 1 and 2. Washington DC, USA: Smithsonian Institute, and Barcelona, Spain: Lynx Edicions. Siddiqui, K. U., Islam, M. A., Kabir, S. M. H., Ahmad, M., Ahmed, A. T. A., Rahman, A. K. A., Haque, E. U., Ahmed, Z. U., Begum, Z. N.T., Hassan, M. A., Khondker, M. & Rahman, M. M., eds. (2008) Encyclopedia of flora and fauna of Bangladesh, 26, Birds. Dhaka, Bangladesh: Asiatic Society of Bangladesh. Temple, S. A. & Wiens, J. A. (1989) Bird populations and environmental changes: can birds be bio-indicators? Amer. Birds 43: 260-270. M. Monirul H. KHAN, Deportment of Zoology, Jahanglrnagor University, Savor, Dhoko 1342, Bangladesh. Email: mmhkhan@hotmail.com Nasim AZIZ, International Resources Group - Bangladesh, House # 68, Road # 1, Block # 1, Banani, Dhaka 1213, Bangladesh. Present address: lUCN-Bangladesh, House # 1 1, Road # 138, Gulshan 1, Dhaka 1212, Bangladesh. Email: nasim.aziz@iucn.org 24 M. MONIRUL H. KHAN & NASIM AZIZ Forktail 28 (2012) Appendix List of bird species recorded in five NSP sites in the breeding season (February-August) during 2005-2008 Distribution: Wl - wide (all NSP sites), L - Lawachara, S - Satchari, RK - Rema-Kalenga, C - Chunati, and T - Teknaf. * Primarily forest species. Species Principal diet Principal foraging guild Status Distribution Blue-breasted Quail Coturnix coromandelica granivoreand herbivore ground fairly common, resident C, T, RK White-cheeked Partridge Arboropbila atrogularis* granivoreand herbivore ground uncommon (in NSP sites), resident L, S, RK Red Junglefowl Gallus gallus* granivoreand herbivore ground common, resident Wl Kalij Pheasant Lophura leucomelanos* granivoreand herbivore ground fairly common, resident Wl Grey Peacock Pheasant Polyplectron bicalcaratum* granivoreand herbivore ground uncommon (in NSP sites), resident T, RK Lesser Whistling-duck Dendrocygna javanica granivoreand herbivore ground common, resident T, RK, C Cotton Pygmy-goose Nettapus coromandelianus granivoreand herbivore ground uncommon (in NSP sites), resident T Barred Buttonquail Turnixsuscitator granivoreand herbivore ground fairly common, resident C, T, RK Eurasian Wryneck lynx torquilla insectivore ground fairly common, winter visitor C, T, RK White-browed Piculet Sasia ochracea* insectivore middle canopy uncommon (in NSP sites), resident S, RK Rufous Woodpecker Celeus brachyurus insectivore middle canopy common, resident Wl Great Slaty Woodpecker Mulleripicus pulverulentus* insectivore upper canopy uncommon (in NSP sites), resident T, C Grey-capped Pygmy Woodpecker Dendrocopos canicapillus* insectivore middle canopy fairly common, resident Wl Fulvous-breasted Woodpecker Dendrocopos macei insectivore middle canopy very common, resident Wl Lesser Yellownape Picus chlorolophus* insectivore upper canopy uncommon (in NSP sites), resident L, C Greater Yellownape Picus flavinucha* insectivore upper canopy common, resident Wl Grey-headed Woodpecker Picus canus insectivore upper canopy uncommon (in NSP sites), resident L, S Black-rumped Flameback Dinopium benghalense insectivore middle canopy very common, resident Wl Greater Flameback Chrysocolaptes lucidus* insectivore upper canopy very common, resident Wl Lineated Barbet Megalaima lineata frugivore upper canopy very common, resident Wl Blue-throated Barbet Megalaima asiatlca frugivore upper canopy very common, resident Wl Blue-eared Barbet Megalaima australis* frugivore upper canopy fairly common, resident L, S, RK, T Coppersmith Barbet Megalaima haemacephala frugivore middle canopy very common, resident Wl Oriental Pied Hornbill Anthracoceros albirostris* frugivore upper canopy fairly common, resident L, S, RK, T Common Hoopoe Upupa epops insectivore ground common, resident C, T, RK Red-headed Trogon Harpactes erythrocephalus* insectivore middle canopy uncommon (in NSP sites), resident L, S, RK, T Indian Roller Coracias benghalensis insectivore ground common, resident C, T, RK Dollarbird Eurystomus orientalis* insectivore upper canopy uncommon (in NSP sites), vagrant (in Bangladesh) L, S, RK, T Common Kingfisher Alcedo atthis carnivore (inch piscivore) ground (water) common, resident Wl Oriental Dwarf Kingfisher Ceyx erithacus* carnivore (inch piscivore) ground (water) uncommon (in NSP sites), vagrant (in Bangladesh) L, S White-throated Kingfisher Halcyon smyrnensis carnivore (inch piscivore) ground (water) common, resident Wl Pied Kingfisher Ceryle rudis carnivore (inch piscivore) ground (water) uncommon (in NSP sites), resident T Blue-bearded Bee-eater Nyctyornls athertoni* insectivore upper canopy uncommon (in NSP sites), resident Wl Green Bee-eater Merops orientalis insectivore middle canopy very common, resident Wl Blue-tailed Bee-eater Merops philippinus* insectivore upper canopy common, resident Wl Chestnut-headed Bee-eater Merops leschenaulti* insectivore upper canopy very common, resident Wl Pied Cuckoo Clamator jacobin us insectivore middle canopy uncommon (in NSP sites), summer visitor Wl Chestnut-winged Cuckoo Clamator coromandus * insectivore middle canopy uncommon (in NSP sites), vagrant (in Bangladesh) L, S, RK Common Hawk Cuckoo Hierococcyx varius insectivore middle canopy very common, resident Wl Indian Cuckoo Cuculus micropterus insectivore middle canopy common, summer visitor Wl Plaintive Cuckoo Cacomantis merulinus insectivore middle canopy common, resident Wl Asian Emerald Cuckoo Chrysococcyx maculatus * insectivore middle canopy uncommon (in NSP sites), summervisitor s Violet Cuckoo Chrysococcyx xanthorhynchus* insectivore middle canopy uncommon (in NSP sites), summervisitor s Drongo Cuckoo Surniculuslugubris* insectivore upper canopy common, resident L, S, RK, T Asian Koel Eudynamys scolopacea insectivore middle canopy very common, resident Wl Green-billed Malkoha Phaenicophaeus tristis* insectivore middle canopy very common, resident Wl Forktail 28 (2012) Bird species diversity in five protected areas of Bangladesh 25 Species Principal diet Principal foraging guild Status Distribution Greater Coucal Centropus sinensis insectivore ground very common, resident Wl Lesser Coucal Centropus bengalensis* insectivore upper canopy common, resident Wl Vernal Hanging Parrot Loriculus vernalis* granivoreand herbivore middle canopy fairly common, resident L, S, RK, T Rose-ringed Parakeet Psittacula krameri frugivore upper canopy common, resident Wl Blossom-headed Parakeet Psittacula roseata* frugivore upper canopy fairly common, resident L, S, RK Red-breasted Parakeet Psittacula alexandri* frugivore middle canopy very common, resident Wl Asian Palm Swift Cypsiurus balasiensis insectivore upper canopy (mid-air) common, resident C, T, RK Fork-tailed Swift Apus pacificus insectivore upper canopy (mid-air) uncommon (in NSP sites), winter visitor T Oriental Scops Owl Otussunia* insectivore middle canopy uncommon (in NSP sites), resident L, S, RK Collared Scops Owl Otus bakkamoena insectivore middle canopy uncommon (in NSP sites), resident Wl Spot-bellied Eagle Owl Bubo nipalensis* carnivore (inch piscivore) middle canopy uncommon (in NSP sites), resident T Dusky Eagle Owl Bubo coromandus* carnivore (inch piscivore) middle canopy uncommon (in NSP sites), resident RK Brown Fish Owl Ketupa zeylonensis carnivore (inch piscivore) middle canopy uncommon (in NSP sites), resident Wl Tawny Fish Owl Ketupa flavipes * carnivore (inch piscivore) middle canopy uncommon (in NSP sites), resident RK, T Brown Wood Owl Strix ieptogrammica* carnivore (inch piscivore) middle canopy uncommon (in NSP sites), resident L, RK Asian Barred Owlet Glaucidium cuculoides* insectivore middle canopy common, resident Wl Spotted Owlet Athene brama insectivore middle canopy very common, resident Wl Brown Hawk Owl N inox scutulata insectivore middle canopy common, resident Wl Large-tailed Nightjar Caprimulgus macrurus* insectivore middle canopy (mid-air) common, resident Wl Rock Pigeon Columba livia granivoreand herbivore ground common, resident Wl Green Imperial Pigeon Ducula aenea* frugivore upper canopy uncommon (in NSP sites), resident L, S, RK Oriental Turtle Dove Streptopelia orientalis* granivoreand herbivore middle canopy uncommon (in NSP sites), resident L, S, RK Spotted Dove Streptopeiia chinensis granivoreand herbivore ground very common, resident Wl Red Collared Dove Streptopelia tranquebarica granivore and herbivore ground common, resident Wl Eurasian Collared Dove Streptopelia decaocto granivoreand herbivore ground common, resident Wl Barred Cuckoo Dove Macropygia unchall* granivoreand herbivore middle canopy uncommon (in NSP sites), resident S Emerald Dove Chalcophapsindica* granivoreand herbivore ground very common, resident Wl Orange-breasted Green Pigeon Treron bicincta* frugivore middle canopy uncommon (in NSP sites), resident L, S, RK Pompadour Green Pigeon Treron pompadora* frugivore middle canopy common, resident Wl Thick-billed Green Pigeon Treron curvirostra* frugivore middle canopy uncommon (in NSP sites), resident L, S, RK Yellow-footed Green Pigeon Treron phoenicoptera frugivore middle canopy common, resident Wl Wedge-tailed Green Pigeon Treron sphenura* frugivore upper canopy uncommon (in NSP sites), resident L, S, RK White-breasted Waterhen Amaurornis phoenicurus insectivore ground fairly common, resident RK, C, T Pintail Snipe Gallinago stenura insectivore ground uncommon (in NSP sites), winter visitor RK, C, T Common Snipe Gallinago gallinago insectivore ground uncommon (in NSP sites), winter visitor RK, C, T Green Sandpiper Tringa ochropus insectivore ground uncommon (in NSP sites), winter visitor RK, C,T Wood Sandpiper Tringa glareola insectivore ground common, winter visitor RK, C, T Common Sandpiper Actitis hypoleucos insectivore ground common, winter visitor RK, C, T Greater Painted-snipe Rostratula benghalensis insectivore ground fairly common, resident RK, C, T Bronze-winged Jacana Metopidius indicus granivoreand herbivore ground fairly common, resident C Little Ringed Plover Charadriusdubius insectivore ground uncommon (in NSP sites), winter visitor C Red-wattled Lapwing Vanellus indicus insectivore ground fairly common, resident RK, C, T Little Tern Sterna albifrons carnivore (inch piscivore) ground (water) fairly common, resident T Whiskered Tern Chlidoniashybridus carnivore (inch piscivore) ground (water) fairly common, winter visitor T Osprey Pandion haiiaetus carnivore (inch piscivore) ground (water) uncommon (in NSP sites), winter visitor T Jerdon's Baza Aviceda jerdoni* carnivore (inch piscivore) upper canopy uncommon (in NSP sites), resident L, S, RK Black Baza Aviceda leuphotes* carnivore (inch piscivore) upper canopy fairly common, resident L, S, RK, T Oriental Honey-buzzard Pernis ptilorhynchus insectivore (while feeding honey-comb) middle canopy fairly common, resident winter visitor Black-shouldered Kite Elanus caeruleus insectivore ground fairly common, resident C, T, RK 26 M. MONIRUL H. KHAN & NASIM AZIZ Forktail 28 (2012) Species Principal diet Principal foraging guild Status Distribution Black Kite Milvus migrans carnivore (inch piscivore) ground fairly common, resident C, T, RK, S Brahminy Kite Haliastur indus carnivore (inch piscivore) ground common, resident Wl White-rumped Vulture Gyps bengalensis carnivore find piscivore) (carrion) ground fairly common, resident RK, C,T Himalayan Griffon Gypshimaiayensis carnivore (inch piscivore) (carrion) ground uncommon (in NSP sites), vagrant (in Bangladesh) RK Crested Serpent Eagle Spilornis cheela* carnivore (inch piscivore) middle canopy very common, resident Wl Shikra Accipiter badius* carnivore (inch piscivore) middle canopy fairly common, resident Wl Besra Accipiter virgatus* carnivore (inch piscivore) middle canopy fairly common, resident Wl Changeable Hawk Eagle Spizaetus cirrhatus* carnivore (inch piscivore) middle canopy uncommon (in NSP sites), resident L, S, RK Common Kestrel Falco tirwunculus insectivore ground fairly common, winter visitor Wl Amur Falcon Falco amurensis insectivore upper canopy (mid-air) uncommon (in NSP sites), winter visitor L, RK Little Cormorant Phalacrocorax niger carnivore (inch piscivore) ground (water) uncommon (in NSP sites), resident T Little Egret Egretta garzetta carnivore (inch piscivore) ground fairly common, resident C, T, RK Cattle Egret Bubulcus ibis carnivore (inch piscivore) ground fairly common, resident C, T, RK Indian Pond Heron Ardeola gray'd carnivore (inch piscivore) ground very common, resident C, T, RK, L Black-crowned Night Heron Nycticorax nycticorax carnivore (inch piscivore) ground uncommon (in NSP sites), resident RK, C,T Malayan Night Heron Gorsachius melanolophus* carnivore (incl. piscivore) ground uncommon (in NSP sites), vagrant (in Bangladesh) L, RK Yellow Bittern Ixobrychus sinensis carnivore (inch piscivore) ground uncommon (in NSP sites), resident T, C Cinnamon Bittern Ixobrychus cinnamomeus carnivore (inch piscivore) ground fairly common, resident RK, C, T Asian Openbill Anastomus oscitans carnivore (inch piscivore) (snail) ground uncommon (in NSP sites), resident RK Lesser Adjutant Leptoptilos javanicus carnivore (inch piscivore) ground uncommon (in NSP sites), resident RK Blue-naped Pitta Pitta nipalensis* insectivore ground uncommon (in NSP sites), resident RK, L, S, T Hooded Pitta Pitta sordida* insectivore ground uncommon (in NSP sites), summer visitor S, L, RK Asian Fairy Bluebird Irena puella* frugivore middle canopy common, resident L, S, RK, T Blue-winged Leafbird Chloropsis cochinchinensis* insectivore middle canopy uncommon (in NSP sites), resident L, T Golden-fronted Leafbird Chloropsis aurifrons* insectivore middle canopy very common, resident Wl Brown Shrike Lanius cristatus insectivore bush and undergrowth common, winter visitor Wl Long-tailed Shrike Lanius schach insectivore bush and undergrowth common, resident Wl Grey-backed Shrike Lanius tephronotus insectivore bush and undergrowth fairly common, winter visitor Wl Common Green Magpie Cissa chinensis* omnivore middle canopy uncommon (in NSP sites), resident S, L,T Rufous Treepie Dendrocitta vagabunda omnivore middle canopy common, resident Wl Grey Treepie Dendrocitta formosae* omnivore middle canopy fairly common, resident L, S, RK, T House Crow Corns splendens omnivore ground fairly common, resident C, T, RK Large-billed Crow Corvus macrorhynchos omnivore ground common, resident Wl Ashy Woodswallow Artamus fuscus insectivore upper canopy (mid-air) fairly common, resident Wl Black-naped Oriole Oriolus chinensis* omnivore middle canopy uncommon (in NSP sites), winter visitor L, S, RK Black-hooded Oriole Oriolus xanthornus omnivore middle canopy very common, resident Wl Maroon Oriole Oriolus traillii* omnivore upper canopy uncommon (in NSP sites), resident L, S, RK Large Cuckooshrike Coracina macei omnivore middle canopy common, resident Wl Black-winged Cuckooshrike Coracina melaschistos* omnivore middle canopy uncommon (in NSP sites), winter visitor L, S, RK Black-headed Cuckooshrike Coracina melanoptera omnivore middle canopy uncommon (in NSP sites), resident RK Rosy Minivet Pericrocotus roseus* insectivore upper canopy uncommon (in NSP sites), resident L, S, RK Ashy Minivet Pericrocotus divaricatus* insectivore upper canopy uncommon (in NSP sites), resident L, RK Small Minivet Pericrocotus cinnamomeus* insectivore upper canopy very common, resident Wl Scarlet Minivet Pericrocotus flammeus* insectivore upper canopy common, resident Wl Bar-winged Flycatcher-shrike Hemipus picatus* insectivore middle canopy fairly common, resident L, S, RK White-throated Fantail Rhipidura ai bicoids insectivore bush and undergrowth common, resident Wl Black Drongo Dicrurus macrocercus insectivore middle canopy very common, resident Wl Ashy Drongo Dicrurus leucophaeus insectivore middle canopy uncommon (in NSP sites), winter visitor Wl Bronzed Drongo Dicrurus aeneus* insectivore middle canopy very common, resident Wl Forktail 28 (2012) Bird species diversity in five protected areas of Bangladesh 27 Species Principal diet Principal foraging guild Status Distribution Lesser Racket-tailed Drongo Dicrurus remifer* insectivore upper canopy uncommon (in NSP sites), winter visitor L, S, RK Spangled Drongo Dicrurus hottentottus* insectivore middle canopy common, resident Wl Greater Racket-tailed Drongo Dicrurus paradiseus* insectivore upper canopy common, resident Wl Black-naped Monarch Hypothymis azurea* insectivore bush and undergrowth common, resident Wl Common lora Aegithina tiphia insectivore middle canopy very common, resident Wl Large Woodshrike Tephrodornisgularis* insectivore upper canopy common, resident L, S, RK Common Woodshrike Tephrodornis pondicerianus* insectivore middle canopy common, resident Wl Blue Rock Thrush Monti co la solitarius insectivore ground fairly common, winter visitor Wl Blue Whistling Thrush Myophonus caeruleus * insectivore ground uncommon (in NSP sites), resident T Orange-headed Thrush loothera citrina insectivore ground uncommon (in NSP sites), resident Wl Red-throated Flycatcher Ficedula parva insectivore middle canopy very common, winter visitor Wl Verditer Flycatcher Eumyias thalassina* insectivore upper canopy fairly common, winter visitor Wl Pale-chinned Flycatcher Cyornis poliogenys* insectivore middle canopy uncommon (in NSP sites), resident RK, L, S Grey-headed Canary Flycatcher Culicicapa ceylonensis insectivore middle canopy common, resident Wl Oriental Magpie Robin Copsychus sauiaris insectivore ground very common, resident Wl White-rumpedShama Copsychus maiabaricus* insectivore ground common, resident Wl Black Redstart Phoenicurus ochruros insectivore bush and undergrowth uncommon (in NSP sites), winter visitor RK, C, T Black-backed Forktail Enicurus immacuiatus * insectivore ground uncommon (in NSP sites), resident RK, T Common Stonechat Saxicoia torguata insectivore bush and undergrowth common, winter visitor RK, C, T Pied Bushchat Saxicola caprata insectivore bush and undergrowth uncommon (in NSP sites), resident C,T Asian Glossy Starling Aplonis panayensis* frugivore middle canopy uncommon (in NSP sites), winter visitor T Chestnut-tailed Starling Sturnus maiabaricus frugivore middle canopy very common, resident Wl Asian Pied Starling Sturnus contra omnivore ground very common, resident Wl Common Myna Acridotheres tristis omnivore ground very common, resident Wl Bank Myna A cridotheres ginginianus omnivore ground uncommon (in NSP sites), resident RK, C, T Jungle Myna Acridotheres fuscus omnivore middle canopy very common, resident Wl Hill Myna Gracula religiosa* omnivore upper canopy common, resident Wl Velvet-fronted Nuthatch Sitta frontalis* insectivore middle canopy fairly common, resident RK, L, S Great Tit Parus major insectivore middle canopy very common, resident Wl Barn Swallow Hirundo rustica insectivore upper canopy (mid-air) common, winter visitor Wl Black-headed Bulbul Pycnonotus atriceps* omnivore middle canopy fairly common, resident Wl Black-crested Bulbul Pycnonotus melanicterus* omnivore middle canopy common, resident Wl Red-whiskered Bulbul Pycnonotus jocosus omnivore middle canopy very common, resident Wl Red-vented Bulbul Pycnonotus cafer omnivore middle canopy very common, resident Wl White-throated Bulbul Alophoixus fiaveolus* omnivore middle canopy common, resident Wl Olive Bulbul lole virescens* omnivore middle canopy uncommon (in NSP sites), resident RK, L, S Ashy Bulbul Hemixos flavala* omnivore middle canopy uncommon (in NSP sites), resident L, RK Grey-breasted Prinia Prinia hodgsonii insectivore bush and undergrowth common, resident Wl Plain Prinia Prinia inornata insectivore bush and undergrowth fairly common, resident C,T, RK Zitting Cisticola Cisticola juncidis insectivore bush and undergrowth common, resident Wl Oriental White-eye Zosterops palpebrosus insectivore middle canopy very common, resident Wl Blyth's Reed Warbler Acrocephalus dumetorum insectivore bush and undergrowth common, winter visitor Wl Striated Grassbird Megaluruspalustris insectivore bush and undergrowth fairly common, resident C, T Common Tailorbird Orthotom us sutorius insectivore bush and undergrowth very common, resident Wl Dark-necked Tailorbird Orthotomus atroguiaris* insectivore bush and undergrowth fairly common, resident T, C Dusky Warbler Phyiloscopus fuscatus insectivore bush and undergrowth fairly common, winter visitor Wl Tickell's Leaf Warbler Phyiloscopus affinis insectivore middle canopy fairly common, winter visitor Wl Yellow-browed Warbler Phyiloscopus inornatus insectivore middle canopy common, winter visitor Wl Greenish Warbler Phyiloscopus trochiloides insectivore middle canopy fairly common, winter visitor Wl 28 M. MONIRUL H. KHAN & NASIM AZIZ Forktail 28 (2012) Principal Species Principal diet foraging guild Status Distributi Blyth's Leaf Warbler Phylloscopus reguloides insectivore middle canopy fairly common, winter visitor L, S, RK Yellow-vented Warbler Phylloscopus cantator insectivore middle canopy fairly common, winter visitor L, RK Golden-spectacled Warbler Seicercus burkii insectivore middle canopy uncommon (in NSP sites), summer visitor L, RK Grey-hooded Warbler Seicercus xanthoschistos insectivore middle canopy uncommon (in NSP sites), winter visitor L, RK White-crested Laughingthrush Gamilaxleucolophus* insectivore bush and undergrowth uncommon (in NSP sites), resident C, T Lesser Necklaced Laughingthrush Garrulaxmoniliger* insectivore middle canopy uncommon (in NSP sites), resident RK, L, C Greater Necklaced Laughingthrush Garrulaxpectoralis* insectivore middle canopy common, resident Wl Rufous-necked Laughingthrush Garrulax ruficollis* insectivore bush and undergrowth common, resident Wl Abbott's Babbler Malacocincla abbotti* insectivore bush and undergrowth very common, resident Wl Puff-throated Babbler Pellorneum ruficeps* insectivore bush and undergrowth common, resident Wl Large Scimitar Babbler Pomatorhinus hypoleucos* insectivore middle canopy uncommon (in NSP sites), resident L, RK, T White-browed Scimitar Babbler Pomatorhinus schisticeps* insectivore bush and undergrowth uncommon (in NSP sites), resident L, S, RK Grey-throated Babbler Stachyris nigriceps* insectivore bush and undergrowth uncommon (in NSP sites), resident C, T Striped Tit Babbler Macronous gularis* insectivore bush and undergrowth common, resident Wl Chestnut-capped Babbler Timalia pileata* insectivore bush and undergrowth uncommon (in NSP sites), resident C,T Yellow-eyed Babbler Chrysomma sinensis* insectivore bush and undergrowth uncommon (in NSP sites), resident C, T Brown-cheeked Fulvetta Alcippe poioicephala* insectivore middle canopy uncommon (in NSP sites), resident L Nepal Fulvetta Atcippe nipalensis* insectivore middle canopy uncommon (in NSP sites), resident RK Rufous-winged Bushlark Mirafra assamica granivoreand herbivore ground common, resident Wl Thick-billed Flowerpecker Dicaeum agile omnivore middle canopy fairly common, resident RK, C, T Yellow-vented Flowerpecker Dicaeum chrysorrheum* omnivore middle canopy uncommon (in NSP sites), resident L, RK, T Orange-bellied Flowerpecker Dicaeum trigonostigma* omnivore middle canopy uncommon (in NSP sites), resident T Pale-billed Flowerpecker Dicaeum erythrorynchos omnivore middle canopy common, resident Wl Plain Flowerpecker Dicaeum concolor omnivore middle canopy common, resident L, RK, T Scarlet-backed Flowerpecker Dicaeum cruentatum* omnivore middle canopy very common, resident Wl Ruby-cheeked Sunbird Anthreptessingalensis * nectarivore middle canopy fairly common, resident Wl Purple-rumped Sunbird Nectarinia eeylonica nectarivore middle canopy fairly common, resident RK, L, C, T Purple-throated Sunbird Nectarinia sperata* nectarivore middle canopy common, resident Wl Purple Sunbird Nectarinia asiatica nectarivore middle canopy very common, resident Wl Crimson Sunbird Aethopyga siparaja* nectarivore middle canopy very common, resident Wl Little Spiderhunter Arachnothera longirostra* nectarivore middle canopy very common, resident Wl Streaked Spiderhunter Arachnothera magna* nectarivore middle canopy uncommon (in NSP sites), resident T House Sparrow Passer domesticus granivoreand herbivore ground common, resident Wl Forest Wagtail Dendronanthus indicus* insectivore ground common, winter visitor Wl White Wagtail Motacilla alba insectivore ground common, winter visitor RK, L, C, T White-browed Wagtail Motacilla maderaspatensis insectivore ground common, resident RK, L, C, T Citrine Wagtail Motacilla citreola insectivore ground uncommon (in NSP sites), winter visitor RK, C, T Grey Wagtail Motacilla cinerea insectivore ground fairly common, winter visitor RK, C,T Paddyfield Pipit Anthus ruficollis insectivore ground common, resident RK, C, T Olive-backed Pipit Anthushodgsoni* insectivore ground common, winter visitor Wl Rosy Pipit Anthus roseatus insectivore ground uncommon (in NSP sites), winter visitor RK Baya Weaver Ploceus philippinus insectivore ground common, resident Wl Indian Silverbill Lonchura malabarica granivoreand herbivore ground uncommon (in NSP sites), resident RK, C, T White-rumped Munia Lonchura striata* granivoreand herbivore ground fairly common, resident Wl Scaly-breasted Munia Lonchura punctulata granivore and herbivore ground common, resident RK, C, T Black-headed Munia Lonchura malacca granivoreand herbivore ground uncommon (in NSP sites), resident RK, C, T FORKTAIL 28 (2012): 29-37 Peat swamp forest birds of the Tuanan research station. Central Kalimantan, Indonesia, with notes on habitat specialists MARY ROSE C. POSA & DAVID ALEXANDER MARQUES The avifauna of tropical peat swamp forest has not been well documented, even though it is an extensive habitat in parts of South-East Asia. We conducted surveys using various methods at the Tuanan research station and surrounding areas in Central Kalimantan, Indonesian Borneo. These observations resulted in a list of 138 bird species and numerous noteworthy records. Although more depauperate than lowland rainforest on mineral soils, peat swamp forest is an important habitat for many threatened and Near Threatened bird species, especially habitat specialists such as Hook-billed Bulbul Setornis criniger and Grey-breasted Babbler Malacopteron albogulare. We also recorded in selectively logged peat swamp several high-profile, globally threatened species such as Crestless Fireback Lophura erythrophthalma, Storm's Stork, Ciconia stormi, Great Slaty Woodpecker Mulleripicus pulverulentus, Black Hornbill Anthracoceros malayanus and Wrinkled Hornbill Aceroscorrugatus. In view of its importance to certain species, peat swamp forest should be afforded more protection, especially in light of the recent rapid loss of this habitat to land conversion and forest fires. INTRODUCTION Borneo is the third largest tropical island in the world and particularly rich in biodiversity, with 630 recorded bird species (Mann 2008). Geopolitically, the island is divided into Brunei Darussalam, the Malaysian states of Sabah and Sarawak, and the four Kalimantan provinces of Indonesia. Most of the ornithological work on Borneo has been conducted in the northern part of the island, and Kalimantan remains poorly studied, with only a few avifaunal lists published specifically for this region (e.g. Holmes & Burton 1987, Holmes 1997). In Central Kalimantan, the second largest province of Indonesian Borneo with a land area of 1 54,564 km2, bird surveys have largely concentrated in a few well-known areas such as Tanjung Puting National Park and Barito Ulu (Bohap & Galidikas 1987, Nash & Nash 1988, Dutson et al. 1991, Wilkinson et al. 1991). The extensive tropical swamp forests dominating the southern lowland plains of Central Kalimantan have largely been ignored and unexplored, even though these habitats constitute one of the seven biogeographic zones of Borneo and have an important influence on species distribution (MacKinnon et al. 1996). Tropical peat swamp forest (PSF) is a unique wetland ecosystem that develops in areas where waterlogging prevents the complete decomposition of plant debris, which over time accumulates as peat soils (Anderson 1983). PSF occurs throughout the tropics but reaches its greatest extent and depth in South-East Asia, especially in the lowlands of Sumatra and Kalimantan (Rieley et al. 1996). PSFs are characterised by periodic flooding, nutrient limitation and high acidity due to the leaching of organic compounds. PSF trees are adapted to tolerate nutrient deficiency, unstable substrate and fluctuating water levels, and in this respect exhibit structural features such as stilt roots and pneumatophores. The overall primary productivity and biodiversity levels in this nutrient- deficient forest type are lower than in lowland forest on mineral soils (Bruenig& Droste 1995). The neglect of PSF by biologists might result either from its relatively depauperate flora and fauna or from the difficult access and working conditions brought about by the boggy soils and dense understorey vegetation, which severely hamper movement and visibility. However, recent research indicates that PSF may harbour a considerable proportion of the South-East Asian fauna (Posa et al. 2011). Thus, there is an urgent need for more information on the flora and fauna of this unique ecosystem, as its destruction has accelerated in recent years. Nearly half of the PSF in Peninsular Malaysia, Borneo and Sumatra has been lost since 1990 (Miettinen & Liew 2010). Many areas have already been converted into oil palm and paper pulp plantations and much of what remains under forest cover has been selectively logged (Miettinen & Liew 2010). Such disturbance renders PSF extremely prone to forest fires, since peat itself is combustible when dry (Page et al. 2009). Fire is now one of the major drivers of PSF loss and conversion to degraded land. In this paper, we present the first avifaunal list for the Tuanan research station in the Mawas Conservation Area and surrounding areas based on field observations, mistnetting and camera trapping conducted in Central Kalimantan from 2009 and 2010. STUDY AREA AND METHODS The Mawas Conservation Area comprises a 3,000 km2 area managed by the Borneo Orangutan Survival Foundation located east of the Kapuas river, about 55 km from Palangkaraya, the capital of Central Kalimantan (Figure 1). Here, the Tuanan research station (2°09,06"S 114°26'26"E) was established in 2003 for long-term Orangutan behavioural studies. It comprises a 9.45 km2 grid-based trail system situated on peat of varying thickness up to 2 m. The forest was subjected to selective commercial logging in the early 1990s, followed by illegal logging (van Schaik et al. 2005). However, there has been no systematic logging since 2002 (Vogel et al. 2009). Despite this disturbance, the forest supports a relatively high density of Bornean Orangutans Pongo pygmaeus wurmbii , Bornean Agile Gibbons Hylobates agilis albibaris and other globally threatened mammals (van Schaik & Brockman 2005, pers. obs.). Central Kalimantan has a humid tropical climate, with very little variability in temperature. The wet season normally occurs during the north¬ west monsoon in November to April: the climate is drier during June to August. There is some climatic variation associated with the El Nino Southern Oscillation cycle, which affects the duration and severity of the seasons. During strong El Nino events, southern Kalimantan can experience prolonged drought conditions (Page et al. 2009). The mean average annual rainfall measured from 2004 to 2007 at the Tuanan research station was 2,678 mm, with an average monthly rainfall of 223 mm (Wartmann 2008). Outside of the Mawas Conservation Area, the PSF has been heavily disturbed by humans. Drainage canals dug in the early 1990s for the Indonesian government’s Mega-Rice Project have disrupted the natural hydrology of the area, making it extremely prone to fire (Page et al. 2009). As a result, large areas have been subjected to one or more fires and are now dominated by ferns and other low-growing plants. 30 MARY ROSE C. POSA & DAVID ALEXANDER MARQUES Forktail 28 (2012) Figure 1 . Map of the ex-Mega-Rice Project area with remaining peat swamp forest cover (in black). Approximate location of the Tuanan research station indicated. Straight solid grey lines are canals that were dug for drainage. Bird observations were made in the intact PSF around the Tuanan research station as well as in surrounding degraded areas using a variety of methods. DM mad e ad libitum observations from November to December 2009, while MRCP conducted standard 10 minute, 25 m radius point-count surveys from August 2009 to July 2010 (Posa 2011). In addition, MRCP also used mistnets and camera traps to survey Tuanan for a total of 2,535 net hours and 3,924 trap nights respectively (details of methodology in Posa 2011). We included in our list only species that were confirmed by visual sightings or distinctive calls, mistnetting and camera trapping. With the exception of Bornean Ground Cuckoo Carpococcyx radiatus (following Collar & Long 1996), nomenclature follows the 2009 checklist of the Oriental Bird Club (sequence of Dickinson 2003) available online at http://orientalbirdimages.org/ new-obc-checklist.html. RESULTS We recorded a total of 138 bird species from all survey methods in the intact and degraded PSF habitats in and around the Mawas Conservation Area (Appendix). Four of these were migratory species, and two were most likely introduced recently to Borneo. Thus, 1 32 resident birds were found in the PSF. Twenty-six species were observed only along canals or rivers and in the non-forested regrowth vegetation in areas that had previously been burned. The rest of the species were observed in logged PSF and remnant forest fragments. Several species of particular interest were observed, including seven globally threatened, 31 Near Threatened species as well as two PSF specialists (Hook-billed Bulbul Setornis criniger and Grey-breasted Babbler Malacopteron albogulare) and three Bornean endemics (Bornean Ground Cuckoo, Bornean Bristlehead Pityriasis gymnocephala and Dusky Munia Lonchura fuscans). We captured a total of 293 birds from 28 species in mistnets (see Appendix), with recaptures (21 individuals) comprising 7.2%. Two species that were recorded only by mistnetting were Blue-eared Kingfisher Alcedo meninting and Oriental Cuckoo Cuculus saturatus. Camera traps took 45 photographs of birds from ten species, of which we were unable to identify three. Three species were detected only through photography, namely Black Partridge Melanoperdix niger , Crestless Fireback Lophura erythrophthalma and Bornean Ground Cuckoo. Significant records Species accounts are given for threatened species, endemics and habitat specialists and a few other notable records. We give the species conservation status (Endangered, Vulnerable, Near Threatened, Least Concern) based on the 2010 Red List by the International Union for Conservation of Nature (IUCN 2010). Black Partridge Melanoperdix niger Vulnerable. A lone female was photographed in July 2010 in the early morning. An uncommon and local resident on Borneo, this species is poorly known, as it is shy and secretive. It has been recorded from swamp forest in Gunung Palung National Park (Laman et al. 1996). CrestSess Fireback Lophura erythrophthalma Vulnerable. Only detected by camera traps in intact forest, but the commonest bird species recorded using this method (29 of 45 photographs of birds). Pictures of individuals or pairs were taken between dawn and dusk, but the majority of photos was taken before 08h00. Other researchers reported occasional encounters on man-made transects, but in general this species is very elusive. Storm's Stork Ciconia stormi Endangered. Recorded three times by camera traps and two seen flying over the research station in the early morning in June 2010. This species has also been reported from other swamp forests, includingpeat swamp (Laman et al. 1996, Danielsen etal. 1997, Page et al. 1997). It is considered very rare throughout its range, but our record and the recent one in Thailand, where it was thought to be extinct (Cutter et al. 2007), indicates that camera trapping is a very useful method in detecting this species’s presence in forested areas. Lesser Adjutant Leptoptilos javanicus Vulnerable. A few to a dozen individuals were spotted at various times in drained and deforested areas while travelling on canals in May-July 2010. Wallace's Hawk Eagle Spizaetus nanus Vulnerable. An adult was observed perched in the PSF interior in December 2009. Cinnamon-headed Green Pigeon Treron fulvieollis Near Threatened. Individuals and a group of three were seen feeding in fruiting trees lining canals in the deforested area in September 2009. This species was not recorded in intact forest, but was possibly overlooked. It has also been recorded from PSF in Sebangau (Page etal. 1997) and was reportedly the most abundant green pigeon in the wooded areas of the Barito' region and southern Kalimantan by Holmes & Burton (1987). Long-tailed Parakeet Psittacula longieauda Near Threatened. One bird was observed in PSF in December and groups of more than a dozen birds were seen on dead remnant trees in degraded areas in August to November 2009. Forktail 28 (2012) Peat swamp forest birds of theTuanan research station, Central Kalimantan, Indonesia 31 Chestnut-bellied Malkoha Phaenicophaeus sumatranus Near Threatened. Surprisingly, a commonly encountered bird in the PSF interior habitat, travelling in singles, pairs or groups of three birds. Reported as uncommon throughout Bornean lowland and hill dipterocarp forests (Mann 2008). Bornean Ground Cuckoo Carpococcyx radiatus Near Threatened. Endemic. One individual was photographed following a Sun Bear Helarctos malayanus in December 2010. Described as a rare resident of lowland forests (Mann 2008). It has been recorded in a few other places in Central Kalimantan in alluvial and swamp habitats (Long & Collar 2002, Fredriksson & Nijman 2004). Black Hornbil! Anthracoceros malayanus Near Threatened. Small groups of up to eight individuals were uncommonly encountered in intact forest. Wrinkled Hornbil! Aceros conugatus Near Threatened. One adult male was observed in PSF in November 2009. Red-crowned Barbet Megalaima rafflesii Near Threatened. The commonest barbet in PSF and disturbed forest around Tuanan with up to eight individuals recorded in a day. Great Slaty Woodpecker Mulleripicus pulverulentus Vulnerable. Noisy conspicuous groups were uncommonly encountered in intact forest and also observed in forest fragments in the degraded area. This species has also been reported from PSF in West Kalimantan (Laman etal. 1996). Bornean Bristlehead Pityriasis gymnocephala Near Threatened. Bornean endemic. Considered to be rare and uncommon on the island (Mann 2008). Individuals and small groups of up to five birds were uncommonly encountered in intact PSF and forest fragments in the degraded area. This species has been reported from PSF in Sarawak (Laman et al. 2006) and the ‘swamp’ forests ofTanjungPuting (Nash & Nash 1988). Smythies (1981) suggested that it may prefer PSF, but Witt & Sheldon (1994) refuted this. Hook-billed Bulbul Setornis criniger Vulnerable. PSF habitat specialist. Fairly common in intact forest, often travelling in small groups. Their call has been described as a rattling series of notes (3.4 kHz, 1 1 notes) or a soft crrrk (Myers 2009) or harsh alarm cuurrk (MacKinnon & Phillipps 2008). These bulbuls are quite easy to detect in the PSF understorey because of these calls, which we were able to record (Figure 2; www.xeno- canto.org catalogue number XC74801). We also caught six individuals in mistnets. This species has been observed in other nutrient-poor forests in Borneo and was described as a local lowland resident (Mann 2008). In Central Kalimantan, it has been recorded in swamp forest in Sebangau (Pag eetal. 1997) and TanjungPuting, as well as frequently encountered in kerangas at Barito UIu (Dutson etal. 1991). It has been suggested that Setornis criniger is intolerant of habitat degradation (Dutson et al. 1991), but our observations indicate that it can persist in large blocks of selectively logged PSF. However, the rapid loss of this habitat will continue to threaten this species. Grey-breasted Babbler Malaeopteron albogularis Near Threatened. PSF habitat specialist. This species is rare except in poor soil habitats such as PSF, kerangas and ultrabasic forests (Sheldon etal. 2001). It is often overlooked because of its skulking habits, absence from mixed-species flocks and infrequent vocalisations. We observed it only on three occasions in the understorey. However, it was the fourth most commonly mistnetted species (19 out of 272 individuals), supporting the suggestion by some authors that this method can reveal its presence (Sheldon 1987, Dutson et al. 1991). Thus it may be overlooked even in habitats where it is fairly common. Birds from Barito Ulu and T anjung Puting are described as having white lores, while those from north Borneo have yellow lores (Dutson etal. 1991, Sheldon et al. 2001), but see Collar (201 1). The lores of the adult birds captured at Tuanan were white but consistently have a few rufous feathers on the edge near the forehead. We captured one juvenile with an inflated gape on 14 June 2010. It had similar coloration to the adult, except for a yellow lower mandible (grey in adult), brown iris (red in adult) and pink legs (grey in adult). Crimson-breasted Flowerpecker Prionochilus percussus Least Concern. Observed on four days in November-December 2009 and on three days in March-June 2010 with a maximum of four individuals. This species is supposed to be a rare lowland resident, especially in northern Borneo (Mann 2008), but identification issues may mask its true occurrence. No Yellow- rumped Flowerpeckers P. xanthopygius were observed in the Tuanan PSF, supporting the statement by Holmes & Burton (1987): ‘ xanthopygius occurs with percussus in upper Barito Sep 1986 but not with it in southern lowlands of Kalimantan’. Scarlet-breasted Flowerpecker Prionochilus thoracicus Near Threatened. Single adult males were observed on two separate occasions in the canopy of the PSF interior. An uncommon and local bird on Borneo (Mann 2008). It seems to prefer poor soil habitats (Sheldon et al. 2001). Figure 2. Sonagram of the crrrk or cuurrk call typical for Hook-billed Bulbul Setornis criniger. 32 MARY ROSE C. POSA & DAVID ALEXANDER MARQUES Forktail 28 (2012) Dusky Munia Lonchura fuscans Least Concern. Bornean endemic. Not uncommon in disturbed forest, forest edge and forest regrowth areas around the Tuanan site. DISCUSSION While only one study (Gaither 1994) has made a direct comparison and shown that bird diversity in PSF is lower than in lowland rainforest on mineral soils, we reach the same conclusion from our survey in this nutrient-poor environment. We observed only 132 of Borneo’s 398 resident bird species in PSF, including only three of at least 41 Bornean endemics, whereas other lowland sites around Borneo show higher species richness — even up to twice those numbers in eastern Sabah where forests are unusually rich (e.g. Lambert 1992, Johns 1996, Cleary etal. 2007, Edwards etal. 2011). Aside from fewer numbers of species and endemics, total abundance of observed birds is also very low. However, if we compare the bird species (102) found in the Tuanan PSF (excluding degraded and riverine areas) with other sites on Borneo containing PSF (Appendix), the numbers of species are roughly similar. Laman et al. (1996) reported 104 resident species from Gunung Palung National Park, which contains about 400 ha of swamp forest in a mosaic with lowland dipterocarp and upland forests. Tanjung Puting National Park, which is composed of 50% PSF in mosaic with freshwater swamp and heath forests, has 111 resident species reported in its ‘swamp forest’ by Nash & Nash (1988), although these authors surveyed peat basin margins and not true PSF. Our list shares 97 species (41%) with the lowland habitats of the Cabang Panti research site in Gunung Palung (61 reported from PSF and an additional 36 reported from lowland dipterocarp forest) and 123 species (56%) with Tanjung Puting, including 104 species reported from swamps by Nash & Nash (1988) with an additional 19 reported by Bohap & Galdikas (1987). Page et al. (1997) reported 150 species of birds observed over three years from various habitats, including both forest and riverine sedge swamp, at the Sungei Sebangau catchment in Central Kalimantan. However, because they did not provide a complete list of species, we cannot directly compare their results with ours. Mistnetting has been conducted in PSF at only a few other sites on Borneo. Gaither (1994) captured 34 species in Gunung Palung but did not provide a complete species list. In Sarawak, sporadic mistnetting from 1996 to 1999 in a previously logged 20-ha patch of PSF at the UNIMAS Campus near Kuching revealed 68 resident species (Tuen & Darub 1999, Rahman & Tuen 2006). However, most of these (31) were represented by one or two captures only, including M. albogulare. They also failed to detect S. criniger, which suggests the site is heavily degraded and fragmented. In PSF at Loagan Bunut National Park, mistnetting during a short 10-day survey yielded 18 resident species, including S. criniger but not M. albogulare; observations produced an additional 12 species (Laman etal. 2006). In Sabah, six-days of mistnetting in primary PSF of the Klias Forest Reserve (Sheldon etal. 2004) yielded 28 species. While it is more difficult to set up mistnets in PSF than in dryland forests, this method is very effective and, thus, useful for studying understorey species, particularly in revealing the presence of M. albogulare. However, with the mistnetting bias towards understorey species and the low number of recorded species compared with point count sampling, researchers should be judicious in the use of mistnetting for rapid assessments of avian biodiversity in PSF (Remsen & Good 1996). Camera trapping of birds has usually been incidental to surveys of terrestrial mammals, but it is starting to be explored as a viable method for sampling large ground-dwelling bird species (O’Brien & Kinnaird 2008). Although low numbers of birds were recorded with this method, it is a valuable method for detecting rare and elusive species such as L. erythrophthalma and C. stormi in addition to the other standardised procedures. Kalimantan’s PSF may represent a stronghold for S. criniger and M. albogulare , as large, albeit disturbed, tracts of this habitat still remain. These species are most likely to be declining in Peninsular Malaysia and Sumatra, where a greater percentage of PSF has been converted to plantations and other non-forest land uses (Miettinen & Liew 2010). Their current status needs to be assessed and monitored in light of the recent rapid loss of PSF habitats. More research is needed on the local distribution of bird species in PSF, and specific microhabitat requirements of these species in PSF needs to be elucidated. Despite low bird diversity in PSF, the occurrence of the PSF specialists M. albogulare and S. criniger together with other threatened and Near Threatened species underlines the urgent need for continued efforts in the Mawas Conservation Area to prevent further habitat loss and hunting. ACKNOWLEDGEMENTS We thank the Borneo Orangutan Survival Foundation (BOS), especially BOS Jakarta and BOS-MAWAS Palangka Raya, Universitas Palangka Raya and Universitas Nasional, Jakarta, for their facilitation and logistical support. 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C. & Sheldon, F. H. (1994) A review of the status and distribution of the Bornean bristlehead. Kukila 7: 54-67. Mary Rose C. POSA, Department of Biological Sciences, National University of Singapore, 1 4 Science Drive 4, Singapore 1 17543; dbspmrc@nus.edu. sg David Alexander MARQUES, Zentralstrasse 6, 8003 Zurich, Switzerland; dmargues@bluemail.ch 34 MARY ROSE C. POSA & DAVID ALEXANDER MARQUES Forktail 28 (2012) Appendix List of bird species recorded at Tuanan and surrounding areas and their occurrence in other sites containing peat swamp forest habitat Abbreviations: VU - Vulnerable; EN - Endangered; NT - Near-threatened; LC - Least Concern; I - introduced to Borneo; M - migrant to Borneo; PSF - intact peat swamp forest; DIS - disturbed and non-forest habitat; RIV - riverine forest. Species marked with x* were recorded in 'swamp' (including peat swamp) habitat in other sites, x - habitat type not specified or recorded from non-swamp habitat, * - caught in mistnet. TanjungPuting, Unimas, Sarawak Central (mistnetting in Kalimantan Berbak, Sumatra Loagan Bunut regrowth, Tuen & Tuanan record (Bohap & Galdikas Gunung Palung, (Silvius & National Park, Darub1999, Status habitat 1987; Nash& West Kalimantan Verheugt 1986, Sarawak Rahman Species (IUCN 2010) (this study) Nash 1988) (Laman etal. 1996) Hornskov 1987) (Gumalefa/. 2008) & Tuen 2006) Black Partridge Melanoperdix niger VU PSF X X* X Crestless Fireback Lophura erythropthalma VU PSF X X X* Storm's Stork Ciconia stormi EN PSF X* X* X X Lesser Adjutant Leptoptilos javanicus VU DIS X X Black-thighed Falconet Microhierax frirgillarius LC PSF, RIV, DIS X* X X X Black-winged Kite Elanus caeruleus LC DIS X X Brahminy Kite Haliastur indus LC DIS, RIV X* X X Crested Serpent Eagle Spilornis cheela LC PSF, DIS, RIV X* X* X X* Crested goshawk Accipiter trivirgatus LC PSF* X* X X X Changeable Hawk Eagle Spizaetus cirrhatus LC PSF X X* Wallace's Hawk Eagle Spizaetus nanus VU PSF X X X* White-breasted Waterhen Amaurornis phoenicurus LC DIS X X X Spotted Dove Streptopelia chinensis LC DIS X X Cinnamon-headed Green Pigeon heron fulvicollis NT DIS X* X Pink-necked Green Pigeon heron vernans LC DIS, RIV X* X X Thick-billed Green Pigeon heron curvirostra LC PSF X* X* X Blue-crowned Hanging Parrot Loriculus galgulus LC PSF, DIS, RIV X* X X X* Long-tailed Parakeet Psittacula longicauda NT PSF, DIS, RIV X* X* X X* Indian Cuckoo Cuculus micropterus LC PSF X* X X X* Oriental Cuckoo Cuculussaturatus LC, M PSF' Banded Bay Cuckoo Cacomantis sonneratii LC PSF Plaintive Cuckoo Cacomantis merulinus LC PSF, DIS X* X X* Violet Cuckoo Chrysococcyx xanthrohynchus LC PSF, DIS X* X X* Drongo Cuckoo Surniculuslugubris LC PSF, DIS, RIV X* X X Black-bellied Malkoha Phaenicophaeus diardi NT RIV X* X* Chestnut-bellied Malkoha Phaenicophaeus sumatranus NT PSF X* X X* Raffles’s Malkoha Phaenicophaeus chlorophaeus LC PSF X* X* X X* Chestnut-breasted Malkoha Phaenicophaeus curvirostris LC PSF X* X* X X* Bornean Ground Cuckoo Carpococcyx radiatus NT, E PSF X Greater Coucal Centropus sinensis LC PSF, DIS X* X* X X Lesser Coucal Centropus bengalensis LC DIS X X Brown Wood Owl Strix ieptogrammica LC PSF X* X* X Savanna Nightjar Caprimulgus affinis LC DIS Silver-rumped Needletail Rhaphidura leucopygialis LC DIS, RIV X* X X* Grey-rumped Treeswift Hemiprocne longipennis LC DIS, RIV X* X X X* Diard's Trogon Harpactes diardii NT PSF, DIS X* X* X* X* Scarlet-rumpedTrogon Harpactes duvaucelii NT PSF X* X* X X* Dollarbird Eurystomus orientalis LC PSF, RIV X Stork-billed Kingfisher Halcyon capensis LC DIS, RIV X* X X X* Forktail 28 (2012) Peat swamp forest birds of theTuanan research station, Central Kalimantan, Indonesia 35 Species Status (IUCN 2010) Tuanan record habitat (this study) Tanjung Puting, Central Kalimantan (Bohap&Galdikas 1987; Nash & Nash 1988) Gunung Palung, West Kalimantan (Laman era/. 1996) Berbak, Sumatra (Silvius & Verheugt1986, Hornskov 1987) Loagan Bunut National Park, Sarawak (Gumal etal. 2008) Unimas, Sarawak (mistnetting in regrowth, Tuen& Darub1999, Rahman &Tuen 2006) Collared Kingfisher Todiramphus chloris LC DIS X X X Oriental Dwarf Kingfisher Ceyxerithaca LC PSF* X* X* X X* X* Blue-eared Kingfisher Alcedo meninting LC PSF, RIV X* X X X X* Blue-throated Bee-eater Merops viridis LC PSF, DIS, RIV X* X X* Bushy-crested Hornbill Anorrhinus galeritus LC PSF X* X* X X Oriental Pied Hornbill Anthracoceros albirostris LC DIS, RIV X* X Black Hornbill Anthracoceros malayarus NT PSF X* X* X X* Wrinkled Hornbill Acer os cor rug at us NT PSF X* X* X X Red-crowned Barbet Megalaima rafflesii NT PSF, DIS, RIV X* X* X X* X* Red-throated Barbet Megalaima mystacophanos NT RIV X X* X X Blue-eared Barbet Megalaima australis LC PSF, RIV X* X* X X* Brown Barbet Caloramphus fuliginosus LC PSF X* X* X X* Rufous Piculet Sasia abnormis LC PSF' X* X* X* X* Sunda Pygmy Woodpecker Dendrocopos moluccensis LC PSF, DIS X X Rufous Woodpecker Celeus brachyurus LC PSF X* X X* X* White-bellied Woodpecker Dryocopus j avert sis LC DIS, RIV X* X X X* Banded Woodpecker Picus mineaceus LC PSF, RIV X X X X* Crimson-winged Woodpecker Picus puriceus LC RIV X* X* X* X* Maroon Woodpecker Blythipicus rubiginosus LC PSF X X* X X* X* Orange-backed Woodpecker Reinwardtipicus validus LC PSF, DIS X* X* X Buff-rumped Woodpecker Meiglyptes tristis LC PSF, DIS X* X X* Buff-necked Woodpecker Meiglyptes tukki NT PSF X* X* X X* X* Grey-and-buff Woodpecker Hemicircus concretus LC PSF, DIS, RIV X* X* X Great Slaty Woodpecker Mulleri picus pulverulentus VU PSF X* X* X Green Broadbill Calyptomena viridis NT PSF X* X* X Black-and-red Broadbill Cymbirhynchus macrorhynchos LC RIV X* X X X X* Banded Broadbill Eurylaimus javanicus LC PSF X* X X Black-and-yellow Broadbill Eurylaimus ochromalus NT PSF X* X X X* Dusky Broadbill Corydon sumatranus LC PSF X* X X* Golden-bellied Gerygone Gerygone sulphurea LC RIV X* Large Woodshrike Tephrodornis virgatus LC RIV X X X* White-breasted Woodswallow Artamus leucorynchus LC DIS, RIV X X Common lora Aegithina tiphia LC PSF, RIV X* X X X X* Green lora Aegithina viridissima NT PSF X* X X* X* Bornean Bristlehead Pityriasis gymnocephala NT, E PSF X* X X* Lesser Cuckooshrike Coracina fimbriata LC PSF, RIV X* X X* Pied Triller Lalage nigra LC DIS X Fiery Minivet Pericrocotus igneus NT PSF, DIS, RIV X* X* X X* Scarlet Minivet Pericrocotus flammeus LC PSF X* X X X* Black-winged Flycatcher-shrike Hemipus hirundinaceus LC PSF, DIS, RIV X* X X X* X* Mangrove Whistler Pachycephala cinerea LC PSF' X* X Long-tailed Shrike Laniusschach LC DIS, RIV X Tiger Shrike Lanius tigrinus LC,M DIS X Dark-throated Oriole Oriolus xanthonotus NT PSF, DIS X* X* X* 36 MARY ROSE C. POSA & DAVID ALEXANDER MARQUES Forktail 28 (2012) TanjungPuting, Unimas, Sarawak Central (mistnetting in Kalimantan Berbak, Sumatra Loagan Bunut regrowth, Tuen & Tuanan record (Bohap & Galdikas Gunung Palung, (Silvius & National Park, Darubl999, Status habitat 1987; Nash & West Kalimantan Verheugt 1986, Sarawak Rahman Species (IUCN 2010) (this study) Nash 1988) (Laman etal. 1996) Hornskov 1987) (Gumal etal. 2008) & Tuen 2006) Bronzed Orongo Dicrurus aeneus LC RIV X* X X X* X* Pied Fantail Rhipidura javanica LC PSF, DIS.RIV X X X, X* Black-naped Monarch Hypothymis azurea LC PSF8 X* X* X X* Asian Paradise-flycatcher Terpslphoneparadisi LC PSP X* X* X X* Pacific Swallow Hirundo tahitica LC DIS.RIV X* X X X* Barn Swallow Hirundo rustica LC, M PSF X* Yellow-bellied Prinia Prinia flaviventris LC DIS X X X X* Dark-necked Tailorbird Orthotomus atrogularis LC PSF, RIV X* X X* X* Rufous-tailed Tailorbird Orthotomus sericeus LC PSP, DIS X* X X, X* Ashy Tailorbird Orthotomus ruficeps LC PSF5, DIS X* X X* X* Arctic Warbler Phylloscopus borealis LC,M PSF Sooty-headed Bulbul Pycnonotus aurigaster LC, 1 RIV Puff-backed Bulbul Pycnonotus eutilotus NT DIS X* X* X X* X* Yellow-vented Bulbul Pycnonotus goiavier LC PSF, DIS X* X X* Olive-winged Bulbul Pycnonotus plumosus LC DIS.RIV X* X X* X* Cream-vented Bulbul Pycnonotus simplex LC PSP X* X* X X* Spectacled Bulbul Pycnonotus erythropthalmos LC PSF X* X* X X* X* Hook-billed Bulbul Setornis criniger VU PSF* X* X* X* X* Black-capped Babbler Pellorneum capistratum LC PSF*, DIS X* X* X X* X* White-chested Babbler Jrichastoma rostratum NT PSP, DIS X* X X* X* Short-tailed Babbler Malacocincla malaccensis NT PSP X* X* X* X* Scaly-crowned Babbler Malacopteron clnereum LC PSF* X* X* X* Rufous-crowned Babbler Malacopteron magnum NT PSP X* X X* Grey-breasted Babbler Malacopteron albogulare NT PSF8 X* X* X* X* Chestnut-rumped Babbler Stachyris maculata NT PSF8, DIS X* X* X X* Black-throated Babbler Stachyris nigricollis NT PSF8 X* X* X* X* Chestnut-winged Babbler Stachyris erythroptera LC PSF8 X* X* X* X* Striped Tit Babbler Macronous gularis LC PSF, DIS X* X X, X* Fluffy-backed Tit Babbler Macronous ptilosus NT PSP X* X X X* X* Asian Fairy Bluebird Irena puella LC PSF, RIV X* X* X X* X* Velvet-fronted Nuthatch Sitta frontalis LC PSF X* X X Common Hill Myna Graculareligiosa LC PSF, DIS X* X* X X* Oriental Magpie Robin Copsychus saularis LC DIS X* X* X X X* White-rumpedShama Copsychus malabaricus LC PSF8, DIS X* X* X* X* X* Rufous-tailed Shama Trichixos pyrropygus NT PSF8 X* X* X* X* Grey-chested Jungle Flycatcher Rhinomyias umbratilis NT PSF8 X* X* X* X* X* Malaysian Blue-flycatcher Cyornis turcosus NT RIV X* X X Greater Green Leafbird Chloropsis sonnerati LC PSF X* X* X X* Lesser Green Leafbird Chloropsis cyanopogon NT PSF, DIS X* X* X X* Yellow-breasted Flowerpecker Prionochilus maculatus LC PSF8 X* X* St X* X* Crimson-breasted Flowerpecker Prionochilus percussus LC PSF8, DIS X X* X Scarlet-breasted Flowerpecker Prionochilus thoraclcus NT PSF X* X* X* Orange-bellied Flowerpecker Dicaeum trigonostigma LC PSF, DIS X* X* X X* X* Scarlet-backed Flowerpecker Dicaeum cruentatum LC PSF X X* Forktail 28 (2012) Peat swamp forest birds of theTuanan research station, Central Kalimantan, Indonesia 37 Species Status (IUCN 2010) Tuanan record habitat (this study) TanjungPuting, Central Kalimantan (Bohap & Galdikas 1987; Nash & Nash 1988) Gunung Palung, West Kalimantan (Lamanefa/. 1996) Berbak, Sumatra (Silvius & Verheugt 1986, Hornskov 1987) Loagan Bunut National Park, Sarawak (Gumal etal. 2008) Unimas, Sarawak (mistnetting in regrowth, Tuen & Darub1999, Rahman & Tuen 2006) Ruby-cheeked Sunbird Anthreptes singalensis LC PSF, DIS X* X* X X* Plain Sunbird Anthreptes simplex LC PSF X* X* X* Brown-throated Sunbird Anthreptes malacensis LC PSF, DIS X* X X* X* Purple-naped Sunbird Hypogramma hypogrammicum LC PSF' X* X* X X* X* Purple-throated Sunbird Nectarinia sperata LC PSF X* X* X X* Olive-backed Sunbird Nectarinia jugularis LC DIS X X X X* Crimson Sunbird Aethopyga siparaja LC PSF, RIV X* X X X* Little Spiderhunter Arachnothera longirostra LC PSP, DIS X* X* X X* X* Yellow-eared Spiderhunter Arachnothera chrysogenys LC PSF X* X* X Eurasian Tree Sparrow Passer montanus LC, 1 DIS Dusky Munia Lonchura fuscans LC, E DIS X* X X X* FORKTAIL 28 (2012): 38-43 An annotated checklist of birds and conservation issues in Salkhala Game Reserve, an isolated Important Bird Area in Azad Kashmir, Pakistan MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE Salkhala Game Reserve (SGR) in Azad Kashmir, Pakistan, lies within an Important Bird Area (IBA) of the Western Himalayas Endemic Bird Area. The conservation status of the reserve and its birds is poorly known due to political instability in the disputed territory of Kashmir and the relative remoteness of the site. The findings of a bird survey undertaken from May 2007 to April 2008 are documented here. In total, 101 species were recorded including 45 resident species, 48 breeding migrants and six winter migrants. There were significant records of the globally threatened Western Tragopan Tragopan melanocephalus, the Near Threatened Pallid Harrier Circus macrourus and European Roller Coraciasgarrulus, and the restricted-range Kashmir Nuthatch Sitta cashmirensis and Spectacled Finch Callacanthis burtoni. Kashmir Flycatcher Ficedula subrubra and Cheer Pheasant Catreus wallichi were not recorded in the IBA, with the latter species now possibly locally extirpated. An annotated checklist of the species recorded is presented along with measures of relative abundance. Habitat fragmentation, degradation and clearance through the collection of fuel and timber, forest fire, livestock grazing, collection of non-timber forest products and unsustainable use of pastures are the major threats to the wildlife of SGR. These conservation issues are discussed briefly along with recommendations for the future management of the reserve. INTRODUCTION Located in the Neelum valley within the Western Himalayas Endemic Bird Area (EBA; Stattersfield etal. 1998), Salkhala Game Reserve (SGR) forms part of the Salkhala Wildlife Sanctuary Important Bird Area (IBA; Chan etal. 2004). It is classified as such owing to the presence of three globally threatened IBA trigger species: Western Tragopan Tragopan melanocephalus , Cheer Pheasant Catreus wallichi and Kashmir Flycatcher Ficedula subrubra (BirdLife International 2011a). All three species are listed as Vulnerable (IUCN 2011). Western Tragopan is distributed in five separate populations in the Western Himalayas of Pakistan and India (BirdLife International 2001). Previous work in the Neelum Valley established its presence in SGR (Mirza et al. 1978, Islam 1982) and recorded it as ‘common’ and at densities of 0.8- 1.6 birds/km2 (Mirza et al. 1978). More recently, it has been recorded as locally rare’ in the region (Hassan 2004). It is found in mixed coniferous forest, often with a dense understorey, from as low as 1,350 m and up to 2,800 m in winter, and from 2,400 m to 3,600 m in summer (Gaston et al. 1983, Islam & Crawford 1987, Ramesh 2003). Cheer Pheasant is patchily distributed, owing to its association with early successional habitats, between 1,200 and 3,000 m throughout the southern foothills of the Himalayas (Gaston et al. 1981, Garson 1983, Kaul 1993). In Pakistan, it is found in the mountains of eastern North-West Frontier Province and Azad Kashmir (Roberts 1991). A previous survey in SGR flushed 20 individuals (Mirza 1978) but, despite a recent record of 126 birds in Jhelum Valley, Azad Kashmir (Awan etal. 2004), there have been no reports of the species in SGR since. Kashmir Flycatcher has a very restricted distribution in northern India and parts of Pakistan, and occurs as a scarce and apparently irregular summer breeding migrant in the side valleys of Kashmir and the Pir Panjal range of northern Pakistan, with one record from Sind, southern Pakistan (BirdLife International 2001). It breeds between 1,800 and 2,300 m where there is predominantly deciduous vegetation (Roberts 1992). In 1983, one breeding pair with newly fledged young was recorded at 2,100 m in SGR (Roberts 1992). In addition to the three IBA trigger species, the site is important for a number of mammal species, including Kashmir Musk-deer Moschus chrysogaster and Kashmir Gray Langur Semnopithecus ajax (both Endangered), Himalayan Black Bear Ursus thibetanus (Vulnerable), and Leopard Panthera pardus and Himalayan Goral Naemorhedus goral (both Near Threatened) (Dar 2006, IUCN 2011). There are six villages with a total population of about 6,000 people adjacent to SGR (Awan 2008). These communities depend on the natural resources of the area, entering the reserve to graze their cattle, cut trees for timber and collect firewood. Trunks of older trees are sometimes partially burnt to make them easier to cut. In addition to the loss of tree cover, these activities cause much damage to the forest understorey of the reserve (Awan 2008). Salkhala Game Reserve is situated at the ceasefire line between Pakistan and India and, consequently, cross-border conflict between 1989 and 2003 prevented the completion of any field studies in the area during that time. This, coupled with its relative remoteness, means there have been few recent ornithological surveys in the reserve (Islam 1982). This survey is the first to consider all bird species in SGR and was conducted to provide a checklist for the site, measures of relative abundance for key species, and a current understanding of the conservation issues in the reserve after a comparatively long period of isolation. METHODS Salkhala Game Reserve (34°33'N 73°50'E), Neelum Valley, is located 80 km north-west of Muzaffarabad in the Himalayan foothills of Azad Kashmir, Pakistan (Figure 1). Covering 810 hectares at 1,320-3,150 m elevation, it was notified as a Game Reserve in 1982 and is classified as an IUCN Category IV protected area (Dudley 2008). The reserve lies within the Himalayan moist temperate ecozone (Roberts 1991) and consists of a range of forest habitats, including coniferous, broadleaf and mixed coniferous- broadleaf forests. These are characterised by the trees Cedrus deodara , Pinus wallichiana , Abies pindrow, Picea smithiana , Taxus wallichiana , Acer caecium , Bettila utilis, Berberis spp., Ouercus spp., Juniper us communis , Vibernum spp., Indigofera gerardiana,Juglans regia and Aesculus indica. It has a mean annual rainfall of 1 25.7 cm, with March and April being the wettest months, and is exposed to heavy snowfall during the winter (Qureshi 1990). We conducted a bird survey in SGR between May 2007 and April 2008 using two methods: dawn and dusk call counts (Gaston Forktail 28 (2012) An annotated checklist of birds and conservation issues in Salkhala Game Reserve, Pakistan 39 1980) for surveying Galliformes; and unlimited radius point counts (Bibby etal. 2000) for surveying all other bird species. Twelve survey points were positioned randomly and approximately 0.5 km apart between 1,377 and 2,970 m elevation (Figure l), which was representative of the altitudinal range and habitats covered by the reserve. Ten points were located in coniferous forest (points 1-10 in Figure 1), and one each in mixed broadleaf-conifer forest and scrub grassland. One point was surveyed during each dawn and dusk survey, and each of the twelve points was surveyed twice per month, once at dawn and once at dusk (total effort = 288 points). Call counts of 60 minutes’ duration were conducted at 04h45- 05h45 (April-September) and 05h30-06h30 (October-March), and 18hOO-19hOO (April-September) and 16h00-17h00 (October-March), with start time varying according to seasonal differences in sunrise/sunset times. All calling Galliformes heard were recorded and mapped. Point counts of 10 minutes’ duration were carried out at the end of each dawn call count and start of each dusk call count. All birds detected were identified and the number of individuals recorded. If a bird group was only detected by call, then a mean group size from visual contacts of that species was used (Lee & Marsden 2008). A checklist for SGR was produced from both sets of survey data. However, the survey methods employed were not appropriate for effectively detecting birds of prey (Marsden 1998) and, consequently, these species are likely to be under¬ recorded in this study. Species encounter rates were calculated based on the number of individuals detected from all points surveyed, and presented as the number of individuals per 100 point counts (± standard error). Encounter rates were converted into ordinal categories of abundance: <5 individuals per 100 point counts = ‘Rare’: 5.1-10 = ‘Uncommon’; 10.1-20 = ‘Frequent’; 20.1-40 = ‘Common’; and >40 = ‘Abundant’ (adapted from Lowen et al. 1996). These simple categories can be used for future monitoring of the abundance of species within the reserve (Robertson & Liley 1998). Mean encounter rates for each species were calculated for each month (24 points/ month), and then a standard error was derived from these sample means. Bird survey data were supplemented by information gathered from interviews with local staff of the AJ&K (Azad Jammu and Kashmir) Wildlife Department (n = 15) and local community members (n = 35); five community members were selected randomly from each of the seven villages around SGR. Specifically, this information was used to help confirm the presence/ absence of Galliformes, and particularly Cheer Pheasant. RESULTS In total 1,959 bird records, comprising 101 species belonging to 38 families, were recorded in the survey. Of these, 45 were resident species, 48 were summer migrants, six were winter migrants and two were passage migrants. Two species were classified as ‘Abundant’, 16 as ‘Common’, 35 as ‘Frequent’, 44 as ‘Uncommon’ and four as ‘Rare’ (Appendix). The survey recorded one of the three IBA trigger species, Western Tragopan (29.1 ± 8.8 individuals/ 100 points; ‘Frequent’), and two Near Threatened species, Pallid Harrier Circus macrourus (6.3 ± 4.2 individuals; ‘Uncommon’) and European Roller (6.9 ±4.9 individuals; ‘Uncommon’) (IUCN 2011), winter and summer migrants, respectively. Cheer Pheasant and Kashmir Flycatcher, the two other IBA trigger species, were not recorded during the survey. Interviews with local communities did not provide any supporting evidence to suggest that Cheer Pheasant is present within the reserve. In addition to the tragopan, two additional restricted-range species (Stattersfield et al. 1998), Kashmir Nuthatch Sitta cashmirensis (21.8 ± 5.7 Figure 1. Map showing the location and land cover types of Salkhala Game Reserve. 74°6‘0"E i 73°54'0"E 73°55'0"E 73°56,0"E 73°57'0"E Legend © Survey Points - Line of control - Major Rivers District Boundary □ Salkhala GR Sill Dense Conifer Forest Medium Conifer Forest Mixed Broadleaved / Conifer Forest feS-Lj Grasses / Shrubs : Agriculture / Cultivated Areas Bare Land (Soil / Rocks) Snow / Glacier N A 74°6'0"E 40 MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE Forktail 28 (2012) individuals; ‘Frequent’) and Spectacled Finch Callacantbis burtoni (12.5 ± 4.0 individuals; ‘Uncommon’), were also recorded. A complete annotated checklist is given in the Appendix. Interviews with local villagers revealed that a number of birds, mainly Galliformes, are hunted to varying degrees within the reserve. Hunting of Galliformes is probably higher in the reserve during the winter months when birds move down to lower altitudes. Western Tragopan is hunted for meat and feathers, and some skins for taxidermy were for sale in local houses. A number of stuffed Himalayan Monal Lopbopborus impejanus were also seen in many homes. Kalij Pheasant Lopbura leucomelanos is hunted locally for food, especially in the winter when pheasants migrate to lower elevations. Koklass Pheasant Pucrasia macrolopha and Chukar Partridge Alectoris chukar are also trapped and hunted for food by local communities. Local villagers occasionally shoot Pallid Harriers because they prey on their domestic chickens. All four species of columbids recorded in the reserve are hunted, primarily by teenagers and younger men, for food. DISCUSSION Salkhala Game Reserve is designated as an IBA due to the presence of Western Tragopan, Cheer Pheasant and Kashmir Flycatcher. This survey recorded 101 species, but included records for only one of the IBA trigger species, Western Tragopan, lor which SGR is an important site, along with Pallid Harrier, European Roller, and Kashmir Nuthatch and Spectacled Finch, two restricted-ranges species of the Western Himalayas EBA (Stattersfield et al. 1998). Kashmir Flycatcher is an irregular and sparse summer migrant to the area, so it is as feasible that it was present but undetected as that it was absent in the reserve during the survey. Ol greater conservation concern is the failure to detect Cheer Pheasant, with its apparent absence from the reserve corroborated in local interviews. For a species with a small and fragmented population (BirdLife International 2011b), this loss from a protected site is a worrying development. The possible local extirpation of Cheer Pheasant from SGR is indicative of a growing human population and an increasing demand on natural resources affecting the conservation status of species and habitats in what is a comparatively small protected area (Awan 2010). Rising human activities are increasing the conservation importance of the reserve in a landscape already heavily impacted, raising concerns about site isolation and the viability of populations of key species. The recent construction of a road within the reserve, and its use for extracting trees that have fallen due to heavy snow or landslides, has now made access to wildlife relatively easy. Conservation threats within SGR include habitat degradation and loss, through the collection of timber, firewood and wild vegetables, hunting and overgrazing. Hunting pressure is particularly high for Galliformes in the reserve, with hunting for food, skins or recreation conducted by local and non-local professional (trophy-hunting) and non¬ professional hunters alike. There is a seasonal shift in the type of hunting pressure within the reserve. During the warmer months of May-September, people from adjacent villages travel with their cattle to higher grazing areas (above 2,400 m) and stay in their summer homes in and around the reserve. At this time, people take the opportunity to collect medicinal plants, vegetables and eggs from pheasant nests, and to hunt wildlife (Qureshi 1990). Owing to difficult terrain in the reserve, dogs are often used to flush birds, especially pheasants, while traps may also be laid (Awan 2010). During the winter months, people and their livestock move back to lower elevations, and any hunting at this time tends to be recreational rather than functional. In addition to the Galliformes, Pallid Harrier, which is a rare winter visitor to SGR, experiences some degree of hunting pressure as local villagers shoot it to protect their chickens from predation. Although commercial tree cutting is prohibited in all protected areas in Pakistan, there is unlawful felling in SGR, especially in the gullies in the north and south of the reserve (MNA pers. obs. 2008). These areas tend to be at lower elevations, but logging activities affect not only the species that occupy those elevations throughout the year but also those that undergo seasonal migration during the winter months. Of these, pheasants are most likely to be affected since they are also hunted for food and trophies. To reduce the impacts of harvesting forest resources, including hunting, Awan (2010) recommended that the reserve be extended south-west to the Gail Nullah area and east to the line of control and, consequently, be better conserved under the protected area system. Adding some form of mixed-use or buffer zone to try to shift pressure away from core areas within what is a small reserve, especially during the summer months when more people are accessing and utilising the reserve, would seem likely to benefit the reserve generally and the Western Tragopan in particular. Man-made forest fires remain a threat to the conservation of wildlife in the reserve, with large areas of forest affected by fires every year (Qureshi 1990). These fires are especially prevalent during the drier summer months, when people spend more time in the forest and make fires for warmth at night and to help bring down standing timber. From 1989 to 2003, cross-border firing between India and Pakistan destroyed areas of natural forest growth in and adjacent to the reserve. A recent community-based awareness campaign was undertaken to support the conservation of key bird and mammal species in the reserve (Awan 2010). This included working with communities, in schools, directly with hunters, and training local wildlife staff. However, there remains a general lack of understanding of the biodiversity importance of the reserve in those communities in and around SGR. Consequently, it is vital that the efforts of this initial programme are built on in a collaborative and constructive manner to help improve the conservation status of SGR and the species within it, while maintaining and supporting local livelihoods. Now that the reserve is more accessible, it would benefit from regular species monitoring to track general trends in species abundance and habitat alteration, which, in turn, will help support effective management of the site. In part, this could be included within the planned surveys for Galliformes in the Western Himalayas of Pakistan, coordinated by the World Pheasant Association-Pakistan and WWF- Pakistan. At a basic level, using the same survey points as this study may be a first step to establishing a bird monitoring scheme in SGR. Although subjective and taking no account of detectability differences between species, the ordinal categories of relative abundance that we have used here may also provide a simple baseline to monitor and detect any large-scale changes in the abundance of individual species within SGR in the future. ACKNOWLEDGEMENTS We are very thankful to Prof. Z. B. Mirza for reviewing an earlier draft oi the manuscript. We are also grateful to Mir Saleem, Supervisor Wildlife, Muzaffrabad, and the wildlife staff of Salkhala Game Reserve for their support during the field study. Thanks are also due to Baseer Qureshi who helped with collecting data in SGR. REFERENCES Awan, M. N. (2008) Community conservation awareness program, Salkhala Game Reserve, Neelum, Azad Kashmir. Unpublished Final Report to the Oriental Bird Club, UK. Forktail 28 (201 2) An annotated checklist of birds and conservation issues in Salkhala Game Reserve, Pakistan 41 Awan, M. N. (2010) Status and conservation of Western Tragopan Pheasant in and around Salkhala Game Reserve, District Neelum, Azad Kashmir, Pakistan. Unpublished Final Report to the Oriental Bird Club, UK. Awan.M.S., Khan, A.A, Ahmad, K.B., Qureshi, M.A., Malik, M. A & Dar, N.l (2004) Population dynamics of Cheer Pheasant ( Catreus wallichii ) in Jhelum Valley , Muzaffarabad, Azad Kashmir, Pakistan. Pakistan J. Biol. Sci. 7: 789-796. Bibby, C. J., Burgess, N. D., Hill, D. A. & Mustoe, S. (2000) Bird census techniques. Second revised edition. London: Academic Press. B i rd Life International (2001) Threatened birds of Asia: the BirdLife International Red Data Book. Cambridge, UK: BirdLife International. BirdLife International (2011a) important Bird Areas factsheet: Saikaia Wildlife Sanctuary. Downloaded on 1 1 May 2011. . BirdLife International (2011b) Species factsheet: Catreus wallichi. Downloaded on 1 1 May 201 1. . Chan, S., Crosby, M. J., Islam, M. Z. & Tordoff, A. W. (2004) Important Bird Areas in Asia: key sites for conservation. Cambridge, UK: BirdLife International (Conservation Series 13). Dar, N. (2006) [Wildlife of Azad Kashmir. Report of the Department of Wildlife and Fisheries, Government of Azad Jammu and Kashmir.] Muzaffarabad, AJK: Al-Sheikh printing press. (In Urdu.) Dudley, N., ed. (2008) Guidelines for applying protected area management categories. Gland, Switzerland: IUCN. Garson, P. J. (1 983)The Cheer Pheasant Catreus wallichi in Himachal Pradesh, Western Himalaya: an update. J. World Pheasant Assoc. 8: 29-39. Gaston, A. J. (1980) Census techniques for Himalayan pheasants including notes on individual species. J. World Pheasant Assoc. 5: 40-53. Gaston, A. J., Garson, P. J. & Hunter, M. L. Jr. (1981 ) The wildlife of Himachal Pradesh, Western Himalaya. Technical Notes No. 82. School of Forest Resources, University of Maine. Gaston, A. J., Islam, K. & Crawford, J. A. (1983) The current status of the Western Tragopan [Tragopan melanocephalus).J. World Pheasant Assoc. 8: 40-49. Hassan, S. A. (2004) Compilation of baseline data for ornithological studies in Machiara National Park. Unpublished Reportto Azad KashmirWildlife Department. Islam, K. (1 982) Status and distribution of the Western Tragopan in north¬ eastern Pakistan. Pp.44-50 in C. D. W. Savage & M. W. Ridley, eds. Pheasants in Asia 1982. Reading, UK: World Pheasant Association. Islam, K. & Crawford, J. A. (1987) Habitat use by Western Tragopan Tragopan melanocephalus (Gray) in northeastern Pakistan. Biol. Conserv. 40: 101 — 115. IUCN (201 1) IUCN Red List of threatened species. Version 201 1 .2. Downloaded on 6 March 2012. . Kaul, R. (1993) Habitat utilization by Cheer Pheasant J. World Pheasant Assoc. 17/18: 84-85. Lee, D. C. & Marsden, S. J. (2008) Adjusting count period strategies to improve the accuracy of forest bird abundance estimates from point transect distance sampling surveys. Ibis 1 50: 315-325. Lowen, J. C„ Bartrina, L., Clay, R. P. & Tobias, J. A. (1996) Biological surveys and conservation priorities in eastern Paraguay. Cambridge, U.K.: CSB Conservation. Marsden, S. J. (1998) Counting single-species. Pp.53-75 in C. J. Bibby, M. J. Jones & S. J. Marsden, eds. Expedition field techniques: bird surveys. London: Expedition Advisory Centre. Mirza, Z. B. (1978) Pheasant surveys in Pakistan. Amer. Pheasant Waterfowl Soc. Mag. 78: 2-6. Mirza, Z. B., Aleem, A. & Asghar, M. (1978) Pheasant surveys in Pakistan./ Bombay Nat. Hist. Soc. 75: 292-296. Qureshi, A. R. (1990) Revised forest management plan for the forests of Neelum Valley. Unpublished Report. Azad Jammu and Kashmir Forest Department, Muzaffarabad, AJK. Ramesh, K. (2003) An ecological study on pheasants of the Great Himalayan National Park, Western Himalaya. Unpublished Report. Wildlife Institute of India, Dehradun, India. Roberts, T.J. (1991) The birds of Pakistan. Vol. I (Non-passeriformes). Karachi: Oxford University Press. Roberts, T. J. (1992) The birds of Pakistan. Vol. II (Passeriformes). Karachi: Oxford University Press. Robertson, P. A. & Liley, D. (1998) Assessment of sites: measurement of species richness and diversity. Pp.76-98 in C. J. Bibby, M. J. Jones & S. J. Marsden, eds. Expedition field techniques: bird surveys. London: Expedition Advisory Centre, Royal Geographical Society. Stattersfield, A. J„ Crosby, M. J., Long, A. J. &Wege, D. C. (1998) Endemic Bird Areas of the world: priorities for biodiversity conservation. Cambridge, UK: BirdLife International (Conservation Series 7). Muhammad Naeem AWAN, Himalayan Nature Conservation Foundation (HNCF), Muzaffarabad, Azad Kashmir, 13100, Pakistan. Email: ajkwildlife@gmail.com Hassan ALI, WWF Pakistan, Ferozepur Road, Lahore, Pakistan. Email: hassanwwf@gmail.com David C. LEE, Division of Biology, University of Glamorgan, Pontypridd, UK. Email: dlee@glam.ac.uk Appendix Annotated checklist of birds recorded in Salkhala Game Reserve IUCN Red List status (IUCN 2011) follows the species name: VU = Vulnerable; NT = Near Threatened. RR after the species name indicates a restricted-range species (Stattersfield et al. 1998). Species encounter rates are per 100 point counts ± standard error (SE). Abundance (ordinal scale), with numbers of individuals encountered in parentheses: A = Abundant; C = Common; F = Frequent; U = Uncommon; R = Rare. Status, with months observed in parentheses: R = Resident; S = Summer migrant; W = Winter migrant; P = Passage migrant; L = Local movement. Species Encounter rate (± SE) Abundance Status (months observed) Altitude (m) Himalayan Snowcock Tetraogallus himalayensis 11.1 ±4.9 U (32) R, L (Feb— Mar, Sep— Oct) 2,880 Chukar Mectoris chukar 39.5 ±12.0 F (114) R, L (Feb— Mar, May-Jun, Sep— Dec) 1,320-2,350 Western Tragopan Tragopan melanocephalus (VU, RR) 33.3 ±10.3 F (96) R, L (Feb— Mar, May-Jun, Sep— Oct) 1,960-2,890 Himalayan Monal Lophophorus impejanus 29.1 ±8.8 F (84) R, L (Feb— Mar, May-Jun, Sep-Oct) 2,130-2,860 Koklass Pheasant Pucrasia macrolopha 58.3 ±17.6 C (168) R, L (Feb— Mar, May-Jul, Sep-Oct, Dec) 2,180-2,840 Kalij Pheasant Lophura leucomelanos 52.0 ±15.8 C (150) R, L (Feb— Mar, May-Jun, Sep— Nov) 1,610-2,270 Himalayan Woodpecker Dendrocopos himalayensis 31.2 ± 11.6 F (90) R (Feb-Jun, Sep-Oct, Dec) 2,590 Scaly-bellied Woodpecker Picus squamatus 29.1 ±7.5 F (84) R (Feb— Mar, May-Jun, Sep-Oct, Dec) 1,970-2,800 Great Barbet Megalaima virens 29.1 ±7.6 F (84) R (Feb— Mar, May-Jul, Sep-Oct, Dec) 2,420-2,770 42 MUHAMMAD NAEEM AWAN, HASSAN ALI & DAVID C. LEE Forktail 28 (2012) Species Encounter rate (±SE) Abundance Status (months observed) Altitude (m) Blue-throated Barbet Megalaima asiatica 6.9 ±4.8 U (20) S (May-Aug) 1,620 Common Hoopoe Upupa epops 24.3 ±16.3 F (70) S (Apr-Aug) 1,560-2,800 European Roller Coracias garrulus (NT) 6.9 ±4.9 U (20) S (May-Aug) 1,500-1,700 Indian Roller Coracias benghalensis 10.4 ±7.2 U (30) S (May-Aug) 1,987-2,700 Common Kingfisher Alcedo atthis 13.1 ±9.0 U (38) S (May-Sep) 1,410-1,570 White-throated Kingfisher Halcyon smyrnensis 37.5 ±11.5 F (108) R, L (Feb— Mar, May-Jul, Sep, Nov) 1,360-1,760 Pied Kingfisher Ceryle rudis 43.7 ±11.2 C (126) R (Feb— Mar, May-Jun, Sep, Nov— Dec) 1,420 Asian Koel Eudynamys scolopacea 8.3 ±5.7 U (24) S (Apr-Aug) 1,800-2,570 Rose-ringed Parakeet Psittacula krameri 29.8 ±20.5 F (86) S (May-Aug) 1,780 Common Swift Apus apus 24.3 ±16.8 F (70) S (Apr-Aug) 1,150-1,600 Fork-tailed Swift Apus pad ficus 17.3 ±11.8 U (50) S (May-Aug) 1,570-1,600 House Swift Apus affinis 1 6.6 ± 11.4 U (48) S (Apr-Jul) 1,550-2,475 Brown Wood Owl Strix leptogrammica 5.6 ±3.8 U (16) S (May-Aug) 1,760 Spotted Owlet Athene brama 24.3 ±6.4 F (70) R (Feb— Mar, May-Jun, Sep— Oct, Dec) 1,450 Rock Pigeon Columbalivia 29.1 ±7.6 F (84) R, L (Feb— Mar, May-Jun, Sep, Nov— Dec) 1,570-2,340 Spotted Dove Stigmatopelia chinensis 17.3 ± 1 1.8 U (50) S (May-Aug) 1,500-2,680 Red Collared dove Streptopelia tranquebarica 24.3 ±16.5 F (70) S (May-Aug) 1,400-2,300 Eurasian Collared Dove Streptopelia decaocto 8.3 ±6.0 U (24) S (May-Aug) 1,440-1,650 Himalayan Vulture Gypshimalayensis 29.1 ±7.8 F (84) R, L (Feb— Mar, May-Jun, Sep— Oct, Dec) 1,600-2,850 Pallid Harrier Circus macrourus (NT) 6.3 ±4.2 U (18) W (Oct-Jan) 2,170-2,380 Common Kestrel Falco tinnunculus 50.0 ±15.2 C (144) R (Feb— Mar, May-Jun, Sep, Dec) 1,320-2,460 Bay-backed Shrike ianius vittatus 6.9 ±5.6 U (20) S (May-Aug) 1,380 Long-tailed Shrike Laniusschach 5.6 ±4.3 U (16) S (May-Aug) 1,500-2,130 Great Grey Shrike ianius excubitor 9.0 ±6.4 U (26) S (May-Aug) 2,130 Yellow-billed Blue Magpie Urocissa flavirostris 31.5 ±8.4 F (91) R, L (Feb— Mar, May-Jun, Sep-Oct, Dec) 1,400-2,615 Rufous Treepie Dendrocitta vagabunda 41.3 ±10.8 C (1 19) R, L (Feb— Mar, May-Jun, Sep-Oct, Dec) 1,350-1,830 Red-billed Chough Pyrrhocorax pyrrhocorax 22.9 ±7.1 F (66) R (Feb— Mar, May-Jul, Sep-Oct) 2,370-2,660 Yellow-billed Chough Pyrrhocorax graculus 14.5 ±4.6 U (42) R (Feb— Mar, May-Jul, Sep-Oct) 2,360-2,530 House Crow Corvus splendens 52.0 ±18.6 C (150) R (Feb— Mar, May-Oct, Dec) 1,340-2,380 Large-billed Crow Conus macrorhynchos 252 ±65.0 A (728) R (Feb— Mar, May-Jul, Sep-Oct, Dec) 1,340-3,040 Eurasian Golden Oriole Oriolus oriolus 17.3 ±12.9 U (50) S (May-Sep) 1,420-2,500 Scarlet Minivet Pericrocotus flammeus 10.4 ±7.7 U (30) S (May-Sep) 1,578-2,520 White-throated Fantail Rhipidura albicollis 24.3 ±6.5 F (70) R, L (Feb— Mar, May-Jul, Sep-Oct, Dec) 1,600-1,800 Black Drongo Dicrurus macrocercus 38.1 ±25.7 F (110) S (May- Sep) 1,350-2,090 Asian Paradise-flycatcher Terpsiphoneparadisi 15.9 ±10.9 U (46) S (Apr-Aug) 1,410-2,110 Brown Dipper Cinclus pallasii 55.9 ±14.5 C (161 ) R, L (Feb— Mar, May-Jun, Sep-Oct, Dec) 1,970-2,360 Blue-capped Rock-thrush Monticola cinclorhynchus 8.3 ± 6.4 U (24) S (May-Aug) 1,460-2,420 Blue Whistling-thrush Myophonus caeruleus 39.5 ±12.0 F (1 14) R (Feb— Mar, May-Jun, Sep— Dec) 1,800-3,000 Dark-throated Thrush Turdus ruficollis 9.0 ±6.8 U (26) W (Oct-Jan) 2,000-2,130 Slaty-blue Flycatcher Ficedula tricolor 10.4 ±8.1 U (30) S (May-Aug) 1,340-2,230 Grey-headed Canary-flycatcher Culicicapa ceylonensis 9.2 ±7.0 U (28) S (May-Aug) 2,640-2,710 Common Redstart Phoenicurus phoenicurus 37.5 ±11.4 F (108) R (Feb— Mar, May-Jun, Sep— Dec) 1,860 Plumbeous Water Redstart Rhyacornis fuliginosus 64.2 ±23.0 C (185) R (Feb-Jul, Sep-Oct, Dec) 2,040-3,050 White-capped Water Redstart Chaimarrornis leucocephalus 22.9 ±16.4 F (66) S (Apr-Aug) 1,880-3,050 Little Forktail Fnicurusscouleri 48.6 ±12.5 C (140) R, L (Jan— Mar, May-Jun, Sep-Oct, Dec) 1,770-1,980 Spotted Forktail Enicurusmaculatus 43.7 ±11.3 C (126) R, L (Feb— Mar, May-Jun, Sep-Oct, Dec) 1,650-1,830 Forktail 28 (2012) An annotated checklist of birds and conservation issues in Salkhala Game Reserve, Pakistan 43 Species Encounter rate (±SE) Abundance Status (months observed) Altitude (m) Common Stonechat Saxicola torquatus 13.8 ±10.2 U (40) S (May-Sep) 1,800-2,180 Pied Bushchat Saxicola caprata 7.6 ±6.3 U (22) S (May-Aug) 2,370-2,510 Brahminy Starling Sturnus pagodarum 13.8 ±10.2 U (40) S (Apr-Aug) 1,565 Common Starling Sturnus vulgaris 13.1 ±10.0 U (38) W (Oa— Jan) 2,320 Common Myna Acridotheres trlstis 89.9 ±23.2 A (259) R (Feb— Mar, May-Jul, Sep— Oct, Dec) 1,500-2,430 Kashmir Nuthatch Sitta cashmirensis (RR) 21.8 ± 5.7 F (63) R, L (Feb— Mar, May-Jun, Sep— Oct, Dec) 1,650-2,640 Chestnut-bellied Nuthatch Sitta castanea 20.8 + 6.4 F (60) R (Feb— Mar, May-Jul, Sep-Nov) 1,350-1,630 Bar-tailed Treecreeper Certhiahimalayana 22.9 ±7.3 F (66) R, L (Jan, Mar, May, Jul, Sep-Nov) 1,690-2,770 Fire-capped Tit Cephalopyrus flammiceps 3.5 ±2.6 R (10) S (May-Aug) 2,485-2,505 Rufous-naped Tit Parus rufonuchalis 43.7 ±11.4 C (126) R, L (Feb-Mar, May-Jun, Sep— Ort, Dec) 1,680-2,720 Spot-winged Tit Parusmelanophus 11.8 ±9.0 U (34) S (May-Aug) 2,140-2,300 Great Tit Parus major 58.3 ±17.6 C (168) R (Mar-Jun, Sep-Oct, Dec-Jan) 1,380-2,670 Green-backed Tit Parus monticolus 13.8 ±9.5 U (40) S (May- Jul) 2,670-2,980 Black-lored Tit Parus xanthogenys 7.6 ±6.3 U (22) S (May- Jul) 1,320-2,940 Barn Swallow Hirundorustica 9.7 ±6.9 U (28) S(May- Sep) 2,330 Himalayan Bulbul Pycnonotus leucogenys 58.3 ±15.1 C (168) R (Feb-Mar, May-Jun, Sep-Oct, Dec) 1,410-2,280 Red-vented Bulbul Pycnonotuscafer 25.6 ± 17.6 F (74) S (Apr-Sep). 1,380-1,710 Black Bulbul Hypsipetes leucocephalus 58.3 ±15 C (168) R (Feb-Mar, May-Jul, Sep-Oct, Dec) 1,410-2,200 Zitting Cisticola Cisticola juncidis 2.8 ±1.9 R (8) S (May-Aug) 1,600 Striated Prinia Prinia criniger 24.3 ±6.4 F (70) R (Feb-Mar, May-Jun, Sep-Oct, Dec) 1,800-2400 Oriental White-eye Zosterops palpebrosus 31 .5 ± 8.4 F (91) R (Feb-Mar, May-Jun, Sep-Oct, Dec) 1,800-2,700 Common Tailorbird Orthotomus sutorius 5.6 ±4.5 U (16) S (Apr-Aug) 1,320-2,050 Common Chiffchaff Phylloscopus collybita 22.9 ±16.2 F (66) W (Sep-Jan) 1,390-2,550 Tickell's Leaf Warbler Phylloscopus affinis 7.6 ±6.0 U (22) S (Apr-Aug) 2,160 Greenish Warbler Phylloscopus trochiloides 7.3 ±5.5 U (22) S (May-Aug) 1,550-2,290 Grey-hooded Warbler Seicercus xanthoschistos 4.9 ±3.9 R (14) ' S (Apr-Jul) 1,540-1,580 Streaked Laughingthrush Garrulaxlineatus 34.0 ±9.8 F (98) R (Feb-Mar, May-Jun, Sep-Oct, Dec) 1,370-2,740 Jungle Babbler Turdoidesstriatus 10.4 ± 3.6 U (30) R (Feb-Mar, May-Sep, Dec) 1,430 Thick-billed Flowerpecker Dicaeum agile 7.6 ±5.8 U (22) S (May-Aug) 2,230-2,840 Purple Sunbird Nectarinia asiatica 6.3 ±4.7 U (18) S (May-Sep) 1,360-1,610 House Sparrow Passer domesticus 54.1 ±28.7 C (156) R (Mar, May-Jan) 1,410-3,000 Russet Sparrow Passer rutilans 34 ± 9.4 F (98) R (Feb-Mar, May-Jun, Sep-Oct, Dec) 2,000 Forest Wagtail Dendronanthus indicus 5.6 ±4.3 U (16) P (Mar-Apr) 1,760-2,310 White-browed Wagtail Motacilla madaraspatensis 6.9 ±4.9 U (20) S (May-Aug) 1,320-2,280 Yellow Wagtail Motacilla flava 7.6 ±5.3 U (22) P (Mar-Apr) 1,470-2,240 Citrine Wagtail Motacilla citreola 9.0 ±6.6 U (26) S (Apr-Aug) 1,530-2,830 White Wagtail Motacilla alba 34.7 ± 24.3 F (100) S (Apr-Aug) 1,340-2,110 Alpine Accentor Prunella collar is 26.7 ±7.1 F (77) R (Feb-Mar, May-Jun, Sep-Oct, Dec) 1,970 Rufous-streaked Accentor Prunella himalayana 9.7 ±6.6 U (28) W (Oct— Jan) 2,100-3,100 Yellow-breasted Greenfinch Carduelis spinoides 41.3 ±10.9 C (119) R (Feb-Mar, May-Jun, Sep— Dec) 1,430-1,610 Common Rosefinch Carpodacus erythrinus 4.9 ±3.9 R(14) S (May-Aug) 1,494 Spectacled Finch Callacanthis burtoni (RR) 12.5 ±4.0 U (36) R (Feb, May-Jun, Sep-Oct, Dec) 2,680-3,100 Rock Bunting Emberiza da 30.5 ±23.4 F (88) S (Apr-Sep) 2,000-3,100 Pine Bunting Emberiza leucocephalos 12.5 ±9.6 U (36) W (Oct— Jan) 2,020-2,700 Chestnut-breasted Bunting Emberiza stewarti 34.7 ±23.6 F (100) S (May-Sep) 1,430-2,570 Crested Bunting Melophuslathami 25.6 ±18.9 F (74) S (Apr-Aug) 1,430-1,530 FORKTAIL 28 (2012): 44-48 A review of the status of Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis SIMON P. MAHOOD & JONATHAN C. EAMES A survey of the globally Endangered Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis was conducted in May 2009 on the Da Lat Plateau, southern Vietnam. Collared Laughingthrush proved to be common throughout broadleaved evergreen forest habitat within its limited altitudinal distribution while the distribution of Grey-crowned Crocias was extended to the south. These results are discussed in the context of the conservation status of these species and it is recommended that both species are retained as Endangered. INTRODUCTION For many species distributed in the tropics, a paucity of basic abundance and distribution data makes the accurate assessment of threat status difficult. There is still much to be learned about the status and distribution of relatively well-known taxa, such as birds, in countries like Vietnam. Vietnam supports over 800 resident bird species (Robson 2008) and, owing to its complex topography, up to six areas which are designated (or would qualify) as Endemic Bird Areas (StattersfieldeTu/. 1998). In recent years the discovery of new species and the revision of species limits has dramatically increased the number of species endemic to Vietnam (or nearly so), revealed a more detailed pattern of endemism, and reinforced the conservation importance of the country (Eames etal. 1999a, Eames etal. 1999b, Eames &: Eames 2001, Eames 2002, Collar 2006). The Da Lat Plateau Endemic Bird Area (EBA 145) is located on an isolated massif in southern Vietnam beyond the southern tip of the Annamite mountains (BirdLife International 2011). The natural vegetation cover is broadleaved evergreen forest with emergent Pinus. The Da Lat Plateau Endemic Bird Area supports more restricted range bird species than any other EBA in mainland South-East Asia. Recent taxonomic revisions have reinforced the status of the Da Lat Plateau EBA as the most important centre of avian endemism in Vietnam. Following taxonomy used in the BirdLife checklist version 4, it supports at least 14 species that qualify as restricted-range species, of which five occur only in this EBA, namely Orange-breasted Laughingthrush Garrulax annamensis , Collared Laughingthrush Garrulax yersini. Grey- crowned Crocias Crocias langbianis , Black-headed Parrotbill Paradoxornis margaritae and Vietnam Greenfinch Carduelis monguilloti. Approximately 40 subspecies of bird are also restricted in range to this EBA (Eames 1995). Of the five species that are restricted to the Da Lat Plateau EBA two are considered globally threatened: Collared Laughingthrush and Grey-crowned Crocias. Both are considered Endangered (BirdLife International 2011). Accurately understanding the distribution and status of species is of utmost importance in assessing their threat status. In Vietnam, there have been few recent field surveys aimed at better understanding species distribution and abundance. In this study we conducted surveys to gather data on the abundance and distribution of the two Endangered passerines that are endemic to the Da Lat Plateau: Collared Laughingthrush and Grey-crowned Crocias. Collared Laughingthrush Garrulax yersini Collared Laughingthrush is part of a species complex that also includes Chestnut-crowned G. erythrocephalus, Assam G. chrysopterus, Silver-eared G. melanostigma, Golden-winged G. ngoclinhensis and Malayan Laughingthrush G. peninsulae. Its habitat, habits and vocalisations are similar to those species, although unlike most of them, it is regarded by some as localised; Robson (2008) describes it as ‘locally common’. Collared Laughingthrush is an understorey bird restricted to high-elevation broadleaved evergreen forest (>1,500 m elevation) on the Da Lat Plateau. Its extent of occurrence (EOO) is only 720 km2 (BirdLife International 2011) and it is known from only five sites: Chu Yang Sin National Park in Dak Lak province, the contiguous Bi Duop Nui Ba National Park and Dam Rong State Forest Enterprise (part of which lies within Cong Troi Important Bird Area), Mt Lang Bian in Dan Kia Suoi Vang Tourism Area, and Nam Ban Police Forest Management Board (part of Tuyen Lam Important Bird Area) (Tordoff 2002). Within these areas populations are fragmented owing to the patchy distribution of suitable habitat, which is often embedded in a matrix of unsuitable pine Pinus forest. Collared Laughingthrush lives in pairs or small parties that forage quietly and unobtrusively in dense understorey and on the forest floor. It is usually detected by vocalisations, which are given infrequently. However, it responds strongly to playback of pre¬ recorded vocalisations. Grey-crowned Crocias Crocias langbianis The Grey-crowned Crocias is known from only four specimens collected at two localities. For a passerine with a continental distribution this paucity of specimen records renders it most unusual. After much intensive effort it was rediscovered in 1994, in Chu Yang Sin National Park (Eames et al. 1995). It has since been found in three parts of Tuyen Lam Important Bird Area, a place known popularly as Ta Nung valley (and near Cam Ly, one of the historical collecting localities) located within Lam Vien Landscape Protection Forest, at Ho Tuyen Lam Tourism Area and in the adjacent Nam Ban Police Forest Management Board (Pilgrim etal. 2007), these constituting two discrete areas of the same forest; at this site the populations may be small and the habitat fragmented. Of the historical collecting localities little is known. At one site, Bjorkegren’s ‘Lang Bian peaks’, which is presumably Mount Lang Bian in Dan Kia Suoi Vang Tourism Area (Eames & Ericson 1996), no suitable habitat within its altitudinal range remains. The location of the other site is imprecisely known but is thought to be within D'Ran Watershed Protection Forest (formerly known as Don Duong Police Forest Management Board) (Pilgrim et al. 2007). There are also recent reports from Bi Doup Nui Ba National Park, including a photograph of an individual of this species purportedly taken there in 2006 by the Vice-Director of the national park, Mr Do Manh Hung (Le Trong Trai pers. comm. 2008). The location at which this photograph was taken has been described variously as ‘the slopes of Mount Lang Bian’ (Pilgrim et al. 2007) and ‘in forest adjacent, along and close to road 723 at 1,450 m and 1,600 m elevation’ (Do Manh Hung pers. comm. 2009). Both locations would place the record within Bi Doup Nui Ba National Park. Forktail 28 (201 2) Status of Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis 45 Plate 3. Hydro-electric dam construction in the Da Nhim Watershed Protection Forest. A male Grey-crowned Crocias could be heard singing from a tree top behind the photographer. (Jonathan C. Eames) Plate 5. Road construction within the altitude range of Grey-crowned Crocias in the Da Nhim Watershed Protection Forest. Note the gully erosion and the tipping of soil over the side of the road. Note also the clearance of pine forest for coffee. (Jonathan C. Eames) The elevational range of Grey-crowned Crocias is poorly understood, because although there are historical records from up to 1,700 m elevation, recent records (excluding those claimed from Bi Doup Nui Ba National Park and the 1 994 record from Chu Yang Sin National Park, which was at c. 1,615 m) are from a narrowband Plate 2. Habitat of Grey-crowned Crocias in the Da Nhim Watershed Protection Forest. (Jonathan C. Eames) Plate 4. Many parts of the Da Lat Plateau now support only an anthropogenic landscape. Evergreen forest degraded by logging and shifting cultivation can be seen in the far distance. A ridge covered by Pinus kesiya plantation in the middle foreground and irrigated rice and market gardening in the foreground. (Jonathan C. Eames) between 910 and 1,130 m. It forages in the canopy and mid-storey of broadleaved evergreen forest, sometimes utilising scrub between forest patches, in pairs or small groups, in monospecific and mixed- species flocks. METHODS Surveys were undertaken in four Forest Management Units with suitable habitat for the target species located within the Da Lat Plateau EBA, as part of a project to locate and map forest of high conservation value (Mahoodef^/. 2009). We focused on sites where the distribution of the target species was poorly known. The Forest Management Units surveyed were Bi Doup Nui Ba National Park, Da Nhim Watershed Protection Forest, D’Ran Watershed Protection Forest and Don Duong Forest Company. Survey locations are defined as Forest Compartments within Forest Management Units. All of these Forest Management Units receive some level of legal protection from illegal logging and encroachment, although in the case of Forest Companies (state owned logging concessions) and especially Watershed Protection Forests legislation is rarely enforced. Bird surveys were undertaken by three teams consisting of an experienced ornithologist and a relevant member of staff from the Plate 1. Expansion of coffee cultivation in the altitudinal range of Grey- crowned Crocias within the Da Nhim Watershed Protection Forest. (Jonathan C. Eames) 46 SIMON P. MAHOOD & JONATHAN C. EAMES Forktail 28 (2012) Plate 6. Grey-crowned Crocias in the Don Duong Forest Company at the southern limit of its range. (Jonathan C. Eames) Forest Management Unit (local guides were also used where required). Surveys were undertaken in the first two weeks of May 2009. Data were collected throughout the day, starting from dawn. If there was moderate or heavy rain, or moderate or strong wind, then the survey was abandoned. The survey teams actively searched for indicator species on foot, using existing roads and trails. Trails were walked slowly (1 — 1 . 5 km/hour) . The survey teams concentrated their effort in habitat at elevations believed to be most suitable for target species. The most suitable habitat for Collared Laughingthrush is believed to be broadleaved evergreen forest with dense undergrowth above 1,500 m, and for Grey-crowned Crocias broadleaved evergreen forest at 900-1,200 m, although this species was also searched for in forest above 1,200 m. In suitable habitat, the survey team broadcast prerecorded songs of the target bird species at 100 m intervals and waited at each broadcast station for three minutes afterwards to detect responding birds. RESULTS Collared Laughingthrush Collared Laughingthrush was recorded at 93% (13 of 14) survey locations where broadleaved evergreen forest reached over 1,500 m elevation. Our records in Da Nhim Watershed Protection Forest are the first for that site, and they fill a gap in the species’s range between contiguous forest in Bi Doup Nui Ba National Park and Dam Rong State Forest Enterprise. Grey-crowned Crocias Grey-crowned Crocias was recorded at 57% (four of seven) survey locations where contiguous forest reached below 1,200 m and 8% (one of 13) survey locations where forest was only found above 1,200 m. Two sites where it was found were in areas where it was previously unrecorded: pairs were seen at two locations at just over 1,500 m along a dirt road in Forest Compartment 62, Da Nhim Watershed Protection Forest and a further seven pairs were found along a 25 km transect in Forest Compartments 326 and 331, Don Duong Forest Company (Plate 6). We also obtained the first modern records of the species in D’Ran Watershed Protection Forest. Three pairs were located in Forest Compartments 314 and 315 in a mosaic of broadleaved evergreen and coniferous forest. Although the survey team spent four person-days surveying suitable habitat for Grey-crowned Crocias in Bi Doup Nui Ba National Park none was recorded. DISCUSSION The survey gathered new data for two globally threatened bird species. These data have allowed us to expand the range of one species (Grey-crowned Crocias) and re-evaluate the abundance of another (Collared Laughingthrush). Based on these data we reassess the global threat status of both of these species and recommend that they be retained as Endangered. Because their altitudinal distributions encompass almost the full elevational range of all of the other restricted-range species that characterise the Da Lat Plateau EBA and because they are reliant on broadleaved evergreen forest, Grey-crowned Crocias and Collared Laughingthrush are effective analogue species for the Da Lat Plateau EBA. Collared Laughingthrush Owing to the distribution of suitable habitat we only found this species in Bi Doup Nui Ba National Park and the south-east corner Forktail 28 (201 2) Status of Collared Laughingthrush Garrulax yersini and Grey-crowned Crocias Crocias langbianis 47 of the north-west sector of Da Nhim Watershed Protection Forest. However, our records suggest that within its small range Collared Laughingthrush is common wherever suitable habitat is found. The population estimate given in BirdLife International (201 1) is precautionarily placed in the band 2,500-10,000 based on the observation that this species is less abundant than Black-hooded Laughingthrush Garrulx milleti , which has been estimated to occur at a density of 70 individuals/km2 in Chu Yang Sin National Park, where it was recorded on 1 6 occasions compared to four encounters with Collared Laughingthrush (Hill et al. 2001). Unlike Black- hooded Laughingthrush, which occurs in demonstrative flocks that vocalise often, Collared Laughingthrush is usually silent and skulking unless provoked. These two species are therefore quite different in habits and detectability. Pre-recorded vocalisations were not used by Hill et al. (2001) to detect birds. We assert that Collared Laughingthrush is more abundant than formerly believed and that population estimates based on the encounter rate in Hill et al. (2001) probably underestimate the population size; the population might number at least 10,000 individuals. In addition, we recorded Collared Laughingthrush at slightly lower than the minimum recorded elevation, at 1,450 m (Robson 2008). The high-elevation forests occupied by Collared Laughingthrush are more secure than low- and mid-elevation forest on the Da Lat Plateau, in part because they are mainly distributed in protected areas which may confer some degree of security to the species, at least in the short to medium term. Outside of the two protected areas (Chu Yang Sin and Bi Doup Nui Ba National Parks) there is relatively little broadleaved evergreen forest above 1.500 m, but because the overall area of habitat within this species range is low, even these areas are of high significance in a conservation context. In Da Nhim Watershed Protection Forest clearance of high-elevation broadleaved evergreen forest, primarily for coffee cultivation, is widespread. This cash crop is growing in popularity with farmers, although much of the coffee produced is robusta, a relatively low-value crop widely used for the production of instant coffee; thus a relatively large area of coffee cultivation is needed in order for farmers to turn a good profit. In 201 1 Nescafe announced plans to build a large instant coffee factory in nearby Dong Nai province, which is likely to increase the attractiveness of coffee as a cash crop on the Da Lat Plateau. Although habitat loss within its elevational range is less severe than for Grey-crowned Crocias, and despite our belief that it is more abundant than previously thought. Collared Laughingthrush must have a relatively small global population, limited as it is to broadleaved evergreen forest on the Da Lat Plateau. We recommend it be retained as Endangered under criterion Bla+b (i,ii,iii,iv,v). Grey-crowned Crocias Our records of Grey-crowned Crocias slightly expand its known range to the south, to the southern limit of the Da Lat Plateau, and bring the number of sites where it has been recorded to seven. Our data indicate that it is uncommon and localised between 900 and 1,200 m and rare above those elevations. Its occurrence in forest at the higher end of this elevational range may be dependent on the persistence of contiguous forest lower down. The sites at which we found the species contain small areas of forest at a suitable elevation, and therefore do not significantly increase the global population estimate. We concur with BirdLife International (2011 ) that the population of this species is likely to be between 2.500 and 10,000 individuals. Don Duong Forest Company, where we found seven pairs along a 25 km transect in logged broadleaved evergreen forest (Plate 6) probably supports one of the largest populations of the species. We have not been able to confirm its occurrence in Bi Doup Nui Ba National Park, where it must be rare and highly localised at best because although there is much apparently suitable habitat it is almost entirely at the higher end of the species’s altitudinal range. We suggest that records from this location be treated as provisional until they are confirmed by an independent survey. Throughout its range most ol the forest at elevations suitable lor Grey-crowned Crocias has already been converted to agriculture or burned and replaced by coniferous forest (Plate 4). This process is ongoing. At the location where we found the species in Da Nhim Watershed Protection Forest, forest was being cleared for a hydropower dam and coffee cultivation (Plates 1 and 3). Throughout the Da Lat Plateau it is likely that forest conversion activities will increase in line with ongoing road construction as new areas are opened up to human colonisation. Because roads are typically built at lower elevations, associated forest clearance inherently affects lower- and mid-elevation forest before it impacts higher-elevation forest. People have already settled alongside a road that was built through Chu Yang Sin National Park in 2010; this road connects to route 722 along which we found Grey- crowned Crocias in DaNhim Watershed Protection Forest (Plates 2 and 5). Suitable habitat for Grey-crowned Crocias is highly fragmented, although its ability to tolerate and even possibly prefer secondary habitats means that, with appropriate land management (which is a long way off in Vietnam), it could potentially recolonise much of its former range. Although it has recently been found in Kon Turn province (Anon. 2012), its distribution there and in intervening forest is extremely poorly known, and it is likely that, overall, it has a very small global distribution within which all suitable forest is rapidly declining in extent and quality, both inside and outside protected areas. Our survey does not indicate a lower level of threat than was previously assumed, and we recommend that Grey-crowned Crocias be retained as Endangered under criteria Bla+b (i,ii,iii,iv,v). It should be noted that although it has the same threat category as Collared Laughingthrush, it is perhaps inherently more vulnerable to extinction owing to its elevational preferences. ACKNOWLEDGEMENTS We thank the Trust Fund for Forests for financially supporting this survey and the management boards of the special-use forests where we conducted fieldwork for their cooperation. Le Trong Trai of BirdLife in Indochina collected invaluable data for this publication. Pham Xuan Nguyen and other staff from Bi Doup Nui Ba National Park were invaluable in arranging logistics and were good company in the field, and Nguyen Cong Mao our driver took us to places we would otherwise have not been able to reach. Craig Robson and an anonymous referee provided comments on a draft of this manuscript. REFERENCES Anon. (2012) Grey-crowned Crocias crops-up in KonTum province, Vietnam. Downloaded from http://birdlifeindochina.org/ on 28 March 201 2. BirdLife International (201 1) Endemic Bird Area factsheet: Da Lat Plateau. Downloaded from http://www.birdlife.org on 08/08/201 1. BirdLife International (201 1) IUCN Red List for birds. Downloaded from http:/ /www.birdlife.org on 08/08/201 1. Collar, N. J. (2006) A partial revision of the Asian babblers (Timalidae). Forktail 22: 85-112. Eames, J. C. (1995) Endemic birds and protected area development on the Da Lat Plateau (Vietnam) Bird Conserv. tnternatn. 5: 593-425. Eames, J. C. (2002) Eleven new subspecies of babbler (Timaliinae) from Kon Turn Province, Vietnam. Bull. Brit. Orn. Club 122: 109-141. Eames, J. C. & Eames, C. (2001) A new species of laughingthrush Garrulax (Passeriformes: Sylviinae: Garrulacinae) from the Central Highlands of Vietnam. Bull. Brit. Orn. Club 121:1 0—23. 48 SIMON P. MAHOOD & JONATHAN C. EAMES Forktail 28 (2012) Eames, J. C. & Ericson, P. G. P. (1996) The Bjorkegren expedition to French Indochina: a collection of birds from Vietnam and Cambodia. Nat. Hist. Bull. Siam Soc. 44: 75-1 1 1 . Eames, J. C., LeTrong Trai & Nguyen Cu (1995) The rediscovery of the Grey- crowned Crocias Crocias langbianis. Bird Conserv. Internatn. 5: 527-537. Eames, J. C., Le Trong Trai & Nguyen Cu (1999) A new species of laughingthrush Garrulax (Passeriformes: Sylviinae: Garrulacinae) from the Western Highlands of Vietnam. Bull. Brit. Orn. Club 1 19: 4-15. Eames, J. C., Le Trong Trai, Nguyen Cu & Eve, R. (1999) New species of barwing Actinodura (Passeriformes: Sylviinae: Timaliini) from the Western Highlands of Vietnam. Ibis 141 : 1 — 1 0. Hill, M„ Eames, J. C., Le Trong Trai & Nguyen Cu (2001) Population sizes, status and habitat associations of forest birds in Chu Yang Sin Nature Reserve, Dak Lak Province, Vietnam. Bird Conserv. Internatn. 1 1 : 49-70. Inskipp, T„ Lindsey, N. & Duckworth, W. (1996) An annotated checklist of the birds of the Oriental region. Sandy, UK: Oriental Bird Club. Mahood, S. P., Le Trong Trai, Tran Van Hung & Le Anh Hung (2009) Identification, planning and management of high-value forests: final consultancy report. Hanoi: B i rd Life International Vietnam Programme. Pilgrim, J. D„ Nguyen Xuan Vinh, Nguyen Xuan Dang, Polet, G.,ThaiTruyen, Tordoff, A. W„ Tran Huy Manh & Peters, J., eds. (2007) Biological Assessment of the Dong Nai River Basin Conservation Landscape, Vietnam. Final report, 1 9 October 2007. Robson, C. R. (2008) A field guide to the birds of South-East Asia. London: New Holland. Stattersfield, A. J., Crosby, M. J., Long, A. J.& Wege, D. C. (1998) EndemicBird Areas of the world: priorities for biodiversity conservation. Cambridge, UK: BirdLife International (Bird Conservation Series No. 7). Tordoff, A. W., ed. (2002) Directory of Important Bird Areas in Vietnam. Hanoi: BirdLife International in Indochina and the Institute of Ecology and Biological Resources. Simon P. MAHOOD, Wildlife Conservation Society Cambodia Programme, House 2 1, Street 21, Sangkat Tonle Bassac, Phnom Penh, Cambodia. Email: simonmahood@gmail.com Jonathan C. EAMES OBE, BirdLife International Cambodia Programme, #9, Street 29 Tonle Basac, Chamkarmon, P.O. Box 2686, Phnom Penh, Cambodia. Email: eames@birdiife.org.vn FORKTAIL 28 (2012): 49-52 Seasonal changes in plumage coloration of Orange Bullfinches Pyrrhula aurantiaca TILLTOPFER Males of the sexually dimorphic Orange Bullfinch Pyrrhula aurantiaca have previously been thought to have a particular subadult body plumage, whereas there is no such plumage stage known in all other Pyrrhula taxa.This particular plumage is characterised by light yellow ventral colour compared to the intensive orange of adult males. It has alternatively been explained as representing geographic variation. In contrast, reconstruction of the hitherto unknown moult cycle using museum specimens shows that birds are coloured richly orange from late autumn to spring and become increasingly yellowish during the summer. The different coloration is therefore most likely a consequence of colour fading because of carotenoid degradation (i.e. photobleaching) during the breeding season. INTRODUCTION The Pyrrhula bullfinch taxa are traditionally grouped on grounds of distinctive plumage coloration, a pattern that is also supported by molecular data (Bianchi 1907, Voous 1949,Vaurie 1956, Topfer etal. 2011). Of the six (Dickinson 2003) or seven (Clement 2010) currently recognised species, four are sexually dimorphic, with the males being the brighter-coloured sex. For the first few months of their life, juveniles of all Pyrrhula taxa are easily distinguished from adults by an overall brownish and comparatively dull body plumage (e.g. see plate 28 in Clement 1993). This plumage is replaced by the adult plumage during the first autumn (Newton 1966, Jenni & Winkler 1994). In contrast, males of the sexually dimorphic Orange Bullfinch P. aurantiaca have been suspected of having a distinctive subadult body plumage, although there is no phenotypically distinct subadult plumage known in any other Pyrrhula bullfinch. This plumage is said to be characterised by a light yellowish ventral colour compared to the intensive orange of adult males (Sharpe 1888, Oates 1890, Roberts 1992, Rasmussen & Anderton 2005). However, the different plumage colours are not always treated as a sign of age (Grimmett et al. 1999), and have alternatively been assumed to represent geographic variation (Bates & Lowther 1952, Roberts 1992). Given the variety of interpretations and the lack of comprehensive field data, I carried out a thorough re-examination of study skins in order to identify the true nature of the light yellow plumage and to understand the moult cycle of Orange Bullfinches. METHODS I examined 47 study skins (32 males and 15 females) of Orange Bullfinches housed in the ornithological collection of the Natural History Museum in Tring, UK (BMNH; Table 1), including the male syntype of P. aurantiaca Gould, 1858 (BMNH 1858.9.7.5). One male is a juvenile, with the characteristic brownish plumage. The birds were collected at 14 localites in the Kashmir area (Pakistan andlndia) during 1857-1937 (Table 1). These specimens represent the most extensive set of Orange Bullfinch skins available in natural history museums (very few specimens exist in other major collections) and clearly outnumber the published records of individuals from the wild. Most of the specimen-based descriptions in the literature probably refer to these particular skins. The series contains five from the Meinertzhagen collection that is notorious for fraudulent data (Rasmussen & Prys-Jones 2003). However, according to their preparation style, these individuals do seem to be reliably labelled (R. Prys-Jones verbally 201 1), so were included in the analyses. Exclusion of these specimens does not change the results reported here. I visually determined the colour of the ventral body plumage of every male, distinguishing five colour classes: orange, yellowish- orange, orange-yellow, yellow, and light yellow. I also compared these with the best colour match in two colour guides (Smithe 1 975, Anon. 2000). I documented the condition of the plumage in each individual regarding signs of wear, feather gloss and abrasion of feather tips, particularly of flight feathers and coverts, distinguishing the stages: fresh, worn and heavily worn. In addition, I recorded several morphometric parameters: wing length; wing tip; tail length and graduation; bill length, depth and width; tarsus length; and sagittal and lateral tarsus diameter. I analysed the measurements of males according to body plumage colour, comparing orange (those classified as orange or yellowish-orange) with yellow (orange -yellow, yellow and light yellow), using unpaired t-tests (with a = 5%). All altitudes of collected specimens were converted from feet to metres in Table 1. RESULTS The occurrence of orange- and yellow-coloured male Orange Bullfinches was confirmed. However, there are individual differences, with some showing intermediate orange/yellowish coloration. Taking into account this individual variation, there is an obvious relationship between plumage colour and season (Figure 1, Table 1). A rich orange fresh plumage appears to be obtained in October-November and remains bright until May, but gradually fades to yellowish-orange and finally to yellow during summer (Figure 2). This is well illustrated by a moulting bird from October (BMNH 1897.12.10.244) that is growing some new feathers on the belly and back that are contrastingly bright orange compared with the other yellowish-orange feathers, while a male from mid- November (BMNH 1858.9.7.5) shows a consistently rich orange body plumage. According to their labels, at least three males from the beginning of July and one male from mid-August (BMNH 1949.25.3775, 3776, 3778 and 1949.Whi. 1.8743) were in full reproductive condition. All four show a yellow or light yellow coloration. Additionally, the labels of two females from the beginning of July and from mid-August (BMNH 1949.25.3779 and 1969.41. 223) confirm this time as the breeding season, since information on the full development of the reproductive tract as well as data on nesting material and nest construction are given. No significant morphological differences were found between orange and yellow individuals: in all ten morphometric dimensions the two groups are statistically indistinguishable (Table 2). There were no signs of moult limits in the greater wing-coverts of any of the specimens and no distinctive differences in size or coloration that allow ageing of first-year and adult Eurasian Bullfinches 50 TILLTOPFER Forktail 28 (2012) Table 1 . List of male Orange Bullfinch specimens examined, indicating locality and elevation, date, ventral body plumage colour as scored by eye and compared with colour guide codes (Smithe 1 975, Anon. 2000), and plumage condition ('h.' = heavily). Specimens are listed chronologically by day and month. BMNH number Locality and elevation Date Ventral colour Smithe (1975) Anon. (2000) Plumage condition 1949.WhU.8739 Liddar Valley, Kashmir, 1,980 m 15.02.1937 orange 16 13-16-7 fresh 1955.6.N13.31 Dachgam, Kashmir, 1,830 m 13.03.1904 orange 116 13-16-8 fresh 1 965-M-1 7967 Kangan, Sind Valley, Kashmir, 1,950 m 31.03.1925 orange 116 13-17-7 fresh 1965-M-17968 Kangan, Sind Valley, Kashmir, 1,950 m 31.03.1925 orange 16 13-0-8 fresh 1 965-M-1 7969 Kangan, Sind Valley, Kashmir, 1,950 m 31.03.1925 orange 16 13-0-8 fresh 1 965-M-1 7970 Kangan, Sind Valley, Kashmir, 1 ,830 m 03.04.1925 orange 16 13-16-8 fresh 1 949. Whi. 1.8743 Gan Nullah, Kishtawar, Kashmir, 2,285 m 04.05.1931 orange 16 13-16-7 fresh 1897.12.10.248 Kashmir, 2,590 m 19.05.1876 yellowish-orange 17 13-0-8 worn 1888.9.12.324 Kashmir, 2,590 m 19.05.1876 yellowish-orange 16 13-0-8 worn 1925.12.23.298 Gund, Kashmir 28.05.1896 orange 16 13-5-8 worn 1887.6.1.1522 Muree 05.1875 orange 16 13-0-8 fresh 1897.12.10.252 Gilgit, Pakistan, 3,350 m 10.06.1876 yellow 18 13-5-7 h.worn 1887.6.1.1517 Gilgit, Pakistan 13.06.1876 orange-yellow 17 13-5-8 h.worn 1889.6.1.1516 Baltal, Sind Valley 20.06.1870 orange-yellow 17 13-4-8 worn 1897.12.10.250 Gilgit, Pakistan, 3,350 m 24.06.1879 orange-yellow 16 13-4-8 worn 1897.12.10.806 Gilgit, Pakistan, 3,350 m 24.06.1879 yellowish-orange 16/17 13-5-8 fresh 1888.9.12.325 Gilgit district, Pakistan, 2,740 m 06.1876 yellow 18 13-4-8 h.worn 1897.12.10.251 Gilgit, Pakistan, 2,740 m 06.1876 yellowish-orange 17 13-5-8 worn 1897.12.10.253 Gilgit, Pakistan, 2,740 m 06.1876 light yellow 153 5-13-8 h.worn 1949.25.3776 Baltal, Sind Valley, Kashmir, 2,880 m 06.07.1929 light yellow 56 13-5-7 worn 1949.25.3778 Baltal, Sind Valley, Kashmir, 2,880 m 06.07.1929 light yellow 57 13-5-7 worn 1949.25.3777 Baltal, Sind Valley, Kashmir, 3,050 m 06.07.1929 light yellow 56 13-5-8 worn 1949.25.3775 Baltal, Sind Valley, Kashmir, 2,880 m 06.07.1929 yellow 17/18 13-5-8 worn 1881.5.1.2248 Sonamarg, Kashmir 16.07 (year unknown) yellow 17/18 13-5-8 worn 1897.12.10.247 Sonamarg, Kashmir, 2,740 m 16.07.1874 yellow 17/18 13-4-8 h.worn 1897.12.10.249 Sind Valley, Kashmir, 2,740 m 18.07.1874 yellow 18 13-5-8 fresh 1887.6.1.1519 Gilgit Valley, Pakistan 07.1876 light yellow 153 5-13-7 h.worn 1949.Whi.1.8742 Thasgam, Kashmir, 3, 200m 16.08.1934 yellow 153 13-5-7 worn 1887.6.1.1520 Kashmir 09.1875 yellow 17/18 13-5-7 h.worn 1897.12.10.244 Astor, 3,350 m 01.10.1880 orange-yellow 17 13-5-8 h.worn 1897.12.10.245 Astor, 3,350 m 01.10.1880 brown (juvenile) 38/39 9-11-6 fresh 1858.9.7.5 Pir Pinjal Mts, Kashmir 10.11.1857 orange 16 13-0-8 worn P. pyrrhula (Jenni & Winkler 1994, Winkler & Jenni 2007). For example, in all birds the tips of the coverts are whitish-brown on the outer vane and yellowish-brown on the inner vane. This pattern does not differ between the juvenile specimen (BMNH 1897.12.10.245) and the other individuals, and is identical on the growing coverts of one specimen from October (BMNH 1897.12.10.244). The correlation between body feather colour and season is also supported by examination of the overall plumage condition, although with greater individual variation. Specimens from February to mid-May show fresh and glossy remiges that subsequently abrade, becoming heavily worn by August-September (Table 1). In some individuals with only slightly worn flight feathers, the uppertail- coverts show small white edges or tips, while the white edging is Figure 1. Seasonal changes in body plumage coloration of male Orange Bullfinches illustrated by selected specimens (ventral view). The rich orange plumage of birds in spring gradually fades to yellow during summer, returning to orange again after moult in late autumn. From left to right: 1 965- M-17967 (March), 1887.6.2.1522 (May), 1897.12.10.247 (July), 1887.6.1.1520 (September), 1858.9.7.5 (November). © The Natural History Museum. Forktail 28 (2012) Seasonal changes in plumage coloration of Orange Bullfinches Pyrrhula aurantiaca 51 Table 2. Morphological comparisons of yellow and orange individuals. Dimension Unpaired t-test Wing length T29= — 1 .061; n.s. Wing tip T2g= 1.547; n.s. Tail length T8 = 0.33; n.s. Tail graduation T20= 0.914; n.s. Bill length T28= -0.468; n.s. Bill width T2, = 0;n.s. Bill depth T20= 1.429; n.s. Tarsus length T28= -1.947; n.s. Tarsus diameter (sagittal) T29= -0.649; n.s. Tarsus diameter (lateral) T29= -1.658; n.s. absent from birds with heavy wear. This pattern holds true also for female Orange Bullfinches that are not subject to such an obvious change in body plumage coloration as males. All of the specimens were taken from three adjacent regions that today belong to the state Jammu and Kashmir in India and the province Punjab and the special territory Gilgit-Baltistan of Pakistan. The sampling localities are spread over an area of about 300 x 250 km (see Wunderlich 1992), with two main concentrations of male specimens: around Srinagar (Jammu and Kashmir, India; n=19) and in the Gilgit area (Gilgit-Baltistan, Pakistan; n=10). Orange and yellowish individuals occur among the birds from each of these regions. From this geographic sampling, there is no evidence for a geographic separation of orange and yellow plumage colours. DISCUSSION Differences in coloration of Orange Bullfinches appear to be unrelated to age or geography, but instead linked to season and caused by fading. The yellowish and orange colours are generated by the carotenoid pigments canary xanthophyll A and B that are known to be subject to fading over time (McGraw 2006). Because carotenoids are usually deposited in feathers in their free, unesterified form, they are probably more prone to light-induced degradation than esterified pigments (through ‘photobleaching’: McGraw 2006). Taking into account the higher levels of ultraviolet radiation in the Orange Bullfinches’ high- altitude breeding grounds compared to that in their much lower wintering habitats (2,400-3,900 m vs 1,550-2,330 m: Ali & Ripley 1974, Clement 1993, 2010, Rasmussen & Anderton 2005), it appears likely that photobleaching may be more extensive during summer. This could also explain the apparent lack of change in the orange coloration of males during winter (Table 1). Seasonal light-induced fading of carotenoid plumage has also been demonstrated in House Finches Carpodacus mexicanus (McGraw & Hill 2004) and Great Tits Parus major (Figuerola & Senar 2005). Although different sets of carotenoids in congeners may be differently susceptible to light-induced degradation, a similar phenomenon to that reported here might be expected for two closely related Himalayan species: Grey-headed Bullfinch P. erythaca and Red-headed Bullfinch P. erythrocephala. Since feather pigments are metabolised from dietary precursors (McGraw 2006), the consumption of carotenoid-poor food during or prior to feather growth could result in a dull yellow instead of rich orange plumage colour. However, there is no evidence for this in the skins studied: all the specimens dating nearest to the time of moult (i.e. in winter) show bright orange plumage. The variability of coloration is most probably not an artefact of long-term colour fading in museum skins (as suggested by Rasmussen & Anderton 2005): as early as 1888, Sharpe, referring to specimens collected within less than 20 years, already remarked that ‘every shade between yellow and deep orange is exhibited by the series in the Museum’. Moreover, Doucet & Hill (2009) found that reflectance spectra of museum specimens were very similar to those from wild birds, including in the UV range. Notably, the variation in reflectance spectra of wild populations might even exceed the variation in museum specimens obtained from different localities and over a century-long period (Doucet & Hill 2009). Although some influence of specimen age on wavelength reflectance cannot be denied (e.g. Armenata etal. 2008, Doucet & Hill 2009), the results presented here are more likely to be connected to season than to specimen age. The specimen BMNH 1897.12.10.244 (collected in 1880) which has some fresh, bright orange feathers within its yellowish-orange body plumage further indicates that proper century-long storage does not necessarily lead to artificial colour fading. It is very likely that seasonal pigment degradation has a significant impact on plumage UV reflectance. Although not tested here, it is possible that the UV reflection spectrum of the orange plumage acts as a signal of sexual maturity, and that there is no need for such recognition during the late stages of the breeding season. Further work using UV reflectance spectrometry would be informative, but is beyond the scope of the present study. In general, the moult cycle of Orange Bullfinches roughly corresponds to that of the European Bullfinch, in which adults have a complete post-breeding moult, while juveniles undergo a partial moult during which they renew the body feathers, but retain the flight feathers until their second summer (Newton 1966, Busse 1984, Roselaar 1994, Jenni & Winkler 1994, Glutz von Blotzheim 1997, Winkler & Jenni 2007). Clement’s statements (1993, 2010) suggesting that first-summer Orange Bullfinch males are deeper orange above and below than females supports the conclusion that males acquire their first orange body plumage during a post-juvenile moult (Fig. 2). Interestingly, Bates & Lowther (1952) reported that male Orange Bullfinches appear ‘clear yellow’ in the field, and mentioned a specimen from March being much ‘deeper and duller in tone’ than three other males from August that are ‘far brighter’, i.e. more yellowish-orange. This observation precisely fits the findings of this work. Unfortunately, other field studies are still lacking. Thus the change in body plumage coloration from orange to yellow occurs in summer without a moult, but appears loosely associated with the timing of abrasion of flight feathers and uppertail-coverts in both sexes. As there is no evidence for a distinct subadult plumage in the Orange Bullfinch, there is also no indication for sexual plumage heterochrony (see Bjorklund 1991. Badyaev & Hill 2003). Feather abrasion, which leads to seasonal changes in plumage colour and patterns in many bird species by revealing previously hidden colours or patterns, can be excluded as a cause for the colour Figure 2. Moult cycle of the Orange Bullfinch, reconstructed from study skins. The colour change in male body plumage is indicated by the colour gradient of the circle. 52 TILL TOPFER Forktail 28 (2012) changes in Orange Bullfinches. Individual body feathers are bicoloured, having a black base and a yellowish to orange outer portion covering about two-thirds of the feather. Thus, even though heavy wear (particularly of flight feathers) can be associated with orange -yellow or yellow plumages (Table l), extensive abrasion of body feathers would not lead to fading but to darkening of the orange coloration because of the underlying black feather bases. However, such extensive abrasion of body feathers was not found in any of the specimens examined (or in specimens of congeners). The lack of statistical differences in measurements between orange and yellow individuals does not necessarily mean that there are not two age classes, because no significant difference exists in wing length between first-year and adult European Bullfinches (Glutz von Blotzheim 1997). However, while morphometric data alone might not permit identification of age classes in Orange Bullfinches, it is obvious from label annotations that yellow males were in active breeding condition and were not immature. This is further supported by the fact that most specimens from the breeding season (late May to late August; Ward 1908, Ali & Ripley 1974, Roberts 1992, Clement 2010) are yellowish or yellowish- orange instead of rich orange. As the pair-bonds form with the break-up of winter flocks in early May (Roberts 1992, Clement 2010), a rich orange coloration may be important for mate choice, although no field data are available to test this. There is also no evidence for a geographic explanation for the different plumage colours. While Roberts (1992) mentions males from the Gilgit area being distinctively ‘much paler saffron yellow’ than those from the Hazara district (today Khyber Pakhtunkhwa, Pakistan), there is no correlation between colour and locality in the specimens I studied. In both of the areas from which specimens were taken (the Srinagar area and Gilgit), there are both orange and yellowish birds, with identical seasonal patterns of colour change. Thus, there is no evidence for a potential subspecific differentiation as suggested by Roberts (1992). ACKNOWLEDGEMENTS I cordially thank Robert Prys-Jones and Mark P. Adams for providing access to the bird collection of the Natural History Museum at Tring and for their helpful discussion of various aspects of this work. My sincere thanks go also to Raffael Winkler and Mats Bjorklund for their useful remarks on the subject and to Harry Taylor for taking the photograph. I also thank the anonymous referees for their thorough review of the manuscript. This work was supported by the Biodiversity and Climate Research Centre (BiK-F), Frankfurt/Main. REFERENCES Ali, S. & Ripley, S. D. (1974) Handbook of the birds of India and Pakistan, 1 0. Oxford: Oxford University Press. Anon. (2000) Michel Farbenfuhrer / Colour guide, 37th edition. Munchen: Schwaneberger Verlag. Armenata, J. K., Dunn, P. O. & Whittingham, L. A. (2008) Effects of specimen age on plumage color. Auk 1 25: 803-808. Badyaev, A. V. & Hill, G. E. (2003) Avian sexual dichromatism in relation to phylogeny and ecology. Ann. Rev. Ecol. Evol. Syst. 34: 27-49. Bates, R. S. P. & Lowther, E. H. N. (1952) Breeding birds of Kashmir. Oxford: Oxford University Press. Bianchi, V. (1907) [Revision of the forms of the genus Pyrrhula, family Fringillidae, their phylogeny and geographic distribution.] Bull. Acad. Imperial Sci. St. Petersburg (5) 25: 1 59-198. (In Russian.) Bjorklund, M. (1991) Coming of age in fringillid birds: heterochrony in the ontogeny of secondary sexual characters. J. Evol. Biol. 4: 83-92. Busse, P. (1984) Key to sexing and ageing of European passerines. Beitr. Naturk. Nieders. 37, Sonderheft: 1-224. Clement, P. (1 993) Finches and sparrows. London: Christopher Helm. Clement, P. (2010) Family Fringillidae (finches). Pp.512-617 in J. del Hoyo, A. Elliott & D. A. Christie, eds. Handbook of the birds of the world, 1 5. Barcelona: Lynx Edicions. Dickinson, E. C. ed (2003) The Howard and Moore complete checklist of the birds of the World. Third edition. London: Christopher Helm. Doucet, S. M. & Hill, G. E. (2009) Do museum specimens accurately represent wild birds? A case study of carotenoid, melanin, and structural colours in Long-tailed Manakins Chiroxiphia linearis. J. Avian Biol. 40: 146-156. Figuerola, J. & Senar, J. C. (2005) Seasonal changes in carotenoid- and melanin-based plumage coloration in theGreatTit Parus major. Ibis 147: 797-802. Glutz von Blotzheim, U. N. (1997) Gattung Pyrrhula Brisson 1 760. Pp.1 1 30- 1181 in U. N. Glutz von Blotzheim, ed. Handbuch der Vogel Mitteleuropas, 14/11. Wiesbaden: AULA Verlag. Grimmett, R., Inskipp, C. & Inskipp, T. (1999) A guide to the birds of India, Pakistan, Nepal, Bangladesh, Bhutan, Sri Lanka and the Maldives. Princeton: Princeton University Press. Jenni, L. & Winkler, R. (1994) Moult and ageing of European passerines. London: Academic Press. McGraw, K. J. (2006) Mechanics of carotenoid-based coloration. Pp.1 77- 242 in G. E. Hill & K. J. McGraw, eds. Bird coloration, I. Cambridge, Mass.: Harvard University Press. McGraw, K. J.& Hill, G. E. (2004) Plumage color as a dynamic trait: carotenoid pigmentation of male house finches ( Carpodacus mexicanus ) fades during the breeding season. Can. J. Zool. 82: 734-738. Newton, I. (1966) The moult of the Bullfinch Pyrrhula pyrrhula. Ibis 108: 41- 67. Oates, E. W. (1890) The fauna of British India, including Ceylon and Burma, Birds, II. London: Taylor & Francis. Rasmussen, P. C. & Anderton, J. C. (2005) Birds of South Asia, 2. Washington and Barcelona: Smithsonian Institution and Lynx Edicions. Rasmussen, P. C. & Prys-Jones, R. P (2003) History vs mystery: the reliability of museum specimen data. Bull. Brit. Orn. Club 1 23A, Supplement: 66- 94. Roberts, T. J. (1992) The birds of Pakistan, 2. Oxford: Oxford University Press. Roselaar, C. S. (1994) Pyrrhula pyrrhula Bullfinch. Pp.81 5-832 in S. Cramp & C. M. Perrins, eds. The birds of the western Palearctic, 8. Oxford: Oxford University Press. Sharpe, R. B. (1888) Catalogue of the birds of the British Museum, 12, pt. 3. London: Trustees of the British Museum. Smithe, F. B. (1975) Naturalist's color guide. New York: American Museum of Natural History. Topfer,T., Haring, E., Birkhead,T. R., Lopes, R. J., Liu Severinghaus, L., Martens, J. & Packert, M. (201 1 ) A molecular phylogeny of bullfinches Pyrrhula Brisson, 1 760 (Aves: Fringillidae). Molec. Phylogen. Evol. 58: 271-282. Vaurie, C. (1956) Systematic notes on Palearctic birds. No. 21 Fringillidae: the genera Pyrrhula, Eophona, Coccothraustes, and Mycerobas. Amer. Mus. Novit. 1 788. Voous, K. H. (1949) Distributional history of Eurasian bullfinches. Condor 51: 52-81. Ward, A. E. (1908) Further notes on the birds of the provinces of Kashmir and Jammu and adjacent districts. J. Bombay Nat. Hist. Soc. 18: 461- 464. Winkler, R. & Jenni, L. (2007) Alters- und Geschlechtsbestimmung europaischer Singvogel. Sempach: Verlag Schweizerische Vogelwarte. Wunderlich, K. (1 992) Pyrrhula aurantiaca Gould. 18th instalment in H. Dathe &W. M. Loskot, eds. Atlas der Verbreitung palaarktischer Vogel, 1 8. Berlin: Akademie Verlag. Till TOPFER, Senckenberg Natural History Collections Dresden, Museum of Zoology, Konigsbrucker Landstrasse 159, 01109 Dresden, Germany; and Biodiversity and Climate Research Centre & Senckenberg Gesellschaft fur Naturforschung, Senckenberganlage 25, 60325 Frankfurt/M. Germany. Email: TiU.Toepfer@senckenberg.de FORKTAIL 28 (2012): 53-56 The use of tree-fall gaps by a forest interior avian frugivore in a tropical evergreen forest D. KHAMCHA & G. A. GALE Previous studies suggest that forest birds commonly use tree-fall gaps owing to the relatively higher availability of fruits and arthropods. However, others have argued that gaps may not be important for food resources but may offer other services, such as increased protection from predators owing to denser vegetation associated with gaps. We examined the use of tree-fall gaps by Puff-throated Bulbul Alophoixus pallidus, an abundant generalist frugivore found in tropical evergreen forests of East Asia. Bulbuls (family Pycnonotidae) are known to be the most important small frugivores in this region (an area with the highest rate of deforestation in the world), and likely to be integral to regional forest maintenance as well as regeneration. This bulbul did not preferentially use small gap interiors (average 1 63 ± 43 SE m2) and probably avoided them, but it appeared to use areas immediately surrounding gaps more than expected particularly during the breeding season. Such areas probably provide increased security for roosting adults and for young fledglings during the post-fledging period. This first detailed study of the movement of forest bulbuls in tropical East Asia suggests that the use of gaps and gap edges by these frugivores is highly variable in space and time (with equally complex reasons for their use or avoidance) but is likely to impact their role in forest maintenance and restoration. INTRODUCTION Differences in age, size or location of tree-fall gaps — defined by Brokaw (1982) as vertical holes in the forest extending from canopy level down through a height of 2 m above ground — can greatly impact the distribution and abundance of animals by influencing the availability of resources (Fogden 1972). Here we focus on gaps created through tree-fall or large branch-fall owing to wind storms or other natural processes, excluding fire, rather than those caused by human factors. Several initial studies suggested that frugivorous birds foraged more frequently in gaps relative to forest interior sites, presumably because gaps provided greater fruit (as well as arthropod) resources. Additional light levels in tree-fall gaps have been suggested to cause understorey shrubs and lianas to produce more fruits; gaps also provide more foliage in the understorey for herbivorous invertebrates and higher near-ground temperatures, which can also be more suitable for a variety of forest arthropods (Blake Sc Hoppes 1986, Levey 1988, 1990, Malmborg& Willson 1988). These studies also suggested that birds may change their foraging behaviour to increase fruit and arthropod foraging in gaps depending on the availability of resources (Blake & Hoppes 1986). However, more recent detailed studies of gaps and food resources suggest that there is a complex relationship between gap age and use by birds, and that many birds do not appear to track resources tightly in gaps at least at the scales measured (Restrepo et al. 1999). Nevertheless, regenerating gaps may provide other services for birds, particularly protection from predators owing to the density of vegetation (Anders et al. 1998, Bowen et al. 2007). The aim of this study was to investigate whether a common frugivorous bird would preferentially use canopy gaps during the course of a year and whether such preferential use was related to food resources or other resources, particularly cover. We focused on a forest bulbul because of the importance of bulbuls as seed dispersers in East Asian forests (Corlett 1998) and because little is known about the movements of forest bulbuls. Furthermore, South-East Asia has the highest deforestation rate in the world (Sodhi & Brook 2006), so understanding the movement patterns of potential seed dispersers such as these will be fundamental in maximising the restoration potential of the region’s forests. We assessed ( 1 ) whether Puff-throated Bulbuls Alophoixus pallidus , a locally abundant forest species, would spend proportionately more time foraging in natural tree-fall gaps or areas immediately surrounding gaps (referred to here as ‘buffer’ areas) compared to areas away from gaps, and (2) whether they would utilise gaps or buffer areas for resources other than food. METHODS The study was conducted in the 30-ha Mo-singto Long-term Biodiversity Research Plot ( 14°26'N 101°22'E), Khao Yai National Park (~2000 km2) in north-eastern Thailand. The plot consists of a series of ridges and valleys with an elevation range of 730-860 m asl. The vegetation is mostly mature evergreen forest dominated by fleshy-fruited trees witli a ~0.25 ha section of secondary forest at the north edge of the plot (Brockelman 1998). The plot is laid out on a 20 m-square grid. All trees with diameter at breast height (DBH) >1 cm have been identified, labelled with unique ID numbers, and mapped using GIS with a precision ±1 m (Brockelman 1998). When the birds are seen, the tree numbers were used to reference their locations precisely. One hundred and sixty-nine species of bird are known to occur on the plot (Round et al. 2011), with Puff-throated Bulbul having the highest density (>3 individuals/ha) (Gale et al. 2009). Puff- throated Bulbul is a common resident of evergreen forest of northern and north-eastern Thailand (Lekagul & Round 1991), and studies from Khao Yai suggest that it is also the commonest avian frugivore (Kitamuraet^/. 2002, Sankamethawee et al. 2011) and consumes the widest variety of f ruits, based on faecal samples (80 plants species from 185 samples) and direct observations of fruiting plants (>100 species between 2003 and 2008) (Sankamethawee et al. 2011). It forages in the lower to middle canopy with an average foraging height of 9.4 m (range 1-15 m) (P. D. Round et al. unpubl. data), in pairs or in groups of 3-7 birds (Sankamethawee et al. 2009), with an average home range of c.2.2 ha (Tanasarnpaiboon 2008). Birds also occasionally join mixed-species bird flocks feeding on insects (McClure 1974, Nimnuan et al. 2004). Their breeding season extends from February to July (Pierce et al. 2004). Colour-banding was initiated in January 2003 such that at the time of this study all family groups (3 1-32 groups) and the majority of individuals (~300 individuals) on the Mo-singto plot could be individually identified. As far as we are aware no quantitative habitat selection data are available for Puff-throated Bulbul, and while generally considered a forest species (Lekagul & Round 1991) it has been observed foraging along forest/ non-forest edges (Chaikuad 2000). Furthermore, Bowen et al. (2007) found that many species generally classified as forest-interior species were frequently observed in forest gaps, often depending on season. Vegetation was classified into three categories: tree-fall gaps, gap buffer areas (0-10 m from the gap edge) and forest interior. 54 D. KHAMCHA & G. A. GALE Forktail 28 (2012) All the gaps in our study plot were created by natural tree-falls. We used a somewhat narrower definition of a tree-fall gap than Brokaw (1982): following Wongs riphuek (2008), who mapped gaps on the Mo-singto plot just prior to the start of this study, we defined gaps as open areas larger than 25 m2 and covered by vegetation shorter than 5 m. Gap edges were marked with plastic stakes at 1 m intervals. Tagged trees on the edge of gaps were used to georeference the gaps. The coordinates were mapped in a GIS (ArcView 3.2a) to create polygons for each gap and to estimate their area. The 1 0-m buffers around these gaps were drawn in ArcView 3.2a using the Buffer Selected Features command in the XTools extension. We selected ten groups of Puff-throated Bulbul which occupied c.20 ha of the core of the 30 ha Mo-singto plot, and precisely mapped their movements using the georeferenced trees. All activities of the ten selected groups were recorded to determine if they spent proportionately more time in gaps, near gaps or away from gaps in the surrounding forest. The ten selected groups were followed two hours twice per month from May 2007 to April 2008 (for logistical reasons no data were collected in September 2007). The observations started when at least one individual in a group was located. The behavioural data to examine the response to gaps were taken from the second hour of each of the two-hour observations to avoid possible biases relating to greater initial detection in gaps. Observations on a particular group were distributed evenly throughout the day to avoid potential biases associated with time of day. Behavioural data were grouped into two seasons: breeding (February-July) and non-breeding (August-January), which included feeding (on both fruit and arthropods), breeding (nest building, incubating, brooding and feeding offspring) and roosting (perching, calling, preening and resting). To test the frequency of gap and non-gap use, as well the frequency of the different behaviours between breeding and non-breeding seasons, Fisher’s exact tests were carried out using R version 2.1 1.0. Total use areas were based on minimum convex polygons drawn around each of the ten groups using the Minimum Convex Polygon command in ArcView 3.2a for estimating the expected use. We used the ratios of observed vs expected observations per group per habitat to examine the overall trends within and among habitat types. To generate the expected values we used the areal proportions of each habitat type for each group mulriplied by the number of observations of each group in each of the three habitat types. RESULTS There were 28 gaps (average 163±43 SE m2) within the home ranges of the ten selected groups of Puff-throated Bulbul (mean 2.8±0.57 SE gaps/group), totalling 0.5 ha. The percentage areas covered by gaps, 10-m buffers and interior forest were 2.09%, 10.05% and 87.86% for the total area occupied by the ten groups. Based on the Mo-singto tree database, as expected most of the older gaps were covered by a dense layer of herbaceous seedlings and saplings, but had few trees with DBH >1 cm (0.1 trees/m2). The 10-m buffer areas were more densely vegetated (0.4 trees/m2) and tree crowns were typically covered with a dense tangle of lianas. A study of lianas on the Mo-singto plot also indicated that the number of lianas was higher in gap areas (gap interiors and gap-edges) than the primary forest (Wongsriphuek 2008). The database also indicated that the forest interior had the highest tree density (0.6 trees/m2). Overall, 84.56% of the observations occurred in the forest interior (n = 2,120), while the proportion of observations of bulbuls in tree-fall gaps was 1.08% (n =27) and in the 10-m buffers 14.36% (n = 360). The use of gaps and 10-m buffer areas tended to vary with time. Of the observations in tree-fall gaps, the largest percentage occurred in June (30%) (n = 7) and none in October- Figure 1. The precentage of observations of Puff-throated Bulbul activities in tree-fall gaps (n = 27), 10-m buffers surounding gaps (n = 360) and the forest interior (n = 2,120) as a percentage of the total number of observations in each particular habitat based on the total number of observations often different groups in each particular habitat over an 1 1 -month period (May 2007— Apri I 2008). December (Fig. 1). The number of observations in tree-fall gaps was more frequent during the breeding season (February-July). The highest percentage of observations in the buffer areas was during March (17%) (n = 61), the lowest in August (3%) (n = 9) (Figure 1). The number of observations in the forest interior fluctuated relatively little during the study period (Figure 1). Overall, there were 1,579 observations obtained during the breeding season and 928 during the non-breeding season. For all groups combined. Puff-throated Bulbuls used the gaps, 10-m buffers and forest habitat differently between breeding and non¬ breeding seasons (Fisher’s exact test, p< 0.001) (Table 1 andbelow). When analysed separately by group, this difference was significant between seasons for only two groups; however, the other groups seem to have similar patterns (Table 1). Approximately 6 5.75%, 32.01% and 2.24% of the behaviours were associated with roosting, feeding and breeding respectively. There were significant differences in behaviour among habitats for all groups combined (Fisher’s exact test, p < 0.001) (see below). When analysed separately by group, behaviours were also different among habitats for four of the ten groups (Fisher’s exact test, p = 0.01-0.03). Nine out of ten groups appeared to use gaps less than expected (ratios were less than one), with three groups having no detections in the gaps (Table 1). There was only one group that appeared to use gaps more than expected (ratio was greater than one), especially during the breeding season. The remaining groups appeared in gaps more often during the breeding season but still lower than expected overall (Table 1). In contrast, seven out of ten groups used the 10-m buffer areas more than expected (ratios were greater than 1), mostly for roosting and feeding, while two groups appeared to use the buffer areas less than expected. Puff-throated Bulbul was detected more often in the buffer areas during the breeding season than non-breeding season (252 vs 108 observations). Although there were more hours of observation during the breeding than the non-breeding season (12 vs 10 hours per group), with an expected ratio of 6:5 for total observations, there were 5.8 and 2.3 times more observations in the gaps and buffer areas during the breeding season than the non-breeding season respectively (Table 1). Fdowever, the magnitude of the overall seasonal difference in gap use was somewhat skewed owing to the relatively frequent use by one group during the breeding season. There was no particular behaviour associated with the use of buffer areas. For the forest interior there were no trends among the different behaviours or between seasons. There was only one group that occurred in the forest interior less than expected (Table 1). Forktail 28 (2012) The use of tree-fall gaps by a forest interior avian frugivore in a tropical evergreen forest 55 Table 1. Use of tree-fall gaps, 10-m buffer zones surrounding gaps and forest interior by Puff-throated Bulbul from May 2007 to April 2008. The breeding season (BR) is during February-July and the non-breeding (NB) August-January. The ratio refers to the ratio of observations per group per habitat to expected observations. Expected observations were generated from the areal proportions of each habitat type for each group multiplied by the total number of observations of each group in all habitats. The total expected observations were generated from the proportions of each habitat type across all home ranges multiplied by the total number of observations of all groups in all habitats. Based on the total number of hours of observation during each season, the expected ratio of breeding to non-breeding season observations was approximately 6:5. Gap Buffer Forest Season Season Season Group BR NB No. of observations Expected no. of observations Ratio BR NB No. of observations Expected no. of observations Ratio BR NB No. of observations Expected no. of observations Ratio 1 2 0 2 4.7 0.42 50 36 86 28.3 3.04 81 67 148 203.0 0.73 2 13 2 15 10.5 1.43 34 17 51 42.5 1.20 121 63 184 197.0 0.93 3 2 0 2 2.5 0.81 16 6 22 14.8 1.48 142 81 223 229.7 0.97 4 1 0 1 1.5 0.65 2 2 4 14.0 0.29 139 75 214 203.2 1.05 5' 2 0 2 2.2 0.92 31 6 37 21.8 1.70 110 69 179 191.8 0.93 6 3 0 3 4.5 0.66 45 8 53 29.5 1.80 100 71 171 193.0 0.89 7 0 0 0 4.2 0.00 18 8 26 27.4 0.95 97 88 185 179.4 1.03 8 0 2 2 2.7 0.73 22 14 36 13.7 2.63 143 92 235 257.6 0.92 9 0 0 0 3.2 0.00 2 4 6 19.4 0.31 204 113 317 300.4 1.06 10* 0 0 0 4.2 0.00 32 7 39 34.2 1.14 167 97 264 264.5 1.00 Total 23 4 27 52.0 0.52 252 108 360 251.0 1.43 1304 816 2120 2,193.0 0.97 'Groups which used habitat types differently between breeding and non-breeding seasons (Fisher's exact test, p< 0.05) Although breeding behaviours did not occur significantly more often in gaps or 1 0-m buffer areas, of the 56 observations associated with breeding, 26 (46%) occurred during the post-fledging period. Eight percent occurred in gaps, 19% in the 10-m buffer areas and 73% in the forest interior. Thus, while more than 25% of the breeding-associated observations were related to caring for fledglings in or adjacent to gaps, the proportion was not significantly different from expected (%2 = 3.7, p = 0.057), although suggestive of greater use of gaps and buffers for fledgling care. Of the seven groups that successfully produced fledglings, three were observed utilising gaps or 10-m buffer areas with their fledglings. In the forest interior the number of observations during the post- fledging period was not significantly different from expected (%2 = 0.37, p = 0.54). DISCUSSION Overall, there were only 27 observations of Puff-throated Bulbuls using gaps out of >2,500 total observations, suggesting there was no preference for gaps, and perhaps gap avoidance, with the frequency of gap use being significantly lower than expected (%2 = 12.02, p < 0.001; Table 1). During the non-breeding season, Puff-throated Bulbuls were detected in gaps much less than expected. Most of the detections in the gaps occurred during the breeding season. Overall only one group out of ten appeared in the gaps more than expected, while the other nine groups used them less than expected, including three which were never observed to use gaps. These results are in agreement with Bowen et al. (2007), who suggested that forest bird use of gaps was seasonal and, depending on species, more or less common than expected. For these bulbuls open gaps may present a greater risk from predators (Willson et al. 1982, Belisle & Desrochers 2002). Although we did not determine the exact ages of the gaps, Puff-throated Bulbuls tended to care for fledglings in older gaps where there was more vegetation cover. Overall, the 10-m buffer areas appeared to be particularly important locations for roosting and feeding year-round, presumably owing to the dense cover and density of food resources. Regenerating gaps and areas immediately surrounding gaps were covered by dense vegetation, especially lianas, at the gap edges rather than in gap interiors. Pull-throated Bulbul prelerred to roost during the day in areas adjacent to gaps where there was probably greater cover and where they seemed to spend more time feeding fledglings as well, presumably to reduce the risk from predators, especially raptors (Anders et al. 1998, Belisle & Desrochers 2002, Bowen et al. 2007). While it is not possible to generalise from one species, this paper and Khamcha et al. (2012) represent the first detailed studies of the movement of forest bulbuls in tropical East Asia, and suggest that the use of gaps and gap edges by bulbuls is probably highly variable in space and time and the reasons for their use or avoidance equally complex and probably dependent on the type of resource that the gap and the surrounding habitat provides. Based on our observations, the Puff-throated Bulbul probably contributes little to forest gap regeneration, but rather to the overall maintenance of forest interiors. Other forest bulbuls are worth investigation, as at least some are highly likely to prefer gaps and therefore may be significant contributors to the regeneration of fragmented forests in the region. ACKNOWLEDGEMENTS We thank the Department of National Parks, Wildlife and Plant Conservation and Khao Yai National Park for granting permission to conduct this study. DK thanks W. Sankamethawee, N. Sukumal, M. Pliusungnoen, J. Khunwongsa, P. Wongsorn, Y. Rattanapongsai, T. Ong-in, the Khao Yai research team and the Conservation Ecology Program students for invaluable assistance in the field. We also thank T. Savini, V. Chimchome and W. Y. Brockelman for advice on the development on this project and manuscript. We especially acknowledge the late David H. Reed for his advice on statistical analysis; we will miss him greatly. The study was funded in part by the Thai Biodiversity Research and Training Program (BRT T_350009). REFERENCES Anders, A. D„ Faaborg, J. & Thompson, F. R. (1998) Post fledging dispersal, habitat use, and home-range size of juvenile wood thrushes. Auk 1 1 5: 349-358. Belisle, M. & Desrochers, A. ( 2002) Gap-crossing decisions by forest birds: an empirical for parameterizing spatia I ly-expl icit, individual-based models. Landscape Ecol. 1 7: 219-231 . 56 D. KHAMCHA&G. A. GALE Forktail 28 (2012) Blake, J. G. & Hoppes, G. W. 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Gale, G. A., Round, P. D., Pierce, A. J., Nimnuan, S., Pattanavibool, A. & Brockelman, W. Y. (2009) A field test of distance sampling methods for a tropical forest bird community. Auk 1 26: 439-448. Khamcha, D„ Savin i,T., Brockleman, W. Y., Chimchome, V. & Gale, G. A. (2012) Influence of food availability and distribution on the movement patterns of a forest avian frugivore, the puff-throated bulbul ( Alophoixus pallidus).J. Trop. Ecol. 28: 1-9. Kitamura, K., Yumoto,T., Poonswad, P., Chualua, P., Plongmai, K., Maruhashi, T. & Noma, N. (2002) Interaction between fleshy fruits and frugivores in a tropical seasonal forest in Thailand. Oecologia 1 33: 559-572. Lekagul, B. & Round, P. D. (1991) A guide to the birds of Thailand. Bangkok: Saha Karn Bhaet. Levey, D. J. (1988) Tropical wet forest tree fall gaps and distributions of understory birds and plants. Ecology 69: 1076-1089. Levey, D. J. (1990) Habitat-dependent fruiting behaviour of an understorey tree, Miconia centrodesma, and tropical tree fall gaps as keystone habitats for frugivores in Costa Rica. J. Trop. Ecol. 6: 409- 420. Malmborg, P. K. & Willson, M. F. (1988) Foraging ecology of avian frugivores and some consequences for seed dispersal in an Illinois woodlot. Condor 90: 173-186. McClure, H. E. (1974) Some bionomics of the birds of Khao Yai National Park, Thailand. Nat. Hist. Bull. Siam. Soc. 25: 99-184. Nimnuan, S., Round, P. D. & Gale, G. A. (2004) Structure and composition of mixed-species insectivorous bird flocks in Khao Yai National Park, Thailand. Nat. Hist. Bull. Siam. Soc. 52: 71-79. Pierce, A. J.,Tokue, K., Pobprasert, K. & Round, P. D. ( 2004) Observations on the breeding of the Puff-throated Bulbul Alophoixus pallidus in north¬ east Thailand. Forktail 20: 100-101. Restrepo, C., Gomez, N. & Heredia, S. (1999) Anthropogenic edges, tree fall gaps, and fruit-frugivore interactions in a neotropical montane forest. Ecology 80: 668-685. Round, P. D., Pierce, A. J., Sankamethawee, W. & Gale, G. A. (201 1) The avifauna of the Mo-Singto forest dynamics plot, KhaoYai National Park, Thailand. Nat. Hist. Bull. Siam. Soc. 57: 57-80. Sankamethawee, W., Gale, G. A. & Hardesty, B. D. (2009) Post-fledging survival of the cooperatively breeding Puff-throated Bulbul ( Alophoixus pallidus). Condor 111: 675-683. Sankamethawee, W., Pierce, A. J., Gale, G. A. & Hardesty, B. D. (2011) Plant- frugivore interactions in an intact tropical forest in northeast Thailand. Integrative Zoology 6:195-212. Sodhi, N. S. & Brook, B. W. (2006) Southeast Asian biodiversity in crisis. Cambridge: Cambridge University Press. Tanasarnpaiboon, S. (2008) Relationships among habitat quality, home range size and group size in the Puff-throated Bulbul ( Alophoixus pallidus ) at Mo-Singto Khao Yai National Park. MSc. dissertation, Mahidol University, Bangkok. Willson, M. F., Porter, E. A. & Condit, R. S. (1982) Avian frugivore activity in relation to forest light gaps. Carib. J. Sci. 18: 1-4. Wongsriphuek, C. (2008) Seed dispersal and seedling recruitment of lianas in seasonal evergreen forest, Khao Yai National Park, Thailand. PhD dissertation, Mahidol University, Bangkok. Daphawan KHAMCHA and George A. GALE, Conservation Ecology Program, School of Bioresources and Technology, King Mongkut's University of Technology Thonburi, 49 Thakham, Bangkhuntien, Bangkok, Thailand. 10150. E-mails: daphawan@hotmall.com and george.and@kmutt.ac.th FORKTAIL 28 (2012): 57-61 Status and distribution of the endangered Baer's Pochard Aythya baeri in Bangladesh SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M.THOMPSON Continued declines in the now endangered Baer's Pochard Aythya baeri imply that this species may well be entering a downward spiral towards global extinction. Knowledge of the drivers of this decline remain fairly anecdotal and even basic data on population trends are hard to come by given the species's similarity to other diving ducks. We conducted a review of past records of Baer's Pochard in Bangladesh, given concerns over the reliability of some past counts, to determine the true magnitude of recent declines, and also present counts from targeted surveys in Bangladesh in the 2010-201 1 winter in addition to some basic autecological data. We conclude that several of the high counts from the period 1 990-2005 may have been in error, but even accounting for these, the population has declined nationally by 99% in just two decades. INTRODUCTION Baer’s Pochard Aythya baeri is a globally threatened diving duck which breeds in Russia and north-eastern China and winters mainly in eastern and southern mainland China, India, Bangladesh and Myanmar. It was uplisted from Vulnerable to Endangered on the IUCN Red List in 2008 in response to evidence of a drastic decline in numbers at known wintering grounds. The principal drivers of this decline are suspected to be habitat loss and degradation plus over-hunting; the most recent global population estimate totalled just 5,000 individuals. It is now absent or rare over much of its former wintering grounds. For example, Thailand held as many as 426 individuals in the late 1980s but now only plays host to single¬ figure counts each winter (BirdLife International 201 1). BirdLife International (2011) estimated the wintering population of Baer’s Pochard in Bangladesh and India to be of the order of 1 ,000-2,000 individuals; however, recent counts from both the countries suggest that this is a significant overestimate of at least one order of magnitude, with a maximum of only c.100 individuals in the last five years. The earlier estimate of 1,000-2,000 is probably based on counts from the 1990s and early 2000s, some of which are of questionable reliability (Li et al. 2009). Given the apparent significance of Bangladesh for this species and a sharp population decline in its wintering areas, we reviewed all past records and counts of the species. In addition, SUC conducted a survey of the wintering population of Baer’s Pochard in Bangladesh in 2010- 2011 that spans a wider spatial and temporal range than previous surveys and complements recent data from Asian Waterbird Censuses (Li et al. 2009). METHODS Review of records In addition to compiling and reviewing past published records of Baer’s Pochards, five senior birdwatchers in Bangladesh were contacted for unpublished records of this species, with counts also obtained from monitoring of a key site — Hakaluki Haor — and information obtained on the reliability of past Asian Waterbird Census (AWC) records. We compiled all records of Baer’s Pochard from Bangladesh based on both published literature and unpublished observations. It is important to note that Baer’s Pochard is often difficult to separate from the sympatrically wintering Ferruginous Duck Aythya nyroca and Tufted Duck A. fuligula (Grimmett et al. 1998, BirdLife International 2001), depending on expertise of the observer, viewing conditions and quality of optical equipment. In order to assess the reliability of old records, previous surveyors were contacted for Baer’s Pochard survey methodology, field descriptions and in rare cases photographic documentation. We treated older records as reliable if the observers were known to be experienced surveyors with previous experience of identifying this species. Some AWC records were treated as unreliable if experienced observers, listed as present, were subsequently found not to have participated in the counts. Waterfowl surveys SUC undertook waterfowl surveys in two haors, Hakaluki and Tanguar, which are large depressions in north-east Bangladesh that are deeply flooded in the wet season and retain water in several beels (shallow waterbodies) in the dry season; historically the majority of Baer’s Pochard sightings in Bangladesh have been located at these haors. A total of 1 8 beels were thoroughly surveyed for Baer’s Pochard and other globally threatened or Near Threatened species such as Ferruginous Duck, Falcated Duck Anas Jalcata and Baikal Teal A. formosa between December 2010 and February 2011. Additional sites which had previously hosted Baer’s Pochards such as Muhuri Dam, Hail Haor and some parts of the Jamuna River were also surveyed. At least two observers conducted surveys together at each site to minimise counting and identification errors. Surveys were conducted both on foot and by boat. Counts were repeated twice on all occasions and the maximum numbers are presented here. Observations were made using 10x42 binoculars and 25-50x spotting scopes. Behavioural observations SUC selected three male Baer’s Pochards to investigate their diurnal time-activity budget, since males were easier to separate while foraging within mixed flocks than females. These three individuals were observed for c.180 minutes between 08h00 and 14h00 over two days. Behaviour categories were established for four main activities: foraging (diving, up-ending), resting (floating motionless on the water with eyes open or sleeping), preening and swimming (moving from one place to another). In each flock-scan, all birds in the flock were checked from a rowing-boat for Baer’s Pochard, working systematically from one side of the flock to the other using a spotting scope. In addition, a GPS unit was used to measure distance between the foraging site and the lakeshore. Water depths were determined using a measuring tape attached to a bamboo stick. RESULTS Globally threatened waterfowl surveys in the 2010-2011 winter During the winter 2010-2011 a minimum total of five Baer’s Pochards were recorded by this survey, comprising four individuals 58 SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M. THOMPSON Forktail 28 (2012) Table 1. Abundance of internationally significant waterfowl counted between December 2010 and February 201 1 during a search for Baer's Pochard in selected key wetlands. Site name Coordinates Survey dates Baer's Pochard Aythya baeri (EN) Ferruginous Duck Aythya nyroca (NT) Falcated Duck Anas falcata (NT) Baikal Teal Anas formosa (VU) No of individuals recorded No of species recorded Chatla Beel (Hakaluki Haor) 24.381 2°N 92.5195°E 22/12/2010 - 46 2 - 1,270 20 Hawa-banna Beel (Hakaluki Haor) 24.3745°N 92.1573°E 23/12/2010 - 52 4 - 640 13 Foot Beel (Hakaluki Haor) 24.3733°N92.1471°E 23/12/2010 - 12 - - 1,655 9 Hingaijur Beel (Hakaluki Haor) 24.3733°N92.147TE 24/12/2010 - 1,450 - - 1,780 11 Haor Khal Beel (Hakaluki Haor) 24.4038°N 92.2543°E 25/12/2010 - 70 2 - 6,400 16 Knna Muina 25.0432°N91.5547°E 19/02/2011 - 270 2 - 788 13 Rowa Beel (Tanguar Haor) 25.0839°N 91.5412°E 20/02/2011 1 248 - - 1,800 8 Rupaboi Beel (Tanguar Haor) 25.0732°N91.4424°E 21/02/2011 - 220 2 - 430 4 Hatirgata Beel (Tanguar Haor) 25.0844°N91.3423°E 22/02/2011 4 120 5 1 9,000 12 Muhuri Dam 22.5050°N 91.2820°E 10/01/2011 - 28 - - 1,290 10 (three males and one female) at Hatirgata Beel and one female at Rowa Beel, both in Tanguar Haor (Table 1). The four individuals (2% of the total number of waterfowl counted in the flock) at Hatirgata Beel were c.200 m from the shoreline at 1.88 m water depth, within a mixed flock of 166 Red-crested Pochard Netta rufrna (83%), ten Gadwall Anas strepera (5%), eight Ferruginous Duck Aythya nyroca (4%), six Common Teal Anas crecca (3%) and six Common Pochard Aythya ferina (3%). The single individual at Rowa Beel was seen c.25 m from the shoreline at 1.26 m water depth and was not accompanied by other waterfowl. In addition, in this winter four individuals were recorded at Pashua Beel during the regular mid-January waterbird census (E. Haquepers. comm. 2011), and in Hakaluki Haor seven were seen at Haor Khal and one male in Nagoa-Loribari during a census of the haor in mid-February, giving a total of 17 Baer’s Pochard for the country (it is unlikely that there was duplication in the counts given the close dates and distances between sites). Three other globally threatened waterfowl species (Table 1) were recorded including the third national record of Baikal Teal and 17 Falcated Duck, which is the highest count since 1993 in Bangladesh (Thompson et al. 1993, Thompson & Johnson 2003). Baer's Pochard diurnal time-activity budget The activity budget data indicated (Figure 1) that Baer’s Pochard spent most of its time between 08h00 and 14h00 resting (58%) and swimming (32%), and considerably less time in foraging (7.4%) and preening (2.6%). Baer’s Pochards were actively foraging and swimming during the early hours of the day, mostly prior to 1 OhOO and primarily stationary (either resting or preening) after 10h30. Certainly, this information is insufficient to draw a firm conclusion on the overall diurnal activity pattern of the Baer’s Pochard, and further detailed observations are needed. Figure 1 . Diurnal time-activity budgets of Baer's Pochard at Tanguar Haor, Bangladesh, observed for c.180 minutes between 08h00 and 14h00 over two days in February 201 1 . llhOO ^ lOhOO •A v. Foraging Swimming H 09h00 ffl Preenins 0 Resting 08h00 ■ 0% 20% 40% 60% 80% 100% Percentage of time Review of records since 1990 We were able to trace 59 reports of Baer s Pochards from Bangladesh in the period 1990-2011, of which 40 records are previously unpublished in detail (including observations from 2011), although aggregate Asian Waterbird Census (AWC) counts have been used in analyses of threat status of this species. Table 2 lists all records known to us. Of these we consider 50 of the 59 records (85%) to be reliable using the criteria outlined above. We consider that the AWC published counts highlighted below and in Table 2 should be treated as inconclusive, and the current national AWC coordinator is of the same opinion (E. Haque pers. comm. 2011). The counts deemed unproven comprise: 510 individuals from Hakaluki Haor and Pasua Beel in 1995; 3,010 individuals mostly from Pashua, Tanguar and Hakaluki Haors in 1996; and 3,580 individuals from Taguar Haor, Pashua Beel and Khana Muair Beel of Sylhet division (Fi et al. 2009) in 2001. These AWC records (D. Fi in lift. 2007) also included names of experienced but non- participatory observers, so there was potentially a dearth of experience within the survey team in 1996 and 2001; hence some of the highest counts of this species which might have played a key role in the current global population estimate (5,000 mature individuals) (BirdFife International 2011) are considered potentially unreliable. Based on Table 2 the peak numbers recorded in a winter were about 700 in January-February 1992 and about 1,700 injanuary- February 1993 (the high estimates at three sites in 1993 are considered reasonable, as very large numbers of duck were present in the haors that winter), and just under 900 in 1994. Between 1995 and 2000 the true status is confused by the unreliable counts and some gaps in coverage of key wetlands. After the last larger count of 70 at Tanguar Haor in 2001 the most important site has been Hakaluki Haor, where an average of about 10 birds has been recorded in each of the last four winters. Table 3 summarises the annual totals and peak counts each year for the two wetland systems most regularly used by Baer’s Pochard — Tanguar and Hakaluki Haors — and also indicates those years for which data are unavailable or unreliable. The average annual count was about 660 during 1991-1995, is unknown during 1996-2000, was 99 during 2001- 2005 and has been only 6 during 2006-2010. DISCUSSION Population decline Records of just five individuals during the targeted survey, an additional 1 2 individuals during regular surveys in the winter 20 1 0- 2011, plus our review of past records, indicate that the wintering population of Baer’s Pochard in Bangladesh has undergone a decline of about 90% in the last ten years, and of 99% since the early 1 990s even after discounting the highest counts that are considered Forktail 28 (201 2) Status and distribution of the endangered Baer's Pochard Aythya baeri in Bangladesh 59 Table 2. Sightings and claims of Baer's Pochard in Bangladesh between 1900 and 2011 Site (name and district) Year Month Day Number Observer/ Reference Zoo Lake, Dhaka 1990 Jan 2 4 AWC Peelkhana Lake, Dhaka 1990 Jan 5 3 AWC Shibaloy, Kamalapur, Bhumiheen Chars, Dhaka 1990 Jan 12 3 AWC Zoo Lake, Dhaka 1991 Jan 4 3 AWC Peelkhana Lake, Dhaka 1991 Jan 5 2 AWC Shibaloy, Kamalapur, Bhumiheen Chars, Dhaka 1991 Jan 6 3 AWC Hakaluki Haor (Chatla & Pingla Beels), Moulavi Bazar 1992 Feb 19 5 D.A. Scott & A. Khan in FAP 6 (1995) (in table prepared by S.M.A. Rashid but not listed in the text written by D.A. Scott) Tanguar-Rauar Beel, Sunamganj 1992 Mar 2 1 D.A. Scott & S.M.A. Rashid in FAP 6 (1995) BanuarHaor.Sunamganj 1992 Mar 2 4 S.M.A. Rashid & A. Khan in FAP 6 (1995) Matian Haor (Palair Beel), Sunamganj 1992 Mar 3 20 D.A.Scott in FAP 6 (1995) Tanguar Haor (Pana Beel), Sunamganj 1992 Mar 3 660 D.A. Scott & S.M.A. Rashid in FAP 6 (1995) Pashua Beel, Sunamganj 1992 Mar 4 2 D.A. Scott & S.M.A. Rashid in FAP 6 (1995) Tanguar Haor (Pana Beel), Sunamganj 1992 Mar 23 6 S.M.A. Rashid & A. Khan in FAP 6 1995 Pashua Beel, Sunamganj 1993 Jan 27 600 S.M.A. Rashid & A. Khan in FAP 6 (1995) Tanguar Haor, Sunamganj 1993 Jan 27 112 P.Thompson in Thompson etal. 1993 Hakaluki Haor (Haorkhal), Moulavi Bazar 1993 Jan 30 300 S. M.A. Rashid in FAP61995 Hakaluki Haor (Chatla & Pingla Beels), Moulavi Bazar 1993 Feb 1 700 S.M.A. Rashid in FAP 6 (1995) Hail Haor (fish ponds), Moulavi Bazar 1993 Oct 24 2 S.M.A.Rashid in FAP 6 (1995) Hakaluki Haor (Haorkhal), Moulavi Bazar 1993 Nov 1 2 S.M.A. Rashid & A. Khan in FAP 6 (1995) Tanguar Haor, Sunamganj 1994 Jan 14 600 AWC Banuar Haor, Sunamganj 1994 Jan 14 4 AWC Pashua Beel, Sunamganj 1994 Jan 15 175 AWC Shanir Haor, Sunamganj 1994 Jan 15 10 AWC Medol Haor, Sylhet 1994 Jan 18 10 AWC Bara Haor, Sylhet 1994 Jan 18 20 AWC Hakaluki Haor, Moulavi Bazar 1994 Jan 20 70 AWC Zoo Lake, Dhaka 1994 Feb 11 1 P.Thompson pers.obs. Hakaluki Haor, Moulavi Bazar 1995 Jan 15 500 AWC - not considered reliable Pashua Beel, Sunamganj 1995 Jan 15 20 AWC - not considered reliable Hakaluki Haor, Moulavi Bazar 1996 Jan 19 800 AWC - not considered reliable Pashua Beel, Sunamganj 1996 Jan 20 1,000 AWC - not considered reliable Shanir Haor, Sunamganj 1996 Jan 20 10 AWC - not considered reliable Tanguar Haor, Sunamganj 1996 Jan 21 1,200 AWC- not considered reliable Tanguar Haor (Pana Beel), Sunamganj 1996 Feb 18 2 D Johnson, R. Haider, P.Thompson, - P.Thompson in litt. 2011 Banua Beel, Matian Haor, Sunamganj 1996 Feb 20 30 D Johnson, R. Haider, P.Thompson, - P.Thompson in litt. 201 1 Muhuri Dam,Feni 1999 Jan 29 4 P.Thompson, E., Haque, R. Haider, W.J. Collis, - P.Thompson in litt. 201 1 Khana Muia Beel 2001 Jan 20 1,000 AWC - not considered reliable Pashua Beel, Sunamganj 2001 Jan 20 80 AWC - not considered reliable Tangua Haor Complex Sunamganj 2001 Jan 21 2,500 AWC - not considered reliable Tanguar Haor (Pana Beel), Sunamganj 2001 Feb 20 4 AWC Tangua Haor, Sunamganj 2001 Feb 24-25 70 E.U. Haque in P.Thompson and Johnson, D.2003 Jamuna River, Gaibanda 2002 Jan 25 28 E. U. Haque in P.Thompson and Johnson, D.2003 Pashua Beel, Sunamganj 2002 Feb 22 24 AWC Tanguar Haor, Sunamganj 2002 Feb 23 95 AWC Khana Muia Beel 2002 Feb 23 6 AWC Tanguar Haor, Sunamganj 2003 Feb 14 30 AWC Shanir Haor, Sunamganj 2004 Jan 11 2 AWC Tanguar Haor (Rowa Beel), Sunamganj 2004 Jan 12 17 AWC Tanguar Haor, Sunamganj 2004 Jan 12 6 AWC Muhuri Dam.Feni 2005 Jan 2 12 AWC Chatidhara (Chatidhora) Beel, Sylhet 2005 Jan 28 2 AWC Tanguar Haor, Sunamganj 2005 Jan 31 202 AWC Hail Haor (Baikka Beel), Moulavi Bazar 2008 Feb 17 2 E.U. Haque and P.Thompson - P.Thompson in litt. 2011 Hakaluki Haor, Moulavi Bazar 2008 Feb 7 10 UNDP via P.Thompson in litt. 201 1 Hakaluki Haor, Moulavi Bazar 2009 Feb 7 14 E.U. Haque in litt.. 2009 Hakaluki Haor, Moulavi Bazar 2010 Feb 7 3 UNDP via P.Thompson in litt. 201 1 Hakaluki Haor, Moulavi Bazar 2011 Feb 20-21 8 P.Thompson, E.U. Haque in litt. 201 1 Pashua Beel, Sunamganj 2011 Jan 7 4 E.U. Haque in litt.. Tanguar Haor, Sunamganj 2011 Jan 20-22 5 S.U.Chowdhury (this paper) 60 SAYAM U. CHOWDHURY, ALEXANDER C. LEES & PAUL M. THOMPSON Forktail 28 (2012) Table 3. Total and highest annual counts of Baer's Pochard in Bangladesh and two main sites. Year Total sightings Tanguar complex (peak) Hakaluki complex (peak) [1990] 10 NS NS 1991 8 NS 0 1992 698 660 5 1993 1,716 112 1,000 1994 890 600 70 1995 0 0 NA [1996] 32 NA NA [1997] 0 NS NS [1998] 0 NS NS [1999] 4 NS NS [2000] 0 NS NS 2001 74 70 NS 2002 151 95 NS 2003 30 30 0 2004 25 23 0 2005 216 202 0 2006 0 0 0 2007 0 0 0 2008 12 0 10 2009 14 0 14 2010 3 0 3 2011 17 5 8 1 NS - Not Surveyed, NA - Not Accepted. 2 Year with square brackets indicate no count or no acceptable count from either of main sites. unreliable (Table 2). Without historical data we cannot refute the potential hypothesis that these early high counts may have been a temporary phenomenon involving displacement of birds from other wintering areas, but we consider this unlikely given that other potential wetland sites were also covered by the AWC, at least within Bangladesh. Moreover, there are few if any unsurveyed sites to which birds might have relocated. Globally, between the period 1998 and 2008, a sharp decline (50-70%) has occurred, leaving an estimated current global population of just 5,000 mature individuals. Considering no double-counting between sites, an estimated c. 1 ,000 Baer’s Pochards were recorded between late 2010 and early 201 1 in the wintering grounds, comprising 760 individuals at Wuchang Hu, Anhui province, eastern China, which appears to be the largest known current population of this species (M. Barter inlitt. 2010); 152 in four sites of Hubei province, central China (J. Lie in litt. 2011) and 1 in Deobali Jalah, Assam, India (P. Baruah pers. comm. 201 1). Current threats in Bangladesh The lowlands of north-east Bangladesh formerly provided a wide variety of habitats for waterbirds; shallow wetlands, seasonally flooded grasslands and swamp forest were ubiquitous just a century ago, but substantial areas have now been converted to agricultural use (BirdLife International 2004). Illegal hunting of waterfowl using poisoned baits (Alpha-Chloralose in boiled rice) in the two major wetland complexes has proved to be a significant problem, particularly at Hakaluki Haor with c.500 ducks poisoned in one recent incident {The Daily Star 2010). The Northern Anas acuta is the commonest victim of the poisoned baits, but Ferruginous Ducks were also reported to be killed ( The Daily Star 2009). Although no evidence was found of trapping or poisoning of Baer’s Pochard in recent years, hunting is still likely to be a significant threat. Changes in attitudes and practices amongst people who presently poison or trap ducks, and the enforcement of laws, including an end to all use of poisoned baits, are urgently needed for future protection of all waterfowl, globally threatened or otherwise, in Bangladesh. Human presence may also cause indirect effects through disturbance. At Tanguar Haor fishing boats regularly pass through most parts of the wetland. As these boats approach closer the ducks abandon where they are feeding or loafing due to the interruption (Muzaffar 2004). This might reduce the foraging efficiency of migratory ducks, considering that the ducks are tolerant of boats only if they are over 30 m from the toraging or roosting spot (Muzalfar 2004). Recent management changes at Tanguar Haor (a Ramsar site) have set rules to permit lishing by local people, but have yet to zone the wetland effectively to provide disturbance- free areas for wildlife. The resultant widespread small-scale fishing may cause more disturbance than the traditional previous practice of mass fishing of large areas once every three years. Moreover, unsustainable fishing practices pose a huge threat to the haor ecosystem and biodiversity as a whole (BirdLife International 2004, Muzaffar 2004). Time-activity budgets have been used extensively elsewhere to provide valuable information on duck habitat use and wintering strategies. However, knowledge of the behavioural ecology of diving ducks (Aythyini) is lagging far behind that of dabbling ducks (Anatini) (Houhamdi & Samraou 2008). Our observations suggest that Baer’s Pochards actively forage between 08h00 and lOhOO at 1.26-1.88 m water depth; recently arrived individuals on the breeding grounds were reported as foraging actively between 08h00-09h00 and diving to depths of 0. 5-2.0 m for c.40 seconds at Xianghai Nature Reserve in China (Gao Jihong et al. 1992b). Further studies on the behavioural ecology of wintering diving ducks including the Baer’s Pochard are important for sound evidence-based conservation planning decisions regarding sustainable wetland management practices in Bangladesh. CONCLUSION This consolidation of both past and recent surveys confirms a dramatic reduction in the wintering population of Baer’s Pochards in Bangladesh, and indicates that the global population may be substantially lower than previously thought. A thorough remote¬ sensing analysis is required to identify potential suitable wetland habitats to be targeted by future surveys for Baer’s Pochard in north¬ east India and Bangladesh. Further studies are needed to determine the potential negative effects on wintering waterfowl of current fishing practices, synthetic agrochemical run-off, and disturbance caused by intensive human activities. Urgent measures must be taken to reduce the pressure of hunting, especially at key wintering sites. Both Hakaluki and Tanguar Haors are recognised as Important Bird Areas and are both considered nationally Ecologically Critical Areas, and the latter also a Ramsar site (BirdLife International 2004). Although both haors have been the subject of recent projects aimed at establishing conservation and wise use through co-management involving local communities, this has not been reflected in recovery of wintering waterfowl populations. Additionally, it is reported that wetland management plans have not been properly implemented in either of these internationally important sites (Muzaffar 2004). In order to maintain a healthy wetland ecosystem to support both human use and biodiversity, appropriate conservation-based wetland management plans and better governance are required. ACKNOWLEDGEMENTS SUC is grateful to the British Ecological Society for funding this survey (Application number: 2776 / 3471). ACL would like to thank CAPES - Coordena^ao de Aperfeiqoamento de Pessoal de Nivel Superior for funding support. SUC is thanklul to Ronald Haider and Farhad A. Pavel for their support in the field. We thank Enam U1 Haque for information on past waterbird counts under the AWC. Forktail 28 (2012) Status and distribution of the endangered Baer's Pochard Aythya baeri in Bangladesh 61 REFERENCES BirdLife International (2001) Threatened birds of Asia: the BirdLife International Red Data Book. Cambridge, UK: BirdLife International. BirdLife International (2004) Important Birds Areas in Asia: key sites for conservation. Cambridge, UK: BirdLife International (BirdLife Conservation Series No. 13). BirdLife International (201 1) IUCN Red List for birds. Downloaded from http:/ /www.birdlife.org on 01/05/201 1 . Gao, J. H„ Ma, J. Z. & Tao, Y. (1992) The behavior of incomplete brood parasitism of two pochard species. Zoological Research 1 3: 327-328 Houhamdi, M. & Samraoui, B. (2008) Diurnal and nocturnal behaviour of Ferruginous Duck Aythya nyroca at Lac des Oiseaux, northeast Algeria. Ardeola 55: 59-69 Li, Z. W. D., Bloem, A., Delany, S., Martakis, G. & Quintero, J. O. ( 2009) Status of waterbirds in Asia - results of the Asian Waterbird Census: 1 987-2007. Kuala Lumpur: Wetlands International. Muzaffar, S. B. (2003) Diurnal time-activity budgets in wintering Ferruginous Pochard Aythya nyroca in Tanguar Flaor, Bangladesh. Forktail 20: 25- 27. Siddiqui, K. U., Islam, M. A., Kabir, S. M. FI., Ahmed, A.T. A., Rahman, A. K. A., Plaque, E. U., Ahmed, Z. U., Begum, Z. N.T., Hassan, M. A., Khondker, M. & Rahman, M. M., eds. (2008) Encyclopedia of flora and fauna of Bangladesh, 26. Birds. Dhaka: Asiatic Society of Bangladesh. The Daily Star (2009) Guest birds in danger: poachers active in Plakaluki Haor. Downloaded from http://www.thedailystar.net/newDesign/ news-details.php?nid=75525 on 28.04.201 1. The Daily Star (2010) Poachers poison to death hundreds of Plakaluki birds. Downloaded from http://www.thedailystar.net/newDesign/news- details.php?nid= 165072 on 28.04.201 1. Thompson, P. M. & Johnson, D. L. (2003) Further notable bird records from Bangladesh. Forktail 1 9: 85-1 02. Thompson, P. M„ Plarvey, W. G„ Johnson, D. L., Millin, D. J., Rashid, S. M. A., Scott, D. A., Stanford, C. & Woolner, J. D. (1993) Recent notable bird records from Bangladesh. Forktail 9: 1 2-44. Sayam U. CHOWDHURY, Department of Environmental Science and Management, North South University, Plot 15, Block-B, Bashundhara, Dhaka 1229, Bangladesh. Email: sayam_uc@yahoo.com Alexander C. LEES, Dept, of Zoology, Museu Paraense Emilio Coeldi, Caixa Postal 399, CEP 66040-170, Belem - Para, Brazil. Email: alexanderlees@btopenworld.com Paul M. THOMPSON, House 32, Road 10, Banani, Dhaka, Bangladesh. Email: paul@agni.com FORKTAIL 28 (20 12): 62-66 Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area ZHIXIN ZHOU, YUE SUN, LU DONG, CANWEI XIA, HUW LLOYD & YANYUN ZHANG We present data on the breeding biology of the largest known colony of Asian House Martin Delichon dasypus, located in the Jiangxi Wuyishan Nature Reserve at 2,1 58 m in the Huanggang Mountains, China. Nest surveys conducted in abandoned buildings in a subalpine meadow during March-August 2007 and 2008 yielded 163 and 132 clutches, from 84 and 82 nests, respectively. Breeding pairs also laid multiple broods and replacement clutches. Average clutch size was 3.0 and 2.6 eggs for first and second broods respectively. Synchronous hatching was detected in 79% of clutches. The proportion of eggs hatching was 0.7 and 0.6 for first and second broods respectively, and the proportion fledging was 0.5 and 0.4 respectively. Nests situated inside buildings were more successful than those situated outside owing to greater protection from severe weather, which was the major cause of breeding failure. Nest losses caused by severe weather were more pronounced later in the breeding season. INTRODUCTION Many bird species raise only one brood per year because of a narrow period of suitable environmental conditions which prohibits multiple breeding attempts (Evans-Ogden & Stutchbury 1996). Others raise multiple broods per breeding season (Verhulst et al. 1997) and the total seasonal fecundity for these species depends on the number of breeding attempts (Geupel & DeSante 1990). One of the principal determinants of multiple broods is the duration of the breeding season, which is often directly related to the abundance and availability of food (Holmes et al. 1992, Morrison 1998). At high elevations, climatic conditions are more severe and unpredictable, with greater daily variation (Lu 2005). Such variable weather may significantly increase the risk of nest failure and may influence parental investment in each clutch and offspring (Bulmer 1984). Consequently, bird species that occupy montane habitats may adjust their reproductive strategy by breeding over shorter periods and producing fewer broods or smaller-sized clutches than species at lower elevations (Sandercock et al. 2005, Bears et al. 2008). We examined the reproductive ecology of the Asian House Martin Delichon dasypus in the high-elevation montane environment of the Huanggang Mountains in south-eastern China. This widespread migrant species is known to occupy a wide elevational gradient spanning both lowland and montane habitats from 800 to 3,000 m (Turner & Rose 1989). In the Huanggang Mountains, birds visit during March-August each year to breed at elevations above 2,000 m, where weather conditions are unpredictable, with severe rainstorms and typhoons. Our aim was to evaluate the reproductive biology and breeding success of this population under such environmental conditions. METHODS Study site Fieldwork was conducted in the Jiangxi Wuyishan Nature Reserve, in the Huanggang Mountains (27°51,38"N 1 17°47/0l"E), Jiangxi province, south-eastern China. Nest surveys were conducted from March to August in 2007 and 2008. The study site represents the highest peak of Wuyishan Cordillera, at an elevation of 2,158 m, with a mean daily temperature of 14.3°C during May and August, and mean annual precipitation of 1,500 mm. From June to August, rainstorms become more severe and weather conditions are extremely poor when typhoons occur. Nest surveys The 3-ha study area is predominately subalpine meadow habitat in which are situated more than 30 abandoned buildings and garages that provide suitable nesting substrate for the breeding martins. The nest of Asian House Martin is a closed cup typical of hirundines, built with tiny mud pellets and then lined with dried moss, sear’ grasses and feathers, leaving only a small entrance hole. We classified two different nest locations: (a) under the eaves outside the buildings, or (b) inside the buildings. Where possible for each nest we recorded the overall size (width x height x depth) and the dimensions of the nest entrance (widest x smallest diameter). Systematic surveys of nests and their contents began on 1 March in each year. Weekly visits were made to determine the arrival dates and record pre-breeding behaviour. Mist-nets (6-8 m long, with 1 x 1 cm mesh) were set 1.8 m from the ground to capture adult birds. For each individual caught we measured mass (g), total body length, wing length, tail length, bill length and tarsus length (all in mm). We banded the birds with coloured plastic rings and also marked each bird (using indelible markers) with a unique pattern on the throat and undertail-coverts to enable individual identification from a distance using binoculars following Massoni etal. (2007). Once nest-building commenced, we recorded details daily. Four observers visited each nest every day to determine the exact timing of egg-laying. All eggs were numbered (in laying sequence) at the pointed end using a waterproof ink pen (Gosler et al. 2005). Egg length and width were measured to the nearest 0.01 mm using plastic vernier calipers, and egg mass was recorded to the nearest 0.1 g using an electronic balance. Egg volume was calculated as length x width2 x 0.000507, following Manning (1979). Clutch volume was calculated as the sum of the volume of each egg laid in a clutch. Clutch size was defined as the number of eggs in the clutch at the onset of incubation. Hatching date was defined as the date on which the majority of eggs in each clutch hatched. Synchronous hatching was defined as occurring when all eggs in the clutch hatched on the same day. Incubation duration was calculated as the time elapsed from the beginning of incubation to the hatching date. During the hatching period, nests were checked every day to monitor the incubation progress and obtain hatching dates. After hatching, we checked nests carefully every day to monitor the growth of nestlings and to determine the exact date they fledged. On each day, nestlings were marked with indelible markers on their legs, and. weighed to the nearest 0.1 g with an electronic balance. The nestling daily mass gain and growth rate were calculated following Ricklefs (1968). Nestlings were not weighed after 18 days to reduce the possibility of premature fledging. Fledging date was defined as the date when Forktail 28 (2012) Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area 63 all nestlings left the nest, and the nestling period was calculated as the duration between the hatching and fledging dates. We determined the length of the breeding season for each year by calculating the number of days between the date when nest-building for the first breeding attempt commenced and the fledging date for the last clutch of the year. Statistical analysis Data were examined for normality using Kolmogorov-Smirnov tests, and we pooled the data of two years for subsequent statistical analysis. Comparison of breeding parameters across successive broods (clutch size, fledging morphology, number of nestlings, etc.) was conducted using non-parametric Mann-Whitney 17-tests. Variation in the duration of incubation and other periods of the reproductive cycle across successive broods was examined using non-parametric Kruskal-Wallis tests. Spearman’s rank correlation was used to detect the relationship between clutch size and the proportion of eggs hatching. Logistic regression was used to model nestling growth. Values are reported as means ± SE with sample size («) and range. Differences were considered significant at P < 0.05. Data were analysed using SPSS 16.0 (SPSS Inc. 2008). RESULTS Nest construction and breeding Asian House Martins were first observed in the study area on 13 March in 2007 and on 10 March in 2008, with nest-site selection and territorial defence by paired birds first observed in early April each year. Asian House Martins apparently preferred re-using existing nest sites: in 2008, all 195 nests remaining from the previous breeding season were re-occupied by the earliest-arriving individuals. Individuals that arrived later constructed new nests. The average external dimensions of nests (»= 37) were 121.9 ± 4.5 mm (70-181.1 mm) wide, 109.3 ± 2.8 mm (73.9-130.9 mm) high, and 101.9 ± 2.6 mm (80-171 mm) deep. The average dimensions of entrance holes (#=3 6) was 37.3 ±1.4 mm (18.3-60 mm) by 22.9 ± 0.6 mm (13.9-30.4 mm). The maximum number of nests on any one day during the season was 628 in 2007 and 675 in 2008. The number of nests increased rapidly during April and May and peaked during June (Figure 1; results for 2007 were similar). The number situated outside buildings subsequently fluctuated and then declined after the end of August, whereas the number of nests situated inside buildings remained relatively stable (Figure 1). Figure 1. Number of Asian House Martin nests located inside and outside buildings each month in 2008. Shaded blocks indicate the timing of severe weather events. 400 The first eggwas laid on 20 April in 2007 and 27 April in 2008, when the majority of pairs had only just begun to construct nests. The last breeding attempt was recorded on 1 August in 2007 and 3 August in 2008. In both seasons, the Asian House Martin population had completely left the area by late August. Clutch size, incubation, fledging and nesting success Pooling both breeding seasons, we monitored 295 clutches totalling 834 eggs from 166 nests. Of these, it was possible to measure and collect data from 571 eggs. Clutch size ranged from 1 to 4 eggs (25, 52, 167 and 51 clutches contained 1, 2, 3 and 4 eggs, respectively), averaging 2.8 ±0.1 eggs. No replacement eggs were laid if any eggs were lost. Eggs were coloured pinkish-white with blood-red spots just after being laid, but gradually faded to a pale white colour during incubation. On average, eggs were 17.9 ±0.1 mm (15.5-21.3 mm, zz = 5 5 8 ) long, 12.9 ± 0.2 mm (1 1 .6- 14.6 mm, 72=558) wide, and 1.5 ± 0.1 g (0.4-2. 2 g, n= 571) at laying. Mean egg volume was 1.5 ± 0.1 cm3 (1.2-2. 2 cm3, 72=558), and clutch volume averaged 4.2 ± 0.1 cm3 (1.3-6. 8 cm3, tz=195 clutches). Incubation by both parents (distinguished by colour rings) lasted 15.3 ± 0.2 days (7-25 days, 72=195 clutches, excluding data from four nests with only one egg since we were unable to determine the exact date of incubation). There was no significant difference in the duration of incubation between clutches of different size (Kruskal-Wallis test, 72=191, /22=5.46, P= 0.07). The hatching proportion among all eggs in both two years was 0.66 (72=834 eggs). At least one nestling hatched in 75% (222) of all clutches recorded, and the mean proportion of eggs hatched in each clutch was 0.62 ± 0.02 (72=295 clutches); this was positively correlated with clutch size (Spearman r= 0.185, P=0.001). Synchronous hatching was detected in 175 clutches (79%), asynchronous hatching in 47 clutches, with the majority of these (87%, 72 = 41) hatching within 48 hours (1-5 days). The number of nestlings in a brood averaged 2.5 ± 0.1 (1-4, 72=222 clutches). Among clutches which led to the fledging of at least one chick (72=157), the nestling period lasted on average 28.2 ± 0.2 days (22- 35 days) and increased with the number of nestlings in a brood (Spearman r=0.213, 72=153, 73=0.008). For these 157 successful clutches, the mean number of fledglings was 2.5 ±0.1 (1-4). The mean proportion of fledged nestlings in the clutches which had at least one nestling averaged 0.69 ± 0.03. Of the 834 monitored eggs, 47% fledged. The mean proportion of fledglings per egg in a clutch was 0.44 ± 0.03 (0-1, 72= 295). The survival rate (to fledging) of nestlings inside buildings was 0.84 ± 0.04 (n=73), which was significantly higher (Mann- Whitney U test, 72=166, P=0.006) than the survival rate of nestlings outside (0.65 ± 0.04, 72=93). Severe weather conditions (rain and wind) caused by a typhoon in June 2008 resulted in the loss of 50 nests inside buildings and 112 nests outside. The total number of broods lost during two breeding seasons was 141 (48%, 72=295). Of these, 72 were lost during incubation, comprising 46 (64%) lost owing to nest damage caused by severe weather conditions, and 26 (36%) owing to egg ejection by parental or other adult individuals for unknown reasons. During the nestling stage, 69 broods failed owing to severe weather (49 clutches, 71%), nestling starvation (10, 14%), eggs falling accidently from the nest (9, 1 3%), and predation (presumed by a small rodent based on bite marks; 1 clutch, 1%). Nestling growth We measured 381 individuals in 2007 and 2008 (199 adults and 1 82 fledglings; Table 1 ). There were significant differences between adults and fledglings in mass (Mann-Whitney U test, 72=333, P<0.0001) and tarsus length (Mann-Whitney U test, 72 = 353, P=0.000l) (Table 1). Mean mass of nestlings at hatching was 1.4 64 ZHIXIN ZHOU etal. Forktail 28 (2012) Table 1. Morphological characteristics of Asian House Martin adults and fledglings in study area (means ± SE, with range in parentheses). Adult Fledgling Mass (g)~ 13.5 ±0.1 (9.3-19.3) o=196 12.7 ±0.1 (11.1-15)0=137 Beak length (mm) 5.4 ±0.0 (3.9-77) n=199 5.5 ±0.0 (4.5-6.8)0=154 Tarsus length (mm)" 12.2 ±0.1 (9.3-15.9)n=199 10.9 ±0.1 (8.7-13.5)0=154 Wing length (mm) 93.4 ±0.2 (84.4-102)0=199 95.8 ±0.3 (57-1 11. 5) 0=172 Body length (mm) 1 11 .6 ± 0.4 (95.4-123.7) 0=199 11 1.0 ±0.5 (100-128) o=133 Tail length (mm) 37.0 ±0.3 (26.4-47.2)0=199 37.4 ±0.3 (27.5-49)0=128 " Significant difference between adults and fledglings, Mann-Whitney U test. clutch (Table 2), except for the duration of both the incubation (Mann-Whitney U test, «=169, P=0.054) and nestling period (Mann-Whitney U test, « = 136, P= 0.93). Of the pairs that failed to rear their first clutch successfully, 39 also laid a replacement clutch. Of the 79 nests which had successfully fledged nestlings 44 clutches failed, for which 1 1 pairs laid further replacement clutches, but only three of these hatched nestlings and none fledged successfully. Replacement clutches for first broods also had significantly lower breeding parameters than the first breeding attempt (Table 2). DISCUSSION Figure 2. Nestling mass (means ± SE) as a function of age. Day 0 corresponds to the hatching day. The horizontal line indicates the mean adult mass. 21 r 18 15 - 12 - 6 L 3 - 1 1 1 0 3 6 9 12 15 18 21 Age (d) ± 0.1 g (1.2-1. 5 g, n = 5). Nestlings reached a peak mass of approximately 18.6 ± 0.8 g at 14 days of age, but then showed a marked decrease to 16.7 ± 0.7 g at 18 days (Figure 2). Fitting the mass as a function of nestling age to a logistic growth curve gave a growth rate constant (if) of 0.210. Multiple clutches After successful fledging of nestlings, a second breeding attempt was recorded for 79 nests (48%, 72 = 1 66). The majority of breeding parameters from the second brood, in particular clutch size, hatching proportion, fledging success and nesting success, were significantly reduced when compared to the first successful Our detailed description of the breeding parameters for Asian House Martins nesting at high elevation (2,158 m) reveals that they lay apparently smaller clutches (2.8 ± 0. 1 vs 3.4 eggs) but with larger eggs (1.5 ± 0.1 vs 1.1 g) compared to colonies at lower elevations (data from Yu etal. 1998 and Mi etal. 2007, at 500 m and 1,290 m, respectively), presumably as a local adaptation to boost reproductive success by investing more in fewer offspring (Lu 2005), although we were unable to test this. The Jiangxi Wuyishan Nature Reserve breeding colony, with over 600 pairs, is the largest described to date. Typically, hirundines breed in smaller colonies (Snapp 1976), and smaller colonies of Asian House Martins are known at a number of lower-elevation sites (<1,800 m) in the Wuyishan mountains. The large colony size may be a consequence of a local shortage of nest sites (Snapp 1976, Shields & Crook 1987), but we were unable to test this. We found that Asian House Martins at the Jiangxi Wuyishan Nature Reserve breed over a five-month period, from April until August, the beginning of which is apparently at least 30 days earlier than the equivalent date for a population at l,290m(Mi etal. 2007) and compared with Northern House Martins D. urbicum breeding at a latitude of 39°N (Saygili & Yigit 2007). We speculate that this earlier initiation of breeding may be an adaptation to the increased risk of nest failure later in the season when extreme weather conditions are more likely. Selection of nest sites offering greater protection from severe weather conditions was of critical importance to the breeding success in this population. We found that fewer pairs used more exposed nest sites outside buildings, and that such nests were subject to more pronounced fluctuations in nesting success than those located inside, owing to nest damage from heavy rain and strong winds. Such differences have not previously been recorded in hirundines, to our knowledge. Asian House Martins produced multiple broods and replacement clutches in an attempt to maximise the number of Table 2. Breeding parameters for successive clutches. Breeding parameters 1st dutch 1st replacement clutch 2nd clutch 2nd replacement clutch Clutch size3 b 3 ±0.1, 1-4,0=166 2.6 ± 0.1, 1-4,0=39 2.6 ±0.1, 1-4, o=79 2.4±0.2, 1-3,0=11 Nestlings3-1" 2.7 ±0.1, 1-4, o=136 2.2 ±0.2, 1-3,0=25 2.2 ±0.1, 1-4,o=58 1.7 ±0.3, 1-2, o=3 Fledglings3 b 2.6 ±0.01, 1-4,0=104 2.1 ±0.2,1-3,0=18 2.3±0.1, 1-4,o=35 0,o=3 Hatching proportion111 0.7 ±0.0, 0-1,0=166 0.5 ±0.1, 0-1, o=39 0.6±0.1, 0-1, o=79 0.2±0.1, 0-1, 0=11 Fledging success3 0.7 ±0.0, 0-1, 0=136 0.7±0.1, 0-1, o=25 0.6 ±0.1, 0-1,o=58 0,o=3 Breeding success16 0.5 ±0.0, 0-1,0=166 0.4±0.1, 0-1, o=39 0.4±0.1, 0-1,0=79 0,0=11 Incubation period6 15.5±0.2, 7-21, o=111 14.7 ±0.4, 9-18, 0=24 15.2 ±0.4, 10-25, 0=58 15 ±0.6, 14-16,0=3 Nestling period6 28.4 ±0.3, 22-34,0=103 26.3±0.6, 22-30, 0=17 28.7 ±0.4, 25-35, o=33 4 = significant difference between first and second clutches (Mann-Whitney U test, P<0.05) b = significant difference between first and first replacement clutches (Mann-Whitney U test, P< 0.05) Forktail 28 (2012) Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area 65 offspring they produce per season. However, mean clutch size, hatching proportion, fledgling success and nesting success declined significantly across successive broods. Declines in mean clutch size and hatching success through the breeding season have also been reported in other hirundine species (Zhao 1981, Sakraoui et al. 2005) and other migrant species (Dhondt et al. 2002). Multiple broods and replacement clutches have been reported for several hirundine species including Northern House Martin (Bryant 1979, Lahlah et al. 2006), Barn Swallow Hirundo rustica (Jones 1987, Sakraoui etal. 2005), Tree Swallow Tachycineta bicolor (Clapp 1 997, Monroe et al. 2008), White-rumped Swallow T. leucorrhoa (Massoni etal. 2007) and Purple Martin Progne subis (Allen & Nice 1952). In one experimental egg-removal study, 41% of female Tree Swallows laid replacement clutches (Rooneem & Robertson 1 997), which is lower than the 63% of female Asian House Martins at the Jiangxi Wuyishan Nature Reserve. The capacity for laying multiple broods and replacement clutches has been shown in other species to be dependent on a number of factors including female age and quality, timing of first clutch, size of first clutch, and local resource abundance (De Laet & Dhondt 1989, Smith & Marquiss 1995, Tinbergen & Verhulst 2000). Significant declines in breeding success across successive broods are often related to parental energy budgets and how many young both parents are able to feed (Lack 1968, Bryant & Westerterp 1980). Whether or not the declines observed for Asian House Martins are related to seasonal variation in prey abundance, or other environmental constraints imposed on the parents, remains to be seen. 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(1989) A handbook of the swallows and martins of the world. London: Christopher Helm. Verhulst, S., Tinbergen, J. M.& Daan, S. (1 997) Multiple breeding in the Great Tit. A trade-off between successive reproductive attempts? Fund. Ecol. 11:714-722. Yu W. D.,Tang S. H„ Chen L., Yu K., Zhou Z. H. & Zhao M. S. (1 998) The ecology of Asian House Martin ( Delichon dasypus ) in WestTianmu Mountain Nature Reserve (II). J. East China Norm. Univ. Supp. Zool: 140-143. (In Chinese.) Zhao Z. J. ( 1 98 1 ) The breeding ecology of the Golden-rumped Swallow in Changbai Shan area, Jilin province. Acta Ecol. Sinica 1: 369-374. (In Chinese.) Zhixin ZHOU, Ministry of Education Key Laboratory for Biodiversity and Ecological Engineering, College of Life Sciences, Beijing Normal University, Beijing, 100875, China, and Key Lab of Animal Ecology and Conservation Biology, Institute of Zoology, Chinese Academy of Sciences, 1 00 101, China. Email: zhixinzhou@gmail.com Yue SUN, Ministry of Education Key Laboratory for Biodiversity and Ecological Engineering, College of Life Sciences, Beijing Normal University, Beijing, 100875, China, and State Oceanic Administration Beijing, 100860, China. Email: sunyuebird@ 1 63.com Lu DONG & Canwei XI A, Ministry of Education Key Laboratory for Biodiversity and Ecological Engineering, College of Life Sciences, Beijing Normal University, Beijing, 100875, China. Emails: tiantai_bird@yahoo.cn & xiacanwei@126.com Hu w LLOYD, School of Science and the Environment, Manchester Metropolitan University, Chester Street, Manchester, UK, Ml 5GD. Email: H.Lloyd@mmu.ac.uk Yanyun ZHANG, Ministry of Education Key Laboratory for Biodiversity and Ecological Engineering, College of Life Sciences, Beijing Normal University, Beijing, 100875, China. Email: zhangyy@bnu.edu.cn (author for correspondence) FORKTAIL 28 (201 2): 67-70 Sex bias in a wintering population of Dunlin Calidris alpina in central Taiwan DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU The sexes of many migratory shorebird species differ in timing and distance of their migrations, but this phenomenon has been little studied for birds in the East Asian-Australasian Fly way (EAAF). Using molecular sexing techniques (but also taking morphometric data), we compared the sex ratio of Dunlin Calidris alpina sampled on the west coast of Taiwan between September 2007 and May 2008. We found an overall sex bias (59.2%) toward females, including juveniles. A discriminant function deduced from three morphological measurements correctly identified 89.0% of Dunlin, but performed better in identifying males than females. Since Taiwan is at the southern end of the Dunlin's East Asian winter range, our results support the 'arrival-time' hypothesis whereby the sex that arrives earlier on the breeding grounds winters closer to them. To maintain stable populations, conservation must take account of this differential use of wintering grounds by the two sexes. INTRODUCTION In many migratory bird species, individuals of different sex or age differ in their routes and timing of annual migration (Cristol et al. 1999). Several hypotheses have been proposed to explain these differential migrations, focusing on differences in territorial behaviour, body size and behavioural dominance. The arrival time hypothesis proposes that certain individuals winter closer to the breeding grounds because competition for mates or territory favours early arrival to the breeding grounds (see Morbey & Ydenberg 2001). The body size hypothesis proposes that the larger-bodied sex can winter at higher latitudes due to larger fuel reserves and better thermal efficiency (see Catry et al. 2005). The dominance hypothesis suggests that the dominant sex will out- compete the less dominant one for resources, forcing the less dominant sex to migrate longer distances or winter in inferior habitats (see Marra 2000). Testing these hypotheses is difficult because they often are not mutually exclusive. An early test of these hypotheses by Myers (1981), using data from several shorebird species, found that the sex that arrived earlier at the breeding grounds was the sex that wintered at higher latitudes. He proposed that, regardless of size and behavioural dominance, difference in arrival time alone is sufficient to predict a latitudinal cline in wintering shorebirds. Dunlin Calidris alpina is a long-distance migratory shorebird, breeding along the Arctic rim in Europe, Iceland, Scandinavia, Siberia, North America and Greenland (Piersma et al. 1996) and wintering in temperate and tropical regions throughout the Northern Hemisphere (Wenink & Baker 1996). Studies on wintering Dunlin in Hong Kong have found two distinct mtDNA haplotypes, Siberian and Alaskan (Wenink & Baker 1996). Significant differences exist in culmen length and mtDNA haplotypes among Dunlin of different breeding populations (Wenink & Baker 1996, Wenink et al. 1996). Dunlin ringed in Taiwan have been recaptured on breeding grounds in both Alaska and Siberia, and in Taiwan we have re-sighted Dunlin which were ringed in Alaska and Sakhalin Island (Lanctot et al. 2009, Fernandez et al. 2010, CC pers. obs.). During the non¬ breeding season, significant numbers migrate from breeding grounds to winter in eastern China, Japan, Korea, Hong Kong and Taiwan (Bamford et al. 2008). Around 950,000 Dunlin use this flyway every year, making the East Asian-Australasian Flyway (EAAF) one of the five major flyways for this species (Bamford etal. 2008). Female Dunlin are larger than males, but it is the males that arrive earlier at their breeding grounds and defend territories each spring (Holmes 1966). According to the arrival time hypothesis, males should winter nearer the breeding grounds than females, whereas the dominance and body size hypotheses predict the opposite. The sex of wintering Dunlin is difficult to identify in the field, even in the hand (Prater etal. 1977). In many studies, sex is assigned by difference in bill length (e.g. Chao et al. 1991, Warnock et al. 1997, Shepherd et al. 2001, Shepherd 8c Fank 2004). However, bill length overlaps to some extent among the sexes (Brennan etal. 1984, Shepherd et al. 2001, Meissner 2005) and geographically distinct groups of Dunlin also vary in bill length (Wenink 8c Baker 1996). Instead, by amplifying fragments of sex-specific DNA sequences, the sex of all individuals of monomorphic bird species like Dunlin can be accurately determined (e.g. Baker 8c Piersma 1999, Fridolfsson 8c Ellegren 1999). Differential migration of Dunlin has been reported in North America (Shepherd et al. 2001), but their migration patterns have not been explored in the EAAF. In this study, we present data on the sex ratio of Dunlin wintering in central Taiwan, one of their southernmost wintering areas (Wei et al. 2009). In addition, using the results of molecular sexing, we derive a discriminant function and examine its accuracy in distinguishing the sex of Dunlin in Taiwan. METHODS Field methods Our study area is Han Pao wetland, Changhua County, Taiwan (23-95°N 1 20. 36°E). Each year, more than 10,000 Dunlin migrate through or winter here (Hu 2005). This area is one of the largest intertidal mudflats in Taiwan (c.38 km2). It has a long history of shellfish and oyster farming, and shorebirds often rest and feed in these habitats (Hu 2005). We mist-netted Dunlin from September 2007 to May 2008, three to four times every month, at night from evening to high tide. Most ringing sites were on agricultural lands with some shallow water. We defined birds netted from September to November 2007 as the autumn population, December 2007 to February 2008 as the winter population, and March to May 2008 as the spring population. We ringed and weighed each Dunlin captured and, for most, measured (to the nearest mm): maximum wing length, tarsus length, total head length and bill length. We then extracted 20- 40 yl of blood by venipuncture from the brachial vein and placed the blood in SET buffer (0.15 M NaCl, 0.05 M Tris, 0.001 M EDTA, pH = 8.0). We aged the Dunlin based on the presence of either buffy-edged inner median coverts (first-years) or white-edged inner median coverts (after first year) (Choi et al. 2010). We could 68 DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU Forktail 28 (2012) identify birds as first-years before February, but coverts became worn and buffy fringes disappeared from March. Molecular sexing We extracted DNA from blood using standard methods (Gemmell & Akiyama 1996). Fragments of the chromo-helicase-DNA binding protein (CFID) gene from the sex chromosomes were amplified by polymerase chain reaction (PCR), using the primers 2550F and 2718R (Fridolfsson & Ellegren 1999). The reaction volume was 20 pi and contained c.10 ng DNA, 0.8 pi dNTP (2.5 mM of each nucleotide), 0.4 pi 2550F primer (10 mM), 0.4 pi 2718R primer (10 mM), 2.0 pi 10 x PCR buffer, 15.92 pi dH,0 and 0.4 U pi Taq DNA polymerase (Protech). The PCR included 3 minutes at 95°C, followed by 35 cycles of 30 seconds at 95°C, 40 seconds at 46°C and 50 seconds at 72°C, and a final extension reaction at 72°C for 5 minutes. The PCR products were electrophoresised in 1% agarose gel, using a Bio- 100™ DNA Ladder (Protech) as size reference. We then dyed the gels in ethidium bromide and examined them under ultra-violet light. We identified the sex of birds by the presence or absence of the female- specific DNA fragment of the CHD-W gene. Statistical analysis We used a chi-square goodness-of-fit test to determine if the sex ratio significantly deviated from 1:1 for both adult and juvenile Dunlin. We also compared the difference of sex ratios within each season with the same test. Mean bill length, total head length, maximum wing length and tarsus length were compared between sexes using the Student’s /-test. We used three measurements (maximum wing length, tarsus length and bill length) to derive a discriminant function using SPSS 14.0 to assign sex. RESULTS From September 2007 to May 2008, we caught and sexed 748 Dunlin in the Han Pao wetland (Figure 1). Excluding recaptured individuals, 693 birds were ringed (Table 1). The sex ratio was significantly different from 1:1 (xr= 23.27, P < 0.001): 410 individuals were female (59.2%) and 283 individuals were male (40.8%). We identified 138 Dunlin as juvenile (<20%), but failed to age some individuals because the buffy-edged inner median Table 1 . Number of Dunlin ringed from September 2007 to May 2008 at Han Pao Wetland, Taiwan. Females Males Total Number of birds ringed 410 283 693 Adult 316 239 555 First-year young 94 44 138 Table 2. Measurements (mean ± SE, with ranges below in parentheses) of Dunlins captured from September 2007 to May 2008 at Han Pao Wetland, Taiwan. Measurement Females /V=325 Males A/=240 f-test (two-tailed) Weight (g) 57.92 ±5.79(41.70-79.90) 53.36 ±5.91 (33.6—75.30) rs6=9.15,p<0.001 Bill (mm) 37.61 ±1.81 33.69 ±1.58 r56 = 26.83, p< 0.001 (31.81-37.61) (28.89-38.26) Head (mm) 62.33 ±1.93 58.09 ±1.81 r56 = 26.50, p< 0.001 (55.97-68.38) (50.57-63.45) Tarsus (mm) 26.95 ±1.06 25.87 ±1.09 fS63= 11.75, p<0.001 (23.80-31.33) (21.14-29.00) Wing (mm) 127.09 ±2.77 123.89 ± 3.14 CS6 = 12.81,p< 0.001 (119.00-134.00) (114.00-134.00) coverts were unclear after midwinter. Among juveniles, 94 were female (68.1%) and 44 were male (31.9%), significantly female- biased (-),12= 18.1, P < 0.001). Combining data from all captures, the sex ratio did not significantly deviate from 1:1 in autumn (73 females, 61 males, zl2 = 1.07, P = 0.3). However, in winter and spring, the sex ratio was significantly female-biased (winter: 238 females, 163 males, %i2 = 14.03, P < 0.001, spring: 137 females, 76 males, xl2 = 17.47, P < 0.001 respectively). Of Dunlin that were both sexed molecularly and measured (n = 565 individuals), females were significantly larger than males. However, these measurements showed overlap between the sexes (Table 2). For discriminant analysis, we excluded total head length in the analysis because it was not independent from bill length and both were highly correlated (Pearson’s correlation coefficient r =0.92, P< 0.001). The best discriminant function included bill length (BL), tarsus length (TL) and wing length (WL). The resulting function was: 140 Figure 1. Number of female and male Dunlin captured from September 2007 to May 2008 at Han Pao Wetland, Taiwan. Month Forktail 28 (2012) Sex bias in a wintering population of Dunlin Catidris alpina in central Taiwan 69 Figure 2. Distribution ofthe D values of the discriminant function in male and female Dunlin captured from September 2007 to May 2008 at Han Pao Wetland, Taiwan. D = 0.515 x BL + 0.05 xTL + 0 .105 x WL - 33.018 (Wilks’ lambda = 0.413). The value of D < 0 was identified as male and D > 0 was identified as female. This function correctly identified the sex of 89.03% of Dunlin, with 92.9% of males and 86.2% of females respectively (Figure 2). This function performed better in identifying males than in females (%2 = 6.46, P = 0.01 1). DISCUSSION The wintering sites of Dunlin in East Asia range from the eastern coast of Liaoning province, China (45°N), Japan, South Korea to Hong Kong (22°N). Few Dunlin are found wintering farther south than Hong Kong (Bamford et al. 2008). As our study site (Han Pao wetland) is located at 23.95°N, we assume it is one of the southernmost wintering sites. Numbers of Dunlin reached their peak from December to February, indicating that most Dunlin used the Han Pao wetland as a wintering ground, instead of as a stopover site. Within the E AAF, at least three species of shorebirds show sex- biased migration: Curlew Sandpiper Calidris ferruginea males winter farther south; and Eastern Curlew Numenius madagascariensis and Bar-tailed Godwit Limosa lapponica males winter farther north (Nebel 2007). We found that the overall sex ratio of Dunlin in central Taiwan is significantly female-biased. Both the body size hypothesis and the dominance hypothesis predict that females, which are larger and therefore presumably more dominant than males, would winter at northern sites. Our data thus support the arrival time hypothesis, which states that the sex that arrives later at the breeding grounds will winter farther south (Morbey & Ydenberg 2001). The arrival time hypothesis also predicts more male Dunlin in the northern wintering grounds, such as Japan, Korea or northern China. With further studies from these regions, this hypothesis can be further tested. In North America, Shepherd et al. (2001) found male-biased populations of Dunlin at higher latitude wintering areas and suggested that there is a latitudinal cline in the sex ratio ofwintering Dunlin populations. Our data are consistent with this. Another possibility for the female-biased sex ratio in our study site comes from local differences in habitat preference between sexes. Shepherd & Lank (2004) reported that male Dunlin preferred agricultural land and females preferred mudflats on wintering grounds in British Columbia. All Dunlin we sampled were caught on agricultural land, but we found more females, suggesting that the findings of Shepherd & Lank (2004) are not generally applicable. Our sampling, however, was conducted during high tides when all Dunlin came into agricultural lands, thus minimising such habitat- based sex bias. We used the results of molecular sexing to develop a discriminant function to identify Dunlin sex by morphological measurements. For sex-size dimorphic species, discriminant function analysis of morphometric data is a common technique to sex birds that are monomorphic in plumage (Brennan et al. 1984, Iko et al. 2004). Previous discriminant function and maximum likelihood function analyses have correctly identified the sex of over 90% of non-breeding Dunlin (Brennan et al. 1984, Shepherd et al. 2001, Meissner 2005), but were all conducted on a single subspecies. In Taiwan, wintering Dunlin come from several geographically distinct subspecies (Lanctot et al. 2009), which differ in body size (Wenink & Baker 1996, Bamford et al. 2008). The mix of subspecies in our study site may result in the lower discriminant power of the function, or might be a result of differing morphological measurements among ringers in our study (Barrett et al. 1989). CONCLUSION We found a significant female -biased wintering Dunlin population in central Taiwan. This is the first study to reveal a differential sex ratio in this species in the EAAF. East Asia has one of the highest human population densities in the world and the fastest population and economic growth, especially in coastal areas (Liu 2009). Shorebird habitats are therefore being rapidly altered (Ge et al. 2007). The loss of Dunlin habitats used heavily by either sex could result in disproportionate population declines (Steifetten & Dale 2006). Identification of sites that are preferred by members of one sex is crucial for conservation (Nebel 2007). Our data alone are not sufficient to test hypotheses for the differential migration of Dunlin in this region. More long-term data from different latitudes in the EAAF, such as eastern China, Korea, Japan and other parts of Taiwan, will help the understanding of the migration ecology of this species. 70 DZUNG-YUN YANG, CHUNGYU CHIANG & YU-CHENG HSU Forktail 28 (2012) ACKNOWLEDGEMENTS We thank J. Pilgrim, N. Warnock and K. Mathot for their insightful comments on the manuscript, Taiwan Bird Banding Center for providing bird rings and the Changhua County Government for issuing a ringing permit. The ringing work could not have succeeded without the involvement of the wader-catching crew of the Taiwan Wader Study Group and the students of Department of Environmental Science and Engineering, Tunghai University. We also thank many volunteers for their hard work with us at night. REFERENCES Baker, A. J.& Piersma.T. (1999) Molecular vs. phenotypic sexing in Red Knots. Condor 101:887-893. Bamford, M. J., Watkins, D. G., Bancroft, W. & Tischler, G. (2008) Migratory shorebirds of the East Asian-Australasian Flyway: population estimates and important sites. Canberra: Wetlands International-Oceania. Barrett, R. T„ Peterz, M., Furness, R. W. & Durinck, J. (1989) The variability of biometric measurements. Ringing & Migration 1 0:1 3-1 6. Brennan, L. A., Buchanan, J. B., Schick, C. T., Herman, S. G. & Johnson, T. M. (1984) Sex determination of Dunlins in winter plumage. J. Field Orn. 55:343-348. Catry, P., Lecoq, M., Araujo, A., Conway, G., Felgueiras, M„ King, J. M. B., Rumsey, S., Salima, H. & Tenreiro, P. (2005) Differential migration of Chiffchaffs Phylloscopus collybita and P. ibericus in Europe and Africa.! Avian Biol. 36: 184-190. Chao, A., Chang, S. H. & Tsung, Y. H. (1991) ['Sexing Dunlins of Kung-Du area by a statistical method.'] J. Chinese Stat. Assoc. 29: 131-143 (In Chinese.) Choi, C„ Hua, N„ Persson, C., Chiang, C. Y. & Ma. Z. (2010) Age-related plumage differences of Dunlins along the East Asian-Australasian Fly way. J. Field Orn. 81: 99-1 1 1. Cristol, D. C., Baker, M. B. & Carbone, C. (1999) Differential migration revisited: latitudinal segregation by age and sex class. Current Orn. 1 5: 33-87. Fernandez, G., Buchanan, J. B, Gill, Jr. R. E., Lanctot, R. & Warnock, N. (2010) Conservation plan for the Dunlin with breeding populations in North America ( Calidris alpina arcticola, C. a. pacifica, and C. a. hudsonia ), Version 1.1. Manomet Center for Conservation Sciences, Manomet, Massachusetts. Available at http://www.whsrn.org/sites/default/files/ file/WHSRN_Dunlin_Conservation_Plan_10_02-28_v1 .1 .pdf. Fridolfsson, A. K. & Ellegren, H. (1999) A simple and universal method for molecular sexing of non-ratite birds. J. Avian Biol. 30: 1 16-121 . Ge Z.-M., Wang T.-H., Zhou X., Wang K.-Y. & Shi W.-Y. (2007) Changes in the spatial distribution of migratory shorebirds along the Shanghai shoreline, China, between 1 984 and 2004. Emu 1 07: 1 9-27. Gemmell, N. J. & Akiyama, S. (1996) An efficient method for the extraction of DNA from vertebrate tissues. Trend. Genet. 1 2: 338-339. Holmes, R.T. (1966) Breeding ecology and annual cycle adaptations of the Red-backed Sandpiper (Calidris alpina) in Northern Alaska. Condor 68: 3-46. Hu W.-Y. (2005) ['Population estimate of Dunlin (Calidris alpina) at Changhua coastal area.'] Taichung: M.Sc. dissertation, Tunghai University. (In Chinese.) Iko, N. M., Dinsmore, S. J. & Knopf, F. L. (2004) Evaluating the use of morphometric measurements from museum specimens for sex determination in Mountain Plovers (Charadrius montanus). Western N. Amer. Nat. 64: 492-496. Lanctot, R. B„ Barter, M., Chiang, C. Y., Gill, R., Johnson, M„ Haig, S„ Ma, Z. J., Tomkovich, P. & Wunder, M. (2009) Use of band resightings, stable isotopes and genetics to understand the migratory connectivity of dunlin ( Calidris alpina) breeding in the Circumpolar Arctic and wintering in the East Asian-Australasian Flyway. Pp.149-164 in L.-Y. Hsieh & M.-L. Hsueh, eds. Proceedings of the International Symposium on Coastal Wetlands and Water Birds Conservation. Taiwan: Endemic Species Research Institute. Liu, G. Q. (2009) Mainstreaming in the UNDP/GEF/SFA project 'wetland biodiversity conservation and sustainable use in China'. Pp. 89-97 in E. D. Ongley & M. H. Allard, eds. Mainstreaming wetland biodiversity conservation: experience and lessons learned in practical applications of mainstreaming. Beijing: UNDP and the State Forestry Administration of China. Marra, P. P. (2000) The role of behavioral dominance in structuring patterns of habitat occupancy in a migrant bird during the non-breeding season. Behav. Ecol. 1 1 : 299-308. Meissner, W. (2005) Sex determination of juvenile Dunlins migrating through the Polish Baltic region.! Field Orn. 76: 368-372. Morbey, Y. E. &Ydenberg, R. C. (2001 ) Protandrous arrival timing to breeding areas: a review. Ecol. Lett. 4: 663-673. Myers, J. P. (1981) A test of three hypotheses for latitudinal segregation of the sexes in wintering birds. Canad. ! Zool. 59: 1 527-1534. Nebel, S. (2007) Differential migration of shorebirds in the East Asian- Australasian flyway. Emu 107: 14-18. van Gils, J. & Wiersma, P. (1996) Dunlin Calidris alpina entry in species accounts for family Scolopacidae (sandpipers, snipes and phalaropes). P. 526 in J. del Hoyo, A. Elliott & J. Sargatal, eds. Handbook of the birds of the world, 3. Barcelona: Lynx Edicions. Prater, T., Marchant, J. & Vuorinen, J. (1977) Guide to the identification and ageing ofholarctic waders. Tring: British Trust for Ornithology. Shepherd, P. C. F. & Lank, D. B. (2004) Marine and agricultural habitat preferences of Dunlin wintering in British Columbia. ! Wildlife Mgmt. 68:61-73. Shepherd, P. C. F., Lank, D. B., Smith, B. D., Warnock, N., Kaiser, G. W. & Williams, T. D. (2001) Sex ratios of Dunlin wintering at two latitudes on the Pacific coast. Condor 103: 352-360. Steifetten, O. & Dale, S. (2006) Viability of an endangered population of ortolan buntings: the effect of a skewed operational sex ratio. Biol. Conserv. 1 32: 88-97. Warnock, N., Page, G. W. & Sandercock, B. K. (1997) Local survival of Dunlin wintering in California. Condor 99: 906-91 5. Wei, D. L. Z„ Bloem, A., Delany, S., Martakis, G.& Quintero. J. O. (2009) Status ofwaterbirds in Asia — results of the Asian Waterbird Census: 1 987-2007. Kuala Lumpur: Wetlands International. Wenink, P. W. & Baker, A. J. (1996) Mitochondrial DNA lineages in composite flocks of migratory and wintering Dunlins ( Calidris alpina). Auk 1 13: 744-756. Wenink, P. W., Baker, A. J., Rosner, H. U. & Tilanus, M. G. J. (1996) Global mitochondrial DNA phylogeography of holarctic breeding Dunlins (Calidris alpina). Evolution 50: 318-330. Dzung-Yan YANG, Institute of Natural Resources, National Dong Hwa University, No. I, Sec. 2, Da-Hsueh Rd. Shou-Feng, Hualien, 97401, Taiwan. Email: dzungyun@yahoo.com.tw Chungyu CHIANG, Department of Life Science, Tunghai University, Taichung, 40704, Taiwan. Email: dec.chiang@gmail.com Yu-Cheng HSU, Department of Natural Resources and Environmental Studies, National Dong Hwa University, No. 1, Sec. 2, Da-Hsueh Rd. Shou-Feng, Hualien, 97401, Taiwan. Email: ycsheu@maiindhu.edu.tw (corresponding author) FORKTAIL 28 (2012): 71-76 Habitat associations of the Manchurian Reed Warbler Acrocephalus tangorum wintering on the Tonle Sap floodplain and an evaluation of its conservation status JEREMY P. BIRD, BERRY MULLIGAN, ROURS VANN, PHILIP D. ROUND & JAMES J. GILROY Sixty tape playback trials and 17 net rides were used to investigate the habitat associations of Manchurian Reed Warbler Acrocephalus tangorum at what is potentially its most significant wintering site, the Tonle Sap floodplain in Cambodia. Fieldwork in March 2006 concentrated at three sites within the inundation zone during the dry season when floodwaters are at their lowest. This corresponds with the non-breeding season, when Palearctic migrant Acrocephalus warblers visit. We analysed cover of different habitat types at two scales: close to and broadly surrounding playback locations. Detections within broad habitat types differed significantly from random (c2 = 32.8, d.f = 5, P <0.001 ) with an apparent bias towards grass habitats represented in the study area by tall (> 1 .0 m) grassland. A principal component analysis (PCA) of the proportionate abundance of different habitats within 10 m of playback locations generated just two PCA axes, correlating strongly with the abundance of grassland and wet habitat features (PCA1 ) and woodland and scrub (PCA2). Logistic regression with both axes as predictor variables revealed a significant effect of PCA1 (z = -2.566, P = 0.010), but no significant effect of PCA2 (z = 0.088, P = 0.419). All sites with detections had a low loading on PCA1 , suggesting a strong association with grasslands. Capture rates were extremely low compared with one wintering location in Thailand, so while our study suggests theTonle Sap is of global importance for the species, we cannot find sufficient evidence to warrant revising the species's IUCN Red List status from Vulnerable to a lower category of threat. INTRODUCTION The Manchurian Reed Warbler Acrocephalus tangorum is a migratory passerine listed as Vulnerable on the IUCN Red List (BirdLife International 2012) because it is inferred to have a small and declining population. The species is only known to breed at a few sites in extreme south-east Russia and north-east China. It is considered rare or threatened in north-east Asia by Gluschenko (1989) and of the three sites in the Chinese breeding range (excluding Inner Mongolia), Alstrom et al. (1991) reported that two did not appear to have much suitable habitat remaining and they ‘doubt that significant numbers of tangorum remain’, but it remained ‘locally common’ at Zhalong. The species’s status in, and even the extent of, its wintering range remains particularly poorly understood. The wintering grounds were unknown until a population was found in Phragmites reedswamp atKhaoSam RoiYot, Thailand, in 1981 (Round 1993, Round & Rumsey 2003). Subsequently, it has been found wintering in sedge beds within deciduous dipterocarp forest at a number of sites: Champasak province, southern Lao PDR (Round 1998), Preah Vihear province, northern Cambodia (Clements et al. 2005) and southern Mondulkiri province, eastern Cambodia (Bird et al. 2007). In 2000 ‘at least 20’ Manchurian Reed Warblers were recorded at Krous Kraoum, KompongThom province, on the Tonle Sap floodplain in Cambodia (Robson 2000, Davidson 2001), and more records have since followed in Kompong Thom and Siem Reap provinces, Cambodia. These observations originated from a variety of habitats, particularly tall grass stands (away from water), sedge beds (both wet and dry), scrub-fringed lotus swamps, and heterogeneous scrub/grass mixes away from water (BirdLife International 2012). To date, the only study examining winter habitat use by the species was conducted within the least disturbed part of a c.50 km2 marsh in Khao Sam Roi Yot National Park, Thailand (Round & Rumsey 2003). This demonstrated a strong positive association of Manchurian Reed Warbler with stands of mature Phragmites. Because Phragmites is rare elsewhere in Thailand, and because a moderate increase in mist-netting in South-East Asia and an improved knowledge of the species’s identification failed (at least before 1997) to produce many records, the Red Data Book asserts that Manchurian Reed Warbler should be considered truly scarce and locally distributed with a small global population (BirdLife International 2001). However, more recent records at wintering; sites away from Khao Sam Roi Yot demonstrate that other wetland vegetation is widely used in the absence o l Phragmites. and it remains undetermined whether this is truly suboptimal habitat (Round & Rumsey 2003). The extent of potentially suitable habitat in the Tonle Sap Lake inundation zone, South-East Asia’s largest freshwater wetland, is vast (P. Davidson in litt .) and therefore, even if population densities in floodplain habitats other than Phragmites should prove to be lower, Manchurian Reed Warbler may be more widespread and numerous than previously considered. At Khao Sam Roi Yot, the best documented (and formerly only known) wintering site for the species, the extent of Phragmites is declining owing to illegal encroachment with plantations of casuarinas, eucalyptus and coconut palms established around the margins of the area, while major prawn farms have been established in the centre, pumping salt or brackish water into previously freshwater areas (BirdLife International 2001). There is now very little Phragmites habitat in Thailand and wetlands in the country are imminently threatened by reclamation and urbanisation (Round 1993). The Tonle Sap Lake floodplain supports large areas of natural and semi-natural seasonally inundated grassland (Evans et al. 2005) that are currently under intense pressure from land- use change, with grassland cover declining since the late 1990s owing to scrub encroachment and, since 2004, an extraordinary intensification of rice cultivation (Gray 2006, Gray et al. 2007). Gray et al. (2009) documented losses of 27% of grassland cover on the Tonle Sap floodplain within 10 grassland blocks between January 2005 and March 2007, primarily as a result of land conversion for intensive dry season rice cultivation. Bengal Florican Houbaropsis bengalensis was uplisted to Critically Endangered in 2007 as a direct result of these land-use changes in the Tonle Sap floodplain (BirdLife International 2012). The impacts of land-use change on Manchurian Reed Warbler remain unknown but, based on the above, a population decline has been inferred in assessing its current IUCN Red List status (BirdLife International 2012). We used tape playback and mist-netting to investigate the habitat associations of Manchurian Reed Warbler on the Tonle Sap floodplain. Our results are discussed in the context of the species’s likely abundance and trends, in order to review its assessed extinction risk. 72 JEREMY P. BIRD ero/. Forktail 28 (2012) METHODS Study area The Tonle Sap Lake in Cambodia is the largest freshwater lake in South-East Asia (Gray et al. 2009). The lake floodplain system is characterised by notable seasonal differences in water level and surface area (Kummu & Sarkkula 2008 in Vastila et al. 2010). The lake expands from some 2,500 km2 in the dry season (December to June) to between 12,500 km2 and 15,000 km2 during the wet season (Campbell et al. 2006). The floodplain landscape is a complex mosaic of flooded forests, wet and dry scrub, cropped grasslands, tall grasslands structurally similar to Phmgmites reedbeds, and areas of rice cultivation radiating out from the lake in roughly concentric rings. During March 2006 we conducted surveys on the northern boundary of the Tonle Sap Lake inundation zone around the Siem Reap and Kompong Thom provincial border (Figure 1 ) in natural and semi-natural habitats and areas of dry season rice production (headponds for irrigation and rice paddy). The study area experiences annual flooding up to 4 m depth for 4-7 months a year (Gray 2007). Three relatively distinct sites were studied during the dry season (Figure 1): (1) an area ofseasonally inundated grassland and short tree scrubland near Stoung (dry at the time of field visits), (2) natural and semi-natural seasonally inundated grassland at Kruos Kraom (fields were dry at the time of fieldwork but headponds remain inundated year-round for irrigation) and (3) permanently flooded forest and seasonally flooded bankside scrub at Boeung Tonle Chhmar, one of the core areas of the Tonle Sap Biosphere Reserve. Tape playback Manchurian Reed Warbler has previously been shown to respond to playback (Alstrom et al. 1991 ). To investigate habitat preferences of the species, relative abundance was compared in seven broad habitat types using 60 separate playback trials. A series of random playback positions was generated and a representative sample chosen for each habitat type. At each playback position, five minutes of continuous vocalisations by Manchurian Reed Warbler was played on a portable Sony CD player through a pair of Philips SBA220 speakers at equivalent volume to reed warbler vocalisations (judged by ear). The vocalisations included song and calls alternating for a minute each for the total five minutes. During the following ten minutes, two observers standing back-to-back recorded all Manchurian Reed Warblers seen and heard from the position of the speakers. An estimation of the total number of individuals was made. Parameters recorded at each playback position were site number, date, time, cloud cover and precipitation. Broad habitat type was subjectively assigned by eye to dry scrub (which is inundated by shallow water during the wet season), permanently flooded forest, wet/flooded scrub (potentially submerged in the wet season), scrub-island within paddy (inundated in the wet season), grass- island within paddy (inundated within the wet season), tall dry grass (>1.0 m average height, inundated in the wet season), or tall wet / flooded grass ( > 1.0 m average height, artificial headponds and natural). Percentage cover within a 10 m radius (measured with a measuring tape) of playback positions of sedge, floating vegetation, cultivated rice, Mimosa sp., tall grass (>1.0 m), scrub, flooded forest, water and mud was estimated by eye. A radius of 10 m was selected by default: no comparable studies were found to indicate the attractant radius of playback on individual reed warblers; to maximise the number of playbacks completed, a manageable radius was selected; and because isolated habitat islands were part of the trial and these pockets were small, a 10 m radius was optimal. Surveys were carried out from 06h00 to lOhOO and again from 14h00 to 18h00, periods of maximum bird activity, between 4 and 24 March 2006. Using R version 2.13 (R Core Development Team 2011), we applied principal component analysis (PCA) to data on the proportionate cover of habitat features within 10 m of each playback location in order to isolate key components of habitat variation. Using ordination plots classified according to the presence or absence of Manchurian Reed Warbler detections, we then performed a visual assessment of the relationship between habitat variation and warbler detection. We used logistic regressions with presence/absence as a response variable to test for significant relationships with extracted values from habitat principal Kampong Svay Stueng Saen lantuk Prasat Balangk Siem Reap Province Chi Kraeng Stoung Kampong Thom Province TONLE SAP LAKE V S' r \ Preah/Mhear i Scale - Kampong Chhnang Province Prasat Sambour Figure 1. Study sites where playback was performed around the Tonle Sap, Cambodia. Forktail 28 (2012) Manchurian Reed Warbler Acrocephalus tangorum wintering on the Tonle Sap floodplain 73 components, as well as overall habitat classes. We also compared detection rates across the broad habitat types at playback sessions using a Chi-square test. Mist-netting Birds were caught during 14 mornings of netting and ringing sessions on 17 separate net rides, each of which was classified under a broad habitat type. Only where there was relatively extensive homogeneous habitat of the same type were net rides established. Nets were operated for up to four hours, mostly between 06h00 and lOhOO. All birds caught were identified, ringed, measured and examined for moult and feather wear. Birds were weighed to the nearest O.lg using Pesola spring balances before being released. Capture rates allow comparison of relative abundance of Manchurian Reed Warbler between sites and with other species. The number of Manchurian Reed Warbler caught per metre-hour (mh) of mist-netting was calculated for different habitat types. Owing to the difficulties of mist-netting in flooded scrub and forest, the survey of Manchurian Reed Warbler was limited to playback in the maze of channels and streams linking Boeung Chhmar and the Tonle Sap Lake. RESULTS An estimated 35 Manchurian Reed Warblers were recorded (based on both calls and direct sightings) during 60 playback sessions. Overall densities of Manchurian Reed Warbler appeared to be relatively low, with detections being made at just 16 playback locations (27% of total). Detection rate across different habitat types was significantly different from random (Table 1, c2 = 32.8, d.f=5, P<0.001), with most detections occurring in grass habitats (Table l). Tall grassland habitats, both wet and dry, appeared to be the main vegetation types supporting Manchurian Reed Warblers within the study area. Multiple encounters also occurred in man¬ made habitats, mostly in tall grasses surrounding artificial wet headponds (used for dry season rice irrigation — an estimated 12 individuals encountered in 6 playbacks), as well as from small ‘islands’ of tall grass habitat in and along the edges of rice fields (an estimated 7 individuals encountered in 10 playbacks). Detection rates in forest and scrub habitats, both flooded and dry, were very low, and no detections were made in extensive areas of permanently inundated scrub and flooded forest in the Boeung Chhmar/Moat Khla IBA. However, logistic regression with habitat type as an explanatory variable revealed no significant differences between habitat types, potentially owing to the small sample sizes in each habitat. Table 1 . Broad habitat classifications of playback locations, together with the numbers of Manchurian Reed Warblers recorded in each habitat type and numbers that would be expected if habitat selection was random. The distribution of records was significantly different from random (%2 = 32.8, d.f = made head-ponds. 5, P <0.001) . Tall wet grass' includes man- tangorum numbers Habitat Playback locations Observed Expected Dry scrub 4 2 2.3 Flooded forest 6 0 3.5 Flooded scrub 9 0 5.2 Small grass island in paddy 11 7 6.4 Scrub island in paddy 10 0 5.8 Tall dry grass 10 14 5.8 Tall wet grass 10 12 5.8 Sum 60 35 Table 2. Factor loadings for each variable in a principal component analysis of habitat variation across playback locations, showing the first two principal components (proportionate eigen values: PCA1 69.2%, PCA2 23.7%). Variable PCA1 PCA2 Sedge -0.030 -0.026 Floating vegetation -0.008 0.001 Rice 0.011 0.073 Mimosa sp. -0.017 0.017 Tall grass 0.824 -0.171 Scrub -0.305 0.511 Flooded forest -0.231 -0.794 Water -0.416 -0.262 Mud -0.032 -0.065 Fig ure 2. Ordination plot showing playback locations, classified according to the presence or absence of Manchurian Reed Warbler in relation to principal component axes explaining variation in habitat characteristics. Axes PCA1 and PCA2, collectively, explain 93% of variation in the habitat characteristics shown. NB 'snaow' is Sesbania spp. PC A of the proportionate abundance of different habitats within 10 m of playback locations revealed that habitat variation was rather simple (Table 2), with 93% ofvariation being explained by just two PCA axes, correlating strongly with the abundance of grassland and wet habitat features (PCA1) andwoodland andscrub (PCA2). Logistic regression with both PCA1 and PCA2 as predictor variables revealed a significant effect of PCA1 (z = -2.566, P = 0.010), but no significant effect of PCA2 (z = 0.088, P = 0.419). Figure 2 shows an ordination plot of playback sample locations against both PCA axes, with sites classified according to the presence or absence of Manchurian Reed Warblers. All sites at which the species was recorded have a low loading on PCA1, suggesting a strong association with locations dominated by grasslands. Manchurian Reed Warblers were absent from all locations which received a high loading for PCA1, suggesting avoidance of habitats associated with wet features such as floating vegetation, open water, mud and sedges. Figure 2 also shows that locations with a high or low loading on PCA2 did not produce detections, suggesting avoidance of wooded or scrub-rich sites. This analysis underlines the importance of tall grassland for Manchurian Reed Warblers within the study area. Grasslands, structurally similar in height and density to Phragmites reedbed, also tended to be the dominant habitat at playback locations at which detection rates were high. Tall grass comprised 76% habitat cover within 10 m of playback locations where Manchurian Reed Warbler was detected versus just 26% at locations where none was detected. The sample of Manchurian Reed Warblers from mist-netting was too small to test statistically, but their distribution was heavily 74 JEREMY P. BIRD era/. Forktail 28 (2012) Table 3. Habitat types and capture rate for Manchurian Reed Warblers (MRW) netted in Kompong Thom and Siem Reap provinces. Broad habitat type No. of net rides Metre- hours netted (mh) No. of MRW caught No. per hundred metre-hours netted (n/IOOmh) Grass/scrub 2 391 1 0.26 Dry scrub 4 1,719 0 0.0 Wet/flooded scrub 3 975 0 0.0 Tall dry grass (>1.0m) 5 1,914 6 0.31 Tall wet grass (>1.0m)/scrub (headpond) 1 819 2 0.24 Tall wet grass (>1.0m) (natural) 2 825 0 0.0 “Pure mature Phragmites 6 2,706 60 2.2 *Data from captures at Khao Sam Roi Yot, Thailand, from Round & Rumsey (2003) skewed towards grassland. All eight of the net rides where individuals were caught were dominated by grass (Table 3). Capture rates (n/ mh netted) for the species were very low in comparison to those recorded by Round & Rumsey (2003) in pure Phragmites karka stands at Khao Sam Roi Yot (Table 3). During 6,643 mh of mist-netting, captures of all Acrocephalus warblers involved 9 Manchurian Reed Warbler, 73 Black-browed Reed Warbler A. bistrigiceps, 3 Blunt-winged Warbler A. concinens , 118 Oriental Reed Warbler A. orientalis and 1 Thick-billed Warbler A. aedon. DISCUSSION Habitat association and abundance A strong positive association of Manchurian Reed Warbler with grassland was demonstrated using playback. The species has been observed in a wide variety of wetland habitats in Cambodia, but within the Tonle Sap inundation zone areas of tall grass (possibly a Saccharum sp.), similar in appearance to Phragmites reedbed, appear to be strongly favoured over alternative habitats. It remains unclear whether the species is wintering in suboptimal habitat on the Tonle Sap floodplain, as no Phragmites reedswamp could be located for direct comparison with earlier studies (e.g. Round & Rumsey 2003). Densities of Manchurian Reed Warbler on the floodplain were low relative to other Acrocephalus species and locations. Despite 6,643 mh mist-netting effort, just nine individuals were captured at a rate of 0.0013 individuals per metre hour, whereas Round & Rumsey (2003) recorded a capture rate of 0.022 in stands of mature Phragmites at Khao Sam Roi Yot, capturing 60 different individuals during April 1995. The high levels at Khao Sam Roi Yot might partly be explained by influxes of migrant individuals and a concentration effect in limited habitat area, but this is conjecture. Inundated grasslands within the Tonle Sap floodplain are of exceptional importance for conservation (Gray et al. 2009). It has been suggested that Manchurian Reed Warbler might have a large wintering population in Cambodia given the extensive available habitat in the Tonle Sap floodplain (Round & Rumsey 2003, P. Davidson pers. comm. 2005). Our findings support this assertion: although densities throughout the floodplain are likely to be low given the habitat associations and densities we observed, the available area of tall grass stands remain vast (but needs quantifying). Furthermore, with a spate of new wintering records in the late 1990s and 2000s redefining the species’s non-breeding range to include much of Cambodia to southern Lao PDR and the Vietnamese border, the availability of tall grass within this range is unlikely to be a limiting factor considering its prevalence in human modified landscapes on the floodplain. Although Manchurian Reed Warbler is now known to be more widespread in the non-breeding season, the Tonle Sap lies at the core of its wintering range, and as South- East Asia’s largest permanent freshwater wetland it probably represents a stronghold for the species. Continuing destruction of wetland habitats in Thailand may further increase the global significance of this population (BirdLife International 2001, Round & Rumsey 2003). Potential threats Alluvial grasslands have been cleared preferentially for agriculture in most parts of East and South-East Asia, so the large areas remaining around the Tonle Sap Lake are of outstanding regional importance (Evans et al. 2005). Drainage and conversion of natural wetlands and grasslands, primarily for rice agriculture, have resulted in a significant decline in the area of available habitat for conservation priority species in Cambodia (Seng et al. 2003) and expansion of dry season rice is probably the most serious threat to the biodiversity of the Tonle Sap grasslands (Evans et al. 2005). It is possible, however, that Manchurian Reed Warbler has been less heavily impacted than other species. The major land-use changes occurring around the floodplain, particularly the shift from traditional wet season rice cultivation to dry season rice cultivation (Chan et al. 2004, Evans et al. 2005), may influence habitat availability for the winteringpopulation. Manchurian Reed Warbler appears to persist around headponds created for dry season rice irrigation, as well as in small blocks of tall grass within paddyfields themselves. This implies that the influence of these land-use changes will not necessarily be negative. It is also likely that traditional management systems are beneficial to the species, as regular grazing and burning help maintain grasslands (both tall and short swards) by preventing successional reversion to scrub and forest (BirdLife International 200 6). Despite the availability of apparently suitable habitat within Cambodia, it is important to emphasise that population densities are likely to be low in most areas. The continuing degradation and inadequate habitat protection at Khao Sam Roi Yot, the only other relatively large area of known habitat to support this species in South-East Asia, highlights the need for conservationists to ensure the maintenance of suitable habitat for this species in Cambodia. Management measures such as retaining tall grass margins may be relatively simple and cost-effective to implement, and should be considered in mid- to long-term management objectives for inundated grassland areas around the floodplain. Another potential threat to the species, not reported in the IUCN Red List account, is trapping. Trapping is widespread in South-East Asia for food and for merit release (the practice of releasing caged wild-caught birds for religious purposes). Trapping has been implicated in rapid declines in Yellow-breasted Bunting Emberiza aureola populations: roosting flocks in reedbeds are disturbed and then caught in mistnets, and are cooked and sold as ‘sparrows’ or ‘rice-birds’ (BirdLife International 2012). This practice has now become widespread and, although it is illegal in some areas, over a million individuals are reportedly killed annually to be sold as snacks (BirdLife International 2012). While Manchurian Reed Warbler is unlikely to be a focus of trapping effort it may well be caught as a bycatch species. We observed passerine trapping while conducting fieldwork. Merit release is now also widespread at tourist sites. JPB observed a tail-less individual of either^. bistrigiceps or A. tangorum in central Phnom Penh that was presumably a merit-released bird. Red List implications The IUCN Red List criterion that has been triggered to qualify this species as Vulnerable, C2a(ii), states that the population is estimated to number fewer than 10,000 mature individuals and is undergoing a continuing decline, with all mature individuals in one subpopulation (IUCN 2001). The population estimate for Manchurian Reed Warbler of 2,500-9,999 mature individuals Forktail 28 (2012) Manchurian Reed Warbler Acrocephalus tangorum wintering on theTonle Sap floodplain 75 (BirdLife International 2012) is derived from the original Red Data Book assessment that it ‘must be presumed to have a fairly small world population’ owing to the scarcity of wintering records and its restricted wintering range (BirdLife International 2001). We recommend revising the text that accompanies the Red List account to reflect that the wintering range is now known to be much larger than when the Red Data Book account was compiled. Based on our findings it seems unlikely that wintering habitat is population- limiting and it should not be used as the basis for the current population estimate. However, it would be premature to revise the population estimate upwards and downlist the species to Near Threatened or Least Concern as a consequence, for two reasons. First, although the species has a broad non-breeding range we cannot definitively say its population is larger than the current estimate. It is possibly noteworthy that increased mist-netting in Phragmites in central Thailand (Bung Boraphet), during which several thousand Acrocephalus warblers were ringed in 2007-2012, yielded fewer than ten Manchurian Reed Warblers (PDR unpubl. data), and very few have been caught on migration in Hong Kong, suggesting the global population is indeed relatively small. Second, the Red Data Book provides sufficient justification for a cautious approach: various problems exist at the three best known breeding grounds of the species, Khanka Lake in Russia and Zhalong and Dalai Hu National Nature Reserves in China, while the two other reported breeding sites in China (excluding Inner Mongolia) are regarded as no longer suitable for the species (Alstrom et al. 1991, BirdLife International 2001). We are unaware whether updated information exists from these sites in the Russian or Chinese literature. If it does, it could provide valuable insights into the conservation status of the species. If not, gathering this information should be a priority for further research. It is also clear from the Red Data Book account that assessing the species’s status in Inner Mongolia is important. After small population size, the second assertion of the current IUCN Red List assessment is that the species’s population is declining. Despite conversion and degradation of habitat in some wintering sites (notably Khao Sam Roi Yot), the Manchurian Reed Warblers ability to exploit a variety of habitats throughout its non¬ breeding range suggests that habitat conversion may not be causing declines on the wintering grounds, and this should be reflected in the Red List text account. However, again we feel it is premature to recommend changing the Red List status owing to uncertainties on the breeding grounds, where habitat conversion may well be causing the population to decline. While the population consequences of documented threats remain unquantified, there is insufficient evidence to infer a rapid population decline (>30% over 10 years or three generations) that could qualify the species as Vulnerable under the A criterion of the IUCN Red List (IUCN 2001). Furthermore, now that the wintering range has been increased substantially through improved knowledge, the species’s estimated extent of occurrence, 490,000 km2 (BirdLife International 2012), is much larger than the threshold triggering the B criterion. Overall we recommend that the status quo, listing the species as Vulnerable under criterion C2a(ii), be maintained. CONCLUSION This study is the first to investigate the wintering habitat associations of Manchurian Reed Warbler in Cambodia since records revealed the country encompasses the majority of its known non-breeding range. While the findings deserve further scrutiny and ideally more prolonged research, they go some way towards outlining Cambodia’s importance for this threatened species. Preliminary evidence suggests Cambodia, and the Tonle Sap Lake floodplain in particular, probably supports a substantial proportion of the global population. Establishing more reliably what that population is likely to be requires considerably more study, and is probably best achieved by concentrating on the breeding sites. Currently there is little evidence that the species is unduly threatened by habitat conversion in Cambodia but we must stress that this conclusion requires confirmation. Additionally, the unregulated trapping of small birds (including reed warblers) through the use of nets is widespread within the species’s non¬ breeding range and its impacts at the population level warrant further investigation. ACKNOWLEDGEMENTS We thank Pete Davidson for helpful advice and information before undertaking fieldwork, Tom Evans and colleagues at the Wildlife Conservation Society in Cambodia for providing essential help and support throughout, and the many donors for sponsoring the research: the Astor of Hever Trust, British Ecological Society, British Ornithologists’ Union, Chris and Liz Shepley, IdeaWild, Opticron, Oriental Bird Club, People’s Trust for Endangered Species, Robert and Elaine Dawson, Royal Geographical Society, Sevenoaks RSPB members’ Group, Sir Phillip Reckitt Educational Trust, University of East Anglia Travel and Expeditions Committee, The Wetland Trust and Wexas International. Also, thanks to Santhosh Kurian for his assistance with the map. Finally we thank two anonymous referees for helpful comments that greatly improved the manuscript. REFERENCES Alstrom, P., Olsson, U. & Round, P.D. (1991) The taxonomic status of Acrocephalus agricola tangorum. Forktail 6: 3-1 3. Bird, J. 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(1998) Wildlife, habitats and priorities for conservation in Dong Khanthung proposed NBCA, Champasak Province, Lao PDR. Final Report. Center for Protected Areas and Watershed Management, & Wildlife Conservation Society, Vientiane. Round, P. D. & Rumsey, S. (2003) Habitat use, moult and biometrics in the Manchurian Reed Warbler Acrocephalus tangorum wintering in Thailand. Ringing & Migration 21 : 21 5-221 . Seng K. H„ Pech B., Poole, C. M„ Tordoff, A. W., Davidson, P & Delattre, E. (2003) Directory of Important Bird Areas in Cambodia: key sites for conservation. Phnom Penh: Department of Forestry and Wildlife, Department of Nature Conservation and Protection, B i rd Life International in Indochina and Wildlife Conservation Society Cambodia Programme. Vastila, K., Kummu, M„ Sangmanee, C. & Chinvanno, S. (2010) Modelling climate change impacts on the flood pulse in the Lower Mekong floodplains. J. Water and Climate Change 1 : 67-86. Jeremy P. BIRD, 36 ThodaySt, Cambridge, Cambridgeshire, CB1 3 AS, UK. Email: Jezbird@gmail.com Berry MULLIGAN, Fauna & Flora International, Cambodia Programme, 19, Street 360, BKK1, Chamkarmorn, POBox 1380, Phnom Penh, Cambodia. Email: Berry. Mulligan@fauna- flora.org ROURS Vann, Wildlife Conservation Society #21 Street 21, Sangkat Tonle Bassac, Khan Chamkarmorn, Phnom Penh, Cambodia. Email: roursvann@gmail.com Philip D. ROUND, Department of Biology, Faculty of Science, Mahidol University, Rama VI Road, Rachadhavi, Bangkok 10400, Thailand. Email: pdround@ksc.th.com James GILROY, Department of Ecology and Natural Resources, Norwegian University of Life Sciences, 1432 As, Norway, james.gilroy 1 @googlemail.com FORKTAIL 28 (201 2): 77-92 The avifauna of Alor and Pantar, Lesser Sundas, Indonesia COLIN R.TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE The status of birds on Alor and Pantar is reviewed based on the literature plus surveys in 1991, 2002, 2009 and 2010 totalling about five weeks and two weeks respectively. A total of 116 birds (86 resident landbirds) were recorded on Alor including 37 new island records (including 27 resident landbirds) and 97 birds were recorded on Pantar including 75 new island records (including 55 resident landbirds). The most significant records were the discoveries on Alor of a population of Timor Bush Warbler Locustella timoriensis which may represent an undescribed subspecies, an undescribed species or subspecies of honeyeater resembling Crimson Myzomela Myzomela kuehnii, and an undescribed subspecies of Timor Stubtail Urosphena subulata. The endemic subspecies of Southern Boobook Ninox boobook plesseni collected historically from Alor was found to be vocally distinct and may be a distinct species; it was recorded on both Alor and (a new population) Pantar. Vocalisations of a presumed scops owl Otus on Pantar are distinct from Moluccan Scops Owl Otus magicus and may also be an undescribed taxon. Alor appears to have an important population of the Critically Endangered Flores Hawk Eagle Nisaetus floris but populations of the heavily traded Critically Endangered Yellow-crested Cockatoo Cacatua sulphured have declined greatly since 1991. The Vulnerable Flores Green Pigeon Treron floris was recorded regularly on Alor, which may be a stronghold. Other notable records are significant range extensions of Mees's Nightjar Caprimulgus meesi to both islands, new island records for Little Cuckoo Dove Macropygia orientalis and Ruddy Cuckoo Dove M. emiliana, and improved knowledge of the montane avifauna of Alor including the first records of Mountain White- eye Zosterops montanus. Our fieldwork shows that the avifauna of these islands contains more globally threatened species than previously thought, and a stronger representation of birds originating from Timor. Alor and Pantar are therefore of greater global conservation significance than previously highlighted. INTRODUCTION The Flores island chain spans more than 1,000 km from Lombok in the west through to Alor in the east: these islands delineate the Northern Nusa Tenggara Endemic Bird Area (EBA) (Stattersfield et al. 1998). Most islands in this chain would have coalesced (joined) or have been briefly isolated during ice-ages and associated sea-level changes during the Pleistocene period (10,000 yrs BP and earlier) (Voris 2000). This global biological hotspot hosts at least 29 restricted-range bird species, including 17 endemic to the EBA (Stattersfield et al. 1998). These numbers are even greater when the recently described Mees’s Nightjar Caprimulgus meesi (Sangster & Rozendaal 2004) and several taxonomic splits including the Critically Endangered Flores Fdawk Eagle Nisaetus floris (Gjershaug et al. 2004) are included. Most of the avian endemics of the Flores island chain are found on the large islands of Flores (at least four island endemic birds and 24 restricted-range birds; 13,540 km2) and Sumbawa (22 restricted-range birds; 15,448 km2), and consequently most historical and modern-day fieldwork has concentrated on these islands (White & Bruce 1986, Butchart et al. 1996, Johnstone et al. 1996, Pilgrim et al. 2000, Hutchinson etal. 2007). Despite recent efforts (Trainor 2002a, 2003, Schellekens et al. 2011), most other islands in the group have been neglected by ornithologists, with little published on the avifauna of Pantar (728 km2) since the nineteenth century (Hartert 1898, Johnstone 1994). Alor Island (2,800 km2, 1,825 m) is rhe fourth largest island in the EBA but has been visited rarely, and consequently its avifauna has also remained poorly known. A. H. Everert visited both islands in March-April 1897, collecting 64 bird species on Alor and 22 species on Pantar (Hartert 1898). The collecting on Alor ‘cannot be regarded as at all an exhaustive one’ because a lack of water and food hampered the visit, causing Everett to become feverish and then injure a leg which required hospitalisation on Sulawesi (Hartert 1898). The collection was enough to determine that Alor has a ‘Flores ornis’ (Hartert 1898) although a few species typical of Timor Island, such as Olive¬ headed Lorikeet Tricboglossus euteles , Bar-necked Cuckoo Dove Macropygia magna and the Alor-endemic subspecies of Southern Boobook Ninox boobook plesseni were collected on Alor, with the latter taxon being described as late as 1929 (Stresemann 1929, White & Bruce 1986). V. von Plessen visited Alor in July- August 1927, collecting 68 bird specimens of 31 bird species (Rensch 1929). A major study ( 1987- 1 993) of the avifauna of the Lesser Sundas and the islands of Maluku by the Western Australian Museum (WAM) and the Indonesian Institute of Sciences (LIPI) included a visit to Pantar and Alor in April-May 1991. This resulted in new information on the shorebirds and seabirds of Pantar and Alor (Johnstone 1994), the discovery and description of a new form of Sunda Bush Warbler Horornis [ Cettia\ vulcanius kolichisi (Johnstone & Darnell 1997a) on Alor, and a redescription of the Alor subspecies of Southern Boobook (Johnstone & Darnell 1997b). Substantial additional information on the landbirds of Alor and Pantar, including many new island records, has remained unpublished. The first published record of a montane bird on Alor (Lesser Shortwing Bracbypteryx leucophrys) by Holmes (1995) was notable. Recent visitors to Alor include limited ornithological observations by Mason (1991, 1993) and Holmes (1995), and a status survey of rhe Critically Endangered Yellow-crested Cockatoo (Setiawan et al. 2000). Following a failed attempt to reach Wetar Island by CRT, birds on Alor were surveyed at several locations over three weeks in April- May 2002, with an account of the birds at one location in the vicinity of Mt Koya Koya and Tanglapui Timur published (Trainor 2005a). A visit to Alor by PV and Veerle Dossche (VD) in June- July 2009, and to Pantar and Alor in September 2009 (PV), resulted in the discovery of a population of Timor Bush Warbler Locustella timorensis on Alor (Trainor et al. 2012), prompting further fieldwork by CRT on Alor and Pantar in December 2009 and January 2010. The aim of this paper is to document the new general ornithological observations of REJ, PV and CRT, particularly significant new island records, and records of globally threatened and restricted-range bird species. Many of the supposed new island records mentioned in Holmes (1995) and Trainor (2005a) had actually been recorded during the WAM-LIPI expeditions, so we take the opportunity to correct the record. Recent discoveries by PV also prompted brief visits to Alor and Pantar by Peter Collaerts (PC) and a BirdTour Asia tour led by James Eaton (JE) and Rob Hutchinson (RH), with significant records from these ornithologists also noted in this paper. 78 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) METHODS We independently visited seven sites on Alor over about 40 field days, including visits to three islets (Kepa, Ternate and Sika off Alor), and five sites on Pantar Island over about 14 days. Details of the sites, survey dates and habitat are provided in Table 1 and Figure 1 . The main sites were in the far east near Tanglapui Timur and Mt Koya Koya, and above Apui (near Subo village and Manmas hamlet): both these locations were surveyed over a total of about 14 field days. To support bird identifications, and descriptions, CRT and PV took photographs using respectively a Canon 7D and a Canon 40D digital camera with a 100-400 mm lens. Sound recordings were made with an Olympus LS-10 recorder (CRT) and an Edirol R09 HR, and Olympus WS 331 M digital recorder and a Sony Minidisk HI- MD MZ-RH 1 (PV) — all in combination with an ME-66 Sennheiser directional microphone, and playback was done with a RadioShack mini amplilier-speaker. A set of recordings will be uploaded to web databases such as xeno -canto and the Avian Vocalization Centre (AVoCet). REJ visited three sites on Alor and two sites on Pantar as part of the WAM-LIPI survey of the Nusa Tenggara islands. Birds were collected by trapping with mistnets and general observations. Specimens are housed at the WAM and LIPI. Taxonomy and nomenclature follows Gill & Donsker (2012) except that we use Common Golden Whistler Pachycephala pectoralis instead of Rusty-breasted Whistler P. fulvotincta and Brown Honeyeater Lichmera indistincta instead of Indonesian Honeyeater L. limbata. ANNOTATED LIST We recorded a total of 116 bird species on Alor including 37 new island records (27 resident landbirds) and a total of 97 bird species on Pantar including75 new island records (55 resident landbirds). Only eight bird species, including one Australian migrant and several visiting waterbirds, appear to have not been recorded on Alor since the historical collections (Australasian Grebe Tachybaptus novaehollandiae , Pacific Baza Aviceda subcristata. Red- backed Buttonquail Turnix maculosa , White-headed Stilt Himantopus leucocephalus, Swinhoe’s Snipe Gallinago megala, Channel-billed Cuckoo Scytbrops novaehollandiae , Savanna Nightjar Caprimulgus affinis and Thick-billed Flowerpecker Dicaeum agile). On Pantar, only two species recorded historically (Australian Hobby Falco longipennis and Sunda Cuckoo Cuculus lepidus ) went unrecorded in the current surveys. The following annotated list focuses on significant resident landbirds, but includes information on a few migrants. Many of the numerous new island records are unsurprising and these are denoted clearly, with the tabulated summary of all records in the appendix. Table 1 . Study site details, survey dates and habitat. Refer also to Figure 1 . Location (elevation range in m) Coordinates Dates Habitat Pantar Batu 8015'16"S124°17'59"E 17—19 April 1991 (REJ) Regrowth forest; Eucalyptus woodland Kabir 8°15'30"S124°13'05"E 20-21 April 1991 (REJ) Regrowth forest; Eucalyptus woodland; mangrove Mt Wasbila/Pantar Timur/ (400-800) 8°2T16"S 124°03'03"E 3,5,6 September 2009 (PV); 11-12 December 2009 (CRT) Regrowth forest; Eucalyptus woodland; grassland Mt Sirung/PantarTengah (500) 8°29,26"S124°07'28'/E 4 September 2009 (PV); 13 December 2009 (CRT) Eucalyptus woodland; grassland Baranusa 8°21'32"S124o05'38"E 2-5 September 2009 (PV); 10-13 December 2009 (CRT) Village; mangrove; Eucalyptus alba woodland Puntaru 8°26'5T'S 124°03'04"E 13-14 December 2009 (CRT) Village; coastal woodland; grasslands and beach Alor Pulau Kepa (c.0.45 km2) Pulau Ternate (c.3.7 km2) Pulau Sika (c. 0.9 km2);also mainland at Mali Wah Wah/Alor Barat Laut (400-800) Mainang/Alor Tengah (400-1,100) 8°16'22"S 124°24'02"E 8°13'21"S 124°22'30"E 8‘>16'22"S124°24'02"E 8°10'32"S 124°30'57"E 8°16'39''S 124°36'38"E 26 April 2002 (CRT);28-30 June 2009 (PV & VD) 27 April 2002 (CRT) 29 April 2002 (CRT) 21 January 2010 (CRT) 9-11 January 2010 (CRT) Coastal woodland; beach Secondary tropical forest; agricultural land Coastal strand; short grass Regrowth forest; tree crops Evergreen forest; Eucalyptus alba woodland Apui-Subo-Manmas/Alor Selatan (700-1,250) 8°17'39"S 124°43'17"E Tanglapui Timur (including Kunggwera, Lipa, Kobra, Mt Koya Koya/Alor Timur (400-1,200) 8°14'56"S 124°59'30"E 22-25 April 1991 (REJ); 1 July 2009 (PV &VD), 31 August 2009 & 9 September 2009 (PV), 4-6 December 2009 (CRT), 14 January 2010 (CRT) 2-7 & 1 1-16 May 2002 (CRT);21-27 June 2009 (PV & VD), 16— 18 January 2010 Mixed tree crops; and regrowth forest; E urophylla woodland Regrowth forest; E. alba woodland Figure 1. Map of Pantar and Alor showing survey sites visited or localities mentioned in the text. Sites: 1 = Puntaru village; 2 = Mt Wasbila; 3 = Baranusa 4 = Mt Sirung 5 = Kabir; 6 = Treweg Island; 7= Batu village; 8= Pura Island, 9= Ternate Island; 10= Buaya Island; 1 1 = Kepa Island, 1 2= Kalabahi; 13= Wahwah; 14= Mali; 1 5= Mainang; 1 6= Sika Island; 1 7= 10 km north of Apui; 18= Apui- Subo-Manmas; 19= Kunggwera; 20= Mount Koya Koya; and, 21 = Tanglapui Timur (Lipa, Kobra hamlets). Source is Google Earth, 10 March 2012. Forktail 28 (2012) The avifauna of Alor and Pantar, Lesser Sundas, Indonesia 79 Yellow Bittern Ixobrychus sinensis Alor: An adult male (WAM A24495) was collected in rice fields at Apui-Subo on 24 April 1991 (REJ). The Yellow Bittern is poorly known in Wallacea. It was thought to be largely migratory, except on Sulawesi and Flores (Coates & Bishop 1997), but there appear to be no confirmed breeding records on Flores (Verheijen 1964, Mees 2006). Dated specimens range from September to April (White 1976). A nest with one chick was discovered on Sumbawa in May 198-8 ( Johnstone et al. 199 6). The number of April, May, June and July records for East Timor, and a recent July record on Sumba (Trainor 2011), suggest that this bird either breeds widely in Nusa Tenggara or that some individuals occasionally oversummer in the region. Eastern Osprey Pandion cristatus Pantar: The first island record was of a single bird observed in flight along the coast at Baranusa village on 10 December 2009 (CRT). Eastern Osprey appears to be uncommon to rare in the central Lesser Sundas (White & Bruce 1986). Everett observed one bird on Alor during April or May (Hartert 1898), and one bird was observed in the hills ofAlor during December 1994(Holmes 1995). During about 300 field-days in Timor-Leste, CRT has only observed a single bird (11 October 2004). These presumably refer to resident birds. Migratory Western Osprey/? haliaetus has been recorded in northern Wallacea (Sangihe, Talaud and Sulawesi) but none has been noted entering Wallacea from Bali (White & Bruce 1986, Germi 2005, 2006) , suggesting that the migratory species is either very rare or does not enter Wallacea along the Continental Flyway (Germi 2009). Short-toed Snake Eagle Circaetus gallicus gallicus Alor: One was observed and sketched over Apui-Subo on 24 April 1991 (REJ). One bird was observed above Apui-Subo on 1 July 2009 (PV and VD); a pair were seen at the same site on 9 September 2009 (PV), and one bird photographed at Mainang on 9 January 2010 (CRT). Pantar: One bird was observed by WAM staff near Kabir on 21 April 1991 (REJ). Singleswere observed daily in September 2009, including one bird photographed, on the slopes of Mt Wasbila (PV) . The resident Lesser Sundas population (Lombok to Timor/Roti) of Short-toed Snake Eagle is isolated by at least 2,500 km from the nearest resident populations in India and Nepal ( Robson 2000). Migrant birds reach Bali at least (Coates & Bishop 1997). Despite the isolation of this population, there are few apparent morphological differences from the nominate subspecies (Mees 2006). Bonelli's Eagle Aquiia faseiata renschi Alor: One bird was observed about 12 km north-east of Mt Koya Koya (on the road to Tanglapui Timur) on 10 May 2002 soaring over tropical forest; a pair was seen on the slopes of Koya Koya on 13 May 2002, and a single bird was observed south of Lipa hamlet on 14 May 2002 (CRT). Two pairs and a single were observed near Mainang on 9 January 2010 flying over Eucalyptus alba woodland and secondary forest, with one pair photographed (CRT). A pair was photographed near Baifui village (5 km south-west of Apui- Subo) on 1 1 January 2009 (CRT). Pantar: One adult was giving a display flight on the slopes of Mt Wasbila on 3 September (PV). The isolated resident Lesser Sunda population of Bonelli’s Eagle ranges from Lombok to the Tanimbar Islands (Debus et al. in press) with the nearest resident populations in north-west Thailand and northern Laos (Robson 2005). The photographs of birds near Mainang confirm the presence of this eagle on Alor. However, records of this eagle from Alor in 2002, and Pantar in 2009, may have been confused with Flores Hawk Eagle Nisaeetusfloris. Flores Hawk Eagle Nisaetus floris Alor: A pair was seen daily giving a territorial display over evergreen forest at Kunggwera (23-25 June 2009) and two birds were photographed in the hills above Apui-Subo on 9 September 2009 (PV). A pair and two singles were subsequently photographed at Apui-Subo on 6 December 2009 with a pair on 14 January 2010 (CRT). One was photographed in flight with a large bird (probably a rail) in its talons (CRT). A pair was photographed at Mainang at 800 m on 9January 2010 (CRT) displaying over secondary tropical forest. One bird was seen in highly degraded forest in the hills near Kalabahi on 3 July 2011 (Collaerts et al. 201 1). This critically endangered species was known from Lombok through to Flores (including Satonda, Sumbawa and Rinca islands) with a population estimated at fewer than 100 pairs (Gjershaug et al. 2004). Flores Hawk Eagle has not been recorded on Komodo Island (Raharjaningtrah & Rahman 2004). However, it was recently recorded in mangrove habitat on an unnamed islet 800 m off Komodo, apparently hunting amongst a fruit-bat colony (probably Large Flying Fox Pteropus vampyrus ) (M. van Buuren pers. comm. 2001). Our Alor records are therefore an important range extension. The use of matrix habitats on Alor including secondary tropical forest (about 10-20% cover) on valleys and slopes, mixed with more extensive Eucalyptus woodland and agricultural land, suggests that the species can persist in highly fragmented forest estates. The only extensive (>20 km2) area of tropical forest on Alor appears to be on the slopes of Mt Koya Koya, but tropical forest remnants are frequent throughout the island’s mountains, especially in gullies and on slopes. The frequency of records on Alor suggests that a good population is likely to present, although repeat observations above Apui-Subo are likely to refer to the same pair. If the home range of birds on Alor is similar to the 38.5 km2 estimated for Flores (Raharjaningtrah & Rahman 2004) then a maximum of 73 pairs could be present on the island (assuming that they can use all available habitat, which is unlikely). Flores Hawk Eagle is likely to be present on Pantar (where similar habitat was surveyed but the species went unrecorded), and perhaps Wetar, Lembata and Adonara. Although the Flores Hawk Eagle is a relatively large and distinctive raptor, in the field we (CRT, PV) initially assumed that these birds were Bonelli’s Eagles, and relied on species identification from photos. Crested Honey Buzzard Pernis ptiiorhynchus Alor: One bird was photographed above Apui-Subo on 6 December at 900 m, and another near Mainang on 1 1 and 1 3 January 20 1 0 at 700 m (CRT). Pantar: One bird was photographed in flight below Mt Wasbila over agricultural land on 10 December 2009 (CRT). Despite the lack of historical specimens (White 1976, White & Bruce 1986) this bird is a common migrant to Nusa Tenggara during the northern winter, with more than 1,000 birds sighted crossing the Lombok Strait during October-November 2004 (Germi 2005) and 7,717 birds sighted in September-November 2005 (Germi 2006). It should be expected in small numbers throughout Nusa Tenggara, and there are Christmas Island and recent Australian mainland records (Gregory 2007). Black-winged Kite Eianus caeruleus hypoleucos Alor: A single bird was observed in Eucalyptus alba woodland in Lipa hamlet on 2 May 2002, and a pair of birds was seen on 7 May 2002, hovering over woodland and perched in an E. alba tree in the same area (CRT). One bird was also seen on 21 June 2009 in coastal/:, alba woodland about 15 km north-west of Lipa (PV and VD). Three birds were also observed by Widodo (2009): one at Tanglapui and two at Tuti Adagae Nature Recreation Park. These appear to be the first Alor records since Everett’s observation of one bird (Hartert 1898). Pantar: One adult perched in a Tamarindus indicus tree at the edge of mangroves at Kabir on 20 April 1991 (REJ). In Nusa Tenggara this species appears to be ‘a local and sparsely distributed resident’ (Coates & Bishop 1997) perhaps with the exception of the grassland and woodland habitats 80 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) on Sumba (Olsen & Trost 2007) and Sumbawa, where seven birds were seen including one sitting on a nest on 7 July 2000 (Trainor 2002b). A nestling has been recorded on Flores during July (Mees 2006). The only recent Lombok records appear to be the four birds seen in 1997 (Myers & Bishop 2005), and it appears to be uncommon and local on Flores (Butchart etal. 1994, Pilgrim 2000, Mees 2006). Peregrine Falcon Falco peregrinus ernesti Alor: The first island records were of a pair displaying over agricultural land at Kunggwera on 5 May 2002 (CRT). Subsequently one bird was photographed above Apui-Subo on 9 September 2009 (PV), one was seen on a tower in Apui-Subo village on 14 January 2010 (CRT) and one bird was photographed in a roadside tree about 4 km north of Mainang on 13 January 2010 (CRT). Pantar: The first island record was of one observed in Baranusa village on 10 December 2009 (CRT). The photographs confirm that these birds are the resident subspecies F p. ernesti. Migrant F p. calidus appears to migrate through Nusa Tenggara in low numbers, with seven birds noted on Bali flying towards Lombok (Germi 2006). Red-legged Crake Rallina fasciata Alor: The trill-like call (grebe-like chatter, see http://www.xeno- canto.org/xc33425) of Red-legged Crake was heard and sound- recorded from tall grass above Apui-Subo on 4 December 2009 (CRT). The first Alor record was of a bird killed by a catapult near Kalabahi on 9 December 1994 (Pdolmes 1995). Eggs were collected from a nest in central Flores on 25 February 1959, and specimens of adults were collected in December (Mees 2006). The eggs had similar dimensions to birds breeding on Java (Mees 2006). On neighbouring Timor, the first records in February and March included observations of breeding (Dymond 201 1, Trainor 2011). These records suggest that Red-legged Crake migrates into the Lesser Sundas during the northern winter (wet season) to breed. Red-legged Crake has also been recorded recently from the east Lesser Sunda islands of Sermata and Romang (CRT unpubl. data). Flores Green Pigeon Treron fioris Vulnerable. Alor: During 2002 there were about 1 0 records mostly of 1-2 birds in tropical forest, or feeding in figs at 580-780 m, and one flock of 40-50 birds feeding in figs in forest-lined ravines at 860 m on the slopes of Mt Koya koya (CRT). In July 2009, Flores Green Pigeon was quite common in groups of up to 15 in fruiting trees about Kunggwera and calling from the forest canopy in ravines (PV and VD). A small party of 5-10 birds were observed feeding in a roadside fruiting tree near Mainang, at 700 m (CRT); one bird called from secondary forest at 1,050 m above Apui-Subo on 14 January 2010 (CRT); and two birds called from a gully 2 km west of Lipa at 720 m on 17 January 2010 (CRT), two birds were seen in tropical forest at about 230 m on the road to Apui on 9 September 2009 (PV). Pantar: The only records are two observations of single birds in flight on 3 September 2009 over tropical forest at c.400 m on Mt Wasbila (PV). Alor appears to be a stronghold for the Flores Green Pigeon, which seems to be common around the sparsely populated Kunggwera village and on the slopes of Mt Koya Koya, which retains some of the most extensive tropical forest on the island. In western Alor our records were less frequent, and populations of the Flores Green Pigeon might be limited by intensive hunting. However, throughout inland Alor there are also many extensive steep forest-lined ravines that are relatively remote from villages. There is little recent information on the status of Flores Green Pigeon from Flores and Sumbawa apart from bird tour reports (Eaton 2009), and the last published Lombok record was from about 1909 (Myers & Bishop 2005). Pied imperial Pigeon Ducula bicolor Alor: A flock of eight birds feeding in a fruiting Garuga tree and later a flock of four in a teak Tectona plantation near Kalabahi on 10 April 1991 (REJ) are the first Alor records. The Pied Imperial Pigeon is a small-island species that possibly visits islands nomadically (Coates & Bishop 1997) to access fruit resources. There are few Nusa Tenggara records, but most of these have come from Komodo, Rinca and West Flores, associated with extensive mangrove landscapes (apparently used for feeding and roosting) during July-November and January and March (Schellekens et al. 2009). It is frequent on the large island of Yamdena in the Tanimbar islands (Bishop & Brickie 1998). Pied Imperial Pigeon is clearly not resident on Alor, but the likely source of these birds is unclear. It typically breeds on small islets, but the islands around Alor (Treweg, Kepa, Pura, Ternate and Buaya) are relatively heavily populated and are unlikely breeding habitat. It might be expected to occur about the extensive mangrove stands on Pantar Island. Green / Pink-headed Imperial Pigeon Ducula rosacea laenea polia Alor: Pink-headed Imperial Pigeon D. rosacea was recorded in small flocks of up to five birds in coastal forest near Kalabahi, with Green Imperial Pigeon D. aenea also moderately common in flocks of up to 18 birds in the same area in April 1991 (REJ). Imperial pigeons were common in forest in Kunggwera at 600-900 m with up to 1 5 Green present in a fruiting fig, and occasional on the slopes of Mt Koya Koya (CRT). Green Imperial Pigeons were also recorded frequently in the forests around Kunggwera in June 2009 (PV). A few imperial pigeons, probably Pink-headed, were recorded a couple of times on Kepa Island and Ternate Island (CRT). Pantar: Green Imperial Pigeons were recorded in small flocks in forest and mangrove at Batu and Kabir in April 1991 (REJ). Pink-headed Imperial Pigeons were moderately common in small flocks up to nine at Batu and in mangrove at Kabir in April 1991 (REJ). Green were recorded relatively commonly in tropical forest on Mt Wasbila in September 2009 (PV) but were heard only a few times in December 2009 over two days at the same site (CRT). Small numbers of imperial pigeons (2-4), most probably Pink-headed, were observed and heard calling on 5 September 2009 in a stand of mature mangrove forests some 5 km from Baranusa (PV). Imperial pigeons appear to have been severely impacted by hunting on Alor: they were relatively common in remote forested areas in the east, but none was recorded at Apui- Subo or Mainang in west Alor, with only a few birds heard on two islets. A total of 13, 9 and 17 Green Imperial Pigeons were recorded at three Alor sites and seven Pink-headed were recorded at one site by Widodo (2009). Imperial pigeons may be more abundant in lowland coastal forests on Alor, especially those dominated by Canarium trees, although this habitat has not been covered in recent surveys. Bar-necked Cuckoo Dove Macropygia magna magna Alor: Th is dove was frequently seen and heard in tropical forest and sometimes adjacent Eucalyptus woodland at Kunggwera in 2002 and 2009 (CRT, PV and VD), and up to five birds were heard calling (and sound-recorded) in tropical forest at 250 m about 10 km north of Apui-Subo, and above Apui-Subo on 1 July 2009 and 9 September 2009 (PV). A few birds were also observed in degraded forest in the hills near Kalabahi on 3 July 2011 (P. Collaerts in litt. 2012). Pantar: Relatively commonly heard and seen (and sound-recorded) in tropical forest on Mt Wasbila (PV) with up to 20 birds calling at dusk from a small area there on 10 December 2009 (CRT). One bird was calling in beach forest at Puntaru (CRT). Bar-necked Cuckoo Dove occurs on the Flores Sea islands of Tanahjampea and Kalaotoa, and in Forktail 28 (2012) The avifauna of Alor and Pantar, Lesser Sundas, Indonesia 81 the Lesser Sundas from Pantar through to Tanimbar (White & Bruce 1 986). Both Alor and Pantar probably host large populations of this dove but these may be impacted by hunting in heavily populated areas. The vocalisations of the different races are distinct, suggesting that at least three species are involved (Eaton & Hutchinson 2011). On Pantar, Alor, Wetar, Atauro, Timor and associated islands At. m. magna gives a two-note call, while At. m. timorlaoensis of the Tanimbar archipelago gives a three-note call (Coates & Bishop 1997, www.xeno-canto.org) and subspecies At. m. longa of the Flores Sea also gives a distinctive call (Eaton & Hutchinson 201 1 ). Little Cuckoo Dove Macropygia ruficeps orientalis Alor: Frequently heard and seen in tropical forest at Kunggwera (PV) and secondary forest above Apui-Subo (500-1,000 m) (PV.CRT), and a few birds heard at Mainangat 700 m (CRT), and a single bird heard calling from secondary forest c.2 km east of Lipa village (CRT). Pantar: A pair was photographed at c.400 m in tropical forest on Mt Wasbila (PV), and heard calling there about 10 times over two days at 300-400 m in December 2009 (CRT). Little Cuckoo Dove occurs widely in South-East Asia and the Greater Sundas, with the endemic subspecies At. r. orientalis reputedly present on the islands from Lombok to Flores, Lembata and Timor, Sumbawa, Flores, Pantar (White & Bruce 1986) and now Alor. It is generally poorly known in Nusa Tenggara, but is a common bird in the mountains of Timor (CRT unpubl. data). The call of birds on Timor, Pantar and Alor is unlike other members of the species and unlike those of the endemic Lesser Sundas subspecies At. r. orientalis , suggesting that it is an undescribed species (Eaton & Hutchinson 2011). An analysis of calls is currently underway (JE in lift. 2012). Ruddy Cuckoo Dove Macropygia emiiiana Alor: A sound recording of this bird was made in tropical forest about 10 km north of Apui on 29 August 2009 (PV). On 23 October 2011 it was observed in flight at least twice in secondary forest at Mainangat c.900-950 m (RH, JE) with afield note at the time of a ‘reddish large cuckoo dove’ (JE in litt. 2012). Ruddy Cuckoo Dove was known from Lombok, Sumbawa, Flores and Paloe in Nusa Tenggara (White & Bruce 1986), with these records the first for Alor. This bird calls rarely on Java and Flores ( JE in litt. 20 1 2) and consequently it is relatively little known in Wallacea. A recorded bird gave a downslurred double note whu-ivhu repeated at about 5-second intervals, which is similar to calls described from Flores (Coates & Bishop 1997). This bird is considered to be generally uncommon on Flores (Verhoeye & Holmes 1999), but locally common in the mountains of Flores (Pilgrim et al. 2000) and rare in lowland forest, but common in upper montane forest on Sumbawa (Johnstone et al. 1996). Yellow-crested Cockatoo Cacatua sulphurea parvula Alor: A loose flock comprising six, eight and four birds was observed over forested gullies at c. 230 m, 1 0 km north of Apui, on 1 July 2009 (PV and VD). Local people said that cockatoos were regularly seen in these river valleys (PV). In 2002, local people at Kunggwera stated that flocks of up to 10 birds were occasionally seen about the village, and that c.20 birds were present in a nearby river valley, but none was seen there during 2002 or 2009 (CRT PV). Pantar: In September 2009, a caged bird was seen in the port of Baranusa (PV) ready to be shipped off the island, and another caged bird, caught on the island, was seen and photographed in the village of Anis Lilo (PV). This latter, a male, was on sale for Rp. 400, 000 (c.$US40), but prices of up to Rp. 1,500,000 (c.$US150) are sought for ‘red-eyed’ female birds on Pantar (PV). The Yellow-crested Cockatoo is currently a rare bird on both Alor and Pantar as a result of intense historical and ongoing captures for trade. On 9 April 1991 a total of 23 adults and 12 immature cockatoos were seen in small cages at Kalabahi ready to be shipped out (REJ). No birds were observed at a Kalabahi market on 9 May 2002 (CRT). No cockatoos were seen by Holmes (1995) during a brief visit to sites in west Alor, but in the late 1990s a total of 80 birds were seen at seven Alor sites (4—30 birds per site) and 29 were seen at two sites on Pantar (Setiawan et al. 1999; no survey dates given in report). The total population was crudely estimated at 678-782 birds for Alor and 444-534 birds on Pantar (Setiawan et al. 1999) based on the extent of preferred riverine forest. No cockatoos were recorded at four Alor sites during a specific search for this species (Widodo 2009; no survey dates given in article). Riverine and coastal swamp forest dominated by Canarium are probably the most important habitats for Yellow-crested Cockatoo (and possibly imperial pigeons) on Alor (Setiawan et al. 1999, Widodo 2009), as these large trees form hollows suitable for nesting and the large nut of this tree is a preferred food item. These habitats were poorly covered during our fieldwork (CRT, PV). No Yellow- crested Cockatoo have been observed during recent visits to the islands of Lembata, Adonara and Solor (Trainor 2002a, c, Schellekens et al. 2011), and the species must now be close to extinction on most Nusa Tenggara islands except Sumba and Komodo, and possibly Timor-Leste (East Timor). Little Bronze Cuckoo Chrysococcyx minutillus Alor: One bird called from coastal dry forest near Kalabahi on 1 1 April 1991 (REJ). This cuckoo was recorded frequently about Tanglapui Timor area in 2002, particularly in Eucalyptus woodland above c. 1,000 m (CRT), but was unrecorded and apparently absent from dense tropical forest at Kunggwera in 2002 and 2009 (CRT, PV). Pantar: The first island records were of several birds photographed and sound-recorded in Eucalyptus alba woodland and coastal scrub in September 2009 (PV). The taxonomic relationships within this species complex remain unresolved, but most authorities prefer to re-unite Little Bronze Cuckoo with Goulds Bronze Cuckoo C. russatus (Mees 2006, Gill & Ddnsker 2012). In the Lesser Sundas this bird occurs on Flores, Timor, Wetar and several Banda Sea islands (White & Bruce 1986). Previously it was listed for Alor as Gould’s Bronze Cuckoo (Trainor 2005a). The lack of records of this bird on Alor during 2009 and 2010 (PV, CRT) was notable, but it may vocalise less duringperiods of high rainfall such as December and January. The new record on Pantar is expected, although surprisingly it is yet to be recorded for Lembata or Adonara (Trainor 2003, Schellekens et al. 2011). The host species on Flores is Golden-bellied Gerygone Gerygone sulphurea (Mees 2006), and probably sunbirds Cinnyris (Coates & Bishop 1997). It appears to be uncommon on Flores with the only recent published records by Pilgrim et al. (2000). On Alor the abundance of Little Bronze Cuckoo in high-elevation Eucalyptus woodland matched the high abundance of its likely host in that habitat, the Golden-bellied Gerygone. Sunda/Oriental Cuckoo Cuculus lepidus/optatus Alor: A single Cuculus cuckoo was observed in flight in forest above Apui-Subo on 9 September 2009 (PV), and a single bird was observed in Eucalyptus woodland south of Mainangat 1,050 m on 23 October 201 1 ( JE in litt. 2012). The distinctive calls of Sunda Cuckoo have not been heard by any recent visitors to Alor (or Pantar) although there is a specimen from Pantar (Hartert 1898) and it is common on the adjacent islands of Atauro (Trainor & Soares 2004, where mentioned as Oriental Cuckoo), Wetar (Trainor et al. 2009) and Timor (Trainor et al. 2008). It would be expected to occur on Alor. The brief observations cannot exclude Oriental Cuckoo, which is one of the commonest non-passerine migrants in Wallacea, with records from September to March (White 1976). 82 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) Asian Koef/Pacific KoeS Eudynamys scolopaceus/cyanocephala Alor: One of the most frequently heard birds in tropical forest on Mt Koya Koya in May 2002 at 600-1,200 m (CRT), heard on 31 August 2009 in highly degraded forest near Kalabahi (PV); common above Apui-Subo in September and December 2009 (PV.CRT), at and below Mainang (350-900 m) and in mangroves at the head of Muitara Bay (CRT). Pantar: Commonly heard in forest on the slopes of Mt Wasbila during September 2009 (PV), and a few birds heard calling on Mt Wasbila at 200-400 m on 12 December 2009 (CRT). A. H. Everett collected koels but reported to Hartert (1898), who identified them as Asian Koel, that ‘it [is not] easy to determine these birds from Alor’. The status of Pacific Koel E. cycanocephala in Wallacea is unclear, with White & Bruce (1986) suggesting that the Australian bird occurs mainly as a migrant with some resident populations likely. On current knowledge Pacific Koel reaches Ashmore Reef (140 km south of Roti Island) as a rare austral migrant (M. Carter in litt. 2012). Alor birds have been considered as Pacific Koel (White & Bruce 1986, Coates 8c Bishop 1997), although December records on Alor were considered by Holmes (1995) to be of resident Asian Koel. Our records suggest that a large population presumably of Asian Koel is present throughout the year, but we observed few birds and took no photos to confirm the taxonomic status of this bird. Southern (Aior) Boohook Ninox (boobook) plesseni Alor: This bird called frequendy from Eucalyptus urophylla woodland and tropical forests in gullies on Mt Koya Koya (700-1,200 m), with up to four birds calling simultaneously and sound-recorded before and at dusk; they also called throughout the night until dawn (CRT). In June 2009, up to five birds were heard to call simultaneously (in forest in the Kunggwera area), and 2-3 birds called, and were photographed, at dusk in a forested gully about 2-3 km from Lipa on 21 June (PV, VD). Pantar: Birds were photographed and sound- recorded in coastal E. alba woodland, and called frequently from mangrove forest adjacent to eucalypt woodland and from evergreen forest on Mt Wasbila up to at least 600 m (PV), and several were heard calling on Mt Wasbila at 600 m (CRT). The subspecies N. b. plesseni was described from a single female collected at c. 1,000 matTanglapoi [Tanglapui] (Stresemann 1929) and was discussed by Mees (1964) and re-described by Johnstone & Darnell (1997b), but there were no field observations of this bird between 1927 and our fieldwork. Using morphological features and colours to distinguish the many subspecies of Southern Boobook has been of relatively limited value and it is likely that several of these subspecies are specifically distinct (Johnstone & Darnell 1997). However, vocalisations are often distinctive. Birds on Aior and Pantar have the same low-pitched call which is repeated rapidly 6- 14 times over c. 1 .5 seconds and which is highly distinctive among all members of the complex, making it a prime candidate for upgrading to species level. Most members of the Southern Boobook complex have similar low-pitched calls, but vocalise with bru-book or bru-brook double notes. We intend to publish further details of the distinctiveness of this taxon elsewhere. Despite the restricted global distribution of the ‘Alor Boobook’, it appears to be under little threat of extinction because it can use both Eucalyptus woodland and tropical forest. This appears to be the only species endemic to Alor and Pantar, although this may change when the distinctiveness of several passerines recently recorded on Alor is resolved. We had no records from west and central Alor during limited (c.4 nights) nocturnal observations (PV, CRT) but presumably it is present throughout much of the island. [Pantar scops owl? Pantar: Calls presumed to belong to a scops owl Otus were heard and sound-recorded after dusk and irregularly throughout the night until at least 0 lhOO at the edge of mangrove forest some 5 km from Baranusa in September 2009 (PV). These were again heard and recorded at the same site on 3 July 201 1 (Collaerts etal. 2011) and on 22 October 2011 during a Birdtour Asia visit (JE in litt. 2012). The calls are reminiscent of those of Moluccan Scops Owl Otus magicus of Flores and Lembata (www.xeno-canto.org, Schellekens et al. 2011) but clearly differ in having a less harsh barking quality. Sound recordings were posted on the Xeno-Canto website (http:/ / www.xeno-canto.org/ asia/ discussion. php ?snd_nr= 1 7 04). When an attempt was made by PC to get closer to the source of the call, it retreated further away. Unfortunately there was not the typical Otus response to playback of flying closer to the playback source. During a visit to the Pantar mangroves in October 2011 there was again no response to playback by the bird, which suggested that the calls may not have originated from a bird (JE in litt. 2012). However, future visitors to Pantar and Alor should remain vigilant for the presence of a scops owl perhaps especially about mangrove habitat and playback of these unidentified calls should be tried to confirm the identity of the sources of the calls.] Mees' s Nightjar Caprimuigus meesi Alor: One bird was seen and sound-recorded after it called in response to playback in Eucalyptus alba woodland near Lantoka (7 km east of Tanglapui Timur) on 27 June 2009 (PV and VD). Pantar: The species was targeted with playback on Pantar because of the likelihood it was present on the island following its discovery on Alor. In open woodland at 100 m about 3 km east of Puntaru on the road to Sirung volcano on 4 September 2009 a nightjar responded strongly to a recording of Mees’s Nightjar, by immediately flying towards the recording and landing on the ground less than 1 m from a motorbike. After it flew off a recording of Savanna Nightjar C. afftnis was played, but the bird did not respond (PV). The bird was not photographed or observed long enough to confirm identification. Playback was used twice, with the same result: no reaction to playback of Savanna Nightjar calls, but immediate reaction to a recording of Mees’s Nightjar recording. The lack of a vocal response makes confirmation impossible, but its behaviour indicates that it was likely to be Mees’s Nightjar rather than Savanna Nightjar. The recently described Mees’s Nightjar (Sangster & Rozendaal 2004) was previously known from Flores and Sumba. These two new island records are an important range extension, and presumably it also occurs on the intervening islands of Adonara, Lembata and possibly Solor, although little nocturnal work has been conducted on these islands (Schellekens etal. 2011). Stork-billed Kingfisher Pelargopsis capensis Pantar: One bird was observed in mangroves near Desa Batu on 19 April 1991 (REJ). The Stork-billed Kingfisher is known from Lombok, Sumbawa and Flores in the Lesser Sundas (Coates & Bishop 1997), with several recent records from mangrove and estuary habitat on Lembata (Trainor 2003). This bird is generally uncommon to rare in Nusa Tenggara, but there are regular records on Flores from mangroves and Lake Rana Mese at 1,200 m (Pilgrim etal. 2000) and occasional records on Sumbawa from the coast to hills up to 870 m (Butchart etal. 1994, Johnstone etal. 1996). Oriental Dwarf Kingfisher Ceyx erythraca ssp. Pantar: Three pairs of Oriental Dwarf Kingfisher were seen about mangroves, coconut and shrub, 3 km north of Baranusa on 22 October 2011, including a pair apparently using a nest-hole on a steep slope (JE in. litt. 2012). This bird occurs widely in South and South-East Asia, and in Nusa Tenggara it occurs on Lombok, Sumbawa, Flores and Sumba (White & Bruce 1986). On Alor, none was noted during two visits (7 December 2009 and 10 January 2010) to extensive mangroves at the head of Mutiara (Kalabahi) Bay (CRT). Recently, the Oriental Dwarf Kingfisher was found to Forktail 28 (2012) The avifauna of Alor and Pantar, Lesser Sundas, Indonesia 83 be locally common in evergreen swamp forest on neighbouring Lembata Island (Schellekens et al. 201 1). There are relatively few Nusa Tenggara records but it is a small and inconspicuous bird (with a high-pitched and near-inaudible call) and is likely to be regularly overlooked. It has been considered ‘rare’ on Flores (Butchart et al. 1994, Coates & Bishop 1997), there are no recent Lombok records (Myers & Bishop 2005), but there have been records from riverine forest on Sumbawa up to 650 m (Johnstone et al. 1996). Sunda Pygmy Woodpecker Dendrocopus moluccensis grandis Alor: One adult male (WAM A2451 1, length 160 mm, weight 22 g) was collected at Apui-Subo on 26 April 1991 (REJ). This bird was recorded once or twice per day in tropical forest and Eucalyptus woodland, usually as single birds, especially along gullies in the Tanglapui Timur area (600-1,200 m) (CRT, PV), above Apui-Subo (PV, CRT) and Mainang (CRT). Pantar: The only record was of two birds on 4 September 2009 in Eucalyptus urophylla woodland on the slopes of Mt Sirung at about 500 m (PV). In Wallacea the Sunda Pygmy Woodpecker occurs from Lombok through to Alor (White & Bruce 1986) and would have been expected to occur on Pantar. It seems to prefer tropical forest on Alor, but also occurs in woodland and plantations when in close proximity to tropical forest. Specimens on Alor average larger than birds on Lombok, Sumbawa and Flores, and were originally described as an endemic subspecies D. m. excelsior (Hartert 1898), but differences were not considered significant by White & Bruce (1986) and this subspecies has not been recognised recently by authorities. Elegant Pitta Pitta elegans concinna Alor: Moderately common, calling with a single bird observed in forest at Apui (650-740 m) and in tropical dry forest at Mali in April 1991 (REJ), with three specimens collected (WAM A24458, A24544 and A24488). Pittas called occasionally at dawn and dusk from tropical forest at Kunggwera and the slopes of Mt Koya Koya (600-950 m) during May 2002 (CRT), and called there frequently in June 2009 (PV). At least three birds called at dusk in degraded forest 15 km east of Kalabahi on 30 June 2009 (PV); two birds called from forested gullies at Mainang on 9 January 20 10 (750 m), and up to four birds called along the road to Wah Wah (4 km north-east of Kalabahi at c.400 m) on 20 January 2010 (CRT). Pantar: One bird was recorded from dense Acacia and shrubs at the edge of mangroves on 19 April 1991 and one bird called from dense stands of the weed Wedelia biflora near Ivabir on 20 April 1991 (REJ). A total of 5-7 birds called from tropical forest on Mt Wasbila during 3 September 2009 (PV) and several birds were sound-recorded in a dry thicket at 120 m and heard at 650- 800 m there on 10 December 2009 (CRT). Elegant Pitta occurs widely in the Lesser Sundas with four subspecies present (White & Bruce 1986). The subspecies P. e. concinna of Lombok-Alor appears to be resident with records throughout the year, but on the drier islands of Roti and Timor the subspecies P. e. elegans (which has a three-note call) is migratory (White & Bruce 1986, Trainor 2005b, Tebb et al. 2008, PV unpubl. data). Surprisingly, there are no published records of Elegant Pitta from Timor-Leste (East Timor), Wetar (Trainor et al. 2009) or Atauro (Trainor & Soares 2004). Recordings of the two-note calls of P. e. concinna from Alor, Pantar and Lembata have been uploaded to the www.xeno-canto website. Wallacean Cuckooshrike Cottacina petrsonata alfrediana Alor: Moderately commonly observed in ones, twos and threes in Eucalyptus woodland at Apui in April 1991 (REJ). Regularly observed in small groups of 3-5 birds in Eucalyptus woodland on Mt Koya Koya (600-1,100 m) and frequently observed, mostly as singles, in Eucalyptus woodland and tropical forest at Kunggwera, above Apui-Subo (PV, CRT) and Mainang (CRT). Pantar: Several pairs were observed in dry tropical forest at the foot of Mt Wasbila at 250 m (PV); several birds were observed in tropical forest and Eucalyptus woodland at 300-600 m (CRT) and one pair was observed in open eucalypt woodland en route to Mt Sirung (PV). Wallacean Cuckooshrike occurs widely in the Lesser Sundas and the Kai Islands (Maluku), with seven subspecies described (White & Bruce 1986). On Alor, C. p. alfredianus was described from six specimens which differed from C. p. floris (Sumbawa-Flores) by being a ‘distinctly paler grey above and below, and the males do not have a black throat and the sides of the head’ (Hartert 1898). A specimen from Lembata was ‘a shade darker, thus pointing towards [...]_ floris’ (Hartert 1898). The presence of Wallacean Cuckooshrike on Pantar is expected: presumably this bird is included with C. p. alfrediana. Based on distribution, the Lembata population is as likely to be C. p. floris as C. p. alfrediana , but plumage characteristics were not noted during recent field surveys (Trainor 2003). Wallacean Cuckooshrike is part of a complex which also includes Black-faced Cuckooshrike C. novaehollandiae. Some taxa within this complex have already been split (e.g. Buru Cuckooshrike C. fortis ) and Mason & McKean (1982) argued for species status for the Timor group subspecies C. p. per sonata but without considering other forms (Mees 2006). A thorough review of this group is needed. There are substantial differences in vocalisations among populations (e.g. Figure 2a, b), and these may help to set taxonomic limits. Figure 2. Song of the Wallacean Cuckooshrike from Apui-Subo, Alor, 5 December 2009 (a) and Hatu Builico, East Timor, 28 April 2010 (b) (both recordings by C. R. Trainor). 84 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) Common Cicadabird Coracina tenuirostris s$p. Alor: A single female was initially heard calling (slow took notes) and then seen in the subcanopy of a candlenut Aleurites moluccana plantation above Apui-Subo village at 950 m on 4 December 2009. The bird had obvious barring on the neck, breast and belly, but no other notes were taken before it flew off (CRT). A single immature male bird was collected on Lembata (Hartert 1896), but there have been no subsequent records on that island (Trainor 2002c, 2003, Schellekens etal. 2011). On nearby Atauro Island a single male bird observed gave ruk ruk notes (Trainor & Soares 2004), and females on Timor also give reasonably similar contact notes (http:// www.xeno-canto.org/32585). The taxonomic affinities of birds on Lembata and Alor are unclear, but on biogeographic grounds (and probably vocalisations) they may be close to C. t. timoriensis. Common Cicadabird is an inconspicuous bird that is generally uncommon throughout its range. Ir is most easily detected by its distinctive vocalisations. There are at least 23 described subspecies of Common Cicadabird (Gil & Donsker 2012) including many distinctive forms (Rheindt et al. 2010). A review of these taxa is needed. Lesser Shortwing Brachypteryx leucophrys Alor: Common in tropical forest along creeks near Apui (650- 800 m) in April 1991 with four specimens collected (REJ). The song of this bird was frequently heard on the slopes of Mt Koya Koya (PV and VD), above Apui-Subo (PV, CRT), particularly in dense vegetation and along streams and gullies. At Mainang, and along the 17 km walk to Apui-Subo, this bird was restricted to watered gullies above about 850 m (CRT). In the village ofTanglapui Timur it occurred in a gully, and was recorded down to 680 m about 2 km west of the village (PV, CRT). This bird was noted also by Holmes (1995) on 10 December 1994 on the road between Mainang and Apui-Subo. Records of this bird given by Trainor (2005a) were confused with Timor Stubtail Urosphena subulata (see below), including the description of a call given which clearly refers to the latter bird. Examination of recordings in 2002 have since confirmed that Lesser Shortwing songs and high-pitched single notes were sound- recorded. Field notes show that three Lesser Shortwings were observed in watered gullies, but perhaps these were also misidentified Timor Stubtail. This bird is patchily common throughout much of Alor above 600-700 m, but is absent from the extensi Eucalyptus woodlands. None was heard on the upper slopes of Mt Wasbila on Pantar (PV, CRT). At least to the human ear, the song of birds on Timor and Alor bird sound similar to birds on Java (www.xeno- canto.com). Mimicry of the song of Timor Bush Warbler by Lesser Shortwing was sound-recorded (www.xeno-canto.com/xcl03157). The subspecies is presumably B. 1. leucophrys. Timor Bush Warbler Loeustella timorensis ssp. Alor: First discovered at Apui-Subo on 9 September 2009, when up to five singing males were heard calling from tall grass and thickets below secondary tropical forest at 1,050 m (PV). Subsequent visits to the same location noted at least 6-8 singing males along a ridge at 900-1,250 m, mostly calling from tall grassland beneath Eucalyptus urophylla woodland (CRT). A second population was located at Mainang with seven males heard singing in grassland below E. alba at 859-930 m. A total of 1 1 birds were heard, with some birds photographed and videographed near Mainang at 928- 1,070 m on 23 October 2011 ( JE and RH in litt. 2012). Details of this discovery were recently published (Trainor et al. 2012) and videos and photos are available online (http://ibc.lynxeds.com/). Timor Bush Warbler is clearly patchily distributed on the island. For example, none was heard singing in the vicinity of Mt Koya Koya in 2002 (CRT) or 2009 (PV) or 2010 (CRT) despite the use of playback (in 2010) in apparently suitable habitat, and none was heard during a 17 km walk between Mainang and Apui-Subo through seemingly suitable Eucalyptus woodland (CRT), indicating that they are local and have specialised habitat requirements. Differences in song characteristics (frequency) suggest that the Alor birds may be subspecifically distinct from the Timor population (Trainor et al. 2012). None was recorded after using playback in apparently suitable habitat on Mt Wasbila at 800 m, but birds may be present elsewhere on Pantar. Chestnut-backed Thrush Geokichla dohertyi Alor: Birds were observed and heard occasionally in forested ravines on the slopes of Mt Koya Koya (CRT ) and one was seen and several heard singing in a forested gully at Kunggwera in June 2009 (PV). This bird was occasionally heard and recorded singing from secondary forest above Apui-Subo village in December and January, with one bird killed with a catapult by a local villager on 4 December 2009 (CRT). Chestnut-backed Thrush occurs on the large Nusa Tenggara islands of Lombok, Sumbawa, Flores, Surnba and Timor, mostly in hill or montane forest above 500 m (White & Bruce 1986). A ‘ Zoothera' thrush was noted by A. H. Everett on Alor but no specimens were collected (Hartert 1898). Its presence on Alor is unsurprising, particularly given the extent of forested montane habitat; by contrast, the absence of records from Pantar was noteworthy (PV, CRT). Chestnut-backed Thrush is an inconspicuous forest-dweller and is easy to overlook if not vocalising. There have been no records from Lembata and Adonara (Trainor 2002a, c, 2004, Schellekens etal. 201 1), Atauro (Trainor & Soares 2004) or nearby Wetar Island (CRT unpubl. data) but more effort is needed in the hills and mountains of these islands. Kamchatka Leaf Warbler Phylloscopus examincmdus Alor: Birds were occasionally observed on Ternate Island on El April 2002, and on Sika Island on 29 April 2002; one bird was observed in a gully in Lipa village at 700 m on 2 May 2002, and six were heard or seen 2-4 km east of Lipa, at 440-620 m, on 1 7 January 2010 (CRT). Pantar: A single bird was heard in coastal vegetation at Puntaru on 14 December 2009 (CRT). The Kamchatka Leaf Warbler was recently split from Arctic Warbler (Alstrom et al. 2011). This bird breeds in south Kamchatka, Sakhalin and north¬ east Hokkaido (Alstrom etal. 2011) and winters throughout South- East Asia with Alor at about the southern limit of its wintering range. The ‘Arctic Warbler’ subspecies examinandus was described from material collected on Flores (Mees 2006), and birds were collected on Alor in April by A. H. Everett, but no details of subspecific identity were noted (Hartert 1898). The 'Arctic Warbler’ was the second most commonly collected passerine migrant to Wallacea (White 1977). There are regular records of Kamchatka/Arctic Warbler on Adonara (Trainor 2002a) and Lembata (Trainor 2002c, Schellekens et al. 2011), but it is very uncommon on Timor (R. Noske in litt. 2012) with no East Timor records (CRT unpubl. data). Sumda Bush Warbler Horornis vulcanius kolichisi Alor: Noted as moderately common in weedy Wedelia biflora shrubs and grass at c.700 m above Apui, with eight specimens collected including the holotype on 24 April 1991 (REJ, Johnstone & Darnell 1997a). In 2002, found to be frequent on the slopes of Mt Koya Koya and Lipa village in weedy tall grassland gullies and stream channels (CRT), and in 2009-2010 it was common in the same habitat at Mainang (700-900 m, CRT), Apui-Subo (800- 1,200 m) and Tanglapui Timur area at 650-1,150 m (PV and VD, CRT). The Sunda Bush Warbler is a poorly known bird which occurs on Sumatra, Borneo, Java and Bali, with Nusa Tenggara populations on the mountain areas of Lombok, Sumbawa and Timor (Coates & Bishop 1997). Like many bush warblers its Forktail 28 (2012) The avifauna of Alor and Pantar, Lesser Sundas, Indonesia 85 plumage is conservative and nondescript (Kennerley & Pearson 2010) but vocalisations are often distinctive. There are recent new observational records for Sumbawa (Johnstone et al. 1996) and Wetar (Trainor et al. 2009). The Alor subspecies H. v. kolichisi differs from both H. v. vulcanius (of Bali, Lombok) and H. v. everetti (of Timor and possibly Wetar) by its smaller size, duller dark brown upperparts lacking the rusty tones of the nominotypical form and the olive tones of H. v. everetti (Johnstone & Darnell 1997a). The bill is also proportionately longer and finer with a more extensive dark zone on the lower mandible (Johnstone & Darnell 1997a). The songs ofNusaTenggara birds are relatively distinctive from birds of the Greater Sundas (PV unpubl. data) and work analysing calls has begun (G. Sangster/« litt. 2011). The Sumbawa bird is presumably similar to those of Lombok (Johnstone & Darnell 1997a) and the song of the Wetar bird sounds close to birds on Timor (CRT, PV unpubl. data). Birds on Timor and Wetar occur from sea-level to the mountains (Noske & Saleh 1996, Trainor et al. 2009) but the Alor bird appears to occur in the hills only (but more lowland surveys are needed on Alor). The Sunda Bush Warbler appears to be absent from Pantar. Timor Stuhtail Urosphena subulata ssp. Alor: This bird was not identified during the 2002 survey, but its high-pitched note was regularly sound-recorded from the slopes of Mt Koya Koya, and these were later (following publication of Trainor 2005a) identified as Timor Stubtail (CRT). Subsequently this diminutive bird has been found to be relatively common and vocal, especially in gullies at Kunggwera (PV and VD), Tanglapui Timur (PV, CRT), above Apui-Subo (PV, CRT), Mainang (CRT) and Wahwah (CRT) at elevations of 660- 1,200 m. During a 17 km walk between Mainang and Apui-Subo it was recorded only in a single well-vegetated gully near Baifui village, and was absent from Eucalyptus alba woodland. This is a significant and unexpected range extension for this essentially ‘Timor-group’ bird. Only recently has the range of this species become better known, with new island records for Wetar (Coates & Bishop 1997), Atauro (Trainor & Soares 2004), Roti (Trainor 2005b) and Romang (CRT unpubl. data). It is generally common in tropical forest throughout its range, although suitable habitat appears to be limited on Roti and Atauro. The closest relatives of the Timor Stubtail are restricted to the mountains of north-east Borneo (Bornean Stubtail U. wbiteheadi ), while Asian Stubtail U. squameiceps is also confined to mountains when wintering in South-East Asia (Kennerley & Pearson 2010). On Timor this bird occurs from sea-level to the mountains, but there is limited data on the elevational limits of this bird on Alor. It may be restricted to the hills, but there has been limited lowland survey effort on Alor. The song has a uniform structure on all islands — a monotone whistled pseeee (Kennerley & Pearson 2010) which varies substantially in length, and less so in frequency. The relationships of these island forms are still unclear, but they are probably distinctive at the subspecific level. Little Pied Flycatcher Ficedula westermanni mayrii Alor: One bird was observed in the forest canopy at Apui on 27 April 1991 (REJ). Noted as occasional in Casuarina junghuhniana and tropical forest-lined gullies, and E. alba woodland (650- 1,120 m) around Tanglapui Timur in 2002 (CRT). A pair was seen in a gully at Tanglapui Timor in 2010 (CRT), and birds were frequent in secondary forest above Apui-Subo in 2009 and 2010 (PV,CRT), but the species was recorded only once on the walk from Mainang to Apui-Subo (CRT ) . One male bird was observed singing c.10 km north of Apui at 230 m (PV). Little Pied Flycatcher is typically a montane bird (above about 800 m) throughout its broad South-East Asian range, but on Timor (Noske & Saleh 1996), Atauro (Trainor & Soares 2004), Wetar (Trainor et al. 2009) and now Alor it occurs from the lowlands to the mountains. This unusual aspect of its habitat use in the Lesser Sundas has been widely overlooked (White & Bruce 1986, Coates & Bishop 1997, Mees 2006). The Little Pied Flycatcher is morphologically conservative with relatively minor differences in plumage throughout its range. Analysis of songs might help to disentangle cryptic diversity within this complex. Three specimens (all females) were collected on Alor by von Plessen (White & Bruce 1986) presumably at an elevation of about 1,200 m (which was given as the Alor elevation range for this bird: Coates & Bishop 1997). YeSlow-breasted Warbler Seircercus montis ssp. Alor: Th is bird was local in well-forested gullies at 800-930 m on the slopes of Mt Koya Koya, with only a few birds seen and recorded during April 2002 (CRT). Several birds were photographed and recorded in dense forest in the hills above Apui at c. 1 ,000-1 ,100m on 9 September, 2009 (PV), but none was recorded there in December or January (CRT). In Nusa Tenggara this bird occurs on Flores (subspecies floris-. compared to S. m. montis of Borneo it has less well developed dark lateral crown-bands to nape) and Timor ( paulinae : compared to S. m. montis it has a brighter rufous crown, brighter olive back, yellower rump, and less developed crown-bands) in Wallacea (White & Bruce 1986). Photos of the Alor bird indicate that it is probably closer to S. m. paulinae with a bright orange crown which appears to lack lateral crown-bands entirely. Although the subspecies described for Nusa Tenggara have been considered to have diverged little from S. m. montis of northern Borneo (White & Bruce 1986) morphological differences shown by photos on www.orientalbirdimages.org, and differences in vocalisations (www.xeno-canto.org) suggest that species-level splits should be expected in these taxa. Birds on Timor and Alor have a song consisting ofa ‘high pitched piercing note’ (Coates & Bishop 1997) similar to the song of birds in the Greater Sundas. This is similar to songs of some Dicaeum flowerpeckers. However, contact notes regularly given on Timor and Alor (a buzzing bresh note given singly, doubly or often repeated) are unlike calls of birds from Borneo and mainland Asian populations archived at http:// www.xeno-canto.org or http://macaulaylibrary.org/. Asian Paradise-flycatcher Tersiphone paradisi floris Alor: There were three observations of this bird in gallery forest at 580-820 m on the slopes of Mt Koya Koya (CRT), and birds were observed daily and considered as frequent in tropical forest at the same sites in 2009 (PV), and in forest above Apui-Subo (PV). Pantar: One bird was observed in vine forest at Desa Batu on 17 April 1991 (REJ). Several birds were observed and sound-recorded in lowland tropical forest and coastal woodland (PV) and one male was observed and sound-recorded at 630 m on Mt Wasbila on 10 December 2009 (CRT). Alor delimits the south-eastern limits of this widespread Asian bird. The subspecies T. p. floris is known from Sumbawa, Flores, Besar, Lembata and Alor, with our Pantar records filling an important distributional gap. Asian Paradise-flycatcher is typically one of the least common resident birds through its Nusa Tenggara range (Coates & Bishop 1997, Mees 2006), although can be locally common on Flores (Butchart et al. 1994, Pilgrim et al. 2000). Females appear to be particularly inconspicuous, and we only observed males on Alor and Pantar, matching historical collections of males from Alor and Lembata (Hartert 1896). All males in Nusa Tenggara are predominantly white above, whereas a high proportion of males in South and South-East Asia are rufous (Owen 1964). Broad-billed Flycatcher Myiagra ruficoilis ruficollis Alor: Singles or pairs were recorded four times in E. alba woodland usually next to streams or a dam at 625-750 m about Tanglapui 86 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) Timur (CRT), and occasional in mangroves near Kalabahi (CRT), degraded lorest near Kalabahi (PV) and coastal strand forest on Sika Island (CRT). Pantar: Regularly recorded in mangroves and E. alba woodland in September 2009 (PV), and once heard from secondary shrubland along the road to the Sirung volcano (CRT). Broad-billed Flycatcher was known from Lembata and Alor in the Flores island chain, with the Pantar record filling a gap in its distribution. In the Lesser Sundas it occurs on Timor and most associated islands, as well as Sumba (Coates & Bishop 1997). It appears to be generally uncommon to rare on Alor except locally near water and in mangroves, and on Pantar it appears to be common in mangroves as on neighbouring Lembata (Trainor 2003) and Timor (CRT unpubl. data). Common Golden Whistler Paeycephala pectoraiis fulvotincta Alor: This bird was common in tropical dry forest, lantana thickets and prickly shrubs at Kalabahi and Mali and in secondary forest and plantations at Apui in April 1991, with eight specimens collected (REJ). It was observed on several occasions in secondary forest above Apui-Subo and in evergreen forest at Kunggwera (PV), and was heard regularly from secondary forest and plantations at 400-900 m near Mainang, and in secondary forest above Apui- Subo to 1,250 m (CRT). Pantar: Moderately common in vine forest at Batu and in mangrove at Kabir in April 1991 with two specimens collected (REJ). Birds were photographed and sound- recorded in coastal woodland (PV), degraded forest patches in the lowlands (<200 m) as well as on the slopes of Mt Wasbila and frequently heard in tropical forest on Mt Wasbila from 100-800 m and coastal shrubland about Puntaru (CRT). The Common Golden/Mangrove Whistler P. pectoraiis/ mela.nu.ra complex, with 66 named populations, is one of the most complex avian examples of geographic variation (Jonsson et al. 2008). The IOC currently treats Alor and Pantar populations as part of a species called Rusty-breasted Whistler P. fulvotincta (of Java, Lesser Sundas, Flores Sea islands: Gill & Donsker 2012) but we prefer to wait until genetic and vocal analyses are published on the complex to support this split. Common Golden Whistler is typically one of the most vocal birds in secondary and primary forest in the Lesser Sundas (Coates & Bishop 1997), although it is inconspicuous and difficult to observe without the use of playback. It appears to be mostly absent from Eucalyptus woodland on Alor and Pantar, with a clear preference for tropical forest. For example, it was only heard once while walking from Lipa to Lantoka village (7 km), and heard once on the walk from Lipa to Naumang (CRT). Cinereous Tit Parus cinereus Alor: This bird was uncommon to moderately common in gardens at' Apui in April 1991, when two specimens were collected (REJ). In 2002, 2009 and 20 1 0 it was occasional to frequent at all sites, in all wooded habitats, from sea-level to 1,250 m (PV, CRT). Pantar: Observed in small numbers in tropical dry forest at 200 m (PV) and common in coastal shrubland about Puntaru village on 14 December 2009, but unrecorded from Mt Wasbila (CRT). Alor delimits the easternmost occurrence of this bird, which ranges from Afghanistan, India, mainland South-East Asia, Greater Sundas and the Lesser Sundas from Lombok along the Flores island chain to Sumba (White & Bruce 1986). Typically this bird is considered a common species along the Flores island chain (Butchart et al. 1 994, Johnstone et al. 1996, Pilgrim et al. 2000, Trainor 2002a, c, Myers <3c Bishop 2005). Thick-billed Ftowerpecker Dieaeum agile Pantar: One bird was collected at Batu (WAM A24474) and several were observed in canopy of woodland on 19 April 1991 (REJ). Thick-billed Flowerpecker is typically uncommon to rare (and inconspicuous with low detectability) throughout its Lesser Sunda range (Lombok, Sumbawa, Flores, Lembata, Alor and Timor) (White & Bruce 1986, Johnstone et al. 1996, Coates & Bishop 1997), with only four specimens known for Flores (Mees 2006). Although collected on Alor by A. FT. Everett (originally thought to have been a specimen of Golden-rumped Flowerpecker D. annae ), there have been no subsequent records on the island. This is the first Pantar island record. Consistent with its general rarity is the single observation of one bird during a recent seven- week survey of Wetar (Trainor et al. 2009). This is an aberrant flowerpecker with recent genetic analyses failing to resolve its generic affinities, although it may actually be a Prionochilus (Nyari etal. 2009). Mountain White-eye Zosterops montanus Alor: Common at c. 700-1, 271 m, particularly in Eucalyptus woodland at Mainang, Apui-Subo and Tanglapui Timur (CRT). Both Mountain White-eye and Ashy-bellied White-eye Z. citrinellus were present in Tanglapui Timur village, but only the former was heard during a 9 km walk from Lipa to Lantoka at 700-750 m. At Mainang this bird mostly occurred in Eucalyptus woodland above 1,150 m, while Ashy-bellied White-eye was more frequent in secondary tropical forest below c. 1,1 50 m. During 2002 only Ashy-bellied White-eye was discriminated (Trainor 2005a), with Mountain White-eye obviously overlooked. Mountain White-eye was known from the large Nusa Tenggara islands of Lombok, Sumbawa, Flores and Timor (Coates & Bishop 1997) but has recently been recorded on neighbouring Atauro Island above 800 m (CRT unpubl. data), and in the mountains ofWetar (CRT unpubl. data). Elevation use by Mountain and Ashy-bellied White-eye remains poorly known on Alor, but the latter was widespread on Pantar from sea-level to c.400 m (PV) and at Batu and Kabir (REJ), and was recorded down to sea-level on Alor including the islets of Kepa (REJ, CRT, PV), Ternate and Sika (CRT). About 20 specimens of Ashy-bellied White-eye were collected at Mali and Apui in 1991 (REJ). Mountain White-eye would be expected to occur on Pantar but none was definitively heard or seen in apparently suitable habitat on Mt Wasbila up to 850 m (CRT). Cryptic taxa are expected to occur within this complex but their vocalisations are unlikely to help unravel the affinities of various island forms (Rheindt & Hutchinson 2007). Photos of Alor birds show they are typical of.Z. m. montanus (as described in Coates & Bishop 1997) with forehead greenish to yellowish, abdomen whitish, without a yellow median stripe, and the undertail-coverts are yellow. Lemon-bellied White-eye Zosterops ehloris/ Oriental White-eye Z. palpebrosus Pantar: Lemon-bellied White-eye was observed and sound- recorded in coastal scrub and secondary forest at 400 m on Mt Wasbila (PV). In Nusa Tenggara this bird occurs on Lombok, Sumbawa, Komodo, Rinca, Flores, Besar and Paloe mostly in the lowlands (White & Bruce 1986). The identity of white-eyes occurring on islands east of Flores remains poorly known. Records of Oriental White-eye on Adonara (Trainor 2002a) were queried by Mees (2006), who considered that Lemon-bellied was more likely to occur on the island. We were unable to confirm Oriental White-eye on Pantar, but it may also be present. Short-tailed Starting Aplonis minor Alor: Locally common in pairs and small flocks of up to 10 birds in gallery forest and fruiting trees at Kunggwera and the slopes of Mt Koya Koya in 2002 (570-1,1 50 m, CRT), with flocks of eight and 12 birds seen in the same area in June 2009 (PV). An immature bird with white below and black streaking on the chest was observed Forktail 28 (2012) The avifauna of Alor and Pantar, Lesser Sundas, Indonesia 87 near Tanglapui Timur on 3 May 2002 (CRT). This bird was uncommon above Apui-Subo, with several voice records and flocks of up to six birds seen (800-1,100 m), but was frequently observed at Mainang in flocks of 2-8 birds (CRT). Pantar: The first island records of Short-tailed Starling were of four and six birds observed in flight near Sirung volcano (PV), and a single voice record from evergreen forest on Mt Wasbila at 750 m (CRT). Short-tailed Starling occurs widely as a typically uncommon forest bird in the Lesser Sundas (Lombok, Suntbawa, Flores, Paloe, Sumba, Timor, Wetar, Mo.a, Romang: Coates & Bishop 1997), and was recently confirmed for Atauro Island (Trainor & Soares 2004). The presence of this bird on Alor and Pantar would be expected, although it has not yet been recorded from Lembata or Adonara (Trainor 2002a,c, 2003, Schellekens et al. 2011). Crimson-hooded (Alor) Myzomela Myzomela ( kuehni ) Alor: A single record of one bird feeding at the flowers of Eucalyptus alba, at 900 m, on the slopes of Mt Koya Koya in April 2002 (CRT), which was thought to be a misidentification at the time, with a brief notes crossed out in a notebook (CRT). In 2009 and 2010 there were occasional records of 1-4 birds feeding at flowers or roosting in E. urophylla. Acacia, Casuarina junghuhniana and the montane shrub Photinia integrifolia above Apui-Subo village at 1,150-1,250 m (CRT, PV). Photographs show that the Alor Myzomela is almost identical in appearance to the nominotypical Wetar bird, except for the extent of red on the head, neck and chest. The red hood on the Alor bird is much reduced and only reaches midway on the hindcrown before sharply changing to grey on lower hindcrown (nape), mantle, back and scapulars, whereas the red on the Wetar bird reaches the back of head (c. 1 cm further than the Alor bird). Alor birds have a red throat, but unlike Wetar birds this does not extend down to the chest. Substantial differences in contact notes and song of Alor birds compared to Wetar suggest that the Alor birds have diverged significantly and represent an undescribed taxon at either subspecies or species level. Alor birds appear to be restricted to ‘upper’ montane habitat, although it was apparently absent from forest at 850-1000 m above Apui where a suite of other montane species were present, and was also unrecorded from extensive £. alba woodlands at 700-1,000 m; by contrast, Wetar birds are common from sea-level to the mountains (CRT unpubl. data). None was heard or seen on Pantar, but no habitat above 900 m was surveyed. It is intended to publish further details on this bird elsewhere. Common Hill Myrsa Gracula religiosa venenata Alor: In April 1991 this bird was moderately common in ones and twos in forest areas around Kalabahi and Mali and in forests at Apui (REJ). It was locally common about Kunggwera village with 15 birds observed in a large fruiting fig tree on 3 May 2002, and pairs and small parties were observed in the surrounding tropical forest (CRT). One flock of six birds was observed in flight at Kunggwera in June 2009 (PV and VD). One bird was heard calling from secondary tropical forest about 2 km north of Mainang on 9 January 2010, and two birds were seen and photographed in the same area on 13 January 2010 (CRT). Pantar: Three specimens were collected from Batu village and one was collected from forest at Kabir in April 1991 (REJ). One bird was sound-recorded on the slopes of Mt Wasbila in September 2009 (PV), and four were photographed on the slopes of Mt Wasbila and 10-11 (some possibly double-counted) were heard on 10 December 2009 (CRT). In Nusa Tenggara the Common Hill Myna occurs from Sumbawa to Alor (White & Bruce 1986). It is uncommon on Sumbawa, but moderately common on Moyo Island (Johnstone et al. 1996), locally common on Flores (Butchart et al. 1994, Pilgrim et al. 2000) and uncommon to rare on Adonara and Lembata (Trainor 2003, Schellekens et al. 2011). Populations have undoubtedly declined precipitously in Nusa Tenggara because of capture for trade during the 1980s and 1 990s, but the residual birds on the islands are still being caught. At a market in Kalabahi during May 2002 nine chicks were available for sale, including two not fully feathered, three fully feathered (selling for Rp 300,000 [c. $US30] each) and four fledged subadults (selling for Rp 250,000 each). On Pantar in 2009, local people said that Common Hill Myna was valued at Rp 200,000- 1 ,500,000 (PV). A total of three individuals including one bird at Tanglapui (Alor) were recorded during an undated survey by Widodo (2010). Despite ongoing trade Alor may still retain some of the best populations of this bird in Nusa Tenggara (Trainor 2005a). Wallacean Drang© Biarurus densus bimaensis Alor: Moderately common in pairs at Kalabahi (one specimen collected), Mali and Apui mainly in forest but also in plantations during April 1991 (REJ). One of the most vocal and frequently observed birds in tropical forest at Kunggwera and Mt Koya Koya at 570-1,250 m (CRT, PV); above Apui-Subo (900-1,250 m) (PV, CRT) at Mainang (300-900 m); present in mangroves at the head of Mutiara Bay, but less common in Eucalyptus woodland (CRT). Pantar: One specimen was collected at Batu village on 17 April 1991 (REJ). Several observations on the slopes of Mt Wasbila at 400 m (PV) and frequently recorded there to 800 m (CRT), and in secondary forest and agricultural land about Puntaru village (CRT). The Wallacean Drongo is typically one of the most vocal and conspicuous birds in forest and secondary habitats in Nusa Tenggara (Coates & Bishop 1997). However, it is worth highlighting that there are substantial morphological and vocal differences between the various Lesser Sunda subspecies, with this subspecies informally named the ‘Lesser Wallacean Drongo’ D. bimaensis and the bird on Timor and associated islands dubbed the ‘Greater Wallacean Drongo’ D. densus (Eaton 2009): The subspecies D. d. bimaensis is moderately common to abundant on Sumbawa (Johnstone et al. 1996), Flores (Butchart et al. 1994, Pilgrim et al. 2000), Adonara (Trainor 2002a) and Lembata (Trainor 2002c), and was recently noted on Solor (Schellekens et al. 2011). It is under no threat of extinction. Eurasian Tree Sparrow Passer montanus Alor: This introduced bird was very common in the large town of Kalabahi during 2009 and 2010 (PV, CRT). Pantar: During September 2009, it was observed and sound-recorded at Baranusa (PV). At least three birds were noted at Puntaru village at the church (CRT). This bird has become widely naturalised throughout Nusa Tenggara over the last 30 years. None was noted on Alor during 1991 (REJ) or 2002 (CRT) but birds are thought to have been present on Flores since about 1955 (Mees 2006). They were locally abundant on Adonara during December 2000 (T rainor 2002a), absent from Lembata in December 2000 (Trainor 2002c, 2003) and abundant there in 2009 (Schellekens et al. 2011). DISCUSSION Our records substantially improve knowledge of the avifaunas of both Alor and Pantar. We expected to add many new island records to the Pantar Island list because it was poorly surveyed historically, with little information published on the island since the nineteenth century, but the addition of 37 (including 27 resident terrestrial) bird species to the Alor list was surprising. The continued accumulation of new records for both islands by ornithologists in 2011 suggests that more species will be added in the future. We also show that Alor and Pantar have greater levels of endemicity than previously known, and the presence of the Critically Endangered 88 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) Flores Hawk Eagle and additional endemic and restricted-range birds elevates the global conservation significance of these islands. Both islands were assumed to be dominated by a ‘Flores ornis’ (Hartert 1898) or possess avifaunas ‘evidently derived from Flores’ (White & Bruce 1986), but our fieldwork shows that the avifaunal origins of these islands is more complex. Alor demarcates the eastern limits of a suite of well-known and ‘typical’ Asian bird species (e.g. Ruddy Cuckoo Dove, Rusty-breasted Cuckoo Cacomantis sepulcralis, Sunda Pygmy Woodpecker, Golden-bellied Gerygone, Cinereous Tit and Asian Paradise-flycatcher), but many species typical of the Timor-group islands were also added to the list (e.g. ‘Little Cuckoo Dove’, Common Cicadabird, Timor Bush Warbler and Timor Stubtail) and other widespread Asian birds which appear to be most closely related to Timor birds (Yellow-breasted Warbler, Little Pied Llycatcher and Lesser Shortwing). The discovery of Timor Bush Warbler on Alor is surprising on biogeographical grounds. Some variation in song characteristics of this species have been published (Trainor et al. 2012) but further advances in determining the affinities of this bird in relation to other members of the complex, including those from Timor, Bali and Java, will rely on the collection of specimens and molecular analyses. Timor Bush Warbler is currently considered Near Threatened, but the Alor population appears under little threat. Only two populations were discovered, highlighting that this bird is patchily distributed in suitable montane habitat. The absence of records during a 17 km walk between the two populations also highlights the high degree of habitat specificity of the species (although significantly this pattern was mirrored by a suite of other passerines that are forest-dependent). Although it requires tall grass and low shrub, this needs to be in close proximity to tropical forest because it avoids extensive Eucalyptus woodland. Interestingly, the opposite was the case on Timor, where the species was recorded from landscapes dominated by Eucalyptus woodland with a grassy understorey on the slopes of Mt Ramelau (Trainor et al. 2012). The presence of an endemic subspecies of Sunda Bush Warbler, and a likely endemic subspecies of Timor Stubtail also highlight the importance of Timor as a source of Alor’s avifauna. The Timor Stubtail is a diminutive bird which is most often seen walking on the forest floor, but new island records for Alor, Roti, Atauro and Romang attest to its strong powers of flight and capacity for island colonisation. The song of this bird has a similar structure on all islands (including Babar: PV and CRT unpubl. data) but these scattered populations are presumably at least subspecifically distinctive. The distinctiveness of endemic subspecies of Lesser Sunda populations of Southern Boobook has been the subject of much speculation (Johnstone & Darnell 1997b, Olsen et al. 2010, Verbelen 2010) but only recently have sound recordings of vocalisations started to help unravel the apparent hidden levels of endemism in this complex. The calls of the Southern Boobook on Alor and Pantar are among the most distinctive in the complex, indicating that this form there should be considered a full species. The ‘Alor Boobook’ occurs in tropical forest and Eucalyptus woodland and probably is widely distributed on both islands, and therefore under little threat of extinction. A study is underway on the vocalisations of this complex including samples from most Lesser Sunda islands (F. Rheindt and JE in litt. 2012). The Otus owls have also harboured substantial hidden diversity in south-east Asia (e.g. Lambert & Rasmussen 1998). We are unable to confirm the presence of an Otus owl on Pantar. More fieldwork on Pantar targeting this bird would be of value. Further work is also needed to determine the affinities of the Myzomela honeyeater recorded on Alor. In appearance it is obviously close to Crimson-hooded Myzomela of Wetar, but its calls and songs are different. This is of biogeographical interest because it may be the only clear example of a direct relationship between the avifaunas of Alor and Wetar. Despite our efforts, the avifaunas of both Alor and Pantar remain poorly known. The status of the Flores Hawk Eagle particularly deserves attention. It would be valuable to produce a population estimate, assess the degree to which it is restricted to forested habitats and determine whether it is also present on islands such as Lembata, Pan tar and Wetar. There has been little attention given to the Yellow-crested Cockatoo in Nusa Tenggara since the late 1990s, despite ongoing illegal bird trade which has probably resulted in further population decline. Further effort on Alor, Pantar and other islands would be of value to provide updated information on its conservation status. A study of the status of the Flores Green Pigeon on Alor, Pantar and other islands in its range would be similarly valuable. The taxonomic status of several bird species on Alor (e.g. ‘Little Cuckoo Dove’, Timor Bush Warbler, ‘Crimson-hooded Myzomela’ and perhaps several other montane species) is poorly understood, with vocal or molecular studies now needed to clarify their affinities, and further fieldwork needed to define their distribution, habitat use and conservation status. ACKNOWLEDGEMENTS CRT thanks Heri Ndolu and Neny Herdianawati for logistical and/or field assistance during the 2002 survey, Ian Cowie (Northern Territory Herbarium) for identifying some plants, Mike Carter for information on the status of Pacific Koel on Ashmore Reef and Richard Noske for information on the status of Arctic Warbler in West Timor. PV thanks Veerle Dossche for companionship during his first Alor visit. REJ thanks Dr Soetikno Wiroatmodjo of LIPI and Dr D. J. Kitchener of WAM for facilitating the fieldwork on Alor and Pantar islands and Mr N. Kolichis whose grants to WAM funded some of the field costs. 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(2009) Status of the Pied Imperial Pigeon Ducula bicolor and Pink-necked Green Pigeon Treron vernans on Flores. Kukila 14: 16-20. Setiawan I., Jati, A, Lesmana, D, Trainor, C. & Agista, D. (2000). Telaah awal status dan penyebaran Kakatua-kecil Jambul-kuning (Cacatua sulphurea parvula) di Pulau Alor, Pantar, Timor Barat, Floresdan Moyo. PKA/BirdLife/ WWF, Bogor, Report No. 12. (In Indonesian.) Stattersfield, A. J., Crosby, M. J., Long, A. J. & Wege, D. C. (1998) Endemic Bird Areas of the world: priorities for biodiversity conservation. Cambridge, U.K.: BirdLife International (Conservation Series 7). Stresemann, E. (1929) Ninox fusca plesseni subsp nova. Orn. Monatsb. 37: 47-48. Tebb, G., Morris, P. & Los, P. (2008) New and interesting bird records from Sulawesi and Halmahera. BirdingASIA 1 0: 67-76. Trainor, C. R. (2002a)The birds of Adonara, Lesser Sundas, Indonesia. Forktail 18:93-100. Trainor, C. R. (2002b) Birds of GunungTambora, Sumbawa, Indonesia: effects of altitude, the 1812 cataclysmic volcanic eruption and trade. Forktail 18: 39-51. Trainor, C. (2002c). Status and habitat use of bird communities on Lembata Island, Wallacea, Indonesia, with particular reference to a simple technique for sampling small-island avifaunas. Bird Conserv. Internatn. 12: 365-381. Trainor, C. R. (2003) The birds of Lembata (Lomblen), Lesser Sundas, Indonesia. Kukila 1 2: 39-53. Trainor, C. R. (2005a) Species richness, habitat use and conservation of birds of Alor Island, Lesser Sundas, Indonesia. Emu 1 05: 1 27-1 35. Trainor, C. R. (2005b) Birds ofTapuafu Peninsula, Roti Island, Lesser Sundas, Indonesia. Forktail 21 : 121-131. Trainor, C. R. (2011) Waterbirds and coastal seabirds of Timor-Leste (East Timor): new site records clarifying residence status, distribution and taxonomy. Forktail 27: 68-77. Trainor, C. R., Imanuddin, Aldy, F., Verbelen, P. & Walker, J. S. (2009) The birds of Wetar, Banda Sea: one of Indonesia's forgotten islands. BirdingASIA 10:78-93. Trainor, C. R„ Santana, F., Pinto, P„ Xavier, A. F„ Safford, R. & Grimmett, R. (2008) Birds, birding and conservation in Timor-Leste. BirdingASIA 9: 16-45. 90 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) Trainor, C. R. & Soares, T. (2004) Birds of Atauro Island, Timor Leste (East Timor) Forktail 20: 41-48. Trainor, C. R., Verbelen, F. & Hoste, S. (2012) Rediscovery of theTimor Bush- warbler Locustella (Bradypterus) timorensis on Alor and Timor, Wallacea: clarifying taxonomic affinities, defining habitat and survey recommendations. Bird Conserv. Internatn. DOI: http://dx.doi.org/ 10.1017/S095927091 1000530. Verbelen, F. (2010) Asian enigma: First field observations of Ninox ( novaeseelandiae ) rotiensis on Roti Island, Lesser Sundas, Indonesia. BirdingASIA 13:85-89. Verheijen, J. A. J. (1964) Breeding season on the island of Flores, Indonesia. Ardea 52: 194-201. Verhoeye, J. & Holmes, D. A. (1999) The birds of the islands of Flores - a review. Kukila 1 0: 3-59. Voris, H. K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations. J. Biogeogr. 27: 1 1 53-1 167. White, C. M. N. (1976) Migration of Palaearctic non-passerine birds in Wallacea. Emu 76: 79-82. White, C. M. N. (1977) Migration of Palaearctic passerine birds in Wallacea. Emu 77: 37-38. White, C. M. N. & Bruce, M. D. (1986) The birds of Wallacea (Sulawesi, the Moluccas & Lesser Sunda Islands Indonesia): an annotated check-list. London: British Ornithologists' Union (Check-list 7). Widodo, W. (2009) Population status of Cacatua sulphurea parvula and Trichoglossus euteles in Alor, East NusaTenggara. S/od/Vers/fos 10:81-87. Colin R. TRAINOR, ENV. Australia, Level 1, 503 Murray St, Perth, Western Australia, Australia. Email: colin.trainor@env.net.au Philippe VERBELEN, Torrekenstraat41 , 9820 Munte-Merelbeke, Belgium. Email: filip_verbelen@yahoo.fr Ron E. JOHNSTONE, Curator, Ornithology, Western Australian Museum, 49 Kew Street, Welshpool, Western Australia 6106. Email: ron.johnstone@museum.wa.gov.au Appendix Birds of Alor and Pantar Status: CR = Critically Endangered; VU = Vulnerable; NT = Near Threatened; rr = restricted-range; R = resident, M = Palearctic migrant, AV = austral visitor/migrant, V = visitor, I = introduced; New = new island record; W&B = White & Bruce (1986); C&B = Coates & Bishop (1997). Species Orange-footed Scrubfowl Megapodius reinwardt Brown Quail Coturnixypsilophora Green Junglefowl Callus varlus Australasian Grebe Tachybaptus novaehollandiae Yellow Bittern Ixobrychus sinensis Rufous Night Heron Nycticorax caledonicus Striated Heron Butorides striata Purple Heron Ardea purpurea Great-billed Hem Ardea sumatrana Little Egret Egretta garzetta Pacific Reef Egret Egretta sacra Red-footed Booby Sulasula Brown Booby Sula leucogaster Little Pied Cormorant Microcarbo melanoleucos Eastern Osprey Pandion cristatus Pacific Baza Aviceda subcristata Crested Honey Buzzard Pernisptilorhynchus Short-toed Snake Eagle Circaetus gallicus Black-winged Kit eElanuscaeruleus Brahminy Kite Haliasturindus White-bellied Sea Eagle Haliaeetus leucogaster Variable Goshawk Accipiterhiogaster Brown Goshawk Accipiter fasciata Bonelli's Eagle Aquila fasciata Flores Hawk Eagle Nisaetus floris Spotted Kestrel Falco moluccensis Australian Hobby Falco longipennis Peregrine Falcon Falco peregrinus Red-legged Crake Rallina fasciata Buff-banded Rail Callirallusphilippensis White-breasted Waterhen Amaurornis phoenicurus Common Moorhen Callinula chloropus Red-backed Buttonquail Turnixmaculosus Alor Island Pantar Island Status New W&B C&B REJ KepaTernate Sika CRT R R R M R R R R R R R R R R R M R R R R R R R CR,R,rr x x x x x R R R M R R R R x x X X X X X X X X X X X X X X X X X X X X X X X XX X X X X X X X X X X X X X XX XX X X XX X X X X X XX X X X X XX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXX XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X Forktaii 28 (2012) The avifauna of Alor and Pantar, Lesser Sundas, Indonesia 91 Species Alor Island Pantar Island Status New W&B C&B REJ KepaTernate Sika CRT Eun £ r- 0 r— g ON On C On Cn On -g O "7^ On -C on rn On . — O PV ^ W&B C&B REJ CRT PV Barred Buttonquail Turnixsuscitator R White-headed Stilt Himantopus leucocephalus R Pacific Golden Plover Pluvialis fulva M Grey Plover Pluvialis squatarola M Malaysian Plover Charadrius peronii NT,R Swinhoe's Snipe Gallinagomegala M Bar-tailed Godwit Limosa lapponica M Whlmhtel Numenius phaeopus M Common Redshank Tringa totanus M CommonGreenshank Tringa nebularia M Grey-tailed Tattler Tringa brevipes M Terek Sandpiper Xenus cinereus M Common Sandpiper Actitis hypoleucos M Ruddy Turnstone Arenaria interpres M Red-necked Phalmpe Phalaropus lobatus M x Australian Pratincole Stiltia isabella AV Gull-billed Tern Gelochelidon nilotica M Greater Crested Tern Thalasseus bergii R Bridled Tern Onychoprion anaethetus M Sooty Tern Onychoprion fuscatus R Sterna sumatrana M Spotted Dove Spilopelia chinensis R Island Collared Dove Streptopelia bitorquata R x Bar-necked Cuckoo Dove Macropygia magna R,rr Little Cuckoo Dove Macropygia ruficeps R,rr x Ruddy Cuckoo Dove Macropygia emiliana R x Pacific Emerald Dove Chalcophaps longirostris R Barred Dove Geopelia maugeus R Flores Green Pigeon Treron floris VU,R,rr Banded Fruit Dove Ptilinopus cinctus R Black-naped Fruit Dove Ptilinopus melanospilus R Pied Imperial Pigeon Ducula bicolor V x Green Imperial Pigeon Ducula aenea R Pink-headed Imperial Pigeon Ducula rosacea NT,R,rr Yellow-crested Cockatoo Cacatuasulphurea CR,R Olive-headed Lorikeet Trichoglossus euteles R,rr Lesser Coucal Centropus bengalensis R Asian/Pacific Koel Eudynamysscolapaceus/cyanocephala R Channel-billed Cuckoo Scythrops novaehollandiae R Little Bronze Cuckoo Chrysococcyx minutillus R x Rusty-breasted Cuckoo Cacomantis sepulcralis R Sunda/Oriental Cuckoo Cuculuslepidus/optatus M x [Otus Otus sp] R Eastern Barn Owl Tyto delicatula R x Southern (Alor) Boobook Ninox boobook plesseni R,rr Mees's Nightjar Caprimulgusmeesi R,rr x Savanna Nightjar Caprimulgus affinis R Glossy Swiftlet Collocalia esculenta R Fork-tailed Swift Apus pacificus M x Oriental Dollarbird Eurystomus orientalis M Stork-billed Kingfisher Pelargopsis capensis R Collared Kingfisher Todiramphus chloris R Sacred Kingfisher Todiramphus sanctus AV Common Kingfisher Alcedo atthis R Oriental Dwarf Kingfisher Ceyxerithaca R Rainbow Bee-eater Merops ornatus M Sunda Pygmy Woodpecker Dendrocopos moluccensis R Elegant Pitta Pitta elegans R x x x x x x x x x x x x x x x x x x x x x x X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXX X X X X X X X X X XXX XXX X X X X XXX X X X X X X X X X X X X X X X X X X X X X X XXX X XX X X X X XX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXX X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X XXX X XXX 92 COLIN R. TRAINOR, PHILIPPE VERBELEN & RON E. JOHNSTONE Forktail 28 (2012) Alor Island Pantar Island Species Status New W&B C&B REJ KepaTernate Sika CRT PV Holmes (1995) Johnstone (1991) Mason (1991) Widodo (2009) New W&B C&B REJ CRT PV Helmeted Friarbird Philemon buceroides R X X X X X X X X X X X X Brown Honeyeater Lichmeralndistincta R X X X X X X X X X X X X X Crimson-hooded Myzomela Myzomela (kuehni) NT.R.rr X X X Golden-bellied Gerygone Gerygonesulphurea R X X X X X X X X White-breasted Woodswallow A rtamus leucorynchus R X X X X X X X X X X X Wallacean Cuckooshrike Coracina personata R X X X X X X X X X X X Black-faced Cuckooshrike Coracinanovaehollandiae M X X X X X Common Cicadabird Coracina tenuirostris R X X White-shouldered Triller Lalagesueurii R X X X X X X X X X X Common Golden Whistler Pachycephala pectoralis R X X X X X X X X X X X X X X X Long-tailed Shrike Laniusschach R X X X X X X X X Black-naped Oriole Orioluschinensis R X X X X X X X X X X X Wallacean Drongo Dicrurus densus R X X X X X X X X X X X X X Arafura Fantail Rhipidura dryas R X X X X X X X X X X X X Black-naped Monarch Hypothymis azurea R X X X X X X X X X X X Asian Paradise-flycatcher Terpsiphoneparadisi R X X X X X X X X X Spectacled Monarch Symposiachus trivirgatus R X X X X X X X X Broad-billed Flycatcher Myiagra ruficollis R X X X X X X X X Large-billed Crow Conus macrorhynchos R X X X X X X X X X X X X Cinereous Tit Parus clnereus R X X X X X X X X X X Horsfield's Bushlark Mirafra javanica R X X Barn Swallow Hirundo rustica M X X X Pacific Swallow Hirundo tahitlca R X X X X X X X X X X X X X Striated Swallow Cecropsis striolata R X X X X X X Tree Martin Petrochelidon nigricans R X X X X X X X X X Sunda Bush Warbler Horornis vulcanius R X X X X X Timor Stubtail Urosphenasubulata R,rr X X X Kamchatka Leaf Warbler Phylloscopus examinandus M X X X X X X X X Yellow-breasted Warbler Seicercus montis R X X X Timor Bush Warbler Locustella timorensis NT,R,rr X X X X X Zitting Cisticola Cisticolajuncidis R X X X X X X X X X X X X X Ashy-bellied White-eye Zosterops citrinellus R X X X X X X X X X X X X X X X Mountain White-eye Zosterops montanus R X X Lemon-bellied White-eye/Oriental White-eye Zosterops chloris/palpebrosus R X X Short-tailed Starling Aplonis minor R X X X X X X Common Hill Myna Gracula religiosa R X X X X X X X X X X X X Chestnut-backed Thrush Geokichla dohertyi NT,R,rr X X X Lesser Shortwing Brachypteryx leucophrys R X X X X X X X Pied Bush Chat Saxicola caprata R X X X X X X X X X X X X Little Pied Flycatcher Ficedula westermanni R X X X X X X Thick-billed Flowerpecker Dicaeum agile R X X X X Black-fronted Flowerpecker Dicaeum igniferum R,rr X X X X X X X X X X X X X X Brown-throated SunbM Anthreptes malacensis R X X X X X X X ? X Olive-backed Sunbird Cinnyrisjugularis R X X Flame-breasted Sunbird Cinnyris Solaris R,rr X X X X X X X X X X X X X X X Eurasian Tree Sparrow Passer montanus 1 X X X X X X Zebra Finch Taeniopygia guttata R X X X X X X X X X X Black-faced Munia Lonchuramolucca R X X X X X X X X X X Scaly-breasted Munia Lonchura punctulata R X X X X X X Five-coloured Munia Lonchura quinticolor R X X X X X X X ■ Pale-headed Munia Lonchura pallida R X X X X X X X X X Red Avadavat Amandava amandava R X X X X Eastern Yellow Wagtail Motacilla tschutschensis M X X Grey Wagtail Motacilla cinerea M X X X X Paddyfield Pipit Anthus rufulus R X X FORKTAIL 28 (201 2): 93-100 Foraging ecology of sympatric White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigantea in northern Cambodia H. L WRIGHT, N. J. COLLAR, I. R. LAKE, BOU VORSAK & P. M. DOLMAN White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigantea are two of the most threatened yet poorly known birds of South-East Asia's dry forests. Anecdotal evidence suggests these species have an intriguing combination of ecological similarities and differences, and as they occur sympatrically there may be an opportunity to safeguard them through joint conservation measures. We compared their foraging ecology and proximity to people in an attempt to unravel their ecological differences and better inform conservation. Landscape-scale habitat use was assessed by recording ibis sightings on journeys through a 75,000 ha dry forest landscape; White-shouldered Ibises were surveyed over 526 journeys (totalling 17,032 km) and Giant Ibises over 349 journeys (1 1,402 km). The ibises showed broadly similar habitat selection, using a range of wetland and terrestrial habitats. Giant Ibises were more often sighted further from settlements than White-shouldered Ibises, with maximum sighting frequency predicted at 9.9 km from villages for the former and 8.3 km for the latter. Giant Ibis may be less tolerant of human disturbance and/or White-shouldered Ibis may be more dependent on traditional land management practices, but the species' differing use of abandoned paddyfield (a habitat typically near settlements) could also be a contributing factor. At waterholes in the dry season foraging Giant Ibis used wetter microhabitats than White-shouldered, suggesting the species occupy different foraging niches. We make preliminary observations regarding Giant Ibis breeding strategy and discuss potential habitat management actions, concluding that, although conservation could address these species simultaneously in dry dipterocarp forest landscapes, their ecological differences must also be taken into account. INTRODUCTION Dry dipterocarp forests of South-East Asia contain a distinctive assemblage of species including megafauna such as Asian Elephant Elephas maximus. Tiger Panthera tigris and Banteng Bosjavanicus, and large-bodied birds, such as three vulture, four stork and one crane species (Baltzer et al. 2001). This biodiversity has suffered various human impacts, namely hunting, habitat loss and degradation, with at least 60 dry forest birds, mammals and reptiles classified as threatened on the IUCN Red List in Cambodia alone (Tordoff et al. 2005, WCS 2009). While conservation resources are being increasingly directed at this ecosystem, the ecology of the forest and much of its wildlife remains poorly or only partially understood (Songer 2006, CEPF 2007). Of the dry forest birds, two species stand out as amongst the most enigmatic, threatened and poorly studied: the Critically Endangered White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigantea. These two dry forest ibises experienced dramatic declines in the twentieth century (BirdLife International 2001) and, although once widely distributed across South-East Asia, their ranges contracted to become almost entirely confined to Cambodia (BirdLife International 2001). Remaining populations are fragmented and only 250 individual Giant Ibises (BirdLife International 2012b), and 731-856 individual White-shouldered Ibises (Wright et al. 2012a) are estimated to remain globally. Conversion of dry forests (for infrastructure, settlement and agriculture, including plantations) and changing local land management are projected to cause further, severe declines in ibis populations (BirdLife International 2012a,b). Conservation action is urgently required to secure these ibises from extinction, but is likely to depend on a scientific understanding of their ecological requirements. White-shouldered Ibis and Giant Ibis exhibit an intriguing mixture of ecological similarities and differences. The species occur sympatrically in much of their current ranges (historically they occurred together, or in close proximity, in Cambodia and southern Laos: BirdLife International 2001), and while their wet-season foraging ecology remains poorly known (Keo 2008b, BirdLife International 2012a), both forage at seasonal wetlands, known as trapaengs, in the dry season (November-May: Keo 2008b, Wright et al. 2010). Both ibises breed solitarily in canopies of dipterocarp trees and no evidence of migration has been found for either species. Despite these similarities these ibises have contrasting breeding strategies, with White-shouldered Ibis nesting in the mid- to late dry season (December-May: HLW unpubl. data) and Giant Ibis in the wet to early dry season ( June-November: Clements et al. in press). Available evidence suggests that breeding White-shouldered Ibises forage in exposed substrates at drying-out trapaengs (Wright et al. 2010). However, why the Giant Ibis breeding season differs so markedly is not yet known, and the habitat use and prey selection of these species have yet to be compared. The ibises’ overlapping ranges and ecology suggest that carefully designed conservation measures could attempt to safeguard both species simultaneously. Conservation that supports local land- management practices may benefit both species, particularly the maintenance of foraging habitat by domestic livestock (Keo 2008b, Wright et al. 2010). However, adopting such a strategy requires research into the compatibility of the ibises’ ecological requirements and their interaction with people. This study compares White¬ shouldered Ibis and Giant Ibis foraging ecology, examining habitat selection in the dry forest landscape and microhabitat and prey use at trapaengs. METHODS Study area The study was conducted in a c.75 ,000 ha area with in Western Siem Pang and Sekong Important Bird Areas (IBA; centred on 14°17'N 106°27'E), northern Cambodia (Figure 1, Seng et al. 2003), an unprotected site with at least 262 individual White-shouldered Ibises and an estimated 80 Giant Ibises (BirdLife International 2012a,b). The area comprises a mosaic of dry dipterocarp forest with patches of grassland {veals), river channels, mixed deciduous and semi-evergreen forest, and active and abandoned agricultural land (rice paddy) close to settlements. The climate is strongly monsoonal with average monthly rainfall reaching 333 mm in the wet season and as low as 0.9 mm in the dry season (Thuon & Chambers 2006). Dry forest understorey is burnt annually in the 94 H. L. WRIGHT etal. Forktail 28 (2012) Thailand Vietnam Map c - Country border Western Siem Pang and Sekong IBAs Kong River Settlement Survey tracks on land Survey tracks by river Agricultural land Veal Forest (all types) Hi Kong River — River tributary ® Trapaeng m Settlement - Main road Survey tracks on land Em Survey tracks by river Figure 1 . Location of Western Siem Pang and Sekong IBAs in Stung Treng province, Cambodia (a) and extent of survey journeys within the IBAs (b). Map c demonstrates the extent of main habitat types within the core section of the study area. 'Agricultural land' includes cultivated/ stubble and abandoned paddyfields. dry season, largely as a result of anthropogenic fires. Trapaengs occur frequently in the landscape and vary in size (0.001-3.4 ha: Wright etal. 2010). Water drawdown in trapaengs and rivers is dramatic in the dry season, exposing substrates with varied moisture conditions. Villages were concentrated in the south and east of the study area (Figure 1) and inhabited by c. 10,000 people (Ministry of Planning 2007). Surveys of ibis habitat use Ibis habitat selection was examined at the landscape scale by recording ibis sightings along journeys through the study site. White-shouldered Ibises were recorded during 526 journeys over 22 months between November 2009 and January 2012; the protocol was then expanded also to record Giant Ibises, which were surveyed during 349 journeys over 1 7 months between March 2010 and January 2012 (a subset of White-shouldered Ibis journeys). Journeys were undertaken systematically as part of travel for wider research and on-site conservation activities, with up to three observers travelling independently per day. Journeys took place along forest tracks and paths, covering 33.9 ±18.9 km per journey- day (mean ± SD) and were made by motorbike at low speed or occasionally by foot where tracks were inaccessible ; 2.4% of journeys were made by boat along main river channels. Journeys were made in both the dry and wet seasons and survey effort (km per journey- day) was similar. Survey routes for each journey were noted on datasheets and recorded using a hand-held GPS. The survey recorded the location (using a GPS), number, activity and habitat use of ibis with each sighting. Ibises on or taking off from the ground were assumed to be foraging and selected for analysis; other activities (such as loafing or preening) may also take place on the ground but are typically interspersed with foraging bouts and occur in the same habitat (HLW pers. obs.). FTabitat was categorised as river channel; trapaeng-, dry dipterocarp forest; veal-, cultivated rice paddy; rice paddy stubbles; abandoned paddyfield (unused tor more than one season) and mixed deciduous/semi- evergreen forest. The placement of forest tracks was largely independent of vegetation or topographical features, making journeys representative of habitats with the exception of denser semi-evergreen forest (rarely used by either ibis in mainland South- East Asia: BirdLife International 2012a,b), river tributaries and isolated areas of wet-season inundation. Although not traversed, tracks were frequently beside trapaengs, allowing them to be surveyed. Much veal habitat originated from historic rice cultivation, but swards are typically taller than at more recently abandoned paddies, so these habitats were considered separately; bunds were more apparent in abandoned rice paddies (typically > 10 cm high) than in veals (typically < 10 cm or absent), allowing these habitats to be distinguished. Fdabitats were mapped with a hand-held GPS during journeys in April 2010. Survey effort per habitat type was quantified in a GIS (ArcMap 9.3, ESRI 2010) by intersecting journey tracks onto the habitat segments they traversed. As tracks went beside trapaengs, survey effort for this habitat was quantified by intersecting tracks through buffers surrounding each surveyed trapaeng. Buffer size was a factor of trapaeng radius and viewable distance (in classes of 0, 20, 40, 60 and 80 m) so that large trapaengs visible from far away Forktail 28 (201 2) White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigonteo in Cambodia 95 accounted for greater survey effort than small trapaengs visible only from close by. The survey protocol was kept simple so that local field staff and villagers (with low technical expertise) could collect consistent data; as distances to observed birds were not recorded, data could not be analysed by a distance-sampling approach. Nonetheless, a preliminary survey did measure the distance from observers to ibises seen on the ground, showing that ibis detectability varied with habitat [F4M = 2.71, P = 0.046, distance square-root transformed). Journey distances per habitat segment were therefore multiplied by a habitat-specific estimated transect strip-width, defined by the average sighting distance (or an approximated distance for habitats in which no ibises were observed), to calculate both survey effort and sighting frequency per km2. Estimated transect widths may slightly underestimate effective strip-width and therefore overestimate sighting density, but this is unlikely to have caused a directional bias in the findings presented, and population densities were not estimated. Survey areas were aggregated by habitat type and half-month time-periods for analysis. Foraging observations at trapaengs Ibis microhabitat and prey use were studied at trapaengs by observing foraging ibises between December and February in two dry seasons (2008-09 and 2009-10) for seven trapaeng- year observation periods: three trapaengs in one year and two trapaengs in each of the two years. Trapaengs were selected to contain a complete gradient of moisture conditions (from pools of water to exposed dry substrate) and maximise the chance of ibis visitation (confirmed by preliminary surveys) to provide foraging data. Trapaengs surveyed in both years were observed in different months under novel habitat conditions. Observations were conducted by one observer using a telescope from hides on trapaeng perimeters; observations were from dawn until dusk lasting for 3.9 ± 0.7 contiguous days (mean ± SD per site). Trapaeng microhabitats were defined by four moisture conditions: pools of water, and saturated, moist or dry exposed substrates. Their extents were mapped at each trapaeng by sketching homogeneous habitat patches (Wright et al. 2010), recording coordinates with a hand-held GPS and measuring dimensions with a laser rangefinder. Maps were georeferenced and digitised in a GIS to calculate patch areas, and area data were aggregated to calculate microhabitat extent as a proportion of each trapaeng. Ibis microhabitat use was measured by instantaneous scan¬ sampling at sLx-minute intervals, recording the activity and location (habitat patch) of all ibises present, the latter aided by markers placed around habitat patch boundaries. Prey type and size class (0- 2.49 cm, 2.50-4.99 cm and > 5 cm) of each item captured was recorded for ibis individuals during replicate six-minute focal watches. Items of < 1 cm were consumed infrequently and therefore excluded from analysis. Prey biomass was estimated using average ash-free dry mass (AFDM) calculated for a set of prey specimens comprising all prey types and size classes (Piersma et al. 1994). Analysis Fandscape-scale habitat selection was examined by log-ratio analysis (Aebischer et al. 1993), comparing proportionate habitat use (from number of sightings) with proportionate habitat availability (from survey effort area) using the half-month period as the unit of replication. Analysis was conducted in Compos Analysis software (Smith 2005) with log-ratios weighted by the square-root of total survey effort area per period. Habitat selection was analysed separately for each ibis species in the dry (White-shouldered Ibis n - 23 half-month periods, Giant Ibis n = 15) and wet (White¬ shouldered Ibis n — 17, Giant Ibis n — 16) seasons. Within the wet season, relative use of trapaengs was compared between ibis species with a chi-squared test, contrasting numbers of sightings at trapaengs versus non- trap aengbzb'nzts by pooling records from journeys made after March 2010 (when both ibis species were surveyed). To examine the effect of proximity to people on ibis occurrence, ibis sightings and journey tracks were split using a GIS into five classes of distance to nearest settlement (0-2.49, 2.5-4.99, 5-7.49, 7.5-9.99 and 10-16 km). The effect of distance to settlement (midpoints of the five distance classes, treated as a continuous variable) on ibis sighting frequency (count per distance-to- settlement class per journey-day) was modelled for each ibis species in GFMs with Poisson-distributed error and log link, with log survey effort area included as an offset. Sample units with a journey distance of less than 2 km were excluded to ensure counts were based on adequate survey effort. Non-linear effects of distance to settlement were tested by square-root transformation. Proximity of individual sightings to the nearest settlement (calculated in GIS) was also compared between the two species using a Mann-Whitney test. The species’ microhabitat and prey use at trapaengs were statistically compared for the trapaeng- year observations in which both ibis species were observed, ensuring comparable survey effort and habitat and prey availability. Species’ use of dry versus saturated substrate, and of water versus other microhabitats combined, were compared using chi-squared tests of the frequency of scan-sampled individuals per microhabitat type. Intake rate (centigrams of AFDM per minute) of two prey groups — ( 1 ) amphibians and small invertebrates, and (2) eels and crabs — were calculated using focal watch data and compared between the two species using a Mann- Whitney test. RESULTS Ibis sighting frequency and flock size A total of 446 White-shouldered Ibis and 66 Giant Ibis sightings were obtained from 17,032 km and 1 1,402 km of survey journeys respectively; 328 White-shouldered Ibis and 5 1 Giant Ibis sightings were of birds seen on or taking off from the ground and assumed to be foraging (‘sightings’ refers to foraging birds henceforth). Sighting frequency of foraging White-shouldered Ibis per journey-day was 0.20 ± 0.40 (mean number of sightings per km2 ± SD) compared with 0.06 ± 0.31 for Giant Ibis, and mean flock size per sighting was larger (Mann-Whitney test FT^32S 51 = 10,142, P = 0.01 1) for White-shouldered Ibis (5.9 ± 15.3 birds, mean ± SD) than for Giant Ibis (1.8 ± 0.8 birds). White-shouldered Ibis flock size was greater (^99 229 = 5939, P < 0.001) in the wet (non-breeding) season (10.1 ± 25.1 birds) than in the dry (breeding) season (4.1 ± 7.3); no such difference was found for Giant Ibis. Landscape-scale habitat selection White-shouldered Ibis and Giant Ibis both foraged in a variety of habitats within the dry forest landscape, and both showed marked differences in habitat selection between the dry and wet seasons (Figure 2). Both ibises preferred trapaengs in the dry season, particularly White-shouldered Ibis which breeds in this season. Giant Ibis also made use of river channels in the dry season, a habitat not used by White-shouldered Ibis in this study. In the wet season, ibises made more equal use of habitats and were found more frequently in terrestrial areas than in the dry season. Both species made use of trapaengs, veals and dry dipterocarp forest, but White¬ shouldered Ibises also used abandoned paddyfields (19.4% of wet- season sightings), a habitat in which Giant Ibises were not observed in either season. Trapaengs accounted for a greater proportion of Giant Ibis sightings in the wet season than for White-shouldered Ibis (47% and 22% of sightings respectively), and use of trapaeng versus non-trapaeng habitat was greater for Giant Ibis than for White-shouldered Ibis (at2! = 4.01, P = 0.045). Sighting frequency 96 H. L. WRIGHT etal. Forktail 28(2012) White-shouldered Ibis a Giant Ibis Forest Aban. yea| Trapaeng River paddy WET Forest Aban. \/eal Trapaeng River paddy Figure 2. Habitat use of foraging White-shouldered Ibis (dark grey) and Giant Ibis (light grey) in the dry and wet seasons. The dry season corresponds with the White¬ shouldered Ibis breeding season and Giant Ibis non-breeding season; the wet season is the reverse. All habitats were surveyed in each season; missing bars indicate no birds recorded. Sighting frequency is the average number of foraging ibis sightings per km2 per half-month; bars indicate standard error. Ibis habitat use (mean log- ratios of use versus availability) did not differ significantly (P < 0.01) between habitats sharing a common letter; river was surveyed too infrequently to analyse its dry- season selection by Giant Ibis. 'Forest' refers to dry dipterocarp forest, 'Aban. paddy' is abandoned paddyfield. of White-shouldered Ibis at trapaengs in the wet season was also 79% lower than in the dry season, compared with only a 34% difference for Giant Ibis. Three habitats appeared unimportant for foraging ibis: one or fewer sightings were gained (for either species) in rice-paddy stubbles and in cultivated paddy despite the large extent of these habitats in parts of the study area; and fewer than three sightings came from mixed deciduous/semi-evergreen forest, perhaps reflecting this habitat’s sparse distribution in the study area and/or poor representation by survey journeys. Effect of distance to settlement Ibis sighting frequency per journey-day was positively related to distance to settlement for both White-shouldered and Giant Ibis (Figure 3), with distance to settlement strongly supported in models for both species; removal of the linear term resulted in Akaike Information Criterion (AIC) increases of > 2 units (63.0 for White¬ shouldered Ibis and 13.0 for Giant Ibis), while removal of the non¬ linear term resulted in large AIC increases (96.6 and 22.5 respectively). Predicted White-shouldered Ibis sighting frequency had a steeper response curve with distance to settlement (Figure 3; n - 1,362, linear term )3 = -0.68 ± 0.17 95% Cl, non-linear term / 3 = 3-93 ± 0.79) than Giant Ibis (n = 904, /3 = -0.85 ± 0.46, non¬ linear term f3 = 5.41 ± 2.34); consequently maximum White¬ shouldered Ibis sighting frequency was predicted at 8.3 km from settlements compared with 9.9 km for Giant Ibis. Further highlighting this difference, the mean distance to nearest settlement of individual White-shouldered Ibis sightings (5.7 ± 3.3, mean ± SD) was significantly less than for Giant Ibis (7.8 ± 3.4, = 5474.5, P < 0.001). White-shouldered Ibis sighting frequency appears to decline after 8.3 km from settlements, but lower sample sizes and overlapping confidence intervals (at 8.3 km versus maximum settlement distance) suggest this result is not robust; using additional classes of 10-12.99 and 13-16 km also did not provide strong evidence for a decline (and resulted in even smaller sample sizes). Microhabitat and prey use at trapaengs White-shouldered Ibis foraging data were obtained in all seven trapaeng- year observations and Giant Ibis data in two, yielding 777 and 1 17 scan-samples (3,101 and 242 individual bird records) for these species respectively; scan-samples comprised tens of White¬ shouldered Ibis individuals and a minimum of eight Giant Ibises. Crude comparison of proportionate microhabitat use shows some apparent similarities between the two ibis species at trapaengs in the dry season (Figure 4); both fed in all exposed substrate types and showed proportionally greater use of both dry and saturated substrates than moist substrate. However, relative to other microhabitats, Giant Ibis made greater use of pools of water than White-shouldered Ibis (a:2| = 81.6, P < 0.001) for which only 0.3% of scan-sampled individuals (across all trapaeng- years) foraged in Figure 3. White-shouldered Ibis (dark grey) and Giant Ibis (light grey) sighting frequency (number of foraging ibis sightings per km2 per journey-day), averaged by class of distance to settlement (a) and predicted by Poisson GLMs (b): White-shouldered Ibis model AIC = 1 585.7, dispersion ratio = 0.77; Giant Ibis model AIC = 376.3, dispersion ratio = 0.31 . Bars indicate standard error and dotted lines indicate 95% confidence intervals. 0-2.49 2.5-4.99 5-7.49 7.5-9.99 10-16 Distance to settlement (km) Forktail 28 (2012) White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigantea in Cambodia 97 Figure 4. Microhabitat use by (a) White-shouldered Ibis and (b) Giant Ibis at trapaengs in the dry season. Proportionate use (grey columns) and proportionate availability (white columns) of moisture conditions were averaged across seven trapaengs for White-shouldered Ibis and two trapaengs for Giant Ibis. Bars indicate standard error. Figure 5. Mean proportionate contribution of prey types to biomass consumed by White-shouldered Ibis (dark grey, at seven trapaengs) and Giant Ibis (pale grey, at two trapaengs ) during foraging bouts. Bars indicate standard error. water. Furthermore, the ibis differed significantly in their use of saturated versus dry substrate (x1^ = 140.7, P < 0.001), with Giant making greater use of the former and White-shouldered greater use of the latter. In terms of proportionate use relative to proportionate availability, White-shouldered Ibis appeared to prefer dry substrate whereas Giant Ibis appeared to avoid it (Figure 4). Focal sampling at trapaengs yielded 89.7 and 3.46 aggregate hours of foraging observation data (from 797 and 40 focal watches) for White-shouldered Ibis and Giant Ibis respectively. Marked differences in diet composition were apparent, with amphibians contributing greatest biomass for White-shouldered Ibis and eels contributing most to Giant Ibis diet (Figure 5). At the two trapaengs where both ibis species were observed, combined intake rate of amphibians and small invertebrates was significantly greater for White-shouldered than for Giant Ibis ( lVi49A0 = 3297, P < 0.001). Conversely, combined intake rate of eel and crabs was significantly greater for Giant than for White-shouldered Ibis {tV40Mc, = 8532, P < 0.00 1 ). Prey biomass estimates for a set of ashed prey specimens suggest the average-sized crab caught by either ibis may hold 2.5 times more AFDM than the average-sized amphibian, and the average-sized eel may hold over 5 times more, indicating that Giant Ibis is likely to be consuming considerably greater prey biomass per item caught than White-shouldered Ibis. DISCUSSION White-shouldered Ibis and Giant Ibis showed some broad similarities in foraging ecology, including habitat use at the landscape scale. Nevertheless, the relative importance of wetland to terrestrial habitats differed between the species, and contrasting microhabitat and prey use at trapaengs suggests these species occupy different foraging niches. Giant Ibises were more often recorded further from settlement than White-shouldered Ibises, suggesting that these species may have different tolerance levels to human disturbance and/or different dependency on traditional land- management practices (assumed to predominate closer to settlements) and habitat types. Conservation could be designed to benefit both ibises simultaneously, but will require careful consideration of their ecological differences. Landscape-scale habitat selection White-shouldered Ibis and Giant Ibis used a mixture of dry- forest habitat types, which varied with season. They showed broadly similar habitat use at the landscape scale in contrast to their markedly different breeding strategies. In the dry season, despite rapidly receding water-levels (Wright et al. 2010), both ibises preferred to forage in wetland habitats, similar to Sharp-tailed Ibis Cercibis oxycerca and Green Ibis Mesembrinibis cayennensis habitat selection in the Llanos of Venezuela (Frederick & Bildstein 1992). Mean White-shouldered Ibis sighting frequency was vastly greater at trapaengs than in any other habitat in the dry season. This species adopts a foraging strategy well adapted to increasing extents of exposed dry substrates at trapaengs in this season (fdLW unpubl. data) and access to trapaengs may be essential for breeding. Habitat use in the wet season was more equitable, with both ibis species foraging in a range of wetland and terrestrial habitats. Trapaengs continued to be used, although to a lesser degree than in the dry season, and the use of open terrestrial habitats (abandoned paddyfield and/or veal) suggests that access to the ground is important, as for most ibis species reliant on terrestrial habitats (del Hoyo et al. 1996). Keo (2008b) also noted the value of veals as a Giant Ibis foraging habitat, observing a high wet-season abundance of earthworms. White-shouldered Ibis was gregarious in the wet season and large congregations of foraging birds (up to 185 individuals) were observed in veals and abandoned paddyfields, indicating the importance of such habitats. Apparent habitat preference and response to settlement proximity may be confounded; abandoned paddyfields were not found more than 4.6 km from settlements, so the absence of Giant Ibis from this habitat may represent avoidance of villages rather than habitat suitability. Conversely, the apparently greater tolerance of White-shouldered Ibis of settlement proximity may reflect its greater use of these habitats rather than a differing response to people per se. Although White-shouldered Ibises were not observed in river channels, rivers appear to constitute important foraging habitat elsewhere in Cambodia and Indonesian Borneo (Sozer & van der Heijden 1997, Timmins 2008). Both species are most likely to forage in river channels with large extents of exposed mud and/or sand (R.J. Timmins in litt. 2012); such habitats were rare along surveyed sections of main rivers, and seasonal tributary channels in the wider landscape, perhaps containing the most exposed substrate, were poorly represented by journeys along forest tracks. More dedicated survey of suitable riverine habitat would improve knowledge of its selection by both ibis. Nonetheless, the lack of wet-season sightings along rivers may genuinely reflect prohibitively high water-levels, an effect seen for White-shouldered Ibis along the Mekong River (Timmins 2008). 98 H. L. WRIGHT eta/. Forktail 28(2012) Proximity to people Of the two species. Giant Ibis appeared more limited by proximity to people. Foraging birds were observed significantly further from settlement for this species and in models the predicted maximum sighting frequency occurred 1.6 km further from settlements than White-shouldered Ibis. Keo (2008b) found that Giant Ibis typically nested more than 4 km from settlements and preferred to forage at trapaengs further from villages, postulating that disturbance and/ or persecution may be greater closer to settlements. White¬ shouldered Ibis appears much more accustomed to people, often roosting and nesting in trees in or around rice paddies — even when in use by people — and is less wary when approached (HLW pers. obs.). Sightings from the early twentieth century, although sparse, also suggest that White-shouldered Ibis was found more frequently in cultivated lands than Giant Ibis (Thewlis & Timmins 1996, BirdLife International 2001). It is possible that White-shouldered Ibis is more opportunistic in its habitat use, while Giant Ibis may make more specialist use of dry- forest landscapes; alternatively, these patterns may reflect an underlying difference in responses to human disturbance. Irrespective of the mechanism, Giant Ibis would appear more vulnerable to human activity and settlement in remote areas, while White-shouldered Ibis’s closer proximity to people and less evasive behaviour may make it more vulnerable to hunting, a factor that most probably contributed to its decline, particularly in Laos and Vietnam (BirdLife International 2001). Unchecked habitat conversion, resource extraction, human population growth and settlement expansion, issues requiring urgent conservation action in South-East Asia (CEPF 2007), will threaten both species. Somewhat contrasting dependencies on traditional land- management practices could also shape these responses to settlement proximity. While both species could benefit from grazing of foraging habitat by livestock (Keo 2008b, Wright et al. 2010), White-shouldered Ibis is plausibly more constrained by habitat availability, being a third smaller (in terms of linear body length: del Hoyo et al. 1996) and considerably shorter in height than Giant Ibis (HLW pers. obs.). Vegetation growth in trapaengs , veals and dry dipterocarp forest is substantial in the wet season (HLW unpubl. data) and may reach more than double White¬ shouldered Ibis body height (Wright et al. 2010), potentially restricting this species’s use of habitats otherwise still accessible to Giant Ibis. More frequent White-shouldered Ibis sightings in areas closer to settlements could reflect a stronger requirement for grazed habitat and bare ground, where livestock densities are highest (Wright et al. 2010). The possible decrease in White-shouldered Ibis sighting frequency beyond 8.3 km from settlements would further support this. The effect of people and livestock on ibis abundance deserves further study to understand the potential trade¬ off between the negative effects of human disturbance (particularly for Giant Ibis) and the positive effects of land-use practices. Examining ibis distribution in a study landscape with a steeper gradient of livestock density and greater maximum distance to settlement than occurred in this study area would help in clarifying this potential effect. Foraging ecology at trapaengs The dry-season foraging ecology of White-shouldered Ibis and Giant Ibis has been examined elsewhere (Keo 2008a, Wright et al. 2010, Wright in press). Having observed only a small number of Giant Ibis feeding at two trapaengs , this study cannot draw major new conclusions on this species’s foraging strategy and we limit the discussion to a preliminary comparison of the two species. Similarities included both species’ use of all exposed substrate types, and shared use of the amphibian resource at trapaengs. Although Keo’s (2008b) study (from January to April) found amphibians contributed a major part of Giant Ibis diet (as is the case for White¬ shouldered Ibis), this study found them to be of only marginal importance. However, our survey took place earlier in the dry season when wetter conditions may have sustained more of the aquatic prey seemingly favoured by this species. The most notable differences in microhabitat use were the avoidance of water and apparent selection of dry substrate by White-shouldered Ibis, contrasting with Giant Ibis’s greater use of water and saturated substrates. Prey use reflected this with the contribution of aquatic prey to Giant Ibis diet (83% of consumed biomass) far exceeding that in White-shouldered Ibis diet (0.4%). Average biomass of prey items consumed by Giant Ibis was 263% greater than of items consumed by White-shouldered Ibis, perhaps explaining the large disparity in body size between these species, or demonstrating the Giant Ibis’s need to consume sizeable prey to sustain its large body size. Despite some similarities in foraging ecology, the contrasting use of wet microhabitats suggests the ibises may occupy different foraging niches at trapaengs , at least in the early to middle dry season. Studies of ibis incidence at trapaengs add further evidence, with Giant Ibis selecting trapaengs with greater extent of wet mud (Keo 2008b), but White-shouldered Ibis showing no such selection (Wright et al. 2010). The ibises’ morphology may also point to a degree of niche separation: the White-shouldered Ibis’s neck is feathered from the body to the top of its hindcrown, whereas Giant Ibis has feathering along only one-third of its neck-length. The latter may be an adaptation for a bird that more regularly submerges its head and upper neck in water or wet mud, although the presence ot bare skin may also aid thermoregulation (Buchholz 1996, Ward et al. 2008). When in shallow water, Giant Ibis forage by probing with bill slightly agape (Eames 2011, HLW pers. obs.), matching the technique used by other ibis (e.g. Kushlan 1979) but not witnessed for White-shouldered Ibis. Giant Ibis breeding strategy The wet-season foraging ecology of these ibises is still very poorly known (Keo 2008b, BirdLife International 2012a); examining the ibises’ relative foraging success for wet season prey types and habitats will inform conservation and shed light on the Giant Ibis’s breeding strategy. Accessing tall and densely vegetated wet-season habitats may be easier for Giant Ibis (given its larger body size) than for White-shouldered, allowing it to reach the prey-rich earthworm mounds found in dry dipterocarp forest and particularly veals. A longer, thicker bill may also be more effective at probing for earthworms. Terrestrial habitats, particularly veals, can become inundated in the wet season and trapaeng water-levels increase substantially (HLW unpubl. data). Greater use of trapaengs in the wet season, a greater tendency to forage in wet microhabitats, and longer legs and bill all indicate that Giant Ibis may be better adapted to foraging in flooded conditions. Giant Ibis may therefore have a broader range of suitable wet-season habitats in which to forage and provision chicks. Implications for habitat management White-shouldered Ibis and Giant Ibis both used a mosaic of habitat types, requiring protection and management of habitats at the landscape scale. Trapaengs are important breeding-season habitats for both species, but a range of terrestrial habitats must also be available, particularly in the wet season. Broadly similar habitat use indicates that conserving a suite of dry forest habitats will benefit these sympatric ibis species simultaneously. The use of open habitats such as veals and abandoned paddyfields suggests these features should be protected and kept open; the latter deserve particular attention as they occur in close proximity to settlement and may be particularly vulnerable to agricultural and urban expansion. Clearings could be created to improve habitat availability in landscapes dominated by dense dry forest, and managed (e.g. by livestock grazing) to ensure they remain accessible. Knowledge of Forktail 28 (2012) White-shouldered Ibis Pseudibis davisoni and Giant Ibis Thaumatibis gigantea in Cambodia 99 the ibises’ wet-season foraging ecology would be valuable to assist the design and management of these open areas. Maintaining the extensive rearing of domestic livestock is likely to be important to both ibis species — a requirement shared with many other threatened species in pastoral and mixed farming systems (Wright et al. 2012b). Domestic livestock keep sward heights low at trapaengs , veals and in dry dipterocarp forest, maintaining habitat suitability for White-shouldered Ibis (HLW unpubl. data). Domestic buffalo are key grazers at trapaengs and may also be useful in creating areas of saturated substrate to benefit Giant Ibis at trapaengs (Keo 2008b). Both ibises will be affected if a reduction in livestock causes long-term ecological succession and trapaeng sedimentation. Further study should investigate whether introducing buffalo or cattle in landscapes where both domestic livestock and wild herbivores are lacking can improve habitat suitability for these species, and whether animal wallowing may contribute to trapaengctcsadow. Annual dry-season fires are another component of traditional dry forest land-management, resulting in reduced sward heights; this could be a benefit to ibises foraging in terrestrial habitats and also deserves further research. While conservation could benefit White-shouldered and Giant Ibises simultaneously, care should be taken to ensure that interventions do not overlook their ecological differences. In particular, safeguarding Giant Ibis is likely to require that large areas of undisturbed habitat are protected from development and human interference, contrasting with the White-shouldered Ibis’s potentially stronger need for habitat management. Fduman disturbance and hunting risk require close attention in interventions that expand, enhance or maintain traditional management practices. Equally, maintaining dry-season water-levels at trapaengs for the benefit of Giant Ibis or other wildlife (Keo 2008b) should not be undertaken to the extent that the exposed substrates (including dry ground) preferred by breeding White-shouldered Ibis become scarce or unavailable. ACKNOWLEDGEMENTS We specially thank Mem Mai, Lourn Bun Paeng, Sum Phearun and Net Norin for assisting data collection and R. J. Timmins for ibis records. Thanks to Sum Phearun, Keo Omaliss, D. Buckingham, T. Evans, J. C. Eames and R. J. Timmins for sharing thoughts on ibis ecology, E. Brooks for GIS assistance and two anonymous reviewers for their comments. For support, we are grateful to H.E Cheng Kim Sun and Men Phymean, Forestry Administration, and H.E. Chay Samith, General Department of Administration for Nature Conservation and Protection, Royal Government of Cambodia. Funding was provided by the British Ornithologists’ Union, Critical Ecosystem Partnership Fund (a joint initiative of l’Agence Framjaise de Developpement, Conservation International, Global Environment Facility, Government of Japan, MacArthur Foundation and World Bank), Mohammed bin Zayed Species Conservation Fund, Natural Environment Research Council and Economic and Social Research Council. REFERENCES Aebischer, N. J., Robertson, P. A. & Kenward, R. E. (1993) Compositional analysis of habitat use from animal radio-tracking data. Ecology 74: 1313-1325. Baltzer, M. C., Nguyen, T. D. & Shore, R. G. (2001) Towards a vision for biodiversity conservation in the forests of the lower Mekong ecoregion complex. Hanoi and Washington D.C.: WWF Indochina/WWF US. Bi rd Life International (2001) Threatened birds of Asia: the BirdLife international Red Data Book. Cambridge: BirdLife International. BirdLife International (2012a) Species factsheet: Pseudibis davisoni . Available from http://www.birdlife.org. Accessed 14/5/12. BirdLife International (2012b) Species factsheet: Thaumatibis gigantea. Available from http://www.birdlife.org. Accessed 14/5/12. Buchholz, B. (1996) Thermoregulatory role of the unfeathered head and neck in male wild Turkeys. Auk 1 1 3: 310-318. Clements, T., Rainey, H. J., An, D., Rours, V.,Tan, S., Thong, S., Sutherland, W. J. & Milner-Gulland, E. J. (in press) An evaluation of the effectiveness of a direct payment for biodiversity conservation: the Bird Nest Protection Program in the Northern Plains of Cambodia. Biol. Conserv. Critical Ecosystem Partnership Fund (CEPF) (2007) Indo-Burma Biodiversity Hotspot, Indochina Region: Ecosystem Profile. Critical Ecosystem Partnership Fund. Arlington, USA: Critical Ecosystem Partnership Fund, del Hoyo, J., Elliott, A. & Sargatal, J., eds. (1996) Handbook of the birds of the world, 3. Barcelona: Lynx Edicions. Eames, J. C. (201 1) Feeding ecology of Giant Ibis Thaumatibis gigantea, Stung Treng province, Cambodia. BirdingASIA 1 6: 98-1 00. Environmental Systems Research Institute (ESRI) (2010) ArcMap 9.3. Redlands, C.A.: ESRI. Frederick, P. C. & Bildstein, K. L. (1 992) Foraging ecology of seven ibis species of Neotropical ibises (Threskiornithidae) during the dry season in the Llanos of Venezuela. Wilson Bull. 104: 1-21. Keo, O. (2008a) The ecology and conservation of Giant Ibis in northern Cambodia. Unpublished Ph.D. thesis, University of East Anglia. Keo, O. (2008b) Ecology and conservation of the Giant Ibis Thaumatibis gigantea in Cambodia. BirdingASIA 9: 100-106. Kushlan, J. A. (1979) Feeding ecology and prey selection in the White Ibis. Condor 81 : 376-389. Ministry of Planning (2007) Commune Database Version 6.0. Phnom Penh: Ministry of Planning. Piersma,T., Verkuil, Y. &Tulp, I. (1994) Resources for long-distance migration of knots Calidris canutus islandica and C. c. canutus: how broad is the temporal exploitation window of benthic prey in the western and eastern Wadden Sea? Oikos 71 : 393-407. Seng, K. H„ Pech, B., Poole, C. M., Tordoff, A. W., Davidson, P. & Delattre, E. (2003) Directory of Important Bird Areas in Cambodia: key sites for conservation. Phnom Penh: Department of Forestry and Wildlife, Department of Nature Conservation and Protection, BirdLife International in Indochina and Wildlife Conservation Society Cambodia Program. Smith, P. G. (2005 ) Compos Analysis, version 6.2 plus. Abergavenny, UK: Smith Ecology Ltd. Songer, M. (2006) Endangered dry deciduous forests of Upper Myanmar (Burma): a multi-scale approach for research and conservation. Unpublished Ph.D. thesis, University of Maryland. Sozer, R. & van der Heijden, A. J. W. J. (1997) An overview of the distribution, status, and behavioural ecology of the white-shouldered ibis in East Kalimantan. Kukila 9: 1 26-140. Thewlis, R. M. & Tim mi ns, R. J. (1 996) The rediscovery of Giant Ibis Pseudibis gigantea with a review of previous records. Bird Conserv. Internatn. 6: 317-324. Thuon, T. & Chambers, M. (2006) Situation analysis: Stung Treng province, Cambodia. Vientiane: Mekong Wetlands Biodiversity Conservation and Sustainable Use Programme. Timmins, R. (2008) Birds. Pp. 53-80 in M. R. Bezuijen, R. Timmins &T. Seng, eds. Biological surveys of the Mekong River between Kratie and Stung Treng towns, northeast Cambodia, 2006-2007. Phnom Penh: WWF Greater Mekong - Cambodia Country Programme, Cambodia Fisheries Administration and Cambodia Forestry Administration. Tordoff, A. W., Tim mins, R., Maxwell, A., Huy Keavuth, Lie Vuthy & Khou Eang Hourt (2005) Biological assessment of the central Indochina Dry Forest Ecoregion. Phnom Penh: WWF Cambodia Programme. Ward, J., McCafferty, D. J., Houston, D. C. & Ruxton, G. D. (2008) Why do vultures have bald heads? The role of postural adjustment and bare skin areas in thermoregulation./. Therm. Biol. 33: 168-173. Wildlife Conservation Society (WCS) (2009) List of animals on the IUCN Red List found in Cambodia. Phnom Penh: Wildlife Conservation Society - Cambodia Program. 100 H. L. WRIGHT etal. Forktail 28 (2012) Wright, H. L. (in press) Synanthropic survival: low-impact agriculture and white-shouldered ibis conservation ecology. Unpublished Ph.D. thesis, University of East Anglia. Wright, H. L„ Buckingham, D. L. & Dolman, P. M. (2010) Dry season habitat use by critically endangered white-shouldered ibis in northern Cambodia. Anim. Conserv. 1 3: 71-79. Wright, H. L„ Collar, N. J., Lake, I. R., Net Norin, Rours Vann, Sok Ko, Sum Phearun & Dolman, P. M. (2012a) First census of white-shouldered ibis Pseudibis davisoni reveals roost-site mismatch with Cambodia's protected areas. Oryx 46: 236-239. Wright, H. L„ Lake, I. R. & Dolman, P. M. (201 2b) Agriculture — a key element for conservation in the developing world. Conserv. Lett. 5: 11-19. H. L. WRIGHT, School of Environmental Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, United Kingdom. Email: hugh.wright@uea.ac.uk N. J. COLLAR, BirdLife International, Wellbrook Court, Girton Road, Cambridge, CB3 0NA, United Kingdom. Email: nigel.collar@birdlife.org I. R. LAKE, School of Environmental Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, United Kingdom. Email: i.lake@uea.ac.uk BOU Vorsak, BirdLife International in Indochina Cambodia Programme, #9, Street 29, Tonle Basac, Chamkarmon, Phnom Penh, Cambodia. Email: vorsak@birdlifecambodia.org P. M. DOLMAN, School of Environmental Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, United Kingdom. Email: p.dolman@uea.ac.uk FORKTAIL 28 (2012): 101-106 Survey of the avifauna at Muong Nhe Nature Reserve, Dien Bien province, Vietnam LE MANH HUNG, MARK B. ROBBINS, NATHAN H. RICE & ERICK A. GARGA-TREJO We documented 1 98 species within the Muong Nhe Nature Reserve in extreme north-western Vietnam during March-April 201 1 .The first Vietnam records were confirmed for three warbler species (Phylloscopidae), including the Sakhalin Leaf Warbler P. borealoides, for which there are very few records for South-East Asia. The status and distribution for a number of other species were clarified for this poorly known area of Vietnam. The Muong Nhe Nature Reserve and the contiguous Phou Dendin NPA in Laos are imperative to the continued presence of the relatively high biodiversity in this region. Not only are these reserves essential to resident species, but they are an important stopover for migrant birds. INTRODUCTION Human pressure on the Indochina landscape continues unabated and much of the forest has disappeared and what little remains is heavily impacted (see Sodhi et al. 2004; also Google Earth). For example, the following rates of deforestation occurred in Vietnam (-99.7%), Laos (-24.1%), and Cambodia (-22.0%) between the periods of 2000-2005 and 2005-2010 (http:// news.mongabay.com/2010/1006-fao_forest_cover.html). Most of the megafauna (elephants, large cats, primates, large hornbills) has been eliminated and the status and distribution of the remaining fauna and flora have been severely altered (Sodhi & Brook 2006). Thus, there is an urgency to document and preserve the remaining biodiversity before it is further eroded. Into the 1970s, the Muong Nhe Nature Reserve, located in the extreme north-western corner of Vietnam abutting Laos and China, still maintained a sizeable Asian Elephant Elephas maximus population, as well as other large mammals (Cox et al. 1992, Tu et al. 2001). However, by the 1980s most of that fauna had disappeared. By 1990, less than 20% of the reserve was forested, and much of that was secondary (Cox et al. 1992, Hill et al. 1997). The reserve now consists of only c.46,000 ha of which c. 26, 000 ha is mature forest that is in a strictly protected zone and c. 20, 000 ha that is considered a forest recovery area (Dien Bien Agriculture Department 2008). Most of the moist evergreen forest that remains is degraded, with the subtropical moist evergreen forest on the steep slopes and ridges being the least impacted vegetation in the reserve. The reserve is highly mountainous, average height c. 1,200 m, with Mount Phu Nam Man reaching 2,124 m (Tu et al. 2001). In part because of its remoteness, the reserve has received minimal biotic assessment. For example, the avifauna has had only two preliminary inventories, one of which was concentrated at the southern end of the reserve (Hill et al. 1997) and the other a very short survey during October 2000 (Tu et al. 2001). In order to provide a more complete inventory of the avifauna we worked the northern sector of the reserve from mid-March to mid-April 2011 when migration was occurring and most of the species were initiating breeding, i.e., period of increased vocalisations, and thus detectability was increased. STUDY AREA AND METHODS At the northern end of the reserve, two sites (22.386°N 1 02.238°E, 850 m; 22.26 1°N 102.224°E, 1,000 m) c. 3 km apart, were surveyed from 18 March- 13 April 2011. The first site was worked from 18 March- 1 April and 12-13 April and the second from 1 April- 1 1 April. The first site was at a guard station along a narrow, recently paved road that transects the reserve and leads to the China border. From the station we worked along the road and various trails into scrub and secondary forest. Agricultural fields, scrub, young secondary forest, and taller, continuous secondary forest (along a ridge c.l km to the south-west) were surveyed. Mist-net effort (12 m nets, maximum of 26 nets at first site; maximum of 25 nets at the second camp; opened for daylight hours only, usually from 06h00 to 17h00), observations, and audio recordings were made daily and were concentrated within 2-3 km of each camp. At the first site, nets were placed in a variety of habitats, including across streams and at edges of agricultural fields and secondary forest, and 1 4 were placed along a ridge in tall, continuous secondary forest. At the second site, a tented camp established along a forest stream, all nets were placed in taller, continuous selectively logged forest, with three nets placed across a closed-canopy forest stream. On 18 March, and for at least the previous three consecutive days, it rained and was unseasonably cool (local people, pers. comm.). As a result of these cool conditions, House Swifts Apus affinis appeared to have gone into torpor with several recovered dead or in severe weakened condition on 17 March at the base of buildings in the town of Muong Nhe and at the reserve headquarters (KUBI 23361-4). On 25-27 March, it rained for periods of several hours (primarily at night) with relatively strong winds that were primarily out of the east. It was overcast through the afternoon of 31 March. As a result of precipitation, most streams had running water. Temperature low for nights at the first camp was 53°F on 27 March. During work at the second camp the days were mostly sunny with a prolonged, mid-morning rain only on 9 April. MBR’s digital audio recordings are available online at the Macaulay Library (ML), Laboratory of Ornithology, Cornell University, USA. Specimens are deposited at KUBI (Kansas University Biodiversity Institute) and data are available online via ORNIS, a distributed database. Taxonomy and nomenclature primarily follow Robson (2008), but we have adopted more recent updates (e.g. Moyle et al. 2012) that clearly provide a more thorough perspective on nomenclature and generic and species relationships. RESULTS AND DISCUSSION We recorded a total of 198 avian species of which 85% were documented with concrete evidence, i.e. specimen, audio recording and/ or photographs (Appendix) . The Muong Nhe Reserve species list prior to our survey (163 species) was based on two inventories that involved sight observations only: one conducted at the south end of the reserve by Hill et al. in 1997 (n=158 species) and the other a short ‘rapid’ survey in October 2000 (Tu et al. 2001; n=102 species). The latter effort added five species to the earlier inventory. Combining results from the two earlier inventories and our survey the Muong Nhe list now stands at 232 species (34 species were recorded during the earlier surveys that we did not encounter). In addition to the 35 new species for the Muong Nhe list, we 102 LE MANH HUNG etai Forktail 28 (201 2) documented the first confirmed Vietnam records for three warbler species (Hartert’s Leaf Warbler Phylloscopus goodsoni, Sakhalin Leaf Warbler P. borealoides and Martens’s Warbler Seicercus omeiensis) and we clarified the status for several other species in this region (see species accounts). During our survey, most species were vocalising; specimen data and behaviour confirmed that many species were in the initial stages of breeding. The facts that our inventory occurred at the beginning of the breeding season, was longer in duration, accessed the northern section of the reserve, and included, albeit briefly, elevations up to c. 1,500 m, probably explain why we recorded a greater number of species than the earlier surveys. Access to a ridge at 1,200-1,500 m revealed submontane taxa such as Rufous-throated Partridge Arborophila rufogularis and Blyth’s Shrike Babbler Pteruthius aeralatus that we did not record elsewhere and which went unrecorded during the earlier surveys. At least 34 species of migrant (defined here where at least some individuals of a species were migrating, although some individuals may be summer or permanent residents at this site) were recorded (Appendix). As a result of a mostly unimpeded view of the horizon at our first camp, we noted small numbers of migrant raptors moving north on several days. Dramatic changes in the relative abundance, undoubtedly related to migration, of Phylloscopus species and the Grey-headed Canary-flycatcher Culicicapa ceylonensis were noted in ridge secondary forest at our first camp. Given the degree of deforestation outside the MuongNhe Reserve, we presume that this site is a now a key stopover for migrants. Despite the common presence of ground-level snares, all three Arborophila species were frequently heard and occasionally flushed. Grey Peacock-pheasant Polyplectron bicalcaratum was frequently heard, especially at the second camp, but only one group of Silver Pheasant Lophura nycthemera was observed (second camp). In addition to snares, we discovered unauthorised mist-nets in the forest and observed people shooting birds (primarily drongos Dicrurus ) with sling-shots at our first camp. Buffalo were ubiquitous at the first site and there were recent signs of this species at the second. No large native mammals were recorded at the first site, but a troop of Assam Macaque Macaco, assamensis were encountered just above 1,200 m at the second camp (ML 169313). Surprisingly, no Turdus or Zoothera thrushes have yet been recorded for MuongNhe, although several species undoubtedly occur at least as migrants if not also as winter residents. Two Zoothera species have been recorded in the contiguous Phou Dendin National Protected Area (NPA) in Laos (Fuchs et al. 2007). We were discouraged that parakeets were non-existent — Red-breasted Parakeet Psittacula alexandri was recorded by Tu et al. (2001) — and only a single hornbill species was recorded and this only at higher elevations in the more remote area that we surveyed; we suspect that these taxa had suffered the combined effects of the removal of large trees and of hunting/ trapping. The absence or low density of large woodpeckers may be a reflection of the lack of large dead trees as such trees were scarce and presumably had been used for firewood near our first camp. Additional inventories are needed as we surveyed only a small portion of the reserve and the following areas were not investigated: the uppermost montane areas (above 1,500 m), lower elevations near the park entrance (c.700 m), and the primary forest near the Laos border where the highly endangered Northern White-cheeked Gibbon Nomascus leucogenys is found. Surveys in those areas and during other seasons are needed to document both resident, migrant and winter residents that use this reserve. CONSERVATION PRIORITIES Muong Nhe Reserve and the adjacent Phou Dendin NPA in Laos are imperative to the continued presence of the relatively high biodiversity in this region. Indeed, we presume proper protection in these reserves is critical to the endangered Northern White¬ cheeked Gibbon. It is likely that the core sections, along the border of each country, of both reserves are the only areas where the gibbon, other large mammals, the Great Hornbill Buceros bicornis and perhaps even the Globally Threatened Rufous-necked Hornbill Aceros nipalensis may persist. This Aceros may still have been present in Phou Dendin NPA during 2004-2005 (Fuchs et al. 2007), and efforts should be made to ascertain if the species still occurs in the upper elevations of the western boundary of Muong Nhe. Not only are these reserves essential to resident species (permanent, summer or winter), but they certainly are an important stopover for migrant birds. As one would expect given that they are contiguous, there is extensive overlap in species composition between MuongNhe and Phou Dendin NPA; however, each reserve also has a unique set of species. These reserves should be treated as a single ecosystem with coordinated international conservation efforts. SPECIES ACCOUNTS Blyth's Kingfisher Alcedo Hercules We recorded at least three individuals (photographed) along the closed-canopy stream at our second site. This species is considered Near Threatened by BirdLife International because of its low population size and narrow habitat requirements. Fuchs et al. (2007) encountered the species in Phou Dendin NPA in abutting Laos and surmised that the population was large and continuous between these two reserves. Rufous-bellied Niltawa Niltava sundara The extreme plumage similarity in both males and females among N. sundara, the Fujian N. davidi and Vivid N. vivida niltavas, coupled with all three species being at least partially migratory, has obfuscated the status and distribution of these taxa in northern Indochina. At Muong Nhe we recorded only sundara (5 males, 2 females). Interestingly, one of the males (KUBI 119665) had somewhat enlarged testes, 5.5 x 2.5 mm and enlarged seminal vesicles, which may indicate breeding, although none of the other specimens had enlarged gonads. No vocalisations were detected, although it is easy to overlook this species’s inconspicuous song and calls. According to recent literature (Dyrcz 2006, Robson 2008), the nearest known breeding range of sundara is in Yunnan and Guizhou. However, there are four audio recordings of song by D. Edwards from Mount Fansipan, Vietnam, during May 2010 (www.xeno-canto.org; xc 65076-9). Moreover, MBR audio-recorded (ML 167779) a territorial male sundara on 29 March 2012 at c. 1,600 m, Van Ban Nature Reserve, Lao Cai province, Vietnam (22.949°N 104.255°E) and specimens from there had enlarged testes (unpubl. data). Pale Blue Flycatcher Cyornis unicolor This species was an uncommon breeder at MuongNhe. At 850 m, a mist-netted female had an unshelled egg in the oviduct on 27 March (KUBI 119363) and at 1,000 m a mist-netted male on 5 April had enlarged testes (KUBI 119648). In addition, males were persistently singing from > 8 m above the ground at c. 1,500 m (ML 169212-3). Neither Dyrcz (2006) nor Robson (2008) lists this as a breeder for West Tonkin. Chestnut-tailed Starling Sturnus malabaricus Flocks ranging in size from 4 to 25 individuals were recorded daily and photographed in agricultural fields and at the edge of secondary forest from the park entrance at c.850 m up to above the roadside guard station ate. 1,000 m. Robson (2008) does not list the species for West Tonkin. Forktail 28 (2012) Survey of the avifauna at Muong Nhe Nature Reserve, Dien Bien province, Vietnam 103 Pied Bushchat Saxicola caprata At least five (four males and one female) were seen and photographed in open areas with grass at the northern boundary of the reserve. Robson (2008) did not list this species for West Tonkin. Ashy Minivet Pericrocotus divaricatus Although this was the least common minivet, it was seen and photographed at both of our sites. Apparently these are the first records for West Tonkin (Robson 2008). Rosy Minivet Pericrocotus roseus Robson (2008) mentioned that the status of this minivet was uncertain in West Tonkin. We found the species to be common, with multiple groups, ranging from 4 to c. 1 0 individuals. Although most were seen in what appeared to be small-sized, monospecific flocks, one male (KUBI 1 19322) had enlarged testes, 8x4 mm, but two females did not have enlarged ova; so whether this species breeds in the region is still unresolved. Phylloscopus and Seicercus Species in these two genera are notoriously difficult to identify. Because of the extreme similarity in morphology among taxa of both genera, often voice and genetics are the only means of confirming identification. During our Muong Nhe fieldwork multiple taxa of these two groups were encountered, but very little song was heard or recorded. Thus, we relied on genetic analyses (sequence data from the the cytochrome b gene; sequences pending on Genbank) from specimens to provide unequivocal identifications (Appendix). As a result of the difficulty in identification both in the field and in the hand, the status and distribution of these taxa are poorly known. This underscores the importance of collecting data-rich specimens to accompany genetic and vocal sampling. Below we summarise our findings from Muong Nhe. Six species of Phylloscopus were recorded, including two that apparently are new for Vietnam: Hartert’s Leaf Warbler Phylloscopus goodsoni (n=4 specimens), and a single Sakhalin Teaf Warbler P. borealoides. Hartert’s and Claudia’s Warblers P. claudiae (n=2 specimens) are recently recognised species of the Blyth’s Warbler P. reguloides complex (Olsson et al. 2005). Phylloscopus goodsoni also has been confirmed, via genetic means, to have occurred during March 2005 in the abutting Phou Dendin NPA (Fuchs et al. 2007), and there is specimen documentation for P. claudiae from Ha Giang province, Vietnam (Vogel et al. 2003). The winter and migration status and distribution of the range-restricted P. borealoides is especially poorly known (Bairlein 2006, Brazil 2009) and our specimen (KUBI 119459) apparently represents the first for Vietnam and one of the very few for South-East Asia as a whole. Finally, our documentation (ML 169180) of territorial Yellow-vented Warbler P. cantator appears to represent the first breeding record for West Tonkin (Robson 2008); Fuchs etal. (2007) did not find this warbler during March surveys in adjacent Phou Dendin NPA. We recorded three species of Seicercus , all of which are known from this general region, but for which specimen documentation is apparently lacking for Vietnam (Martens et al. 1999, 2003, Bairlein 2006). Of ten Seicercus netted, eight were Martens’s Warbler S. omeiensis , one (plus another individual audio recorded, ML 1691 62) Bianchi’s Warbler S. valentini, and one Grey-crowned Warbler S. tephrocephalus. The latter two have been reported for Vietnam based on audio recordings deposited on xeno-canto. ACKNOWLEDGEMENTS We are grateful to Muong Nhe Reserve personnel, especially Tran Xuan Tarn, Nguyen Viet Cuong, Dao Cong Tien and Nguyen Quoc Tang. We thank Charles Linkem for his assistance and camaraderie. Pete Hosner and Arpad Nyari kindly provided sequence data for identifying Phylloscopus and Seicercus specimens. Craig Robson identified several species in audio recordings. Paul Sweet reconfirmed identification of Ha Giang province Phylloscopus and two anonymous reviewers provided valuable comments on the manuscript. REFERENCES Bairlein, F. (2006) Family Sylviidae (Old World warblers). Pp.492-709 in J. del Floyo, A. Elliott & D. A. Christie, eds. Handbook of the birds of the world, 1 1 . Barcelona: Lynx Edicions. Brazil, M. (2009) Birds of East Asia. Oxford: Oxford University Press. Cox, C. R., Vu Van Dung & Pham Mong Giao (1992) Report of a management feasibility study of the Muong Nhe Nature Reserve (November/ December 1991). Flanoi: World Wildlife Fund/Ministry of Forestry. Dien Bien Agriculture Department (2008) A revision of the investment plan for the period from 2008-2020 of Muong Nhe Nature reserve. Technical Report to Dien Bien Agriculture Department. Dyrcz, A. (2006) Family Muscicapidae (Old World flycatchers). Pp. 148-163 in J. del Floyo, A. Elliott & D. A. Christie, eds. Handbook of the birds of the world, 1 1 . Barcelona: Lynx Edicions. Fuchs, J., Cibois, A., Duckworth, J. W., Eve, R., Robichaud, W. G., Tizard, T. & Gansberghe, D. V. (2007) Birds of Phongsaly province and the Nam Ou river, Laos. Forktail 23: 22-86. Hill, M„ Hallam, D. & Bradley, J. (1997) Site study; Muong Nhe Nature Reserve, Lai Chau Province, Vietnam. SEE-Vietnam Research Report 4. London: Society for Environmental Exploration and the Institute of Ecology and Biological Resources. Martens, J., Eck, S., Packert, M. & Sun, Y. H. (1999) The Golden-spectacled Warbler Sericercus burkii - a species swarm (Aves: Passeriformes: Sylviidae). Part I .Zoo/. Abh. Mus. Tierk. Dresden 50: 281-327. Martens, J., Eck, S., Packert, M. & Sun, Y. H. (2003) Methods of systematic and taxonomic research on passerine birds: the timely example of the Seicercus burkii complex (Sylviidae). Part 2. Bonn. Zool. Beitr. 51 : 1 09-1 1 8. Moyle, R. G., Andersen, M. J., Oliveros, C. H., Steinheimer, F. D. & Reddy, S. (2012) Phylogeny and biogeography of the core babblers (Aves: Timaliidae). Systematic Biology 61 : D0l:10.1093/sysbio/sys027. Olsson, U., Alstrom, P„ Ericson, P. G. P. & Sundberg, P. (2005) Non-monophyletic taxa and cryptic species - evidence from a molecular phylogeny of leaf- warblers ( Phylloscopus , Aves). Molec. Phylogen. Evol. 36: 261-276. Robson, C. (2008) A field guide to the birds of South-East Asia. London: New Holland. Sodhi, N. S. & Brook, B. W. (2006) Southeast Asian biodiversity in crisis. Cambridge, UK: Cambridge University Press. Sodhi, N. S„ Koh, L. P„ Brook, B. W. & Ng, P. K. L. (2004) Southeast Asian biodiversity: an impending disaster. Trends EcoL Evol. 19: 654-660. Tu, Nguyen Due, Le Trong Trai, & Le Van Cham (2001 ) A rapid field survey of Muong Nhe Nature Reserve, Lai Chau Province, Vietnam. BirdLife International, European Union, FIPI. Vogel, C. J., Sweet, P. R„ Le Manh Hung & Hurley, M. M. (2003) Ornithological records from Ha Giang province, north-east Vietnam, during March-June 2000. Forktail 19:21-30. LE Manh Hung, Insitltutue of Ecology and Biological Resources, 18 Hoang Quoc Viet, Cau Giay, Hanoi, Vietnam. Email: Hungniltava@gmail.com Mark B. ROBBINS, University of Kansas Biodiversity Institute, 1345 Jay hawk Blvd., Lawrence, Kansas, 66045. U.S.A. Nathan H. RICE, Academy of Natural Sciences, 1 900 Benjamin Franklin Parkway, Philadelphia, Pennsylvania 19103. U.S.A. Erick A. GARCIA-TREJO, Museo de Zoolog la Alfonso L. Herrera', Departmento de Biologla Evolutiva, Apartado postal 70-399, Mexico D.F. 045 1 0, Mexico 104 LE MANH HUNG etoi Forktail 28 (2012) Appendix Bird species recorded in Muong Nhe Reserve, 1 8 March-1 3 April 201 1 English name Scientific name Relative abundance Documentation Comments English name Scientific name Relative abundance Documentation Comments Rufous-throated Partridge Arborophila rufogularis U V Large-tailed Nightjar Caprimulgus macrurus F S Scaly-breasted Partridge Arborophila chloropus F * Indian Nightjar Caprlmulgus asiaticus R S Red Junglefowl Callus gallus U V Oriental Turtle-dove Streptopelia orientals U V.P Silver Pheasant Lophura nycthemera X s Spotted Dove Streptopelia chinensis u S.P Grey Peacock-Pheasant Polyplectron bicalcaratum F V Barred Cuckoo-dove Macropygia unchall R S.P Speckled Piculet Picumnusinnominatus U * Emerald Dove (halcophapsindica R * White-browed Piculet Sasia ochracea U * Thick-billed Green-pigeon Treron curvirostra F V.P Grey-capped Pygmy Woodpecker Dendrocopos canicapillus U V.P Mountain Imperial-pigeon Ducula badia R S.P Rufous Woodpecker Micropternus brachyurus X V Black-tailed Crake Porzana bicolor R s Greater Yellownape Plcus flavinucha R S,P Oriental Honey-buzzard Pernisptilorhynchus U S.P migrant Grey-headed Woodpecker Plcus canus R s Crested Serpent-eagle Spilornis cheela U V,p Greater Flameback Chrysocolaptes lucidus R s Crested Goshawk Accipiter trlvirgatus R p migrant Pale-headed Woodpecker Gecinuius grantia R V Grey-faced Buzzard Butastur indlcus U S.P migrant Bay Woodpecker Blythipicus pyrrhotis F * Himalayan Buzzard Buteo burmanicus R S.P migrant Blue-throated Barbet Megalalma asiatica F V.P Common Kestrel Falco tinnunculus R S.P Golden-throated Barbet M. frankllnii F V.P Chinese Pond-heron Ardeola bacchus X s Oriental Pied Hornbill Anthracoceros albirostris R V Little Heron Butorides striatus R S.P Common Hoopoe Upupa epops X S migrant Silver-breasted Broadbill Serilophuslunatus F * Red-headedTrogon Harpactes erythrocephalus U * Long-tailed Broadbill Psarisomus dalhousiae F * Blyth's Kingfisher Alcedo hercules U P Asian Fairy-bluebird Irena puella F V,p Common Kingfisher Alcedo atthis U * Blue-winged Leafbird Chloropsis cochinchinensis F V,P White-throated Kingfisher Halcyon smyrnensls U S.P Orange-bellied Leafbird Chloropsishardwickii F V Crested Kingfisher Megacerylelugubris X s Brown Shrike Lanius cristatus U S.P migrant Blue-bearded Bee-eater Nyctyornls athertoni U p Burmese Shrike Lanius collurioides R S.P Large Hawk-cuckoo Hlerococcyx sparverioides U * Long-tailed Shrike Lanius schach R S.P migrant Hodgson's Hawk-cuckoo Hierococcyx nisi color U V Grey-backed Shrike Lanius tephronotus R S.P migrant Indian Cuckoo Cuculus micropterus U V Common Green Magpie Cissa chinensis R V Eurasian Cuckoo Cuculuscanorus X V migrant Grey Treepie Dendrocitta formosae R * Banded Bay Cuckoo Cacomantis sonneratii X V Large-billed Crow Corvus japonensis U V.P Plaintive Cuckoo Cacomantismerulinus X S.P AshyWoodswallow Art am us fuscus U S.P Asian Emerald Cuckoo Chrysococcyx maculatus R S Maroon Oriole Oriolus traillii F V.P Drongo Cuckoo Surniculuslugubris F V.P Large Cuckooshrike Coracina macei X V Green-billed Malkoha Phaenicophaeus tristis u V.P Black-winged Cuckooshrike Coracina meiaschistos F V.P Greater Coucal Centropus sinensis u V.P Rosy Minivet Pericrocotus roseus C * migrant Himalayan Swiftlet C ollocalla brevirostris c S.P Ashy Minivet Pericrocotus divaricatus U S.P migrant Asian Palm-swift Cypsiurus balasiensis R s Scarlet Minivet Pericrocotus speciosus c V.P Silver-backed Needletail Hirundapus cochinchinensis F s Bar-winged Flycatcher-shrike Hemipus picatus u * House Swift Apus afflnis C * White-throated Fantail Rhipidura albicollis F * Mountain Scops Owl Otus spilocephalus R V Ashy Drongo Dicrurus leucophaeus c * Collared Scops Owl Otus lettia R V,* Bronzed Drongo Dicrurus aeneus F * Brown Wood-owl Strix leptogrammica R s Lesser Racket-tailed Drongo Dicrurus remifer U * Collared Owlet Claucidium brodiel U s Spangled Drongo Dicrurus hottentottus u S.P Asian Barred Owlet Glaucidlum cuculoides U V.P Greater Racket-tailed Drongo Dicrurus paradiseus u * Brown Boobook N inox scutulata R s Black-naped Monarch Hypothymis azurea F * Forktail 28 (2012) Survey of the avifauna at Muong Nhe Nature Reserve, Dien Bien province, Vietnam 105 Relative Relative English name Scientific name abundance Documentation Comments English name Scientific name abundance Documentation Comments Asian Paradise-flycatcher Terpsiphoneparadisi C * Grey-eyed Bulbul lolepropingua R S Common lora Aegithina tiphia U V Ashy Bulbul Hemixos flavala F * Large Woodshrike Tephrodornisgularis U V Black Bulbul Hypsipetes leucocephalus F * Brown Dipper Cincluspallasii R * Rufescent Prinia Prinia rufescens F * Blue Rock-thrush Monticolasolitarius R s migrant Hill Prinia Prinia superciliaris R S,P Blue Whistling-thrush Myophonus caeruleus R * Grey-breasted Prinia Prinia hodgsonii F * Lesser Shortwing Brachypteryx leucophrys U * Asian Stubtail Urosphena squameiceps U * migrant Brown-breasted Flycatcher Muscicapa muttui U * Dark-neckedTailorbird Orthotomus atrogularis U V Ferruginous Flycatcher Muscicapa ferruginea X s migrant Common Tailorbird Orthotomus sutorius U * Red-throated Flycatcher Ficedula parva u S,P migrant Dusky Warbler Phylloscopus fuscatus U v,p migrant White-gorgeted Flycatcher Ficedulamonileger u * Yellow-browed Warbler Phylloscopus inornatus U * migrant Snowy-browed Flycatcher Ficedula hyperythra R * Claudia's Warbler Phylloscopus claudiae 7 * migrant Little Pied Flycatcher Ficedula westermanni R * Hartert's Warbler Phylloscopus goodsoni 7 * migrant Slaty-blue Flycatcher Ficedula tricolor X * migrant Sakhalin Leaf Warbler Phylloscopus borealoides 7 * migrant Large Niltava Niltava grandis R s Yellow-vented Warbler Phylloscopus cantator R V Verditer Flycatcher Eumyias thalassinus U * Grey-crowned Warbler Seicercus tephrocephalus 7 * migrant Rufous-bellied Niltava Niltava sundara U * Bianchi's Warbler Seicercus valentini 7 * migrant White-tailed Flycatcher Cyornis concretus F * Martens's Warbler Seicercus omeiensis 7 * migrant Hainan Blue Flycatcher Cyornishainanus C * Rufous-faced Warbler Abroscopus albogularis R V Pale Blue Flycatcher Cyornis unicolor U * Yellow-bellied Warbler Abroscopus superciliaris F V,p Hill Blue Flycatcher Cyornis banyumas C * Chestnut-capped Babbler Timaliapileota X * Grey-headed Canary-flycatcher Culicicapa ceylonensis C * migrant Pin-striped Tit-babbler Mixornis gularis C * Yellow-bellied Fantail Chelidorhynx hypoxantha X s Rufous-fronted Babbler Cyanoderma rufifrons F * Siberian Rubythroat Luscinia calliope u * migrant Golden Babbler Cyanoderma chrysaeum R s Oriental Magpie-robin Copsychussaularis ' u * Streak-breasted Scimitar Babbler Pomatorhinus ruficollis X s White-rumped Shama Copsychus malabaricus R Orange-billed Scimitar Babbler Pomatorhinus ochraceiceps U * White-capped Water-redstart Chaimarrornis leucocephalus U * Large Scimitar Babbler Megapomatorhinus hypoleucos R V Plumbeous Water Redstart Rhyacornis fuliginosus U * Grey-throated Babbler Stachyris nigriceps F * White-tailed Robin Myiomela leucura F * Collared Babbler Gampsorhynchus torquatus U * Slaty-backed Forktail Enicurusschistaceus U * Rusty-capped Fulvetta Schoeniparusdubius U s White-crowned Forktail Enicurusleschenaulti R * Puff-throated Babbler Pellorneum ruficeps U * Pied Bushchat Saxicola caprata U S,P Spot-throated Babbler Pellorneum albiventre R V Siberian Stonechat Saxicola mourn U S,P migrant Buff-breasted Babbler Pellorneum tickelli C * Grey Bushchat Saxicola ferrea U V migrant Eyebrowed Wren Babbler Napothera epilepidota U s Chestnut-tailed Starling Sturnus malabaricus F V,p Streaked Wren Babbler Napothera brevicaudata R * Chestnut-bellied Nuthatch Sitta cinnamoventris X V Brown-cheeked Fulvetta Alcippepoioicephala C * Velvet-fronted Nuthatch Sitta frontalis U V,P David's Fulvetta Alcippe davidi F * Japanese Tit Parus minor F -* White-crested Laughingthrush Garrulaxleucolophus F * Sultan Tit Melanochlorasultanea X V Lesser Necklaced Laughingthrush Garrulaxmonileger R V Barn Swallow Hirundo rustica U s migrant Chinese Hwamei Garrulax canorus X V,p Red-rumped Swallow Hirundo daurica u s migrant Black-throated Laughingthrush lanthocincla chinensis U * Striated Swallow Hirundo striolata c * migrant Blue-winged Minla Actinodura cyanouroptera R s Black-crested Bulbul Pycnonotus flaviventris u * Chestnut-collared Yuhina Yuhinatorqueola F V Red-whiskered Bulbul Pycnonotus jocosus c * Black-chinned Yuhina Yuhina nigrimenta U V Sooty-headed Bulbul Pycnonotus aurigaster c * Oriental White-eye Zosterops palpehrosus R * Puff-throated Bulbul Alophoixuspallidus u * Japanese White-eye Zosterops japonicus F * 106 LE MANH HUNG etal. Forktail 28 (2012) English name Scientific name Relative abundance Documentation Comments White-bellied Erpornis Erpornis zantholeuca F * Blyth's Shrike-babbler Pteruthius aeralatus F V Yellow-vented Flowerpecker Dicaeum chrysorrheum X S Plain Flowerpecker Dicaeum concolor C V,P Scarlet-backed Flowerpecker Dicaeum cruentatum F s Mrs Gould's Sunbird Aethopyga gouldiae R * Black-throated Sunbird Aethopyga saturata F * Crimson Sunbird Aethopyga si pa raj a F * Purple-naped Sunbird Hypogramma hypogrammicum R s Streaked Spiderhunter Arachnothera magna F * Eurasian Tree Sparrow Passer montanus F s English name Scientific name Relative abundance Documentation Comments White Wagtail Motacilla alba F S,P migrant; a Iboides subspecies Grey Wagtail Motacilla cinerea F * Olive-backed Pipit Anthushodgsoni F * migrant White-rumped Munia Lonchura striata C * Common Rosefinch Carpodacus erythrinus U * Yellow-breasted Bunting Emberiza aureola X S migrant; 2 females, 29 March 2011 Crested Bunting Melophus iathami F V,p Relative abundance criteria:C = common; more than 20 individuals/day; F = fairly common; 5-20 individuals/day; U = uncommon; present in small numbers (<5 individuals/day); R = rare; only occasionally encountered in small numbers; X= single record. Documentation: Collected = *; P = photographed; V = voice recorded; S = sight record only FORKTAIL 28 (2012): 107-112 The avifauna of Lambusango Forest Reserve, Buton Island, south-east Sulawesi, with additional sightings from southern Buton T. E. MARTIN, D. J. KELLY, N.T. KEOGH, D. HERIYADI, H. A. SINGER & G. A. BLACKBURN Lambusango Forest Reserve occupies a large area of south-central Buton, the largest attendant island of Sulawesi, Indonesia. Buton is located off Sulawesi's south-eastern peninsula and remains poorly known ornithologically. Bird surveys were undertaken in the reserve over eight eight-week long research seasons between June and August in 1999, 2001-2003, 2005, and 2008-2010. Variable radius circular- plot point counts were the primary census method, conducted as part of a long-term biodiversity monitoring programme in the reserve, although data were also collected from 840 mist-netting hours and approximately 2,560 hours of observational data. In total, 79 species were detected in the reserve, including 37 regional endemics (46.8% of the total avifaunal community) and four species considered by the IUCN to be globally threatened or Near Threatened. Additionally, a further 60 species (including two more Near Threatened species) were recorded in various habitats around southern Buton that were not detected in Lambusango Reserve, giving a total of 1 39 species records for the island. We believe that 51 of these species represent previously unpublished records for Buton. We present here a full account of our records from Lambusango Reserve and southern Buton, with additional details provided for threatened and Near Threatened species and new records of endemics. INTRODUCTION Buton (formerly referred to as Butung) is the largest of Sulawesi’s attendant islands, located c .6 km (at the nearest point) off the mainlands south-eastern peninsula, between the Flores and Banda Seas. The island covers an area of c. 560, 000 ha and is around 100 km long and 42 km wide at its broadest point. Altitude varies from 0 to 200 m in coastal areas to around 400 m along the island’s central spine, with isolated peaks reachingup to 1 ,000 m (O’Donovan 200 1 , Whitten et al. 2002). Precipitation follows a tropical monsoon climate with a June-September dry season and a November- April wet season. Mean annual rainfall ranges between 1,500 and 2,000 mm, peaking between April and June. Mean annual temperatures range between 25°C and 27°C (Whitten et al. 2002). The geology of the island is complex; much of the lowlands consist of uplifted karst and other limestone formations, while the mountainous interior is more varied, with sandstone, chert and ultramafic soils overlying ophiolitic rock. A large (70,000 ha) asphalt deposit, one of the most significant in South-East Asia, underlies a 60 km north-south strip of southern Buton (Whitten et al. 2002). The natural vegetation cover of much ol the island is seasonal lowland tropical forest, with mangroves occurring in coastal areas. No comprehensive survey ol the island’s flora has been completed but, as with mainland Sulawesi, the diversity of tree species is high, with no single family being predominant (O’ Donovan 2001, Whitten et al. 2002). Extensive cultivation occurs in cleared areas in the lowlands, particularly in the south of the island. Major agricultural crops include rice, maize, sweet potatoes, cassava and plantations of cashew nuts, cocoa and coconut (Priston 2005). Alang-alang’ Imperata cylindrica grassland and rough scrub occur in areas of abandonedcultivation. Elowever, large tracts of relatively undisturbed forest remain in the island’s interior. The island remains a stronghold for two species of Endangered endemic bovid: Lowland Anoa Bubalus clepressicornis and Mountain Anoa B. quarlesi. This is one of only two locations where the ranges of both species are known to overlap, with approximately 1 0% of the global population of both species believed to occur on Buton (Burton et al. 2005). Despite their ecological importance, the forests of Buton have undergone significant clearance in recent years: agricultural expansion, logging and asphalt mining activities led to a reduction in the remaining forested area by over 13% (27,809 ha) between 1991 and 2002 (Seymour 2004). Lambusango Forest Reserve (5°10'-5°24'S 122°43,-123°07'E) is a 65,000 ha expanse of uninhabited tropical monsoon forest, encompassing much of south-central Buton. The reserve was established in 1982 as part of a nationwide policy by the Suharto administration to found new national parks and reserves across Indonesia. Lambusango was selected as one of these new reserves based on its ‘potency of flora and fauna’. Its area is divided into a 28,510 ha strict forest reserve, managed by the Indonesian Natural Resources Conservation Agency (BKSDA), where all commercial, recreational and agricultural activities are nominally prohibited, and a 35,000 ha production forest, regulated by Buton District Forestry Office, where some sustainable activity such as rattan extraction is allowed (Singer & Purwanto 2006) (Figure 1). The avifauna of Buton remains poorly described. Several previous expeditions have taken specimens from the island, although few have involved any detailed survey work. The earliest account of the island’s avifauna we can find comes from the Figure 1. Lambusango Forest Reserve, showing locations of study transects. Inset shows the study area's location on Buton Island. 108 T. E. MARTIN etal. Forktail 28 (2012) specimen collections of Heinrich Kuhn, made at the turn of the twentieth century (Hartert 1903). Further specimens were collected on the island by G. L. den Haan in 1948 (van Bemmel & Voous 1951), and further species were added to the island’s list by White & Bruce (1986), Schoorl (1987) and Coates & Bishop (1997). An extensive atlas survey of the island was completed by M. Catterall during 1996-1997. This was commissioned by the research organisation Operation Wallacea to provide an initial rapid assessment ot Buton’s avitauna. The findings were compiled as an internal report which was formerly available on Operation Wallacea’s website but is no longer publicly accessible (Catterall undated. Operation Wallacea 2011). METHODS Records ol Lambusango Reserve’s avifauna were made in two ways: formal survey work; and informal observations and net captures. Formal surveys were carried out as part of the long-term biodiversity monitoring programme run by Operation Wallacea, and involved conducting 50 m-radius circular plot point counts (Bibby et al. 2002) along transects spread throughout the reserve. A total of 24 transects were used within the reserve, in six clusters of four transects each. Transects within each cluster were spaced 1 km apart. These transects were located between 100 and 700 m above sea- level and spanned a range of forest ecosystems and successional stages, including relatively undisturbed primary forest, well¬ regenerated secondary forest, disturbed secondary forest and forest- edge environments. An additional cluster of four transects was located in agricultural land around the reserve’s periphery. Land- uses here included rice-paddies, coconut, cocoa, cassava and teak plantations, and recently abandoned land. Each transect contained seven sample sites at 150 m spacing, giving a total of 196 sample points. Each point was repeated once per season, giving a total of 1,568 individual point-counts over the eight research seasons. Surveys were led by DJK and HAS during the 1999-2003 research seasons, HAS having several years’ experience with the island’s avifauna prior to this study period. Survey work in 2005 was led by HAS and TM, while surveys in the 2008-2010 seasons were led by TM and DH, who had several months’ prior field experience with avifauna in Lambusango. Three or four Operation Wallacea volunteers would usually assist with survey work. Sampling was conducted each morning between 06h00 and 08h00. Point counts lasted 10 minutes, beginning on immediate arrival at each sampling site, with all birds seen and heard within the radius being recorded during this time. Point counts were not carried out in rain or heavy mist. Informal observations were recorded whenever sightings were made outside the survey work. These included sightings made when travelling to and from survey sites and, on non-survey days, records made from casual exploration of a range of habitats inside the reserve and beyond its borders. Habitats explored in this way included all forest habitats described above, cultivated land, rough scrub, towns, rivers, mangroves and beaches, as well as the ocean passage between Buton and the Sulawesi mainland. Mist-netting was also conducted in these habitats on a casual basis whenever possible. Three 2.6 m x 20 m x 36 mm mesh mist-nets were used, which were checked every 20 minutes after opening. Netting was conducting in morning, afternoon and evening periods, with nets typically being closed around three hours after opening. A handful of records were also reported as by-catch from the mist-nets used by the bat survey team working in the same areas. Once survey work was completed, species recorded in the reserve were tabulated, with the endemism and conservation status of each species being noted. Endemism levels were classified following Stattersfield etal. (1998) and Clements (2007). Sulawesi endemics were classified as those found only on mainland Sulawesi and its satellite islands, including the Talaud, Sangihe, Togian, Banggai, Sulu and Wakatobi (Tukangbesi) island groups (Coates & Bishop 1997). Wallacean endemics were classified as those found only in the Wallacea region, bordered by Wallace’s Line in the west and Lydekker’s Line in the east (Coates & Bishop 1997). National endemics were defined as those only occurring in Indonesia. Conservation status follows the 2010 IUCN Red List (IUCN 2010). Taxonomy follows Inskipp et al. (2001). Categorical abundance estimates for each species were based on frequency of sightings. The designated categories were: abundant (usually recorded multiple times each day in suitable habitat); common (usually recorded at least once per day); fairly common (typically recorded at least once per week); locally common (usually recorded daily, but restricted to specific habitats or spatially small areas); uncommon (recorded less than five or six times in a season); and rare (known only from one or two records). We identified species as being new records for the island if no records of their presence appeared in any of the published accounts of previous visitors (Hartert 1903, van Bemmel & Voous 1 95 L Schoorl 1987), regional field guides (White & Bruce 1986, Coates & Bishop 1997), comprehensive species checklists (Clements 2007), or international databases (BirdLife International 2010, IUCN 2010, Internet Bird Collection 2011). RESULTS Our total survey effort in the study area amounted to 2,560 hours of observational data (based on formal and informal survey work) and 840 mist-netting hours (based on the number of mist-nets used multiplied by the number of hours the nets were open). A total of 79 species was recorded in Lambusango Forest Reserve, of which 37 were endemic to Wallacea (46.8% of the total avifaunal community) and a further three were Indonesian endemics (3.9%). Four species considered globally threatened or Near Threatened (IUCN 20 1 0) were detected in the reserve. An additional 60 species were recorded in various habitats around southern Buton that were not detected in Lambusango Reserve, including another eight Wallacean endemics, four Indonesian endemics, and two Near Threatened species. This gives a total of 139 species and 45 Wallacean endemics (30.9% of the total avifaunal community) recorded for southern Buton. The Appendix summarises species recorded by both formal and informal survey effort. The following annotated list provides further information concerning observations of globally threatened and Near Threatened species, and endemic species previously unrecorded on the island. Grey-headed Fish Eagle lehthyophaga ichthyaetus A widespread species found in suitable habitat across much of South and South-East Asia. Considered Near Threatened (IUCN 2010) owing to habitat loss and over-fishing. A rare species in the study area; individuals were occasional sighted along river systems and over open farmland. This species has not been reported from Buton before and our records may represent an extension of its known distribution (del Hoyo etal. 1994). Small Sparrowhawk Accipiter nanus Endemic to Sulawesi and a few offshore islands. Considered Near Threatened (IUCN 2010). A rare species in the study area, known from two positively identified individuals: one that flew into a window in the coastal village of Kakenauwe in the 2010 season; and another mist-netted in Labundo-bundo village in 2009. These are interesting records as most previous publications have considered this a bird of upland forest (500-2,000 m) (White & Bruce 1987, Coates & Bishop 1997), while both our records come Forktail 28 (2012) The avifauna of Lambusango Forest Reserve, Buton Island, south-east Sulawesi 109 from at or near sea-level. Catterall (undated) also recorded this species near sea-level during his atlas survey of the island. These are significant altitudinal records, and suggest that the species inhabits a wider range of habitats than previously thought. Our records represent an eastern range extension for this species, which has previously been reported only from mainland Sulawesi (del Hoyo et al. 1994, Coates 8c Bishop 1997). Isabeiline Bush-hen Amaurornis isabellinus Endemic to Sulawesi and a few of its attendant islands. Fairly common in the study area. Individuals recorded regularly in cultivated land, particularly paddies and irrigation systems, as well as forest edge. Previously recorded from mainland Sulawesi. Sulawesi Ground Dove Gallicolumba tristigmata Endemic to Sulawesi. A shy and inconspicuous species that is difficult to observe. Known from a single live bird found captured in a snare in 2009. This individual showed noticeably different plumage from other descriptions, having extensive green coloration on the neck, which is absent in all subspecies described by Coates & Bishop (1997). A photograph of this bird was taken on a mobile phone camera but this was unfortunately lost at a later date. This record represents a range extension for the species, which was previously known only from mainland Sulawesi (Coates & Bishop 1997, del Hoyo et al. 1997, Whitten et al. 2002). Given the presence of other locally endemic subspecies on Buton and nearby Kabaena island (Robinson-Dean et al. 2002), the taxonomy of the population on Buton may be worthy of further investigation. White-bellied imperial Pigeon Ducula forsteni Endemic to Sulawesi. Found almost exclusively in relatively undisturbed interior forest, where it is fairly common. Rare in forest edge and non-forest habitats. Usually encountered in small flocks of three to six birds, although much larger groups of up to 30 birds have been encountered around fruiting figs. Occasionally observed in mixed flocks with other imperial pigeon species. This species has previously been reported only from mainland Sulawesi and from Taliabu and Mangole in the Sulu Islands (Coates & Bishop 1997, del Hoyo et al. 1997). Yellow-crested Cockatoo Cacatua sulphurea Indonesian endemic. This species is listed as Critically Endangered owing to the threat posed by trapping for the pet trade, habitat destruction and population fragmentation. This species is rare in the study area. A single pair was observed each year between 2006 and 2009, flying between a stand of trees near coastal mangrove and the forest edge near Labundo-bundo. It is unknown whether these birds are truly wild or escaped cagebirds. This is the only Critically Endangered species in our study area. It has been previously reported as occurring on Buton (Coates & Bishop 1997). Pygmy Hanging Parrot Loriculus exilis Endemic to Sulawesi. Less common than Sulawesi Hanging Parrot L. stigmatus. More frequently observed in forest edge and adjacent cultivation than forest interior, where it is uncommon. Usually encountered singly or in pairs. This species has previously only been reported from mainland Sulawesi (Coates 8c Bishop 1997, del Hoyo etal. 1997). Sulawesi Hawk Cuckoo Hierococcyx crassirostris Sulawesi endemic. A common species frequently detected in forest and forest-edge habitats by its distinctive call, which is repeated at regular intervals throughout the night, early morning and occasionally later in the day, although it is extremely secretive and hard to observe. Previous records class this bird as an upland species occurring primarily between 500 and 1,400 m (Coates & Bishop 1997, del Hoyo etal. 1997). However, our records show it also occurs in lowland habitats, with many observations below 100 m. These new records from Buton represent a significant range extension for the species, which was previously known only from montane forest in northern and central Sulawesi (del Hoyo etal. 1997). Black-billed Koel Eudynamys melanorhyncha Endemic to Sulawesi. Apparently quite rare. Observed most frequently in the eastern side of the reserve, where flocks of up to six birds were recorded. Individuals were also encountered at locations scattered throughout the reserve. These records represent a range extension for the species, which was previously reported only from mainland Sulawesi (del Hoyo et al. 1997). Sulawesi Owl Tyto rosenbergii Endemic to Sulawesi. Rare and infrequently detected. Occasional records from edge habitats and around villages. One individual was captured in a mist-net in farmland near forest edge during the 2008 survey season. These observations represent a range extension for the species, which was previously known only from mainland Sulawesi and the Sangihe islands (del Hoyo etal. 1999). Sulawesi Scops Owl Otus manadensis Endemic to Sulawesi. The most frequently encountered owl in the study area. Fairly common in forest and forest-edge habitats and along roadsides, although it can be quite local. A single individual was also mist-netted in 2001. This species was previously known only from mainland Sulawesi (del Hoyo etal. 1999). Sulawesi Nightjar Caprimutgus ceiebensis Endemic to Sulawesi. Recorded infrequently, although probably under-recorded owing to insufficient surveying of nocturnal birds. This species was most often detected by voice in agricultural land and forest edge, with a few records coming from the forest interior. We believe this is a new record for Buton; it has previously been listed as ‘probable’ on Buton (del Hoyo etal. 1999). Moluccan Swiftlet Collocalia infuscata A Wallacean endemic occasionally seen flocking with other swiftlets over settlements and cultivated land. Previously known from mainland Sulawesi and the Moluccas (Coates & Bishop 1997). Green-backed Kingfisher Actenoides monaehus Endemic to Sulawesi. A rare species known only from a single mist- net capture in the reserve’s interior in 2005. This bird (subspecies capucinus) was netted along a river course at about 600 m altitude. This record represents a range extension for the species, which was previously known only from mainland Sulawesi (del Hoyo et al. 2001). Sulawesi Dwarf Kingfisher Ceyxfallax Endemic to Sulawesi. Considered Near Threatened (IUCN 2010). An uncommon species most frequently observed at night roosting in overhanging branches along forested river systems. Most individuals seem to utilise the same roosting spots each night. The species has previously been reported only from mainland Sulawesi and Lembeh Island (del Hoyo etal. 2001). Pied Cuckooshrike Coracina bicolor Endemic to Sulawesi. Listed as Near Threatened (IUCN 2010) owing to habitat degradation and fragmentation. A common species found throughout the reserve’s interior and forest edge, and in places locally abundant. Rarely observed in non-forest habitats. Lambusango Reserve appears to support a large population of this species. Previously recorded as occurringon Buton (Coates & Bishop 1997). 110 T. E. MARTIN etal. Forktail 28 (2012) Red-backed Thrush Zoothera erythronota Endemic to Sulawesi. A cryptic and rarely observed understorey forest species. Known from several observations of lone individuals and pairs within the reserve’s interior, and in forest-edge habitats. Previously known from mainland Sulawesi and Kabaena Island (Coates & Bishop 1997, Robinson-Dean et al. 2002). Rufous-throated Flycatcher Ficedula rufigula Endemic to Sulawesi. Considered Near Threatened (IUCN 2010). A shy, fairly cryptic species. It was rarely recorded during point- count surveys or casual observations but individuals were captured quite regularly in mist-nets in areas of mature and disturbed forest. Presumed to be severely under-recorded by our formal survey methods. The species has been previously reported only from mainland Sulawesi (Coates & Bishop 1997, del Hoyo et al. 2006). Ivory-breasted Woodswallow Artamus monachus Endemic to Sulawesi. Much less common than White-breasted Woodswallow A. leucorynchus , although still observed fairly frequently in similar habitats, particularly forest clearings. The species has been previously reported from mainland Sulawesi and the Lembeh, Banggai, and Sula island groups (Coates & Bishop 1997, del Hoyo et al. 2009). Pale-headed (Vlunia Lonchura pallida Endemic to Wallacea. A rare species that has been very occasionally observed in paddyfields and other cultivated land. It has been widely reported from other parts of Sulawesi and the Lesser Sundas (Coates & Bishop 1997, del Hoyo etal. 2010). DISCUSSION We believe our survey work in Lambusango Reserve and its surrounds represents the most detailed account of Buton’s avifauna in print, with 49 previously unpublished records for the island, each representing important range extensions of these species’ known distributions. However, while these are new published records, it is important to acknowledge that the majority of these records were also identified by Catterall (undated), based on his atlas survey of the whole island. Only five of our new records were not reported in this earlier report (Slaty-legged Crake Rallina eurizonoid.es, Sulawesi Ground Dove, Sulawesi Nightjar, Rufous-throated Flycatcher and Scarlet Myzomela Myzornela sanguinolenta). We also acknowledge that, while we have provided a detailed assessment of Lambusango Reserve and its immediate surrounds, the bulk of our survey effort has been focused on forest and forest-edge habitats. Further survey work on the island should concentrate on other habitats, such as agricultural areas, abandoned farmland, coastal areas and small offshore islands, as these could produce further unrecorded species. It may be, for example, that some of the small island species noted by Kelly & Marples (2010) in the nearby Wakatobi archipelago may also occur on Buton’s numerous small offshore islets. It would also be valuable to conduct survey work outside the June-August research period used every year in this study, allowing the recording of wintering birds and passage migrants from Australia and the Palearctic. Further survey work targeting groups that were likely under-recorded by our point-count primary survey methods, such as nocturnal birds and cryptic, non-vocal species, may also produce further valuable records. Greater effort towards surveying of nocturnal birds and researching their habitat associations may represent particularly important future priorities, as the taxonomy and conservation status of nocturnal birds remains poorly understood in Wallacea, as it generally does in tropical island ecosystems globally (Sekercioglu 2010). ' It is interesting to note, however, that some of our secondary survey methods, notably mist-netting, appear to be very limited in their effectiveness on Buton, and particularly within Lambusango forest. Despite spending on average just over 100 hours mist-netting in the forest interior each season, only around 10-15 birds were typically caught during this time : a very poor ratio of 6.6- 1 0 netting hours for every capture. Capture rates were considerably greater in forest edge, villages and agricultural land — typically between six and nine birds per four-hour morning session in these habitats. Poor netting results in the forest interior could result from a dominance of canopy-level species in the Lambusango avifauna — 53.2% of species detected within the reserve are described by Coates & Bishop (1997) as being associated mainly with canopy-level habitat strata — or from a low population density of understorey species that would be in a mist-net’s effective capture range. A more complete study exploring the relative effectiveness of mist-nets compared to other survey methods in Lambusango Reserve is planned for the near future. A final notable finding from survey work across Lambusango Reserve is the apparent adaptability of many endemic species to a wide range of forest habitats. While few Wallacean endemics were found in any abundance in cleared, non-forest habitats, only a small number of endemic species (most notably White-bellied Imperial Pigeon Ducula forsteni) were largely confined to undisturbed primary forest in the reserve’s interior. Most endemic birds described in the literature as ‘forest’ species were encountered with similar frequency in primary, old growth secondary, and disturbed secondary forest, as well as forest-edge habitats. Indeed some endemic species, such as Sulawesi Pygmy Woodpecker Dendrocopos temminckii and White-rumped Cuckooshrike Coracina leucopygia, were encountered far more frequently in degraded forest habitats than primary or old secondary growth, and may be useful indicators of disturbed habitats. This has been discussed in detail previously (Martin & Blackburn 2010), and demonstrates the importance of secondary forests as important habitats for many of the region’s range-restricted species, especially when these secondary forests are in close proximity to or contiguous with tracts of relatively undisturbed forest. ACKNOWLEDGEMENTS Our bird surveys were conducted as part of the Lambusango Forest Research Project, implemented by Operation Wallacea and sponsored by the Wallacea Foundation, Kementerian Negara Riset dan Teknologi (RISTEK) and the Indonesian Institute of Science (LIPI). We thank Dr Tim Coles, Dr Phillip Wheeler, and the Operation Wallacea scientific and logistic teams, especially Mr Tasman, for their support in this project. We also acknowledge the valuable assistance of the Operation Wallacea volunteers who assisted with survey work, especially Tom Wilkinson and Sean Kelly. Finally, we express thanks to the residents of Labundo-bundo for their hospitality. REFERENCES van Bemmel, A. C. V. & Voous, K. H. (1951) On the birds of the islands of Muna and Buton, S.E. Celebes. Treubia 21 : 27-1 04. Bibby, C. J„ Burgess, N. D„ Hill, D. A. & Mustoe, S. H. (2002) Bird census techniques. Second edition. London: Academic Press. Bi rd Life International (2010) BirdLife International Datazone. Accessed 1 5/ 1 1/10 from http://www.birdlife.org/datazone Burton, J. A., Hedges, S. & Mustari, A. H. (2005)The taxonomic status, distribution and conservation of the Lowland Anoa Bubalus depressicornis and Mountain Anoa Bubalus quarlesi. Mammal Rev. 35: 25-50. Catterall, M. (undated) Bird survey of Buton island 1 996-1997. Unpublished Operation Wallacea report. Forktail 28 (2012) The avifauna of Lambusango Forest Reserve, Buton Island, south-east Sulawesi 1 1 1 Clements, J. F. (2007) The Clements checklist of the birds of the world. Ithaca: Cornell University Press. Coates, B. J. & Bishop, K. D. (1997) A guide to the birds ofWallacea. Alderley: Dove Publications. Hartert, E. (1 903) On the birds collected on theTukang-Besi Islands and Buton, south-east of Celebes, by Mr Heinrich Kuehn. Novit. Zool. 10: 18-38. del Floyo, J., Elliott, A. & Christie, D. A., eds. (2006) Handbook of the birds of the world, 1 1. Barcelona: Lynx Edicions. del Floyo, J., Elliott, A. & Christie, D. A., eds. (2009) Handbook of the birds of the world, 14. Barcelona: Lynx Edicions. del Floyo, J., Elliott, A. & Christie, D. A., eds. (2010) Handbook of the birds of the world, 15. Barcelona: Lynx Edicions del Floyo, J., Elliott, A. & Sargatal, J., eds. (1994) Handbook of the birds of the world, 2. Barcelona: Lynx Edicions. del Floyo, J., Elliott, A. & Sargatal, J., eds. (1 997) Handbook of the birds of the world, 4. Barcelona: Lynx Edicions. del Floyo, J., Elliott, A. & Sargatal, J., eds. (1999) Handbook of the birds of the world, 5. Barcelona: Lynx Edicions. del Floyo, J., Elliott, A. & Sargatal, J., eds. (2001 ) Handbook of the birds of the world, 6. Barcelona: Lynx Edicions. Inskipp, T., Lindsey, N. & Duckworth, W. (2001) Checklist of the birds of the Oriental region. Sandy, UK: Oriental Bird Club. International Union for Conservation of Nature (2010) The IUCN Red List of Threatened Species. Accessed 12/1 1/10 from www.redlist.org Internet Bird Collection (2011) Internet Bird Collection. Accessed 24/01/11 from http://ibc.lynxeds.com Kelly, D. & Marples, N. (2010) ['Bird populations of the Wakatobi.'] Pp.149- 1 70 in J. Clifton, R. K. F. Unsworth & D. J. Smith, eds. Marine research and conservation in the Coral Triangle: the Wakatobi National Park. Flauppauge, NY: Nova Science Publishers. Lee, D. C. & Marsden, S. J. (2008) Adjusting count period strategies to improve the accuracy of forest bird abundance estimates from point transect distance sampling surveys. Ibis 1 50: 315-325. Martin, T. E. & Blackburn, G. A. (2010) Impacts of tropical forest disturbance upon avifauna on a small island with high endemism: implications for conservation. Conservation & Society 8: 1 27-1 39. O'Donovan, G. (2001 ) Report on the botanical and ecological status of the Kakenauwe and Lambusango Nature Reserves on Buton Island, Sulawesi. Operation Wallacea report. Operation Wallacea (201 1 ) Operation Wallacea. Accessed on 1 9/01/1 1 from http://www.opwall.com Priston, N. (2005) Crop raiding by Macaca ochreata brunnescens in Sulawesi: reality, perception and outcomes for conservation. PhD Thesis, University of Cambridge. Appendix Checklist of bird species recorded in Lambusango Reserve and vicinity during 1999-2010 research seasons Species marked in bold are those which we believe are new records for Buton Island, albeit previously recorded in Catterall (undated). Species in bold and marked # = new records; those marked * = Wallacean endemics, 4 = endemic to Indonesia, t = assessed as threatened or Near Threatened by IUCN (2010). Abundance estimates are denoted as follows: A = abundant; C = common; F = fairly common; L = locally common; U = uncommon; R = rare. Species marked are seasonal migrants to the study area. Species marked X in the final column have been recorded within Lambusango Reserve. All taxonomy follows Inskipp et at. (2001). Common name Scientific name Abundance Lambusango Common name Scientific name Abundance Lambusango Great Frigatebird Fregata minor U/R Osprey Pandion haliaetus F Lesser Frigatebird Fregata ariel C Jerdon'sBaza Aviceda jerdoni R Little Pied Cormorant Pbaiacrocorax melanoleucos u Barred Honey-buzzard Perris celebensis F X Purple Heron Ardea purpurea F Brahminy Kite Haliastur Indus C X White-faced Heron Ardea novaehollandiae U White-bellied Sea Eagle Haliaeetus leucogaster F Great Egret Casmerodius alba A tGrey-headed Fish Eagle Ichthyophaga ichthyaetus R Intermediate Egret Mesophoyx intermedia A ^Sulawesi Serpent Eagle Spilornis rufipectus C X Little Egret Egretta garzetta A Spotted Harrier Circus assimilis L Little Heron Butoridesstriatus C "Sulawesi Goshawk Accipiter griseiceps R Pacific Reef Egret Egretta sacra u t* Small Sparrowhawk Accipiter nanus R Cinnamon Bittern Ixobrychus cinnamomeus Fc "Spot-tailed Sparrowhawk Accipiter trinotatus C X Black Bittern Dupetor flavicollis U Black Eagle Ictinaetus maiayensis F X Woolly-necked Stork Ciconia episcopus F Rufous-bellied Eagle Hieraaetus kienerii R Schoorl, J. W. (1987) Notes on the birds of Buton (Indonesia, South-East Sulawesi). Bull. Brit. Orn. Club 107: 165-168. Sekercioglu, C. FI. (2010) The mystery of nocturnal birds in tropical secondary forests. Anim. Conserv. 13: 12-13. Seymour, A. (2004) Monitoring forest degradation and animal populations in the forests of central Buton: preliminary results from the pilot study. Operation Wallacea report. Singer, FI. A. & Purwanto, E. (2006) Misteri kekayaan hayati hutan Lambusango. Program konservasi hutan Lambusango (PKHL). Baubau, Indonesia: Operation Wallacea Trust. Stattersfieid, A. J., Crosby, M. J., Long, A. J. & Wege, D. C. (1998) Endemic Bird Areas of the world: priorities for biodiversity conservation. Cambridge, UK: BirdLife International. White, C. M. N. & Bruce, M. D. (1986) The birds ofWallacea (Sulawesi, the Moluccas and Lesser Sunda islands, Indonesia). London: British Ornithologists' Union. Whitten, T., Mustafa, M. & Henderson, G. S. (2002) The ecology of Sulawesi. Singapore: Periplus Press. Thomas Edward MARTIN, Lancaster Environment Centre, Lancaster University, Lancaster, Lancashire, LAI 4YQ, UK, and Operation Wallacea Ltd, Hope House, Old Bolingbroke, Spilsby, Linconshire, PE23 4EX, UK. Email: tom_martin_20 1 0@yahoo.co.uk David James KELLY, Operation Wallacea Ltd, Hope House, Old Bolingbroke, Spilsby, Linconshire, PE23 4EX, UK, and Department of Zoology, School of Natural Sciences, Trinity College, Dublin 2, IRELAND. Niall Thomas KEOGH, Operation Wallacea Ltd, Hope House, Old Bolingbroke, Spilsby, Linconshire, PE23 4EX, UK, and School of Biology & Environmental Science, University College Dublin, Belfield, Dublin 4, IRELAND. Dani HERIYADI, Operation Wallacea Ltd, Hope House, Old Bolingbroke, Spilsby, Linconshire, PE23 4EX, UK. Henry Ali SINGER, Operation Wallacea Ltd, Hope House, Old Bolingbroke, Spilsby, Linconshire, PE23 4EX, UK. George Alan BLACKBURN, Lancaster Environment Centre, Lancaster University, Lancaster, Lancashire, LA 1 4YQ, UK. 112 T. E. MARTIN eta/. Forktail 28(2012) Common name Scientific name Abundance Lambusango Common name Scientific name Abundance Lambusango ^Sulawesi Hawk Eagle Spizaetus lanceolatus F X Uniform Swiftlet Collocalia vanikorensis F X 1 Spotted Kestrel Falco moluccensis L Grey-rumped Treeswift Hemiprocne longipennis A X Oriental Hobby Falco severus U ’Green-backed Kingfisher Actenoides monachus R X Wandering Whistling-duck Dendrocygna arcuata F ’Black-billed Kingfisher Halcyon melanorhyncha L T Sunda Teal Anas gibberifrons U Ruddy Kingfisher Halcyon coromanda U X Philippine Scrubfowl Megapodius cumingii U X Collared Kingfisher Todiramphus chloris A X Blue-breasted Quail Coturnix chinensis R Sacred Kingfisher Todiramphus sancta C Red Junglefowl Gallus gallus F X t’Sulawesi Dwarf Kingfisher Ceyx fallax U X Barred Buttonquail Turnixsuscitator R X Common Kingfisher Alcedo atthis F X #Slaty-legged Crake Rallina eurizonoides R X Rainbow Bee-eater Merops ornatus L Buff-banded Rail Gallirallus philippensis F ’Purple-winged Roller Coracias temminckii F X Barred Rail Gallirallus torquatus U ’Sulawesi Hornbill Peneiopides exarhatus F X ’Isabelline Bush-hen Amaurornis isabellinus F ’Knobbed Hornbill Aceros cassidix C X White-breasted Waterhen A maurornis phoenicurus F X ’Sulawesi Pygmy Woodpecker Dendrocopos temminckii R Common Moorhen Gallinala chloropus C ’Ashy Woodpecker Mulleripicus fulvus F X Whimbrel Numeniusphaeopus C Red-bellied Pitta Pitta erythrogaster U X Common Sandpiper Actitis hypoleucos C Barn Swallow Hirundo rusdca C X Red-necked Phalarope Phalaropuslobatus L Pacific Swallow Hirundo tahidca C X Bridled Tern Sterna anaethetus F t’Pied Cuckooshrike Coracina bicolor C X Black-napedTern Sterna sumatrana R ’White-rumped Cuckooshrike Coracina leucopygia U/F X Great Crested Tern Sterna bergii R ’Sulawesi Cicadabird Coracina morio C X Lesser Crested Tern Sterna bengalensis F ’White-rumped Triller Lalage leucopygiaiis R Spotted Dove Streptopelia chinensis A X TWhite-shouldered Triller Lalage sueurii R Brown Cuckoo Dove Macropygia amboinensis F X Spangled Drongo Dicurus hottentottus A X ’White-faced Cuckoo Dove Turacoena manadensis F X Black-naped Oriole Oriolus chinensis A X Stephan's Dove Chalcophapsstephani R X Slender-billed Crow Conus enca C X ff’Sulawesi Ground Dove Gatiicolumba tristigmata R X ’Piping Crow Corvus typicus F X Pink-necked Green Pigeon Treron vernans R ’Sulawesi Babbler Trichastoma ceiebense A X TGrey-cheeked Green Pigeon Treron griseicauda F X ’Red-backed Thrush loothera erythronota R X Black-naped Fruit Dove Pdlinopus melanospila C X Pied Bushchat Saxicola caprata F ’White-bellied Imperial Pigeon Duculaforsteni F X Golden-bellied Gerygone Gerygone sulphurea C Green Imperial Pigeon Ducula aenea A X Zitting Cisticola Cisticola juncidis U Pied Imperial Pigeon Ducula bicolor L Bright-headed Cisticola Cisticola exilis U ’Silver-tipped Imperial Pigeon Ducula luctuosa F X #t*Rufous-throated Flycatcher Ficedula rufigula R/U X ’Ornate Lorikeet Trichoglossusornatus R X Black-naped Monarch Hypothymis azurea A X ttVellow-crested Cockatoo Cacatuasulphurea R X Citrine Canary Flycatcher Culicicapa helianthea A X ’Golden-mantled Racquet-tail Prionituruspiaturus C X White-breasted Woodswallow Artamus leucorynchus A X Azure-rumped Parrot Tanygnathus sumatranus F X ’Ivory-breasted Woodswallow Artamus monarchus U X * Sulawesi Hanging Parrot toriculusstigmatus F X Asian Glossy Starling Aplonis panayensis A X ’Pygmy Hanging Parrot Loricuius exilis U X ’Sulawesi Myna Basilornis celebensis U X ’Sulawesi Hawk Cuckoo Hierococcyx crassirostris C X ’White-necked Myna Streptocitta albicoliis F X Oriental Cuckoo (uculussaturatus R ’Finch-billed Myna Scissirostrum dubium L X Plaintive Cuckoo Cacomantis merulinus U X #ScarletMyzomela Myzomela sanguinoienta U Rusty-breasted Cuckoo (acomantis sepulcralis U X Brown-throated Sunbird Anthreptes malacensis F X Gould's Bronze Cuckoo Chrysococcx russatus R Black Sunbird Nectarinia aspasia A X Drongo Cuckoo Surniculuslugubris C X Olive-backed Sunbird Nectariniajugularis A X ’Black-billed Koei Eudynamys melanorhyncha R X Crimson Sunbird Aethopyga siparaja F X ’Yellow-billed Malkoha Phaenicophaeuscalyorhynchus F X ’Yellow-sided Flowerpecker Dicaeum aureolimbatum F X ’Bay Coucal Centropus celebensis C X ’Grey-sided Flowerpecker Dicaeum celebicum F X Lesser Coucal Centropus bengalensis F TLemon-bellied White-eye losterops chloris F X ’Sulawesi Owl Tyto rosenbergii R X ’Pale-bellied White-eye Zosterops consobrinorum C X ’Sulawesi Scops Owl Otus manadensis C X Eurasian Tree Sparrow Passer montanus A Ochre-bellied Hawk Owl Ninox ochracea U X f Black-faced Munia Lonchura molucca C Great eared Nightjar Eurostopodus macrods L? Scaly-breasted Munia Lonchurapunctulata C #*Sulawesi Nightjar Caprimulgus celebensis U? X Black-headed Munia Lonchura matacca F Glossy Swiftlet Collocalia esculenta A X ’Pale-headed Munia Lonchura pallida R ’MoluccanSwiftlet Collocallia infuscata U FORKTAIL 28 (2012): 1 13-120 A nesting pair of Gecinulus woodpeckers in a likely zone of intergradation between Pale-headed Woodpecker G. grantia and Bamboo Woodpecker G. viridis PHILIP D. ROUND, JOHN M. HOBDAY, RUNGSRIT KANJANAVANIT & JAMES S. STEWARD A nesting of a pair of Gecinulus woodpeckers in a possible zone of intergradation between the parapatric taxa Pale-headed Woodpecker G. grantia and Bamboo Woodpecker G. viridis is described. While the male looked like a more or less typical G. viridis the female bore plumage characters that appeared intermediate between G. grantia and G. viridis. Additionally a specimen labelled as G. grantia indochinensis, collected in Thailand in 1964 (the only record for that country), also appeared atypical, showing characters somewhat intermediate between G. grantia and G. viridis. It is likely that a narrow hybrid zone between G. grantia and G. viridis exists where the two come into contact in northern Thailand and, presumably, northern Laos. Recommendations for further surveys are made in order to determine the extent of postulated hybridisation, and additionally to investigate the ecological and taxonomic relations of these two taxa. INTRODUCTION Gecinulus woodpeckers are medium-sized, three-toed woodpeckers that occur in intimate association with large-culm bamboos. The five or six accepted taxa are either treated as constituting two allospecies (Kingct^/. 1975, Robson 2008), or as one polytypic species (Short 1982, Dickinson 2003). If the former treatment is followed, two subspecies of G. viridis (Bamboo Woodpecker) are distributed in East and South Myanmar and most of Thailand ( G . v. viridis ), and Malaysia and adjacent southern Thai provinces ( G . v. robinsoni). The (mostly) more northerly distributed G. grantia (Pale-headed Woodpecker) ranges along the Himalayas from eastern Nepal, north-east India, to (mainly north and west) Myanmar (nominate grantia)-, Fujian and Guangdong, south-east China ( G . g. viridanus)-, Yunnan, Lao's, marginally northern Thailand (a single record, mentioned below); and Vietnam, from Tonkin south to (probably south) Annam ( G . g. indochinensis). A further subspecies, G. g. poilanei , described by Deignan (1950) from Cochinchina, southern Vietnam, is doubtfully distinguishable and was regarded as a synonym of indochinensis by Short (1982). Nowhere within this large, aggregated range of the various taxa is there indisputable evidence of sympatry between birds in the viridis and grantia species groups. We here report on a nesting pair of Gecinulus, observed in Chiang Rai province, northern Thailand, in which the female showed plumage characters intermediate between those of G. viridis and G. grantia. We were concerned to conduct a review of the distribution of both species where their ranges approach each other, and to determine whether there were any other indications that the taxa G. v. viridis and G. g. indochinensis might intergrade in their narrow zone of contact. STUDY AREA The field observations were made at Ban Saen Jai, Mae Fah Luang district, Chiang Rai province, 20°12'N 99°46'E, c.12 km west- north-west of the town of Mae Jan, and some 65 km due west of the collection site of Thailand’s only G. g. indochinensis specimen. The habitat was farm and plantation in steep hilly country at c.600 m elevation. The area has long supported villages of the Akha, a Tibeto-Burman ethnic minority group of (traditionally) pioneer shifting cultivators, but in recent years large tracts have been bought by urban landowners. While most of the area is deforested, and planted with hill-rice and corn, a c.20 ha community forest, preserved according to Akha land-use tradition, lies adjacent to Ban Saen Jai village. Additionally, ribbons of secondary forest and bamboo along steep gullies (some spring-fed) maintain connectivity among wooded fragments in the otherwise near-totally deforested landscape. During the period of the study the afternoon temperature in the general surroundings varied between a low of 28°C in mid-March and a high of 37°C in mid-April. The temperature on the floor of the shaded, woody gullies was noticeably (c.2°C) cooler than that of the immediate surroundings. METHODS Intermittent observations were made on a single nesting pair of Gecinulus at Ban Saen Jai, whenever one or more observers was present, during 10 March (when the nest was discovered) to 18 April (when the young fledged). Additionally we sought specimens and sight records of G. viridis and G. grantia in northern Thailand and northern Laos, focusing particularly on the details of the Lao range of G. grantia , since Laos is the only country other than Thailand where the ranges of G. grantia and G. viridis approach closely and, indeed, may overlap. We did not attempt any review of specimens from Myanmar where G. g. grantia is known from the south-west, west, centre and north, and G. v. viridis from the south and east (Robson 2008). It is not clear whether this apparent discontinuity in the distributions of the two in Myanmar is genuine or merely an artifact of sampling. While the Mekong River, some sections of which delineate the national boundary between Thailand and Laos, flows generally north to south, in places it also flows west to east (or even briefly south to north). In the context of this paper, the terms ‘east of the Mekong’ and ‘north of the Mekong’ can be used interchangeably, as can west/ south of the Mekong. In discussion of specimens, the following abbreviations are used: BMNH Natural History Museum, Tring, UK; CTNRC Centre for Thai National Reference Collections, Bangkok; FMNH Field Museum, Chicago; MCZ Museum of Comparative Zoology, Harvard University; USNM National Museum ofNatural History, Smithsonian Institution, Washington, D.C. RESULTS Distribution and vocalisations Gecinulus grantia is found widely throughout northern, central and southern Laos, in both primary and degraded semi-evergreen, dry evergreen and mixed deciduous forest (Thewlis et al. 1998, 114 PHILIP D. ROUND etal. Forktail 28 (2012) A G. grantia sight record or handled; ▼ G. viridis specimen; V G. viridis sight record. Localities mentioned in the text are identified by number: Huai Mae Salaep (1), Lo-Tiao (2), Nam Kan (3), Ban Naten (4), Ban Khomen (5), Phou Dendin (6), Ban Moung Liap (7), Kok Kawdinpiang (8), Sang Thong (9), Nam Mang (10). Duckworth etal. 1 999, Evans 200 1 ). Three specimens from Bokeo province, at Lo-Tiao, c.20°28'N 100°22'E (Figure 1), comprise two males, MCZ 267140 and MCZ 267142 collected on 6 and 7 January 1939 respectively, and a female, MCZ 267141, collected on 6 January 1939 (Figure 7). Inexplicably, the account in Delacour & Greenway (1940) implies that only a single specimen (‘un exemplaire’) was collected at Lo-Tiao. A further male specimen was collected from Phongsaly province, probably Ban Khomen, Pongsaly district, at 21°39,N 102°08'E (Bangs & Van Tyne 1931), on 28 April 1929 (FMNH 78170). Neither specimens nor photographs are available for further reported G. grantia in northern Laos, which consist of: one handled in Nam Kan National Biodiversity Conservation Area (NBCA), Bokeo province, probably c.20°28'N 100°48'E (Pasquet 1997); sight records beside the Nam Mang in Phou Khaokhoay NBCA, Vientiane province, c.l8°3TN 103°12'E (Thewlis etal. 1998);and in Phongsaly province at Phou Dendin NPA, c.22°09'N 102°22'E (identification recorded as provisional) and at Ban Naten, 21 °20'N 101°52'E (Fuchs etal. 2007). The lack of any further records known to us probably reflects the paucity of survey in much of northern Laos rather than indicating a genuine scarcity there. The sole record of G. grantia for Thailand is a female specimen, USNM 534656, labelled G. grantia indochinensis, collected by B. King at Chiang Khong, Chiang Rai province (20°17.7,N 100°23.5,E) on the south (west) bank of the Mekong, where the river forms the national boundary, on 26 April 1964 (King 2007). Gecinulus viridis is widespread but uncommon in Thailand, in evergreen and deciduous forests where large-culm bamboos are present, up to an elevation of c. 1,400 m (Lekagul & Round 1991). The only historical record of G. viridis from Laos is a specimen, BMNH 1955.1.2505, from Ban Moung Liap, on the Mekong River, Xaignabouli province, c.18°29'N 101°40'E (Robinson & Kloss 1931). Present-day Lao maps give the village name as Ban Muangliap while the name in today’s official government use is Ban Phaliap (J. W. Duckworth in litt.). As already discussed by Duckworth (1996), the basis for Delacour’s (1951) statement that the specimen probably came from the west bank (‘rive droite’) of the Mekong may have been nothing more substantial than the supposition that the east bank (‘rive gauche’) would support G. grantia indochinensis , presuming that the two species would be unlikely to occur together. Its origin in this respect should therefore be treated as uncertain. Recent surveys have, in fact, extended the Lao range of G. viridis south and east, the species having been widely found up to at least 20 km north of the north bank of the Mekong, in Sangthong district, west of Vientiane (several individuals, including pairs with young: Duckworth 1996). There is also an intervening record somewhat north-west of this, also well inland of the Mekong: a single sighting (of a male and an unsexed individual on 3 April 2010) at Kok Kawdinpiang (in Phou Gnouey Production Forest Area, Vientiane province, at about 18°18.TN 101°46.8'E) (SUFORD in press). The presence of grantia on the south bank of the Mekong, and of viridis on the north, evidently indicates that this major river does not act as a complete boundary separating these two taxa, and therefore that northern Laos and northern Thailand should encompass a zone of contact between them. In spite of this, however, there are no reports that directly indicate their co¬ existence at any site. Relatively little is known of the biology of either species. The vocalisations of the two are extremely similar. These include a dry undulating cackle, somewhat reminiscent of one of the calls of Bay Woodpecker Blythipicus pyrrhotis (‘rattle call’ in Short 1973), and piercing even-toned kweep notes. Both species drum (Winkler & Christie 2002) and the pattern of drumming described for G. Forktail 28 (201 2) Gecinulus woodpeckers in a likely zone of intergradation between G. grantia and G. viridis 115 grantia (‘...initially very rapid and clearly and gradually decelerates. ..start rate 30, end rate 15 taps/s’, of roughly 1.5 s duration: Rasmussen & Anderton 2005) is similar to that of G. viridis (PDR recording from southern Thailand, deposited with Avian Vocalizations Center, Michigan State University). Progress of observations Intermittent sightings of Gecinulus woodpeckers were made by JMH on and near his farm at Ban Saen Jai from August 2009 onwards. On 10 March 2010, at 09h00, a Gecinulus woodpecker was revealed as the source of a loud, insistent tapping, suggesting the excavation of a cavity, near the vicinity of a small pond at the bottom of a steep wooded gully. The nest cavity itself was discovered by JMH a little after 09h30 that day, when he flushed a woodpecker at close range from a dense clump of large-culm bamboos. The female woodpecker was again seen in the vicinity at c. 1 lhOO while, at 1 5h00, a male woodpecker, heard tapping from within the nest cavity, was seen when its head protruded from the nest-entrance, revealing red on the crown. PDR joined JMH at the site during 13-15 March, when both male and female were seen with heads protruding from the nest cavity on different occasions. Bouts of drumming were heard and there were long periods when tapping could also be heard, apparently emanating from within the nest-cavity. Observations were kept to a minimum so as to avoid disturbing the nesting pair, then assumed to be either in the process of laying, or already in the early stages of incubation. On 15 March, JMH watched the female enter the nest at 17h30. JMH continued observations intermittently, observing the head of the female protruding from the nest cavity on two occasions during 23-25 March. During observations of the nest from a blind, 20 m distant, on 3 April, 14h30-17h30, RK secured photographs of both breeding adults and observed both sexes removing faecal sacs from the nest, indicating that the young had hatched. JMH watched the nest further during 4-1 1 April and 14-18 April, and was joined by both PDRand JSS during 14-17 April. The young could be heard calling from within the nest from at least 8 April onwards, and both adults were highly vocal in the vicinity of the nest, giving chattering and kweep notes. Drumming was intermittently heard. Recordings of the calls of adults, made in March, and of chicks, during April, were deposited with the Avian Vocalizations Center, Michigan State University. On 16 April the female was caught in a 12 m superfine mist- net erected in front of the nest cavity as she was exiting the nest, c. 07h30, after having fed the nestlings. She was examined, photographed, measured and ringed. Two feathers were retained for possible future DNA assay. Only one nestling, the presumed female (see below), remained in the nest on 17 April (its presumed male sibling was heard calling nearby but could not be located) . The female chick was also thought to have left the nest later that day, and by the morning of 18 April neither adults nor young could be detected anywhere in the vicinity. The section of bamboo containing the empty nest was removed later that day in order to examine the nest contents and dimensions. JMH observed presumably the same pair of woodpeckers (the female was ringed) at a recently excavated cavity in an adjacent stem in the same bamboo clump, during 11-16 June 2010. On two occasions in the early morning the female was seen with head protruding from the nest cavity, and on the first of these, when the female exited, the male promptly entered, suggesting a possible repeat nesting attempt. However, no further observations were made after 16 June and no firm conclusions could be drawn. Winkler & Christie (2002) specifically remark that daytime frequenting of roost-holes by woodpeckers may, particularly in the tropics, be misinterpreted as breeding behaviour. The birds were again searched for, but not found, by JMH in the following year (2011). Nest site The nest site was situated at the north-eastern edge of a clump of mainly dead or senescent flowering bamboos on the steep flank of a deep gully that supported a narrow (c.60 m wide) band of dense remnant, secondary, semi-evergreen vegetation. The canopy cover was estimated at 70%. In the gully bottom a seasonal stream flowed into a small, dammed pond, which holds water year- round. During the period of observation, the height of the dry season, the stream had dried up, although its bed remained moist. At the pond the gully joined another wooded gully, forming part of a continuous ribbon of dense vegetation that drained to the north. A path along the northern side of the gully, half-way upslope, passed within 6 m of the nest cavity, which was slightly above head height. The steep bank immediately above the path was densely covered with small trees, bushes and herbage, providing a vantage point where a blind was constructed at a distance of 20 m from the nest, and looking down on to it, so as to observe the birds without disturbance. The path was seldom used except by occasional hunters, bamboo cutters and wandering cattle herders. Almost daily in the late afternoon herders brought their cattle to the pond below the nest for water. There was no evidence that this significantly disturbed the birds. Nest description The entrance hole was towards the upper margin of the 11th internode section of the stem of dead bamboo, Gigantochloa apus (Schulz) Kurtz. (Gramineae, Bambusoideae), c.4 m above the ground. The bamboo stem, which contained the nest hole, had been cut at the base earlier in the year and left, dead, in situ, by bamboo cutters, and was angled at roughly 60°. The top of the hole was 8.0 cm from the lower edge of the upper node and its base was 49.8 cm above the top edge of the lower node. The external circumference of the bamboo stem measured at the centre of the hole was 34.8 cm and the internal diameter of the nest cavity c. 9 cm. The entrance hole was hexagonal in shape with the vertical axis longer than the horizontal axis (the apex of the hexagon to its lowest point was 6.8 cm and the parallel sides of the nest-entrance were 3.9 cm apart). The lower rim of the nest-entrance was highly abraded (Figure 2). The internal height from the bottom of the nest cavity to the lowest point of the cavity entrance (the distance the young would have to climb to be fed at the nest entrance) was 47 cm. The interior wall of the bamboo was smooth above the nest hole, but vertically scored and shredded from the level of the hole to the floor of the cavity, and some of this shredded bamboo fibre apparently contributed to a 6 cm deep layer of black, soft, fine, fibrous vegetable matter, infested with small (c. 1 cm) , thin, white maggot-like insect larvae, on the cavity floor. The contents of the nest were preserved in alcohol for further analysis. The site of the second cavity, found in June, was in a similar bamboo stem, which was dead (after flowering) but had not been cut. The nestlings The heads of the two nestlings were seen protruding from the nest- hole on 1 5 April when a presumed male nestling could be seen to have a red mid- and hind-crown, lacking in a presumed female nestling. Appearance of the breeding pair The male bird appeared like a more or less typical G. viridis with greenish body plumage. However, the crown was not solidly red and did not extend fully onto the nape. The tail appeared unmarked when seen from above, but the primaries and secondaries had 116 PHILIP D. ROUND etal. Forktail 28 (2012) A ( Figure 2. Male Gecinulus at nest, Ban Saen Jai, 3 April 2010. (Rungsrit Kanjanavanit) Figure 3. Female Gecinulus at nest, Ban Saen Jai, 3 April 2010. (Rungsrit Kanjanavanit) indistinct pale bars, with a slight rufous tinge evident at times (Figure 2). The female differed markedly from typical G. viridis females in showing rufous-tinged secondaries, and prominent broad whitish barring on the primaries, secondaries and all rectrices (Figure 3). The conspicuous broad, sharply contrasted pale barring on remiges and rectrices was easily visible in the field, both at rest and in flight. Description of female in the hand (Figures 4, 5) Throat and forecrown unmarked, pale brownish. Mid-crown, hind- crown and ear-coverts yellowish-olive. Mantle and lower back bronze-olive (olive-green); upperwing-coverts concolorous dull bronze-green. Rump feathers extensively tipped (maroon) reddish and uppertail-coverts dull bronze-olive. Underparts (breast, belly and undertail-coverts) dull, dark olive. Prominent white spotting/ Figure 4. Lateral view female Gecinulus in the hand, Ban Saen Jai, 16 April 2010. Note the extensively barred primaries and secondaries, and rufous-tinged secondaries. (P. D. Round) Figure 5. Rump, uppertail-coverts and spread tail of female Gecinulus in the hand, Ban Saen Jai, 16 April 2010. Note the prominent barring on inner and outer webs of rectrices 1-5. (P. D. Round) Forktail 28 (2012) Gecinulus woodpeckers in a likely zone of intergradation between G. grantia and G. viridis 117 transverse barring on both outer and inner webs of all primaries (brighter on inner webs). Bright white spotting/transverse barring on all secondaries (less distinct on outer webs). Outer webs of all secondaries rufous-tinged, forming a slightly rufous panel on the closed wing. Rectrices dark olive-brown, with rufescent-olive outer webs. Rectrices 1-4 with four clear white bars, visible on both webs; rectrix 5 with three white bars, visible on the inner web only. Rectrices 1-5 were modified with pointed tips and stiff shafts. Rectrix 6 was short, unstiffened and unmarked, less than half the length of the central pair, as is more or less typical for woodpeckers. Iris ruby- red; narrow grey orbital ring; bill bluish-white, legs and feet olive-green. Wing length 131 mm (maximum chord), tail 91 mm, bill (to skull) 28.2 mm, tarsus 26.9 mm, weight 72.9 g. Secondary 6 right wing was old, unmoulted, as were secondaries 7 and 8 on the left wing. Comparison with specimens Detailed comparison of photographs of the Ban Saen Jai nesting pair was made by PDR with four male (or male-plumaged) specimens and one female specimen of G. v. viridis in CTNRC (Figure 6). The photographs were also compared with five male and two female Thai and Tenasserim G. v. viridis specimens; a further 18 female G. v. viridis specimens from elsewhere in the range; and with specimens of G. v. robinsoni from the Thai-Malay Peninsula, and three taxa of G. grantia (excluding ‘ poilanei ’). The latter comparisons were made by JSS at BMNH, and by PDR and JSS together from photographs. Neither males nor females of any G. viridis specimens examined showed any rufous cast on the secondaries or elsewhere, nor any clearly visible tail barring when the tail was examined from above. Tail barring was restricted to small white spots on the inner webs of rectrices 2-5, with the central pair of rectrices either unmarked or with one or two small white spots on the basal portion of the inner web. A pattern of vague barring on the primaries and secondaries in G. v. viridis specimens was never as contrasted as in the Ban Saen Jai bird, and mainly restricted to white spots on the inner webs. Faint barring, usually visible on the outer webs in females, was never as prominent as on the Ban Saen Jai bird. Prominent wing and tail barring is characteristic of G. grantia. Fiowever, the pale bars are strongly rufous rather than whitish in that species, and are broader, more than half as broad as the intervening dark brown bars. In addition, the mantle and wings in G. grantia are strongly chestnut-red, the sexes scarcely differing in hue. While G. g. indochinensis is slightly less intensely reddish than the nominate race it nevertheless remains strongly chestnut-rufous (Figure 7). The southern Chinese G. g viridanus is dark rufous, less strongly chestnut on the upperparts, which have some greenish feathers mixed in, but it retains prominent wing and tail barring in which the pale bars are rufous (Figure 8). No specimens of any taxon of either G. viridis or G. grantia examined in collections precisely resembled the Ban Saen Jai female. The latter appeared more or less intermediate between the two: in overall plumage tones more akin to viridis than grantia, yet with a pronounced rufous cast on the secondaries, and clear, broad whitish, well-contrasted bars on primaries, secondaries and tail feathers that were not shown by any other viridis specimen. Photographs of King’s female specimen from Chiang Khong (USNM 534656) also revealed that it is somewhat intermediate in appearance. It differs from any other G. g. indochinensis or G. g. viridanus specimen in being markedly and evenly green-tinged on Figure 6. Dorsal view of four Thai-taken male/male-plumaged specimens, and one female specimen, of Gecinulus viridis. Note the restricted areas of red on the hind-crown on the right-hand-most red-crowned individual, CTNRC 53-3344, from Mae Jan, Chiang Rai. (P. D. Round/Centre for Thai National Reference Collections) 118 PHILIP D. ROUND etai Forktail 28 (2012) Figure 7. Dorsal view of three specimens of 6. grantia indochinensis from Lo-Tiao, Bokeo, Laos. From right to left MCZ 267140 (male); MCZ 267141 (female), MCZ 267142 (male). (Jeremiah Trimble, Museum of Comparative Zoology, Harvard University /© President and Fellows of Harvard College) the mantle, recalling the Saen Jai bird, although it possesses rufous- tinged, rather than whitish, bars on the folded wing. The tail-bars, however, are whitish rather than rufous-tinged and neither as broad nor as boldly contrasted as in any G. grantia (Figure 9). Gecinulus viridis and G. grantia also differ in the patterning of red on the crown of males. In G. viridis the mid-crown and hind- crown and nape are solidly red. In G. grantia the red on the crown is less extensive, pinkish-red, broken on the hind-crown and does not extend to the nape (Figures 7, 8). In this respect, the Ban Saen Jai male was unusual among G. viridis in that the red on the hind- crown was less extensive than is typical for the species. Of four male- phimaged specimens in the CTNRC collection, three (two from Kanchanaburi, south-west Thailand, and one, market-purchased, provenance unknown) have extensive and solidly red crowns. A fourth (specimen no. 53-3344; second from right. Figure 6) lacks solid red on the hind-crown. This specimen, labelled as a female, probably in error (the label reported the gonads as small), was collected at Fluai Mae Salaep, Mae Jan district, Chiang Rai, (c.20°l 1 'N 99°42'E), only a few kilometres from Ban Saen Jai. In terms of its weak wing and tail-feather barring and olive-green body coloration, the specimen looked typical for G. viridis. DISCUSSION The existence of a female Gecinulus , clearly outside the normal range of variation of Bamboo Woodpecker, somewhat intermediate in plumage between G. viridis and G. grantia, and the existence of another female Gecinulus (USNM 534656, labelled G. g. Figure 8. Dorsal view of two specimens of G. grantia viridanus, BMNH 1 900.1 .1 8.328 (male, left) and BMNH 1905.1 2.24.423 (female, right). (J. Steward/ © Natural History Museum) indochinensis') from the same general area (Chiang Rai province) which differs markedly from topotypical G. g. indochinenis from further north and east in Indochina, suggests that viridis and grantia may intergrade in this region of northern Thailand and possibly adjacent northern Laos. The coincidence of reduced red on the crown in two Chiang Rai male G.viridis, the Ban Saen Jai nesting bird and CTNRC 53-3344 from nearby Huai Mae Salaep, may possibly also be significant. Is reduced red on the crowns of males typical for Chiang Rai/ northern Thailand G. viridis ? Might this, in fact, be further evidence of intergradation between G. viridis and G. grantia ? The only other G. viridis specimen from the Thai- Lao border region (the Ban Moung Liap bird, BMNH 1955:1.2505) seems also to possess a less solidly red hind-crown, although in other respects it appears typical for G. viridis. Although both species occur in northern Laos there appear to be no reports of them occurring in close proximity at the same location (Figure 1). The few specimens of Pale-headed Woodpecker in Laos closest to the areas supporting Bamboo Woodpecker for which photographs were examined are typical chestnut-backed G. grantia indochinensis , with strongly and broadly barred wings and tails, lacking any intermediate characters. The most significant, since they were collected only an estimated 20 km north of the site of King’s presumed hybrid (albeit on the opposite bank of the Mekong), were the three MCZ specimens from Bokeo province, at Lo-Tiao (Figure 7). Since Gecinulus woodpeckers are relatively shy and hard to approach and observe, the similarity of the vocalisations of the two species may mean that fleeting sight records collected during faunal surveys within the zone of contact or sympatry may not be FOH.KIf", ,v cw*4. Forktail 28 (2012) Gecinulus woodpeckers in a likely zone of intergradation between G. grantia and G. viridis 119 Figure 9. USNM 534656 (dorsal view and lateral view), collected Chiang Khong, Chiang Rai, northern Thailand, 26 April 1964, by B. King. Note the extensively greenish mantle which is atypical for any subspecies of Gecinulus grantia. (J. Dean/© National Museum of Natural History, Smithsonian Institution) assignable as to species with 100% confidence. (So far, purely aural records are not known to have provided the basis for any northern Lao reports of either species: J. W. Duckworth/w litt.). If G. grantia and G. viridis do intergrade widely, then intermediates might be expected to show a highly variable mix of characters, and those individuals with only subtle differences from either parent species might easily be overlooked. On the other hand, if both occur sympatricaliy without intergradation in their zone of contact, such sympatry might remain undetected if one species was rare, and the other relatively common at any given site. Information on the extent of ecological differences between these two taxa is scant. Since both are associated with large-culm bamboos, most if not all nests may be expected to be situated in cavities in bamboos. The only nest described for G. viridis, from the Thai-Malay Peninsula, was excavated in the bamboo Gigantochloa scortechinii Gamb. (Wells 1999), while both Short (1973) and PDR have seen holes presumed to have been excavated by G. viridis in large-culm bamboos at Thai localities where the species is present. There appear to be no nest records of G. grantia anywhere in its Indochinese range, and the only nests described for G. grantia by Baker (1927), from the northern Indian subcontinent, were apparently in tree-stumps. Too few nests of either species have been found to know whether reported differences are typical, or whether nest-sites in either or both species may be situated in either tree stumps or bamboos, depending on availability. However, Baker’s descriptions of nests of G. grantia may be questionable, since his written work contains inconsistencies and discrepancies from that of other workers, and many of his findings have been either discounted or questioned (Rasmussen & Anderton 2005). Until convincing evidence is presented that refutes this, it should be assumed that G. grantia and G. viridis are very similar in their ecology. Efforts are needed in northern Thailand and northern Laos to discover how frequent intermediate-plumaged Gecinulus woodpeckers are, and investigate the ecological and taxonomic relations of G. grantia and G. viridis. Chiang Rai province, north and east of the area of the present sighting, is an obvious priority area for survey, as are sites in Laos where the ranges o i grantia and viridis approach closely: Bokeo province; Vientiane province and municipality; and northern Laos west of the Mekong (Xaignabouli province). Arguably, however, almost all of northern Laos, where relatively few surveys have been implemented, and in which the status of Gecinulus woodpeckers remains largely unknown, would repay survey. Gecinulus grantia and G. viridis presumably diverged from a common ancestor during a previous period of forest fragmentation. The presence of apparent plumage intergrades suggests that these taxa have since come into renewed contact before isolating mechanisms between them have been fully developed. A review of hybrid zones in birds is provided by Price (2008) and, indeed, hybrid zones may prove to be relatively frequent among parapatric taxa in the tropics. Manakins Manacus offer well-studied examples from the Neotropics (Brumfield et al. 2001, Stein & Uy 2006), while hybridisation is also documented among some Melanerpes woodpecker species (Short 1982) and in North American flickers Colaptes (Short 1965, Moore & Price 1993). Given their marked divergence in plumage patterns, Pale¬ headed Woodpecker and Bamboo Woodpecker qualify as species 120 PHILIP D. ROUND eta/. Forktail 28 (2012) using Helbig e/^/.’s (2002) criterion for assigning taxonomic rank (hybridisation is rare, making it unlikely that their gene pools will ever merge). But if intermediate-plumaged birds, apparently caused by interbreeding, prove to be frequent within the zone of contact, a re-examination of their taxonomic status might be necessitated. Even so, provided that the postulated hybrid zone is narrow in relation to the total ranges of the taxa, indicating barriers to gene flow, the two would probably still continue to be treated as species or (following Helbig et al. 2002) semi-species. A simple scoring system based on phenotypic characters, applied to all Gecinulus sight and photographic records, trapped birds and museum specimens from within the likely contact zone, might help to elucidate the extent and pattern of introgression between the two. ACKNOWLEDGEMENTS James Dean in USNM, Mary Hennen at FMNH, and Jeremiah Trimble at MCZ kindly photographed specimens of G. grantia on our behalf. BMNH and CTNRC kindly allowed access to their holdings, and we thank Robert Prys-Jones at the former institution, and Surachit Waengsothon at the latter, and their respective staff's, for their kind assistance, especially Hein van Grouw at BMNH for providing label data and locating and photographing an additional Gecinulus specimen on our behalf. The Harrison Institute, Kent. UK (supported by the Darwin Initiative), and the Wildlife Conservation Society, New York, provided copies of some key references and we are particularly grateful to Paul Bates and Kerry Prendergast, respectively. Warren Brockelman, James Dean, Edward Dickinson, Will Duckworth, Ben King, Joe Tobias and David Wells commented on drafts of this manuscript. Will Duckworth, in particular, drew our attention to some key Lao references and made innumerable suggestions that improved this paper. We also thank two anonymous referees. Jerome Fuchs and Eric Pasquet provided further information on Lao Gecinulus records, as did Le TrongTrai, Simon Mahood, John Pilgrim and Jack Tordoff for those from Vietnam. J. F. Maxwell, Chiang Mai University Herbarium, identified the species of bamboo containing our woodpecker nest. Perawit Insuan kindly prepared the map. Philip Round is supported by The Wetland Trust. REFERENCES Baker, E. C. S. (1927) The fauna of British India including Ceylon and Burma: birds, 4. London: Taylor & Francis. Bangs, O. & Van Tyne, J. (1 931 ) Birds of the Kelley-Roosevelts expedition to French Indochina. Publ. Field Mus. Nat. Hist. (Zool. Ser.) 18: 33-1 19. Brumfield, R. T„ Jernigan, R. W., McDonald, D. B. & Braun, M. J. (2001) Evolutionary implications of divergent dines in an avian ( Manacus : Aves) hybrid zone. Evolution 55: 2070-2087. Deignan, H. G. (1950) Five new races of birds from south-eastern Asia. Zoologica 35 (2): 1 27-128. Delacour, J. (1951) Commentaires, modifications et additions a la liste des oiseaux d'lndochine Franpais. Oiseau 21 : 1-32. Delacour, J. & Greenway, J. C. (1940) Liste des oiseaux recueillis dans la province du Haut-Mekong et le royaume de Luang-Prabang. Oiseau et R.f.O. 10:25-59. Dickinson, E. C. ed. (2003) The Howard & Moore complete checklist of birds of the world. Third edition. Princeton: Princeton University Press. Duckworth, J. W. (1996) Bird and mammal records from the Sangthong District, Vientiane Municipality, Laos, in 1 996. Nat. Hist. Bull. Siam Soc. 44: 217-242. Duckworth, J. W., Salter, R. E. & Khounboline, K. compilers (1999) Wildlife in Lao PDR: 1999 status report. Vientiane: lUCN-The World Conservation Union/ Wildlife Conservation Society/Centre for Protected Areas and Watershed Management. Evans, T. D. (2001 ) Ornithological records from Savannakhet Province, Lao PDR. Forktail 17: 21-28. Fuchs, J., Cibois, A., Duckworth, J. W., Eve, R., Robichaud, W. G„ Tizard, T. & van Gansberghe, D. (2007) Birds of Phongsaly province and the Nam Ou river, Laos. Forktail 23: 22-86. Helbig, A. J., Knox, A. G., Parkin, D. T„ Sangster, G„ & Collinson, M. (2002) Guidelines for assigning species rank. Ibis 144: 518-525. King. B. (2007) Some 1960s additions to the list of Thailand's birds. Nat. Hist. Bull. Siam Soc. 55 (1 ): 1 05-1 1 9. King, B., Dickinson, E. C. & Woodcock, M. W. (1975) Field guide to the birds of South-East Asia. London: Collins. Lekagul, B. & Round, P. D. (1991) A guide to the birds of Thailand. Bangkok: Saha Karn Bhaet. Moore, W. S. & Price, J. T. (1993) Nature of selection in the northern flicker hybrid zone and its implications for speciation theory. Pp. 196-225 in R. G. Harrison, ed. Hybrid zones and the evolutionary process. Oxford: Oxford University Press. Pasquet, E. (1997) Oiseaux, Pp.1-4 in Rapport Museum NHNP-ForespaceAvril 1997. Paris : Museum National d'Histoire Naturelle. Price, T. D. (2008) Speciation in birds. Greenwood Village, USA: Roberts & Co. Rasmussen, P. C. & Anderton, J. C. (2005) Birds of South Asia: the Ripley guide. Washington, D. C. and Barcelona: Smithsonian Institution and Lynx Edicions. Robinson, H. C. & Kloss, C. B. (1 931) Some birds from Siam and Laos (Middle Mekong). Ibis 13(1): 319-341. Robson, C. (2008) A field guide to the birds of South-East Asia. London: New Holland. Short, L. L. (1965) Hybridization in the flickers ( Colaptes ) of North America. Bull. Amer. Mus. Nat. Hist. 1 29: 307-428. Short, L. L. (1973) Habits of some Asian woodpeckers (Aves: Picidae). Bull. Amer. Mus. Nat. Hist. 1 52: 332-336. Short. L. L. (1982) Woodpeckers of the world. Cinnaminson, New Jersey and Dordrecht, Holland: Foris Publications. Stein, A. C. & Uy, J. A. C. (2006) Unidirectional introgression of a sexually selected trait across an avian hybrid zone: a role for female choice? Evolution 60: 1476-1485. SUFORD (Sustainable Forestry for Rural Development) project (in press) Preliminary biodiversity assessment and management recommendations of SUFORD-AF Production Forest Areas. Vientiane: SUFORD project, Department of Forestry. Thewlis, R.M., Timmins, R. J., Evans, T. D. & Duckworth, J. W. (1998) The conservation status of birds in Lao PDR. Bird Conserv. Internatn. 8 (suppl.): 1-1 59. Wells, D. R. (1999) The birds of the Thai-Malay Peninsula, I . Non-passerines. London: Academic Press. Winkler, H. & Christie, D. A. (2002) Family Picidae (woodpeckers). Pp.296- 555 in J. del Hoyo, A. Elliott, & J. Sargatal, eds. Handbook of the birds of the world, 7. Barcelona: Lynx Edicions. Philip D. ROUND , Regional Representative, The Wetland Trust, and Assistant Professor, Department of Biology, Faculty of Science, Mahidol University, Rama 6 Road, Bangkok 10400, Thailand. Email: philip.rou@mahidol.ac.th John M. HOBDAY, 210/3 Mu 1, Soi Kiengdoi, T. Chang Pheuak, A. Muang, Chiang Mai 50300, Thailand. Rungsrit KANJANAVANIT, Lanna Bird and Nature Conservation Club, 76/1 Mu 14, Soi 5 Suthep Road, A. Muang, Chiang Mai 50200, Thailand. James S. STEWARD, Department of Biology, Faculty of Science, Mahidol University, Rama 6 Road, Bangkok 10400, Thailand. FORKTAIL 28 (2012): 121-128 Zappey's Flycatcher Cyanoptila cumatilis, a forgotten Chinese breeding endemic PAUL J. LEADER & GEOFF J. CAREY The Blue-and-white Flycatcher Cyanoptila cyanomelana is a summer visitor to north-east Asia. A review of museum material demonstrates that the present treatment of two subspecies (nominate and cumatilis ) is untenable as (a) intermedia, although not currently recognised, is considered valid, and (b) the name cumatilis is currently incorrectly ascribed, being restricted in reality to central China (outside of the published range of Blue-and-white Flycatcher). Populations of all three taxa were studied on the breeding grounds in Russia, China and Japan and their songs recorded. Using criteria (Tobias etal. 2010) that grade morphological and vocal differences between allopatric taxa, cumatilis readily achieves the threshold for species status. The English name Zappey's Flycatcher is proposed in honour of the collector of the type specimen. Two subspecies of Blue-and-white Flycatcher are recognised, nominate and intermedia. INTRODUCTION The Blue-and-white Flycatcher Cyanoptila cyanomelana is a summer visitor to the forests of north-east Asia. Two subspecies are currently recognised, nominate cyanomelana Temminck, 1 829, which breeds in the southern Kuril Islands, Japan and South Korea, and cumatilis Thayer & Bangs, 1909, which breeds in north-east China south to Hebei, south-east Russia and North Korea (Clement 2006). Although originally described as a full species based on the Very different’ plumage of males (Thayer & Bangs 1909), cumatilis has long been treated as a subspecies of cyanomelana (e.g. Vaurie 1954, 1958). A further subspecies, intermedia Weigold, 1922, is not currently recognised and has long been treated as a synonym of cumatilis (e.g. Hartert & Steinbacher 1934, Vaurie 1954, 1958). In this paper the validity of intermedia and the relationship between the three taxa are reviewed based upon an examination of museum specimens and fieldwork conducted during the breeding season in China, Japan and Russia. METHODS Museum specimens were examined at the Natural History Museum, Tring, United Kingdom (BMNH), the Yamashina Institute for Ornithology, Tokyo, Japan (YIO), the Institute for Zoology, Chinese Academy of Sciences, Beijing, People’s Republic of China (IOZ), and the Museum of Zoology of the Far-Eastern National University, Vladivostok (MZFENU). The following biometrics were taken: wing (maximum chord), tail length (to base of tail measured under the undertail-coverts) and bill width at distal edge of nostrils. Measurements taken accord with standard procedures (Redfern & Clark 2001) and a sample of 20 males each from central China, south-east Russia and Japan was measured. Plumage differences of males and females from different populations were examined in detail, with consideration given to any differences attributable to age (first-summer birds being readily aged owing to retained juvenile greater coverts). Although no type specimens were examined, specimens from the type localities of both intermedia (Vladivostok) and cyanomelana (‘Japan’) were examined and photographs of the type of cumatilis (type locality: Mafuling in Fangxian County, north-west Hubei), which is housed at the Museum of Comparative Zoology, Massachusetts (MCZ), were obtained. During fieldwork conducted on the breeding grounds, sound recordings of males in song were obtained from Russia (Amurskaya Oblast and Primorskiy Kray), Japan (Honshu and Hokkaido) and China (Beijing) (Figure 1). Additional recordings were obtained from China (Beijing, Hebei and Shaanxi) from other sources. Recordings were made using a Telinga Pro 5 with either a Sound Devices 722 or an HHB Portadisc MDP 500, and a Sony PCM- M10 with a Sennheiser ME66. Spectrograms were produced and analysis of various parameters carried out using Raven Pro 1.3 (Cornell Laboratory of Ornithology 2003-08). Contrast was adjusted for each recording to ensure all elements (defined as any continuous line on a sonagram) were retained, while minimising reverberation. Measurements were made using a spectrogram window size of 5 1 2. A total of 889 song strophes was analysed, comprising 209 from eight male cumatilis , 461 from 15 male intermedia and 219 from nine male cyanomelana (see Table 2). Analysis of parameters of each strophe was based on those proposed by Tobias et al. (2010), and comprised: begin and end times (from which duration was calculated); lowest and highest frequency (from which frequency range was calculated); centre frequency (the frequency dividing a strophe into two intervals of equal energy); peak frequency (the frequency at which peak power occurs); pace (calculated by dividing strophe length by number of elements). In addition, in order to bring out a consistent feature that was apparent on listening to the song of each, namely differences in the degree of variation in frequency and structure of elements within each strophe, a further parameter was analysed: the highest number of times that any individual element was repeated. A repeat was determined aurally, rather than via the sonagram, on which minor differences between elements can be seen that are not detected by the human ear. Where appropriate, phrases were identified; these are defined as a discrete group of more than one element within a strophe. For each individual, we calculated the mean of each parameter; we then used the mean and standard deviation of all individuals per taxon to calculate Cohen’s d values. In order to review species limits between cumatilis , intermedia and cyanomelana we applied the quantitative scoring system proposed by Tobias etal. (2010) to assess the degree of phenotypic difference between allopatric taxa. These criteria were summarised by Collar (201 la, b) thus: an exceptional difference (a radically different coloration or pattern) scores 4, a major character (a pronounced and striking difference in the colour or pattern of a body part, or in measurement or vocalisation) 3, a medium character (clear difference reflected, e.g., by a distinct hue rather than a different colour) 2, and a minor character (a weak difference, e.g. a change in shade) 1. Tobias etal. (2010) set a threshold of7 to allow 122 PAUL J. LEADER & GEOFF J. CAREY Forktail 28 (2012) Plate 1 . Dorsal view of specimens of male cumatilis, IOZ, Beijing (Paul J. Leader). Plate 4. Dorsal view of specimens of male intermedia (Paul J. Leader / © The Natural History Museum, Tring). Plate 2. Ventral view of specimens of male cumatilis, IOZ, Beijing (Paul J. Leader). Plate 5. Ventral view of specimens of male intermedia (Paul J. Leader / © The Natural History Museum, Tring). Plate 3. Lateral view of specimens of male cumatilis IOZ, Beijing (Paul J. Leader). Plate 6. Lateral view of specimens of male intermedia (Paul J. Leader / © The Natural History Museum, Tring). Forktail 28 (2012) Zappey's Flycatcher Cyanoptila cumatilis, a forgotten Chinese breeding endemic 123 ft Plate 7. Dorsal view of specimens of mal e cyanomelana (Paul J. Leader / © The Natural History Museum, Tring). Plate 8. Ventral view of specimens of male cyanomelana (Paul J. Leader /©The Natural History Museum, Tring). Plate 9. Lateral view of specimens of male cyanomelana (Paul J. Leader / ©The Natural History Museum, Tring). for species status, stating that only three plumage characters, two vocal characters, two biometric characters and one behavioural or ecological character may be counted. Vocal and biometric characters were assessed for effect size using Cohen’s d using the online calculator at http://www.uccs.edu/~faculty/lbecker/, where 0.2- 2 is minor, 2-5 medium, 5-10 major and >10 exceptional. RESULTS Morphological differences between populations Examination of museum specimens revealed that there are consistent plumage differences between populations from central China, Japan and south-east Russia. Males from central China are highly distinct and show pronounced differences compared to specimens from Japan and south-east Russia. Males from Japan and south-east Russia, whilst more similar to each other than males from central China, are also consistently different. Males from populations that breed in central China are distinct from specimens of other populations in being blue or blue-green across the breast, throat and ear-coverts, and in having black or blackish restricted to the lores (Plates 1-3). Many specimens from central China show extensive fine blackish vertical streaking across the breast and throat, and most exhibit a well-defined black or blackish line between the lower border of the breast and the rest of the underparts, which are white (Plate 2). The upperparts are typically blue-green, often with extensive fine black streaking across the mantle, scapulars, rump and uppertail-coverts (Plate 1). The similarity in the colour of (a) the breast, throat and ear-coverts and (b) the upperparts results in little, if any, contrast between these two areas. The throat, breast, ear-coverts and lores of males from Japan are typically pure glossy black (although a small number of specimens have narrow bluish tips to the breast and throat feathers) and the upperparts are a bright, rich blue; streaking on the upperparts is rare and, if present, restricted to the scapulars (Plates 7-9). Males from Russia are matt-blackish on the throat, breast and ear-coverts (only very rarely pure black), and usually show a bluish wash or distinct blue tones to the throat and breast (Plates 5-6). The upperparts are blue, although never as richly blue as Japanese birds, being intermediate in colour between Japanese specimens and those from central China; streaking on the upperparts is extremely rare and, when present, is restricted to the central part of the mantle (Plate 4). Males from both Japan and south-east Russia show marked contrast between the throat/breast/ ear-coverts and the upperparts. Subtle differences in the plumages of females were also noted; however, as it was not possible to compare specimens of females directly from all three regions, this issue requires research and is not taken further here. Whilst a white centre to the tail (rather than the sides) is considered a feature of ''cumatilis by Clement (2006) and Brazil (2009), none of the specimens examined showed anything other than white bases to the sides of the tail. Only the specimens from central China matched the type of cumatilis , and whilst specimens from south-east Russia (including Vladivostok, the type locality of intermedia ) were closer overall to those from Japan (i.e. cyanomelana), they differed consistently, as described above, and thus the name intermedia needs to be reinstated for populations in north-east mainland Asia. This treatment is followed hereafter and the name cumatilis is used only for the central Chinese population. With longer wing and tail measurements cumatilis averages larger than both intermedia and cyanomelana, while on average cyanomelana is longer-winged than intermedia, but both have similar tail lengths (Table 1). Bill width values are very similar (average width of 5.81 mm for intermedia, 5.80 mm for cyanomelana and 5.83 mm for cumatilis). 124 PAUL J. LEADER & GEOFF J. CAREY Forktail 28 (2012) Table 1. Average wing and tail lengths (mm) and Standard Deviation (SD) of male cumatilis, intermedia and cyanomelana (n = 20 for each taxon). cumatilis intermedia cyanomelana Mean SO Mean SD Mean SD Wing 95.4 2.10 92.8 1.96 94.3 2.14 Tail 66.9 2.10 64.6 2.65 64.0 1.75 Vocalisations The songs of all three taxa are loud and usually uttered from the most prominent trees in the territory, especially those at the top of wooded slopes. Territories appear to be relatively large, and the volume of the song and choice of prominent perch when singing reflect this. For all taxa, the song comprises a regular series of discrete strophes (separated by distinct pauses normally longer than the strophe) , each of which contains a differing number of elements, one or more of which was repeated to a varying degree, often in the terminal section of each strophe in the case of cumatilis and cyanomelana. Representative song strophes for each taxon are illustrated in Figures 2-4, while samples of recordings, including the actual strophe illustrated in the figures, have, where possible, been placed on the online database at XenoCanto (www.xenocanto.org/asia) with catalogue numbers provided after each strophe in Figures 2-4. The typical song of cumatilis is rather simple and repetitive and of a relatively lower pitch overall, and the frequency range compared to the other taxa is notably narrower, largely due to a lower mean high frequency (Table 3 and Figures 2-4). In addition, the centre and peak frequencies are both approximately 10% or more lower. Not only is the pitch lower, however, but the delivery is slightly more measured and slower. Generally speaking, in cumatilis each strophe consists of a measured repetition of similar elements, with or without an introductory series of 1 -5 notes, creating a song of little variety or, usually, melody. The exception to this was a male recorded in Shaanxi province, whose minor variation in pitch of certain elements in each strophe imparted a distinctive rhythm compared to cumatilis males at Beijing. The lack of diversity in elements is indicated by the mean highest number of repeated elements being distinctly higher than the equivalent figures for the other two taxa. Compared to cumatilis the typical song of intermedia sounds less measured and contains more variation in pitch within both individual elements and each strophe as a whole, and in structure of elements. Phrases (a discrete group of more than one element within a strophe) are more clearly defined, as a result of the elements in each strophe being less regularly spaced than those of cumatilis , and the frequency range is relatively wide (Figure 3). The rapid repetition of short elements nearly producing a trill is almost absent from the songs of cumatilis and cyanomelana. For 84% of intermedia strophes, the first element is higher in pitch than the second and, usually, most of the remainder of the strophe. A bias toward a higher-pitched first element is also shown by both cumatilis and cyanomelana , but it is not so marked (57% and 36% respectively). The song of cyanomelana contains more repetition of elements at the same pitch than is the case with intermedia , but not to the same extent as cumatilis. Within-strophe variation in pitch is greater than in cumatilis , although not as marked as in intermedia. Figure 1 . Locations of cumatilis, intermedia and cyanomelana recordings used in this study and locations of all known breeding season records of cumatilis. ▲ ♦ □* ▲ ▲ cyanomelana recordings ♦ ★ □ 0 250 500 intermedia recordings cumatilis recordings Type locality of cumatilis Location of other cumatilis records (see main text) 1,000 1,500 N S 2,000 H Kilometers Forktail 28(2012) Zappey's Flycatcher Cyanoptila cumatilis, a forgotten Chinese breeding endemic 125 / - 6 - 5- 4 3- 9 - kHz •VAiM i H s 2:1 5 2:16 2:17 2:1 8 2:19 2:20 b) XC1 03900 7 6 5- 4 -| 3 2 1 H kHz d) XC 103899 7- 6 c 4 3 2- 1 : kHz zoo" i*nm — i - 1 - 1 - 1 - 1 - 1 - 1 - 1 - 1 — 13 14 15 16 17 iiihin 2:01 2:02 2:03 2:04 Figure 2. Representative song strophes of cumatilis: (a) Shaanxi, China, (b), (c) and (d) Beijing, China. Note the relatively simple structure, lacking sharp or significant frequency variations within element and strophe, and the generally lower pitch compared to intermedia and cyanomelana. Recording (a) made by P. Alstrom, others by G. J. Carey. Reference numbers refer to XenoCanto catalogue number. a) XC1 03906 7 - 6 - 5 - 4 - 3 - 2 1 - kHz 1 :09 1 1 □ " 1:11 ' 1 :1 2 ' 113 d) kHz 33 l i i i « *+ r .1 — i — 34 35 36 37 Figure 3. Representative song strophes of intermedia: (a), (b), (c) Primorskiy Kray, Russia, (d) Hebei, China. Note the greater variation of both pitch within each strophe and structure of individual elements. The series of very short elements in (c) is more typical of this taxon. Recording (d) made by P. Alstrom, others by G. J. Carey. Reference numbers refer to XenoCanto catalogue number. a) XC1 03902 . 6 - 5 - 4 - 3 - ^ k i * » it i 1 i 1 1 b) XC1 03903 j - 6 - 5 - 4- 3 - kHz 1 - — i 3 1 ! - ’ - 1 - 1 - 1 - 1 - 1 32 33 34 35 T kHz 1 : s 1 43 1 44 1:45 1:46 1 47 1:48 d) XC103905 7 J 6 - 5 - 4 - 3 - 2 - 1 - 1 7 1 8 1 9 20 21 Figure 4. Representative song strophes of cyanomelana: (a) Honshu, Japan, (b), (c), (d) Hokkaido, Japan. The repetition of elements is more similar to cumatilis, while the change in pitch is more similar to intermedia. Compared to cumatilis, pitch is generally higher and frequency range greater. Recordings (a) and (b) made by G. J. Carey, recordings (c) and (d) by P. J. Leader. Reference numbers refer to XenoCanto catalogue number. 126 PAUL J. LEADER & GEOFF J. CAREY Forktail 28 (2012) Table 2. Locations of recordings, number of males and number of strophes analysed. Taxon Location No. of males No. of strophes cumatilis Shaanxi, China 1 19 cumatilis Beijing, China 7 190 intermedia Hebei, China 3 101 intermedia Amurskaya Oblast, Russia 1 34 intermedia Primorskiy Kray, Russia 11 326 cyanomelana Honshu, Japan 1 26 cyanomelana Hokkaido, Japan 8 193 Total 32 889 Table 3. Mean and standard deviation (SD) values of parameters (see text) selected for analysis of cumatilis, intermedia and cyanomelana. cumatilis mean SD intermedia mean SD cyanomelana mean SD Low Freq (Hz) 2,283 265 2,347 292 2,554 330 High Freq (Hz) 4,892 624 6,220 937 6,508 1,108 Freq Range (Hz) 2,609 638 3,873 901 3,953 1,053 Centre Freq (Hz) 3,425 348 3,915 430 3,931 303 Peak Freq (Hz) 3,614 488 4,041 567 4,009 451 Length (s) 2.68 1.05 2.37 0.71 2.18 0.59 No. of elements 8.73 3.68 8.55 3.30 7.98 2.45 Pace (elements/s) 3.26 0.46 3.67 1.12 3.73 0.92 Highest count repeated elements 5.63 2.39 2.59 2.11 2.80 1.75 Taxonomic and geographical delimitations Based on these findings, cumatilis is restricted to central China, breeding north to Beijing, west to Shaanxi, and south to north¬ west Hubei. Thayer & Bangs (1909) described cumatilis on the basis of seven specimens (five males and two females) collected by W. R. Zappey in Hubei (Hupeh), China, between 1 1 and 25 May 1907. Altitudes are available for four of these specimens (MCZ online database: http://www.mcz.harvard.edu/collections/ searchcollections.html, accessed January 2012) and are approximately 1,500-1,700 m (‘5000-5500ft’). Given the dates, latitude, altitude and number of individuals involved — including both sexes — it seems reasonable to assume that these were breeding birds. As such, cumatilis has a breeding range almost entirely south and west of that published in much of the modern literature for Blue-and-white Flycatcher, and the type locality is c. 1,000 km south of the range published in Clement (2006), although Dementiev & Gladkov (1954) map the breeding distribution of Blue-and-white Flycatcher south to the Yangtze River, and Cheng (1987) questions whether the species breeds as far south as Hebei. Confusingly, Zheng (2011) lists both cumatilis and cyanomelana as breeding in north-east China (Liaoning, Jilin and Heilongjiang). We have no evidence that cumatilis and intermedia intergrade in the Beijing/Hebei area where they breed within at least 300 km of each other. Therefore, the range currently attributed to cumatilis in much of the modern literature is extensively occupied by intermedia. The correct breeding distribution of the three taxa is considered to be as follows: cumatilis-. a Chinese breeding endemic occurring in central China, north to Beijing, west to Shaanxi (P. Alstrom in litt. 2011) and south to north-west Hubei; presently known from only a small number oflocations (Figure 1); cyanomelana: southern Kuril Islands (specimens examined) and Japan (Kyushu, Tsushima, Shikoku, Honshu and Hokkaido) (Brazil 1991); intermedia : north-east China (Heilongjiang south to eastern Hebei) (Cheng 1987), south-east Russia (Amurskaya Oblast and Primorskiy Kray) (Dementiev & Gladkov 1954) and the Korean peninsula. Both intermedia and cyanomelana are stated to occur in the Korean peninsula (Clement 2006), with intermedia in the north and cyanomelana in the south, with the two reportedly intergrading (Mayr & Cottrell 1986); however, this seems unusual for what are otherwise mainland-breeding ( intermedia ) and island-breeding (i cyanomelana ) taxa. The morphology of birds breeding in South Korea (N. Moores in litt. 2012) clearly fits intermedia and they are treated here as such; however, further research is required to clarify the situation. Species limits Characters of male cumatilis, intermedia and cyanomelana selected for comparison based on Tobias et al. (2010) were assessed (Table 4) . In the absence of clearly independent biometric characters only wing length was assessed and Cohens d values for cumatilis compared to both intermedia and cyanomelana were within the range of 0.2-2 and hence qualified as a minor difference. In terms of vocalisations, spectral differences for cumatilis compared to intermedia and cyanomelana were higher than temporal differences (Cohens d values for all parameters in Table 3 are provided in Table 5) . No behavioural or ecological differences were observed on the breeding grounds. Overall, cumatilis easily achieves the threshold for species status (a score of 7) set by Tobias et al. (2010), scoring 10 when compared against intermedia and 1 1 against cyanomelana (Table 4). Differences between intermedia and cyanomelana (score 3) fell short of the threshold but are viewed here as supporting the treatment of intermedia as a valid subspecies. Table 4. Characters of male cumatilis, intermedia and cyanomelana selected for comparison based on Tobias etal. (2010), with score (see text) in brackets. Character cumatilis vs intermedia cumatilis vs cyanomelana intermedia vs cyanomelana Plumage Underparts Differences in breast, throat and lores combine to render it highly distinctive (3), extensive black streaking (2) Differences in breast, throat and lores combine to render it highly distinctive (3), extensive black streaking (2) Differences in breast, throat and lores due to a difference in shade (1) Upperparts Entire upperparts a different shade (1) Entire upperparts a distinctly different hue (2) Entire upperparts a different shade (1 ) Vocal High frequency (Cohen's d) 3.52(2) 3.69(2) 0.22(1) Pace (Cohen's d) 1.81(1) 1.63(1) 0.06(0) Biometric Wing length (Cohen's d] 0.26(1) 0.48(1) 0.19(0) Total score 10 11 3 Forktail 28 (2012) Zappey's Flycatcher Cyanoptila cumatilis, a forgotten Chinese breeding endemic 127 Table 5. Cohen's d values of vocal parameters of cumatilis, intermedia and cyanomelana selected for analysis. Parameter cumatilis vs intermedia cumatilis vs cyanomelana intermedia vs cyanomelana Low frequency 1.89 3.05 0.88 High frequency 3.52 3.69 0.22 Frequency range 2.92 3.04 0.04 Centre frequency 3.10 4.55 0.25 Peak frequency 2.59 3.45 0.14 Length 0.17 0.35 0.30 Number of elements 0.19 0.03 0.25 Pace 1.81 1.63 0.00 Based on these results the following taxonomic treatment is proposed: Zappey’s Flycatcher Cyanoptila cumatilis Thayer and Bangs, 1909 Blue-and-white Flycatcher Cyanoptila cyanomelana Temminck, 1829 subspecies intermedia Weigold, 1922. male from Hong Kong in October 2008 (Holmes 2010). However, owing to past taxonomic confusion with intermedia , published records of cumatilis cannot be taken at face value: recent examples of published records of cumatilis which clearly refer to intermedia include Shigeta (2003) and Peterson (2006). Lei et al. (2007), Sangster et al. (2010) and Zuccon & Ericson (2010) all concluded that Verditer Flycatcher Muscicapa thalassina is closely related to Cyanoptila cyanomelana and that both should be placed in either Niltava or a sister genus to Niltava. However, none of these studies stated the subspecies or the geographical origin of the Cyanoptila samples and, in view of this, we suggest that more comprehensive genetic work is required, including analysis of material from cumatilis , intermedia and cyanomelana (especially given that some plumage characters are shared between cumatilis and M. thalassina and which may imply a closer relationship than with intermedia or cyanomelana ), so that the exact relationship between the three Cyanoptila taxa, and between each of these and M. thalassina , can be determined before any such changes are adopted. ACKNOWLEDGEMENTS DISCUSSION The English name for cumatilis honours Walter R. Zappey, the collector of the type specimen. The current status of Zappey’s Flycatcher requires further research. It is currently known as a breeding bird from only a small number of locations in central China (see Table 6); however, this includes the type locality, from which it would appear to be unrecorded since 1907. On present knowledge it breeds between 1,000 and 1,700 m in temperate mixed coniferous and deciduous forest from Shaanxi to the mountains of Beijing. This distribution corresponds with the Shanxi Mountains Endemic Bird Area (EBA), and although as yet there are no records from Shanxi, it seems most likely that it occurs there. Two other species breed only in the Shanxi Mountains EBA, Brown Eared- pheasant Crossoptilon mantchuricum and Grey-sided Thrush Turdus fea (BirdLife International 2012a), both breeding at similar altitudes to Zappey’s Flycatcher although the former occurs up to 2,600 m. Both are currently listed as Vulnerable owing to habitat loss to agriculture and urban development and habitat degradation caused by logging and livestock-grazing (BirdLife International 2012b), and it seems likely that Zappey’s Flycatcher would similarly qualify as globally threatened. It is poorly represented in the collections examined (except IOZ) and only two specimens, both migrants, are in BMNH (an adult male from Malaysia dated December 1919 and a first-winter male from Hainan in May 1899, i.e. before cumatilis was formally described). There is a recent field record (photographed) of an adult We would like to thank the following for their invaluable assistance in the field: Alexi Antonov, Mark Brazil, Sergei Surmach, Yoshimirsu Shigeta and David Stanton. Liu Yang helped in locating Mafuling, the type locality of cumatilis. The followingkindly provided additional recordings: Per Alstrom, Mark Brazil, Xia Canwei, Will Duckworth, Paul Holt, Bo Petersson and Mathias Ritschard. Access to museum collections was kindly facilitated by Oleg A. Burkovsky (MZFENU, Vladivostok), Yoshimitsu Shigeta (YIO), Fu Min-Lei (IOZ) and Mark Adams (BMNH). Per Alstrom and Phil Round provided very helpful feedback which improved the clarity of the paper, Yee Lai prepared Figure 1 and Catriona Leven assisted with statistical analysis. Referees Joe Tobias and Nigel Collar provided valuable comments. Finally, Jeremiah Trimble (MCZ) provided photographs of the type of cumatilis. REFERENCES BirdLife International (2012a) Endemic Bird Area factsheet: Shanxi mountains. Downloaded from http://www.birdlife.org on 1 2/06/201 2. BirdLife International (2012b) IUCN Red List for birds. Downloaded from http://www.birdlife.org on 13/06/2012. Brazil, M. A. (1 991 ) The birds of Japan. London: Christopher Flelm. Brazil, M. A. (2009) Birds of East Asia. London: Christopher Flelm. Cheng, Tso-Fisin (1987) A synopsis of the avifauna of China. Beijing: Science Press. Clement, P. (2006) Blue-and-white Flycatcher species account. P.147 in J. del Fioyo, A. Elliott & D. A. Christie, eds. (2006) Handbook of the birds of the world, 1 1 . Barcelona: Lynx Edicions. Table 6. Known breeding-season locations of Cyanoptila cumatilis. Location Coordinates Comment Xiaolongmen, Beijing, China 40°00'N 1 15°28'E Specimens examined (IOZ) and studied in the field Houzhenzi, Shaanxi, China 33°51'N 107°51'E P. Alstrom in lift. 2011; photos examined, recordings analysed for this study Pingli,Shaanxi, China 32°18'N 109°18'E Photos of specimens examined Taibai, Shaanxi, China 34°05'N 107°19'E Photos of sped mens examined Ningshan, Shaanxi, China 33°26'N 108°22'E Photos of specimens examined Foping, Shaanxi, China 33°34'N 107°59'E P. Alstrom in litt. 2011; photos examined Fangxian, Hubei, China 32°03'N 110°44'E MCZ online database: http://www.ma.harvard.edu/collections/searchcollections.html accessed January 2012. Not examined (nor photos seen) by the authors Mafu Ling, Hubei, China 32°03'N 110°44'E Type locality. Photos of type specimen (an adult male) examined 128 PAUL J. LEADER & GEOFF J. CAREY Forktail 28 (2012) Collar, N. J. (201 la) Species limits in some Philippine birds including the Greater Flameback Chrysocolaptes lucidus. Forktail 27: 29-38. Collar, N. J. (201 1 b) Taxonomic notes on some Asian babblers (Timaliidae). Forktail 27: 100-102. Dementiev, G. P. & Gladkov, N. A., eds. (1 954) Birds of the Soviet Union. Vol. 5. Moskva: Sovetskaya Nauka. Hartert, E. & Steinbacher, F. (1934) Die Vogel der palaarktischen Fauna. Erganzungsband. Berlin: R. Friedlander und Sohn. Flolmes, J. (2011) Blue-and-white Flycatcher Cyanoptila cyanometana cumatilis on Po Toi. The first record of this taxon in Flong Kong. Hong Kong Bird Report 2007-2008: 302-303. (In English and Chinese.) Lei, X., Lian, Z.-M., Lei, F.-M., Yin, Z.-H. & Zhao, FH.-F. (2007) Phylogeny of some Muscicapinae species based on cyt b mitrochondrial gene sequences. ActaZool. Sinica 53: 95-105. Mayr, E. & Cottrell, G. W. (1986) Check-list of birds of the world, 1 1. Cambridge, Mass.: Museum of Comparative Zoology. Peterson, A.T. (2006) Taxonomy is important in conservation: a preliminary reassessment of Philippine species-level bird taxonomy. Bird Conserv. internatn. 16: 155-173. Redfern, C. P. F. & Clark, J. A. (2001 ) Ringer's manual. Thetford, U.K.: British Trust for Ornithology. Sangster, G., Alstrom, P, Forsmark, E. & Olsson, U. (2010) Multi-locus phylogenetic analysis of Old World chats and flycatchers reveals extensive paraphyly at family, subfamily and genus level (Aves: Muscicapidae). Molec. Phylogen. Evol. 57: 380-392. Tobias, J. A., Seddon, N„ Spottiswoode, C. N., Pilgrim, J. D„ Fishpool, L. D. C. & Collar, N.J. (2010) Criteria for species delimitation based on phenotype. Ibis 1 52: 724-746. Thayer, J. E.& Bangs, O. (1 909) Descriptions of new birds from central China. Bull. Mus. Comp. Zool. 52: 141. Shigeta, Y. (2003) The first authentic record of Chinese Blue-and-white Flycatcher Cyanoptila cyanomelana cumatilis for Japan. J. Yamashina Inst. Orn. 34: 309-313. (In Japanese with English abstract.) Vaurie, C. (1954) Systematic notes on Palearctic birds. No. 1 2 Muscicapinae, Hirundinidae and Sturnidae. Amer. Mus. Novit. 1 694. Vaurie, C. (1958) The birds of the Palearctic fauna. Passeriformes. London: FI. F. & G. Witherby. Zheng, G„ ed. (201 1 ) A checklist on the classification and distribution of the birds of China. Beijing: Science Press. (In Chinese.) Zuccon, D. & Ericson, P. G. P. (2010) A multi-gene phylogeny disentangles the chat-flycatcher complex (Aves: Muscicapidae). Zoologica Scripta 39: 213-224. PaulJ. LEADER and Geoff J. CAREY, AEC Ltd., 127 Commercial Centre, Palm Springs, New Territories, Hong Kong, Email pji@aechk.hk FORKTAIL 28 (2012): 129-135 Lophura hatinhensis is an invalid taxon ALAIN HENNACHE, SIMON P. MAHOOD, JONATHAN C. EAMES & ETTORE RANDI The Vietnamese Pheasant Lophura hatinhensis was described in 1 975 from one male specimen which was superficially similar to Edwards's Pheasant L. edwardsi but for four white (instead of dark metallic blue) tail feathers. Like L. edwardsi it is poorly known and highly threatened in the wild. Its status as a species has rarely been questioned despite its curious distribution and dubious morphological distinctiveness. To elucidate the taxonomic status of L. hatinhensis we examined the morphology of captive birds of both taxa and analysed mitochondrial DNA. These lines of evidence demonstrated that birds exhibiting the L. hatinhensis phenotype probably represent inbred L. edwardsi. Thus L. hatinhensis should be removed from the IUCN Red List and other checklists of valid extant bird species. Its apparent recent appearance alongside wild populations of L. edwardsi might be taken as evidence that wild populations of this species are also highly inbred and possibly close to extinction. INTRODUCTION The Vietnamese Pheasant Lophura hatinhensis was described by Vo Quy (1975) in his book Chim Viet Nam (translation: ‘Birds Vietnam’) and has been widely recognised as a species ever since (Sibley & Monroe 1990, Inskipp etal. 1996, BirdLife International 200 1 , 20 1 1 , Dickinson 2003). However, owing to its close similarity to Edwards’s Pheasant L. edwardsi it has been considered a subspecies of that species (e.g. del Hoyo et al. 1994, Johnsgard 1999), a species inquirenda (Vuilleumier etal. 1992; also BirdLife International 2001), not recognised at all (Johnsgard 1986) or treated ambiguously (Madge & McGowan 2002, Hennache & Ottaviani 2005). BothZ. hatinhensis andZ. edwardsi are extremely rare denizens of low-lying broadleaved evergreen forests in the Annamite Mountains of central Vietnam, and remain very poorly known in the wild (BirdLife International 2001). Both were classified as Endangered until early 2012 when L. edwardsi was uplisted to Critically Endangered (BirdLife International 2012a). Lophura hatinhensis records derive primarily from the area to the north of the distribution of L. edwardsi, although there is one record from Thua Tien Hue province on the southern limit of the range of L. edwardsi (BirdLife International 2001). Male L. hatinhensis and L. edwardsi are morphologically very similar. The type description of L. hatinhensis (Vo Quy 1975) diagnoses the species as (our translation): Lophura hatinhensis sp. nov. Male (adult) : white crest with black at the tip. Black underparts (belly). Head, neck, breast, upperparts and rump (uppertail) are black with glossy purplish- blue. Wing-coverts are dark blue; upperparts and tail-coverts black with black lines at the tip; four central tail feathers pure white, other tail feathers black; wing feathers black, facial skin and legs red, bill black. Measurements (male holotype): wing 245, tail 270, leg 89, bill 30 mm. Weight 1 , 1 00 g. In comparison with closely related pheasants like L. imperialis, L. edwardsi in Vietnam, L. inornata in Sumatra and L. swinhoei in Taiwan, the new species is closer to L. edwardsi. The only difference is that the new species has a darker colour, no shiny green and four white tail feathers. Other authors have noted additional differences between the taxa, reporting that L. hatinhensis is larger than L. edwardsi with a slightly downcurved tail with pointed central tail feathers and longer tarsus, and that both species have pronounced metallic green wings, except in the breeding season when L. hatinhensis develops a distinct reddish-purple colour on the wings (Dang Gia Tung & Le Sy Thuc 1996, Hennache etal. 1999). Mitochondrial DNA analyses, using 15 samples of L. hatinhensis and six of L. edwardsi, suggested that the two taxa are each other’s closest relatives (Randi et al. 1997, Scott 1997, Hennache et al. 2003) and that they diverged within the last 100,000 years (Scott 1997, Hennache et al. 1999). Although their phylogenetic relationships could not be accurately determined, Scott (1997) proposed that they should be considered evolutionary significant units. Based on these data Hennache et al. (1999) recommended that they should not be allowed to interbreed in captivity. Despite the widespread acceptance of L. hatinhensis as a species, there is considerable uncertainty regarding its diagnosis. Here we present the results of the first thorough investigation into the validity of L. hatinhensis, bringing together genetic and morphological data. The histories of the captive populations of the two taxa are of relevance to any discussion of their morphology, and these are therefore documented here. We present previously unpublished genetic data and synthesise morphological data that suggest that individuals that are phenotypically classifiable as L. hatinhensis probably represent L. edwardsi-, we propose that inbreeding is the most likely mechanism for this phenomenon. We believe that/,, hatinhensis has no taxonomic standing, and therefore that all records of this taxon are attributable to L. edwardsi. MATERIALS AND METHODS The type description of L. hatinhensis apparently involved a single male individual (see above), but the author did not assign it to a particular specimen, nor did he indicate a specimen number, place of deposition of the specimen, or the place and date of its collection. The distribution of the species was given as ‘areas of mountainous forest in Ky Anh district, Ha Tinh province’ and its status as ‘rare in our country’ (Vo Quy 1975). Confusion surrounds this description, since according to BirdLife International (20 1 2b) the species was discovered in 1964 and described by ‘Vo Quy & Do Ngoc Quang (1965)’. Yet this reference does not appear in the BirdLife reference list and we have only been able to trace one paper by these authors in 1 965 whose subject is a collection of birds made in Cao Bang and Lang Son provinces in northern Vietnam (and thus far from Ha Tinh province, which is in central Vietnam) (Vo Quy & Do Ngoc Quang 1965). Rozendaal (1991) also reported thatZ. hatinhensis was described (in Vietnamese in a publication both difficult to obtain and unclearly referenced) from a single male specimen, preserved in the Institute for Ecology and Biological Resources, Hanoi, collected on 26 January 1964 by the late Do Ngoc near Ky Son (Ky Anh district, Nghe Tinh province [name since reverted to Ha Tinh province] c.17°59,N 106°10T, while a second male was taken in 1974 by Truong Van La at the same locality but was only partially preserved; Robson etal. (1989) gave briefer, similar evidence but reported the type locality as ‘Song 130 ALAIN HENNACHE etal. Forktail 28 (2012) Table 1. Samples used in the genetic analysis. 'Origin unknown' refers to wild-caught individuals lacking information on collecting location. Sample ID Species Sample Origin Captive/wild Sex Comments LED29 L. edwardsi Blood Jersey Zoo C F Studbook number 502 LED55 L. edwardsi Toepad MNHN C (FI) U Skin number 878. Born in Cleres, France, died 1931 LED56 L. edwardsi Toepad MNHN - Cam Fo, Quang Tri province W M Skin number 922. Collected 29/1 2/1923 by J. Delacour LED57 L. edwardsi Toepad MNHN - Thua Tien Hue province w F Skin n°2882. Collected December 1927 by Delacour LED58 L. edwardsi Feather Phuong Dien district, Thua Thien Hue province w M Trapped in August 1996 LED59 L. edwardsi Feather Phuong Dien district, Thua Thien Hue province w M Trapped in August 1996 LED74 L. edwardsi Feather Hanoi Zoo - Huong Hoa district, Quang Tri province w M Trapped on 23/11/1996 LED107 L. edwardsi Feather Hai Fang district, Quang Tri province w M Trapped in 2000 and died shortly after LHA1K L. hatinhensis Feather Hanoi Zoo - Minh Hoa district, Quang Binh province w M LHA2 L. hatinhensis Blood Hanoi Zoo w F Origin unknown LHA3K L. hatinhensis Feather Cleres (France) C (FI) M On breeding loan from Hanoi Zoo LHA4K L. hatinhensis Feather Cleres (France) C (FI) F On breeding loan from Hanoi Zoo LHA5 L. hatinhensis Blood Hanoi Zoo W U Origin unknown LHA6 L. hatinhensis Blood Hanoi Zoo w U Origin unknown LHA7 L. hatinhensis Feather Hanoi Zoo w F Origin unknown LHA8 L. hatinhensis Feather Private collection, Thailand w M Origin unknown LHA9 L. hatinhensis Feather Private collection, Thailand w F Origin unknown LHA10 L. hatinhensis Feather Private collection, Thailand C (FI) U LHA11 L. hatinhensis Feather Private collection, Thailand C (FI) U LHA12 L. hatinhensis Feather Hanoi Zoo w U Origin unknown LHA13 L hatinhensis Feather Hanoi Zoo w U Origin unknown LHA14 L. hatinhensis Feather Hanoi Zoo C (FI) M Origin of parents Minh Hoa District, Quang Binh province LHA15 L hatinhensis Feather Hanoi Zoo - Huong Thuy district, Thua Tien Hue province w M Trapped in 1999 when it walked into a farmer's house Tung’. We were unable to make direct comparisons with the holotype in the preparation of this paper, but one of us (JCE) has previously examined and photographed it (Fig. 13 in Rozendaal 1991). Morphological analyses Morphological data were collected from adults, mainly concerning the colour of the neck, mantle and wing-coverts, the number, form and colour of the tail feathers, and the age at which white tail feathers (if any) are developed. These are derived from personal observations by AH (adult A. hatinhensis : four wild-caught birds and at least 12 captive-bred birds; adult A. edwardsi: one wild- caught male, three museum skins of wild-caught birds and at least 16 captive-bred birds) and by others (in pers. comms. to AH) on the plumage of wild-caught A. hatinhensis held at Hanoi Zoo and on captive-bred birds of both taxa held at Hanoi Zoo, in European zoos and in private collections. The history of the captive populations was reconstructed using the international studbooks of A. edwardsi (Hennache 2003) and A. hatinhensis (Hennache 2008). The A. edwardsi studbook was resurrected in 1994 and carefully maintained by AH until 2009. DNA extraction, amplification and sequencing Total DNA was extracted from 95% ethanol-preserved tissue (skin or toe-pad) or feather root samples, using procedures described by Randi & Lucchini (1998). The 5’ domain of the mitochondrial DNA control region (mtDNA CR) was PCR-amplified and sequenced as previously described (Randi & Lucchini 1 998, Randi et al. 2001). CR sequences were obtained from living birds that were identified from morphological features as A. hatinhensis (n=15; comprising ten wild-caught birds and five FI generation captive-bred birds derived from wild birds), birds which showed morphological features of A. edwardsi (n=8; comprising two birds collected during the 1920s and 1930s, four wild birds collected since 1996, one modern captive-bred individual and one captive individual born in the 1930s from wild-caught parents) (Table 1) and A. swinhoei (Swinhoe’s Pheasant, n=l), a closely related outgroup. The CR sequences were aligned using CLUSTAL X with the default options (Thompson etal. 1997). Phylogenetic analyses were performed using the software PAUP* (Swofford 1998) by: (1) a maximum-parsimony procedure (Swofford 1998), excluding all uninformative nucleotide positions, with unordered and equally weighted characters; (2) the neighbour-joining algorithm (Saitou & Nei 1987), with Tamura & Nei’s (1993) DNA distances. Robustness of the phylogenies was assessed by bootstrap percentages (BP: Felsenstein 1985), with 1,000 random resamplings with replacement. Details on phylogenetic analyses are given in Randi et al. (2001). RESULTS Morphological analyses The A. edwardsi studbook revealed that the captive stock is derived from 28 specimens, of which only 6-8 were females, collected between 1924 and 1930, and never subsequently supplemented with wild birds (Ciarpaglini & Hennache 1997). It is therefore highly inbred, particularly in America where birds are derived from an even smaller subset of lounders imported from France and England before World War II. The A. hatinhensis studbook is more Forktail 28 (2012) Lophura hatinhensis is an invalid taxon 131 recent: the first record of the taxon in captivity was in 1 990 when Hanoi Zoo obtained six wild-caught A. hatinhensis (four males and two females) from hunters. These were reportedly caught in Minh Hoa district (QuangBinh province), but further information about the location of their capture is unfortunately unavailable. Two males and one female died shortly afterwards from injuries sustained during their capture. In 1991 Hanoi Zoo purchased an additional female so that it had two pairs of A. hatinhensis for captive breeding. During the following seven years nearly 50 chicks were hatched in Hanoi Zoo from these two pairs and their offspring, and a few additional wild birds were purchased to augment the population. In 1996 the first A. hatinhensis (two male and two female FI generation captive-bred birds) to be exported from Vietnam were received at Cleres, France, where they bred the following year. The descendants of this pair were distributed widely in Europe, thus establishing the European captive stock. There are reportedly no A. hatinhensis in the USA and no importation there is documented. Observations of captive-bred and wild-caught birds have indicated that the plumage ofL. hatinhensis is unstable and exhibits more variation than the type description and other sources would suggest (Corder 1 996, Dang Gia Tung & Le Sy Thuc 1 996, Davison 1996). There is variation in the number, distribution and morphology of white tail feathers and in the timing of their development. Observations of captive male A. hatinhensis have shown that the ‘diagnostic’ white tail feathers normally develop after the first adult moult, when the bird is 1 8 months old, although in some individuals they appear earlier (at 15 months) or do not appear until the bird is 24 or even 30 months old (Dang Gia Tung & Le Sy Thuc 1996, AH pers. obs.). The number of white tail feathers exhibited by A. hatinhensis is variable and ranges from one to six; moreover, they are often distributed asymmetrically (Dang Gia Tung & Le Sy Thuc 1996, AH pers. obs.). Their morphology varies individually (AH pers. obs.). The feathers may be entirely white or partially white with brownish streaks and patches. For instance, a male A. hatinhensis which died on 10 November 1999 aged 30 months at Cau Dien Breeding Centre (Vietnam) had one white tail feather to the left of the centre of the tail and two feathers (one to the left and one to the right of the centre of the tail) which exhibited a mix of brown and white patches, one of which was entirely brown except the white tip (Plate 1). Moreover, the occurrence of white in the plumage of A. hatinhensis is not always limited to the central tail feathers: a male A. hatinhensis (identified by its white tail feathers) caught near Hue in 1999 and subsequently retained in Hanoi Zoo developed white tertials after its first moult in captivity (Plate 2). The female A. hatinhensis is very similar to that of A. edwardsi, as Rozendaal (1991) showed: body plumage and wing-coverts chestnut, head and neck tinged grey; remiges dark brown, vermiculated with chestnut on the inner vane, outer web pale brown; tail incomplete, three outer pairs of rectrices blue-black, the outermost rectrix with brown basal half of outer web; presumably at least two central pairs of rectrices with more brown. Bill dark horn, orbital skin and feet scarlet, iris dark brown. Thus it appears not to differ substantially from female A. edwardsi , except perhaps for the warmer tone to the underparts. Although some authors have claimed that it has more reddish-chestnut plumage (Dang Gia Tung & Le Sy Thuc 1996, Hennache etal. 1999) there is considerable individual variation in captive individuals. Some captive female A. hatinhensis are indistinguishable on plumage from female A. edwardsi , whilst others possess 1-4 central tail feathers which may be entirely white or, more often, brown with white borders and streaks. Morphological features thought to be unique to A. hatinhensis have arisen in pure-bred captive lines of A. edwardsi and in some individuals typical A. edwardsi plumage features have been lost Plate 1 . Tail feathers of a male L hatinhensis which died in Cau Dien Breeding Center in 1999 showing variable pattern of white on tail feathers. (Alain Hennache) Plate 2. Dorsal view of a male L. hatinhensis caught near Hue in 1999 and subsequently retained in Hanoi Zoo that developed white tertials after its first moult in captivity. (Alain Hennache) (Corder 1996). Towards the end of 1999 at Cleres, France, a six- year-old male A. edwardsi with a history well documented in the international studbook (its parents were traced back to four different bloodlines) developed three white tail feathers on the centre-left and two on the centre-right of its tail: morphologically it had become indistinguishable from A. hatinhensis (Plate 3). In 1998 a female A. edwardsi held at a collection in Germany developed one white central tail feather when it was three years old (Plate 4). In 1997 a male A. edivardsi held at a collection in Alabama, USA, developed 132 ALAIN HENNACHE etal. Forktail 28 (2012) Plate 3. Tail of male L. edwardsi born in France 1993 (international studbook number 586) showing two partially white tail feathers that developed after six years. (Alain Flennache) white tail feathers (Plate 5). Inbreeding has led to a number of other morphological changes in the captive population of A. edwardsi: in the 1970s the crest was reduced or absent on some birds (Lovel 1979) and in 1999 birds imported from the USA were on average a third smaller than European captive birds (AH pers. obs.). The other morphological features mentioned by Vo Quy (1975) as unique to A. hatinhensis , namely a lack of shiny green feathers and darker plumage, have on examination of a larger number of captive-bred and wild-caught birds been shown to be invalid (Dang Gia Tung &LeSyThuc 1996, Hennache er ml. 1999). Genetics The relationships of the A. hatinhensis and A. edwardsi mtDNA CR sequences are described by the neighbour-joining tree (Figure 1). Sequences of A. hatinhensis are very similar to those of A. edwardsi with a low level of sequence divergence (mean 0.6%, max 1 %). Lophura hatinhensis shows eight mtDNA haplotypes of which one clusters within edwardsi (LHA3). Lophura edwardsi shows three haplotypes of which one (LED 107) clusters within one group of A. hatinhensis. With the exception of LED 107, all the recent and historical wild A. edwardsi have the same haplotype, which differs slightly from the captive stock (LED 29 and LED 55). The neighbour-joining tree represented in Figure 1 is based on c.700 nucleotides. Bootstrap values were typically very low. DISCUSSION Lophura hatinhensis is a species with a very short history. It was discovered in 1964 and described in 1975; fewer than 50 individuals have been recorded in the wild with any degree of certainty and of Plate 4. Tail of female L. edwardsi born in Germany 1 995 (international studbook number 882) showing one white tail feather. (Alain Hennache) Plate 5. Male L. edwardsi born in Alabama, USA, which developed white central tail feathers in 1997. This bird phenotypically resembles L. hatinhensis. (Michel Klat) those 22 were already dead and at least seven were taken into captivity (BirdLife International 2001, AH pers. data). Our data demonstrate that L. hatinhensis is characterised by little genetic differentiation from L. edwardsi. Morphological analysis has shown that individual L. edwardsi of known pedigree can develop plumage features that ostensibly render them indistinguishable phenotypically from L. hatinhensis. We have also documented variation within the A. hatinhensis tail phenotype and shown that it is unstable and does not conform neatly to the ‘middle four white tail feathers’ described by Vo Quy (1975). Other plumage features Forktail 28 (2012) Lophura hatinhensis is an invalid taxon 133 Figure 1. Neighbour-joining tree showing phylogenetic relationships of the sequenced mitochondrial DNA control regions of L. edwardsi, L. hatinhensis and L. swinhoei (LED, L. edwardsi; LHA, L. hatinhensis ; LSW, L. swinhoei). 0 LHA1K LHAAK LHA6 LHA7 LHA14 — LED107 - LHA12 LHA15 — LHA2 LED29 _ LHA8 LHA9 LHA10 _ LHA1 1 LHA13 _ LHA5 I LED 59 LED58 LED57 LED56 _ED74 LED55 — 0 001 substttutions/site LSW1 previously described as unique to A. hatinhensis have already been shown to be irrelevant (Dang Gia Tung & Le Sy Thuc 1996, Hennache etal. 1 999). Taken together these findings demonstrate that A. hatinhensis has no taxonomic standing. We therefore suggest it be removed from the IUCN list of threatened species, and all other relevant extant bird checklists. This reduces the number of Vietnamese endemic Lophura to one: L. edwardsi. The other enigmatic Vietnamese endemic Lophura , L. imperialism has already been shown to represent a hybrid between L. edwardsi and Silver Pheasant/,, nycthemera, based on genetic and morphological evidence and captive-breeding experiments (Hennache etal. 2003). Although our morphological study relies almost entirely on captive birds, L. edwardsi is so poorly known in the wild that data are insufficient for a thorough analysis. In addition, captive birds are of known heritage and can therefore reveal details about morphology that observations of wild individuals of unknown parentage could not. If an individual L. edwardsi were to develop white tail feathers in a wild state it would just be assumed to represent L. hatinhensis ; indeed this is probably what happened to the male caught near Hue (within the range of L. edwardsi and far to the south of previous records of L. hatinhensis ) in 1999. Albinism in birds is thought to result from the expression of recessive alleles that disrupt melanin pigmentation at feather development (Bensch et al. 2000). These alleles are usually only expressed when the population is highly inbred. This phenomenon is well known to breeders who have reared birds over many generations without change to the bloodlines. Even when populations are highly inbred, characters which arise owing to inbreeding do not usually become fixed; for instance, in an isolated population of Great Reed Warblers Acrocephalus arundinaceus with a small founder stock partial albinism was only recorded during the first five years of the colony’s existence (Bensch et al. 2000). In contrast, the presence of white tail feathers in L. hatinhensis has become at least partially fixed, and this is perhaps because of the prevalence of a white (or buff) tail feathers in the genus Lophura. Lophura edwardsi and Siamese Fireback L. diardi are unique among Lophura pheasants in that they do not possess any white or buff tail feathers, the presence of which otherwise characterises the genus. Mutations expressed in captive Lophura of other species as a result of inbreeding have resulted in birds with additional white tail feathers. For instance, in Australia where the captive population of L. swinhoei is highly inbred, several males developed a second pair of white central tail feathers and one male developed five white tail feathers and a larger white crest (Weber 1992). The/,, edwardsi captive stock is highly inbred; indeed all captive-bred birds analysed by Randi et al. (1997) have the same nucleotide sequence at the mtDNA control region compared with five wild-caught L. edwardsi which exhibited nucleotide substitutions, a result which is perhaps unsurprising since the captive-reared birds were all derived from a single female. The prevalence of white tail feathers in Lophura perhaps explains why it is this feature that is the primary visual manifestation of inbreeding in L. edwardsi. An alternative explanation for the expression of white tail feathers in captive L. edwardsi would be that these birds represent hybridisation with L. hatinhensis. However, the timing of the birth of the three European captive L. edwardsi that developed white tail feathers precludes any chance that they are the result of such hybridisation, since they were hatched before L. hatinhensis was first exported from Vietnam. Unfortunately it was not possible to compare genetic and morphological data presented in this study with the type specimen of L. hatinhensis. However, it is unlikely that any of our conclusions would have changed as a result of this. Although some of Vo Quy’s (1975) measurements of the type specimen are larger than all L. edwardsi measured by Oustalet (1896) and Delacour (1977) (Table 2), this bird is only marginally larger and some of the more striking differences, especially tarsus length, may be the product of differences in methods for taking measurements. With a larger sample size (Vo Quy only measured one L. hatinhensis ) the measurements might be found to overlap with those for A. edwardsi. All L. hatinhensis examined by AH show no differences in size, colour or intensity of gloss from captive L. edwardsi. Therefore the only morphological feature that can be used to identify L. hatinhensis is the presence of one or more white or partially white tail feathers, and our data indicate that these can arise in lines of inbred pure-bred L. edwardsi. Lophura hatinhensis and L. edwardsi exhibit shallow genetic differentiation at a level that does not support their species-level separation. Research has shown that pheasant species pairs typically show genetic divergence of at least 2% (Randi et al. 2001). For instance, L. edwardsi and L. swinhoei differ by 2.5%, L. leucomelanos (Kalij Pheasant) and L. nycthemera by 2.8% (or 2.5%: Moulin et al. 2003), A. diardi and A. ignita (Crested Fireback) by 4.4% (Randi et al. 2001) and Tragopan species pairs by 3. 6-5. 9% (Randi et al. 2000). Our application of the mtDNA genes uncovers considerable differentiation among closely related pheasants (see Figure 2 in Hennache et al. 2003). Sampling was insufficient to determine 134 ALAIN HENNACHE eta/. Forktail 28 (2012) Table 2. Published measurements in mm of male L. edwardsi and L. hatinhensis. Numbers constitute means unless more than one number is given. Taxon No. Birds Length(head-body) Wing Tail Culmen Tarsus Source L. edwardsi 3 580 230 220 30 83 Oustalet 1896 L. edwardsi 14 580-650 220-240 240-260 30 75 Delacour 1977 L. hatinhensis 1 - 245 270 30 89 ('feet') Vo Quy 1975 whether the nine mtDNA haplotypes apparent in the neighbour¬ joining tree represent geographic differentiation, although this seems very unlikely. Data in Hennache etal. (2003) indicated that, when compared with each other, A. hatinhensis showed five unique alleles and A. edwardsi eight. However, all A. edwardsi examined were captive birds from a highly inbred line, and these data are therefore thought to indicate a loss of alleles from the captive population of A. edwardsi rather than evidence of unique alleles in the population of A. hatinhensis. With the exception of the one bird found in Thua Tien Hue province (mentioned above), located to the south of the range of A. edwardsi , all records of A. hatinhensis derive from the area to the north of the range of A. edwardsi. Were it not for the Thua Tien Hue bird it would be plausible that A. hatinhensis represents a northerly subspecies of A. edwardsi. However, to support the subspecies theory the bird from Thua Tien Hue province must be considered an aberrant A. edwardsi whose appearance coincidentally matches that of true A. hatinhensis. Moreover, the appearance of captive A. edwardsi superficially resembling A. hatinhensis in inbred lines would then have to be explained as a coincidence and the unstable phenotype of A. hatinhensis ignored. The improbability of these circumstances is so high that the burden of proof must now be on those who would seek to uphold the validity of this taxon, whether as a species or a subspecies. Based on the occurrence of white tail feathers in inbred captive populations of A. edwardsi we propose that inbreeding might be the mechanism that has caused the occurrence of the A. hatinhensis phenotype in the wild; this was first proposed by Hennache & Ottaviani (2005). Following this theory, we suggest that records of wild birds with the A. hatinhensis phenotype have been made on the northern and southern periphery of the range of A. edwardsi , suggesting that at least outside of the core range (where there have been no records since the late 1990s) the wild population is fragmented and possibly very inbred. It has taken approximately 35 generations for the captive population of A. edwardsi to develop the A. hatinhensis phenotype, despite originating from a tiny founder population. This indicates that the processes that have led to the dominance of the A. hatinhensis phenotype in some wild populations of A. edwardsi have been acting since its discovery, and probably long before. Lophura edwardsi is now very rare in the wild: there have only been two unequivocal records since 2000, a male trapped in Hai Lang district, Quang Tri province (which later died) and a male found in a farmer’s cage in Quang Tri province in 2009 (Dan Tri 2009). It is even conceivable that there are now no remaining wild populations of A. edwardsi ( Babbler 39 [November 201 1]: 41). Any remaining populations may either exhibit the A. hatinhensis phenotype or have not yet developed it, but like the captive populations they may already be so inbred that the appearance of such a phenotype is only a matter of time. Whilst the captive population is known to derive from a very small founder stock the genetic diversity of the wild population is unknown. In addition to showing white tail feathers, inbred birds might possibly exhibit physiological characteristics, such as reduced fertility or higher mortality rates, which might mean that populations showing the A. hatinhensis phenotype are unlikely to persist in perpetuity. As an example, at the end of the 1960s, A. edwardsi was increasingly difficult to breed reliably in the United Kingdom and many eggs laid were infertile (Lovel 1979). The low genetic diversity of the captive population of A. edwardsi , and the recent appearance in it of birds which could be classified as A. hatinhensis , serve as a warning that this population is not an adequate safety net for restocking areas where wild populations have become extinct. Even captive populations that have not yet developed the A. hatinhensis phenotype might yet do so, and care should be taken in managing the captive population to maximise genetic diversity. Although our research has brought some clarity to an enigmatic taxonomic situation, it also indicates that A. ediuardsi may be closer to extinction and more difficult to rescue than previously thought. If wild populations of the inbred A. hatinhensis phenotype can be found it might be prudent, after breeding experiments, to introduce a small number of genetically pure captive-bred birds which still show the A. edwardsi phenotype, in the hope that since they were derived from birds collected many years ago they may introduce some lost genetic diversity into the wild population and rescue them from possible inbreeding depression. ACKNOWLEDGEMENTS We would like to thank Le Sy Thuc (Hanoi Zoo) and Professor Vo Quy (CRES, Hanoi) for providing us with blood samples from wild-trapped birds and Claire and Jean-Fran^ois Voisin (MNHN, Paris) who provided us with museum samples. Dang Gia Tung must be thanked for his assistance in Hanoi in helping us to check captive/,, hatinhensis, and Truong Van La shared information about records of wild L. edwardsi. Michel Klat, Bernard Liauzu and Edouard Jelen sent us photographs or living specimens of inbred captive/,, edwardsi with white tail feathers that were vital for this publication. Le Trong Trai provided us with vital information regarding the holotype and a translation of the type description, and John Sherwell supplied us with some difficult-to-obtain publications at short notice. Two anonymous reviewers provided us with invaluable comments on an advanced draft of this manuscript. REFERENCES Bensch, S., Hansson, B., Hasselquist, D. & Nielsen, B. (2000) Partial albinism in a semi-isolated population of Great Reed Warblers. Hereditas 133: 167-170. B i rd Life International (2001) Threatened birds of Asia: the BirdLife International Red Data Book. Cambridge, UK: BirdLife International. 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(1986) The pheasants of the world: biology and natural history . First edition. Washington, DC: Smithsonian Institution. Johnsgard, P. A. (1999) The pheasants of the world: biology and natural history . Second edition. Washington, DC: Smithsonian Institution. Lovel.T. W. I. (1 979) Studbook for the Edwards's pheasant Lophura edwardsi. Internatn. Zoo Yearbook 19: 281-283. Madge, S. & McGowan, P. (2002) Pheasants, partridges and grouse. London: Helm Identification Guide. Moulin, S., Randi, E.,Tabarroni, C. & Hennache, A. (2003) Mitochondrial DNA diversification among the subspecies of the Silver and Kalij Pheasants ( Lophura nycthemera and L. leucomelanos) (Galliformes, Phasianidae). Ibis 145: El -El 1. Oustalet, M. E. (1896) Description de cinq nouvelles especes d'oiseaux appartenant au Museum national d'Histoire naturelle et provenant de Chine et d'lndo-Chine. Bull. Mus. Paris 1896 (6) : 314-317. Randi, E. & Lucchini, V. (1998) Organisation and evolution of the mitochondrial DNA control-region in the avian genus Alectoris. J. Molec. Evol. 47: 449-462. Randi, E., Lucchini, V. & Hennache, A. (1997) Searching for mtDNA genetic diversity in captive Edwards's Pheasants. Pp.1 17-123 in A. Hennache, coord. Conservation et Studbook International du Faisan d‘ Edwards Lophura edwardsi. The international studbook for the Edwards's Pheasant Lophura edwardsi and its conservation. Paris: Ed. Service du Patrimoine Naturel, MNHN (Patrimoines naturels 30: 1 -254). Randi, E., Lucchini, V., Armijo-Prewitt, T., Kimball, R.T., Braun, E. L. & Ligon, D. (2000) Mitochondrial phylogeny and speciation in the tragopans. Auk 117: 1007-1019. Randi, E., Lucchini, V., Hennache, A., Kimball, R.T., Braun, E. L. & Ligon, J. D, (2001) Evolution of the mitochondrial DNA control-region and cytochrome b genes, and the inference of phylogenetic relationships in the avian genus Lophura (Galliformes)./Wo/ec. Phylogen. Evol. 1 9: 1 87- 201. Robson, C. R., Eames, J. C., Wolstencroft, J. A., Nguyen Cu & Truong Van La (1989) Recent records of birds from Viet Nam. Forktail 5: 71-97. Rozendaal, F. (1991 ) Notes on Vietnamese pheasants with a description of female Lophura hatinhensis. Dutch Birding 13: 12-15. Saitou, M. & Nei, M. (1987) The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molec. Biol. Evol. 4: 406-425. Scott, E. A. (1997) Systematic relationships of endemic Vietnamese Pheasants. Pp. 26-30 in A. Hennache, coord. Conservation et Studbook International du Faisan d'Edwards Lophura edwardsi. The International Studbook for the Edwards's Pheasant Lophura edwardsi and its conservation. Paris: Ed. Service du Patrimoine Naturel, MNHN (Patrimoines naturels 30: 1-254). Sibley, C. G. & Monroe, B. L. (1990) Distribution and taxonomy of birds of the world. New Haven: Yale University Press. Swofford, D. L. (1998) PAUP*: phylogenetic analysis using parsimony (and other methods) Version 4.0b2a. Sunderland MA: Sinauer Associates. Tamura, K. & Nei, M. (1993) Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molec. Biol. Evol. 10: 512-526. Thompson, J. D„ Gibson, T. J., Plewniak, F„ Jeanmougin, F. & Higgins, D. G. (1997)The CLUSTAL-windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 24: 4876-4882. Vuilleumier, F., LeCroy, M. & Mayr, E. (1992) New species of birds described from 1981 to 1990. Bull. Brit. Orn. Club 1 12 A: 267-309. Vo Quy & Do Ngoc Quang (1965) Ket qua suu tarn chim o vung Bao Lac, Trung Khanh (Cao Bang) va vung Mau Son (Lang Son). Sinh V-t HIc T-p (‘Biological Studies') IV. (In Vietnamese.) Vo Quy (1975) Chim Viet Nam, 1 . Hanoi: Nha Xuat Ban Khoa Hoc Va KyThuat. (In Vietnamese.) Weber, R. (1992) The Swinhoe's pheasant. WPA News 37: 29-30. Alain HENNACHE, 91 route de la vallee, 76890 Saint Victor lAbbaye, France. Email: alain.hennache@wanadoo.fr Simon P. MAHOOD, I/I /CS Cambodia, #21 Street 21, Sangkat Tonle Bassac, P.O. Box 1620, Phnom Penh, Cambodia. Email: s.mahood@wcscambodia.org Jonathan C. EAMES OBE, BirdLife International Cambodia Programme, #9, Street 29 Tonle Basac, Chamkarmon, P.O. Box 2686, Phnom Penh, Cambodia. Email: eames@birdlife.org.vn Ettore RANDI, ISPRA Italy, Laboratory of Genetics, Via Ca Fornacetta, 940064 Ozzano Emilia (BO), Italy. Email: ettore.randi@isprambiente.it 136 SHORT NOTES Forktail 28 (2012) Bar-winged Wren Babbler Spelaeornis troglodytoides : a first record for Vietnam, with speculation for 1 7 further avifaunal additions SIMON P. MAHOOD, DAVID P. EDWARDS & FELICITY A. EDWARDS The first record for Vietnam At 10h30 on 4 May 2010, SPM and DPE found a Bar-winged Wren Babbler Spelaeornis troglodytoides just below the summit camp on Mount Fan Si Pan, in the Hoang Lien Son National Park, Sapa, West Tonkin. Mount Fan Si Pan reaches 3,143 m, making it the highest peak in the Hoang Lien Mountains and in Vietnam. The bird was in a rocky gully vegetated with Rhododendron, stunted trees and dense Arundinaria bamboo at c.2,800 m elevation. It was found when it responded to the broadcast of a pre-recorded cut of Gould's Shortwing Brachypteryx stellata. When the cut was played the wren babbler flew up out of the gully, perching openly on a bamboo stem 4 m from the observers, and sang. DPE obtained a number of audio recordings (xc65028, xc65029, xc65030) and FAE obtained a series of photographs (Plates 1 and 2). Although immediately recognised as a Bar-winged Wren Babbler (confirmed in Plates 1 and 2 by the combination of white throat, rich rufous breast-sides and belly-sides, heavily barred wings and relatively long, barred tail), we were aware that this species had not previously been recorded in Vietnam (Robson 2008). It is distributed in the eastern Himalayas, from western Bhutan to northern Myanmar and Yunnan province, China, and in three disjunct populations within central China (Collar & Robson 2007). The Hoang Lien Mountains of Vietnam are the south-easternmost extension of the Himalayas and thus its occurrence in this region is not altogether unexpected. Since wren babblers are not given to vagrancy, it can be safely concluded that our record represents a breeding range extension of some 550 km from the nearest known population in north-west Yunnan, China. A new subspecies? Seven subspecies of Bar-winged Wren Babbler have been described (Collar & Robson 2007). They differ mainly in tone and pattern of plumage, and possibly also in vocalisations, although vocal variation is not well documented. Plumage variation within recognised subspecies is confounded by apparent sexual dimorphism (Rasmussen & Anderton 2005), unusual in the Timaliidae. Comparison of Plates 1 and 2 with published descriptions of recognised subspecies (Rasmussen & Anderton 2005, Collar & Robson 2007) and photographs of known subspecies (OBC image database) indicates that the Vietnam bird has a number of unique plumage features, including: (1) blackish lores that lack contrast with the crown, (2) dark grey cheeks that contrast strongly with the ear-coverts, and (3) crown coloration that lacks a rufous tinge. It is also perhaps brighter orange-rufous above and below than other subspecies, and more extensively white below, although it approaches S. t. sherriffi (of Bhutan and adjacent Arunachal Pradesh, India) in underpart pattern and coloration. We therefore suggest that the Vietnamese population of Bar-winged Wren Babbler may represent an undescribed subspecies. However, our observations and photographs concerned a single damp individual and comparison has not been made against specimens. Moreover, wren babblers can show fairly extreme intra¬ species variation, even within populations of species (see, e.g., Eames & Mahood 201 1). No firm conclusions should therefore be drawn regarding the taxonomic distinctiveness (or otherwise) of the Vietnamese population of Bar-winged Wren Babbler without the greater use of specimen evidence. The species as a whole is in need of taxonomic review owing to significant plumage variation across its wide range; this should incorporate vocal and, if possible, genetic data. Bar-winged Wren Babbler is nowhere regarded as abundant, making it likely to be an uncommon resident in the Hoang Lien Mountains. Our record in Vietnam falls within the published altitudinal range of Bar-winged Wren Babbler in the Indian Subcontinent (2,400-3,355 m; Rasmussen & Anderton 2005) and Myanmar (2,440-2,895 m; Robson 2008), butabovethat published for China (1,600-2,440 m; Collar & Robson 2007). If the elevational Plates 1 and 2. Bar-winged Wren Babbler Spelaeornis troglodytoides : Mount Fan Si Pan, Vietnam, at c.2,800 m elevation, 4 May 2010. Both photos are of the same individual. Note the blackish lores and dark grey cheeks lacking contrast with the blackish crown, and also the extensive white and bright orange-rufous below. (Felicity A. Edwards) Forktail 28 (2012) SHORT NOTES 1 37 Table 1. Species with a similar distribution to Ban from Robson (2005). -winged Wren Babbler that have not been recorded in Vietnam. Data on habitat and elevation Species Habitat Elevation (m) Speckled Wood Pigeon Columba hodgsonii Broadleaved evergreen forest 1,675-2,565 Spotted Nutcracker Nucifraga caryocatactes Open coniferous and mixed broadleaved and coniferous forest 2,285-3,600 Slaty-backed Flycatcher Ficedula hodgsonii Broadleaved evergreen forest, secondary growth 600-2,750 Indian Blue Robin Lusciniabrunnea Bamboo, secondary growth, thickets, broadleaved evergreen forest 1,480-2,040 Bar-tailed Treecreeper Certhiahimalayana Coniferous and broadleaved evergreen forest 2,135-3,000 Rufous-vented Tit Parus rubidiventris Broadleaved evergreen and coniferous forest 2,745-3,600 Coal Tit Parus ater Coniferous and mixed coniferous and broadleaved forest 2,745-3,445 Grey-crested Tit Parus dichrous Coniferous and broadleaved evergreen forest 2,745-3,200 Grey-sided Bush Warbler Cettia brunnifrons Scrub and grass bordering broadleaved evergreen and mixed broadleaved/coniferous forest >2,100 Moustached Laughingthrush Garrulaxcineraceus Edge of broadleaved evergreen and mixed broadleaved/coniferous forest, scrub and grass, bamboo 2,220-2,500 Green Shrike-babbler Pteruthiusxanthochlorus Broadleaved evergreen forest 1,700-2,800 Great Parrotbill Conostoma oemodium Open broadleaved evergreen forest, bamboo, scrub 2,775-3,660 Brown Parrotbill Paradoxornis unicolor Bamboo, open broadleaved evergreen forest 2,135-3,660 Fulvous Parrotbill Paradoxornis fulvifrons Bamboo, edge of broadleaved evergreen forest 2,895-3,660 Vinaceous Rosefinch Carpodacus vinaceus Edge of broadleaved evergreen forest, secondary growth, scrub, bamboo 1,830-2,745 Crimson-browed Finch Propyrrhula subhimachala Broadleaved evergreen and coniferous forest, thickets 1,830-3,050 Collared Grosbeak Mycerobas carnipes Broadleaved evergreen and mixed broadleaved and coniferous forest, secondary growth 2,500-3,655 range of the Vietnam population is similar to these others (i.e. 1,600-3,143 m) then there are relatively few remaining suitable locations where this species might occur. Within the Hoang Lien Mountains habitat conversion to agriculture using fire and habitat degradation via clearance of the understorey for cardamom cultivation have been widespread, even within protected areas. Furthermore, severe habitat alteration is continuing apace. It is possible that the species might also occur on high peaks in EastTonkin, outside of the Hoang Lien range, such as those within Tay Con Linh Nature Reserve (which reaches 2,616 m elevation). The avifauna of north-east Vietnam is under-collected and poorly known, as evidenced by the relatively recent discovery of a new taxon in the genus Jabouilleia (Vogel etal. 2003) and observations that extend the range of bird species whose Vietnamese distribution was previously thought to be limited to West Tonkin (Vogel et al. 2003, Mahood etal. in prep.). Hoang Lien Mountains: their brief ornithological history and future Ornithological study of the Hoang Lien Mountains began in late 1 923 when J. H. Stevens spent a few months in northern Tonkin. He collected 333 specimens of 219 species, many from 'Ngai Tio', 30 km north of Sapa (Kinnear 1929). J. Delacour spent three months collecting on Fan Si Pan and around nearby Sapa in late 1929 and early 1930, with the aid of the ethnic Hmong amassing c.3,000 bird specimens (Delacour 1930). He was followed by Bjorkegren, who was based in Sapa from December 1 938 to February 1 939, where he collected 582 specimens that he purchased from the same hunters as Delacour (Eames & Ericson 1996). Delacour described numerous subspecies and one new species endemic to the Hoang Lien Son — White-throated Wren Babbler Rimator pasquieri (Delacour & Jabouille 1930, 1931, Collar 2006). Small-bodied birds are, however, under¬ represented in these collections, owing to the collectors' dependence on match-lock muskets, which generally either miss small birds completely or damage them too badly to preserve. The Hoang Lien Mountains were out of bounds to the wider ornithological community for much of the rest of the twentieth century. Once the area was reopened to visitors, surveys conducted by Frontier-Vietnam and visits by birders increased the number of montane species recorded on Mount Fan Si Pan. For example, Ashy Woodpigeon Columba pulchricollis was not recorded in the Hoang Lien Son until 1998 (Tordoff 2002) and Rufous-belled Niltava Niltava sundara was not recorded until the early 2000s (J. C. Eames unpubl. data). Short surveys of poorly known sites were conducted in the Hoang Lien Mountains, such as Van Ban Nature Reserve and Mu Cang Chai Species and Habitat Conservation Area (SHCA; Tordoff etal. 2001, 2002), during the preparation of the Vietnam Important Bird Area Inventory. Although these surveys did not make use of pre-recorded vocalisations, they demonstrated that the avifauna of these sites was comparable with (if not more intact than) that on Mount Fan Si Pan. Mount Fan Si Pan in Hoang Lien Son National Park is the only readily accessible forested site in the Hoang Lien Mountains. Forest there is, however, heavily degraded and habitat quality continues to deteriorate. Other than at the highest elevations, all readily accessible forest is under cardamom cultivation, with the canopy thinned by 30-70% via the felling of mature trees and with all native understorey vegetation removed. Forest at the highest elevations has lost most trees and much Rhododendron scrub to fire (often started accidentally by tourists visiting the peakof Fan Si Pan). This may account for the paucity of recent records of Darjeeling Woodpecker Dendrocopos darjellensis, a species that was regarded as 'common' by Delacour, but for which there are few recent records (Delacour 1 930). Some of these higher-altitude areas are little more than scrubby grassland, although there are still large areas dominated by dense Arundlnarla bamboo, such as the area where the Bar-winged Wren Babbler was found. Van Ban Nature Reserve, south of Hoang Lien Son National Park, still retains a relatively large area of at least superficially intact forest (although cardamom cultivation is reportedly widespread), has a peak reaching 2,91 3 m (just 230 m lower than Fan Si Pan) and still retains forest at middle and perhaps even lower elevations (Tordoff et al. 2002). Mu Cang Chai SHCA, south of Van Ban Nature Reserve has fairly intact forest from 2,000 m to its highest peak at 2,51 2 m, with many mature Foklenia hodginsli and large areas where cardamom has not yet been cultivated. Recent surveys there indicate that past survey effort has been incomplete; new species for the site, such as Lesser Rufous-headed Parrotbill Paradoxornls atrosuperciliaris, are easily found (SPM pers. obs.). Both these protected areas, difficult of access, still support large-bodied species that are apparently absent elsewhere: both have Temminck's Tragopan Tragopan temminckii, while Mu Cang Chai is one of only two locations in Vietnam (the other being Pu Mat National Park) retaining a population of Rufous-necked Hornbill Aceros nipalensis (Le Trong Dat 2009, Le Trong Dat & Luong Van Hao 2008). 138 SHORT NOTES Forktail 28 (2012) Table 1 lists 1 7 species that have similar global ranges to Bar¬ winged Wren Babbler, and which therefore could conceivably be resident in the Hoang Lien Mountains. All these species are resident in the eastern Himalayas of north-east India, northern Myanmar, and Yunnan and Sichuan provinces of China. Species only rarely recorded in northern Myanmar (e.g. Rufous-breasted Accentor Prunella strophlata ) are excluded, as are those that do not occur in Sichuan (e.g. Grey-sided Laughingthrush Garrulax caerulatus and Cachar Wedge-billed Babbler Sphenocichla roberti), although of course such species might also conceivably occur in Vietnam. Similarly, species that share a similar distribution to another rare Fan Si Pan resident — Red-winged Laughingthrush Garrulax formosus — but currently only occur in Sichuan and Yunnan are excluded, because they do not occur in north-east India and northern Myanmar. Also excluded are species that would probably only occur in Vietnam in the non-breeding season, such as Fire- capped Tit Cephalopyrus flammiceps. We believe that with sufficient survey effort, some of our 17 listed species will be found. Most of them are regarded as uncommon within their core range (Robson 2008) and, therefore, could have escaped notice in the poorly surveyed Hoang Lien Mountains. Those dependent on high-altitude coniferous forest, such as Spotted Nutcracker Nucifraga caryocatactes, Bar-tailed Treecreeper Certhla himalayana and the three Parus tits, may be least likely to occur. Two other species, Green Shrike-babbler Pteruthlus xanthochlorus and Indian Blue Robin Luscinia brunnea, also seem unlikely to occur since they are considered common within their ranges, and might therefore be expected to have been recorded already. By contrast, low-density species can easily be missed and we urge greater attention be directed at this neglected but important area of Indochina. Acknowledgements Jack Tordoff and Craig Robson provided invaluable comments a draft of this manuscript. We thank Craig Robson, John D. Pilgrim and James Eaton for comments on this record, and Jonathan C. Eames for providing a number of useful papers and discussion on the habits of the early collectors in Vietnam. References Collar, N. J. (2006) A partial revision of the Asian babblers (Timalidae). Forktail 22:85-112. Collar, N. J. & Robson, C. (2007) Family Timaliidae (babblers). Pp. 70-291 in J. del Hoyo, A. Elliott & D. A. Christie, eds. Handbook of the birds of the world, 12. Barcelona: Lynx Edicions. Delacour, J. (1930) On the birds collected during the fifth expedition to French Indochina. Ibis (12)6: 564-599. Delacour, J. & Jabouille, P. (1930) Description de trente oiseaux de I'lndochina Fran^aise. L'Oiseau 1 1: 393-408. Delacour, J. & Jabouille, P. (1931) Les oiseaux de I'lndochine frangaise, Tome III. Paris: Exposition Coloniale Internationale. Eames, J. C. & Ericson, P. G. P. (1996) The Bjorkegren expeditions to French Indochina: a collection of birds from Vietnam and Cambodia. Nat. Hist. Bull. Slam Soc. 44: 75-1 1 1 . Eames, J. C. & Mahood S. P. (201 1 ) Little known Asian bird: White-throated Wren-babbler Rimator pasqueri: Vietnam's rarest endemic passerine? BirdingASIA 15: 58-61. Kinnear, N. B. (1929) On the birds collected by Mr. H. Stevens in northern Tonkin in 1 923-1 924. Ibis 1 2(5): 1 07-1 50. Rasmussen, P. C. & Anderton, J. C. (2005) Birds of South Asia: the Ripley guide. Washington, D.C. and Barcelona: Smithsonian Institution and Lynx Edicions. Robson, C. R. (2005) A field guide to the birds of South-East Asia. London: New Holland. Robson, C. R. (2008) A guide to the birds of South-East Asia. London: New Holland. Tordoff, A. W. (2002) Ashy Woodpigeon Columba pulchricollis: a new species for Vietnam. Forktail 18: 147-148. Tordoff, A.W., LeTrong Dat & Hardcastle, J. (2001) A rapid biodiversity survey of Che Tao Commune, Mu Cang Chai district, Yen Bai province, Vietnam. Hanoi: Bi rd Life International Vietnam Programme. Tordoff, A. W., Le Manh Hung, Nguyen Quang Truong & Swan S. R. (2002) A rapid field survey of Van Ban district, Lao Cai province, Vietnam. Hanoi: Birdlife International Vietnam Programme. Vogel, C. J., Sweet, P. R., Le Manh Hung & Hurley M. M. (2003) Ornithological records from Ha Giang province, north-east Vietnam, during March- June 2000. Forktail 1 9: 21-30. Simon P. MAHOOD, Wildlife Conservation Society Cambodia Programme, House 2 1, Street 21, Sangkat Tonle Bassac, Phnom Penh, Cambodia. Email: s.mahood@wcscambodia.org David P. EDWARDS, Centre for Tropical Environmental and Sustainability Science (TESS) and School of Marine and Tropical Biology, James Cook University, Cairns, Smithfield, Queensland 4878, Australia. Email: dave.edwards@jcu.edu.au Felicity A. EDWARDS, Institute of Integrative and Comparative Biology, University of Leeds, Leeds, LS2 9JT, United Kingdom Long-tailed Duck Clangula hyemalis and Red-breasted Goose Branta ruficollis : two new birds for Sichuan, with a review of their distribution in China ZHU LEI, ZHANG JUN, QIU JING, WEI QIAN, DONG LEI & SUN YUE-HUA Long-tailed Duck One Long-tailed Duck in non-breeding plumage was seen by J. Zhang at Yazihe Reserve, Guanghan, on 31 January 2006 around 14h00. This individual was found in mixed group with Common Coots Fulica atra and Ferruginous Pochards Aythya nyroca on the river surface near the Chengdu to Mianyang highway bridge. Long-tailed Duck is a distinctive small diving duck. Although the individual found at Yazihe lacked the typical long tail for an adult male, the white forehead and black patch around the neck as well as mostly white breast revealed this to be a young male in its winter plumage (MacKinnon & Phillipps 2000). This species is widespread in the Holarctic, breeding mainly above the Arctic Circle (Zhao 1995). In eastern Asia its wintering range extends from the northern Bering Sea on pack-ice south along coasts, commonly to northern Japan, rarely to the Korea Peninsula and very rarely in eastern China (Brazil 2009). La Touche (1 934) reported his specimen collection of Long-tailed Ducks from Chinwangtao, east Chihli (now Qinhuangdao, Hebei) and Foochow (now Fuzhou, Fujian). He also stated that the scarcity of records of this species was probably due to lack of observation (La Touche 1 934). Shaw (1936) reported his collection from Xin'an, Hopei (now An'xin, Hebei). Guan etal. (1963) reported a specimen found at Dongting Lake, Hunan, in December 1959, which turned out to be the first record for the province. Cheng (1987) evaluated this species's status as very rare in China, being a migrant known to occur in Heilongjiang and Lushun, Liaoning. Han etal. (1994) reported an observation of wintering Long-tailed Ducks at Dalian Bay during surveys from 13 to 15 January in 1990. On 28 February 2010, two Forktail 28 (2012) SHORT NOTES 139 Figure 1. Long-tailed Duck Clangula hyemalis at Yazihe Reserve, Guanghan. (Photo by Zhang Jun) females in non-breeding plumage were seen around Cheniushan Island, near the coast of Lianyungang, Jiangsu, which adds an additional record to the eastern China coastal area (Lu era/. 201 0). Records of Long-tailed Duck from inland China are much rarer. Li (1996) published a checklist of waterbirds of the Qinghai-Tibet Plateau wetlands which includes this species but without details of when and where. Holt (2008) noted an observation of a female near Golmud, Qinghai, on 1 9-29 November 1991, by J. Hornskov. Zhao (2008) and Li (2009) reported the Long-tailed Duck in their dissertations, which focused on waterbird studies in Inner Mongolia and Gansu respectively. Table 1 details records of this duck from all over China. Red-breasted Goose J. Qiu and his companion observed one adult Red-breasted Goose on 3 January 201 1 at about 12h00 on the Yazihe river, just near the famous archaeological site of Sanxingdui, Guanghan. At times it associated with a small flock of 30 wintering Ruddy Shelducks Tadorna ferruginea, but usually foraged on the riverbank alone. J. Zhang, Q. Wei and L. Zhu observed it on 16 February 201 1, and considered it in good condition. The goose and Ruddy Shelducks occasionally chased each other. The last sighting was on 22 February (ZJ). It could not be found on 2 and 5 March (Zhao Y. D. and Song Y. pers. comm.). The Red-breasted Goose has a highly distinctive plumage, and is slightly smaller than Ruddy Shelduck (Zhao 1 995, pers. obs.).The adult only shows two distinct white bars on the closed wing, while a juvenile bird usually has 4-5 instead (Svensson etal. 2009). Accordingly, we considered the individual adult. The breeding area of this goose is up into the Arctic tundra of Russia, between 67°N and 76°N, and between 67°E and 1 1 6°E, with the majority of the population nesting on the Taimyr, Yamal and Gydan peninsulas to the east of the Ural Mountains (Cranswick et al. 201 0). The main winter range lies along the Black Sea coast of western Ukraine, Romania and Bulgaria, with just five sites holding 90% of the wintering population of the species (Cranswick et al. 2010). There was no record of this goose in China prior to 1 960, when Cheng reported a juvenile bird collected from Dongting Lake on 1 5 January in Hunan (Cheng 1 960). After that records remained scarce. In 1 992, Liu Y. Z. photographed an individual in Poyang Lake, perhaps the first record from Jiangxi (Shi 2000). A survey from January 1 991 to January 1 992 at the Dongting Lake wetlands did notfind Red-breasted Goose (Liu et a/.1995). Similarly, a thorough survey from May 1996 to July 2003, which focused on rare, endangered and nationally protected waterbirds in Hubei, failed to provide any new information on this species. In Wetland International's Asian Waterbird Census, from 1987 to 2007, only one individual was recorded in 2004 at Poyang Lake (Li etal. 2009). Zhong (2007) reported one record in the Dongting Lake wetlands, the most recent from this site. There were two records of six individuals in total found in 2005 and 2006 respectively at Poyang Lake (Xu et al. 2009). Since 2007, records of Red-breasted Goose in China have been increasing, very likely because of raised observer awareness (see Table 2 for details). Discussion Newton (2008) defined vagrant or accidental as 'a bird seen outside its regular range and migration route'. Both of the Long-tailed Duck and Red-breasted Goose records in Sichuan fit this definition, but the accumulating evidence suggests that both species may now be too frequently encountered to be considered vagrants in China. Long-tailed Duck, with two independent observations within only c.16 km in January 2006 and December 2007, as well as one sighting of two individuals in Mianyang in February 2008, plus the latest record from Dayi, Chengdu, in February 2010 (see Table 1), may now prove to be a very rare but regular winter visitor to Sichuan. Records in Table 2 indicate that the Red-breasted Goose has occurred almost annually in China since 2004, mainly in the lower Yangtze River wetlands. At the IOZ, Beijing, we discovered one adult specimen of this goose (no. 38022) not previously mentioned by any authors. According to the label, it was collected at Yueyang, Hunan, in January 1960. Guan etal. (1963) reported survey results of wintering waterfowl in southern China from December 1959 to January 1960 and October 1961 to February 1962, and they only found the goose at Dongting Lake, Yueyang, Hunan, the site which provided the juvenile specimen (IOZ no. 38021 ) mentioned by Cheng Figure 2. Red-breasted Goose Branta ruficollis flanked by Ruddy Shelducks Tadorna ferruginea in Yazihe Reserve, Guanghan. (Photo by Dong Lei) 140 SHORT NOTES Forktail 28 (2012) Table 1. Records of Long-tailed Ducks Clangula hyemalis in China Site Location Elevation (m) Date Number/Gender Source Qinhuangdao, HB 7 7 5-6 April 1916 1 juv M, 1 F La Touche 1934* Fuzhou, FJ 7 7 November 1918 IF La Touche 1934* Baiyangdian Lake, An'xin, HB 38°50'N 115°50'E 0 10 December 1934 IM Shaw 1936*; No. 495** Dongting Lake, HuN 29°18'N 112°59'E 15 December 1959 7 Guanefs/. 1963* Dawangjia Island, Changhai, LN 39°25'N-39°36'N 122°55'E-123°06'E 0 13— 15 January 1990 2,480 Han etal. 1994 Golmud, QH 36°24'N94°54'E 2,823 19-29 November 1991 IF Holt 2008 Beidagang Reservoir, TJ 38°45'N 117°23'E 0 November 1998 7 Zhang eta/. 2004* Beidagang Reservoir, TJ 38°45'N 117°23'E 0 April 1999 7 ditto Summer Palace Park, BJ 39°59'N 116°15'E 49 28 March 2004 IM Liu & Zhang 2005* Yellow River Reserve, HN 34°5TN 1 12°38'E 113 1 January 2005 2 F Niu 2007* Mangrove Reserve, SZ, GD 22°30'N 1 14°01'E 0 23 January 2006 1 CBR 2006, 2007* Yazihe Reserve, Guanghan, SC 30°58'N 104°18'E 460 31 January 2006 1 juv Zhang J., pers. obs. * Sailimu Lake, XJ 44°35'N80°59'E 2,080 29 October 2006 1 Holt 2008* Huayuankou, Zhengzhou, HN 34°53'N 1 1 3°40'E 92 23 December 2006 IF CBR 2006, 2007 Ebinur Lake, XJ 44°51'N 82°44'E 195 29 October 2007 1 Ma & Mei 2007 Miquan, XJ 44°02'N87°37'E 530 5-7 November 2007 2 ditto Dalai Nur, IM 49°00'N 1 1 7°23'E 550 24 April 2007 3 Zhao Gerelt 2008* Hangzhou Bay, ZJ 30°18N 120°45'E 0 August 2007 7 Jiang et al. 2011 Danghe Reservoir, GS 39°56'N94°19'E 1,429 28 October 2007 1 M, 1 F Li 2009* Jing Lake, Deyang, SC 31°07'N 104°23'E 492 31 December 2007 1 F, 1 juv CBR 2007, 2008 Dazhanhe Reserve, HU 48°23'N 127°13'E 420 7 7 Zhang et al. 2009 Qionghai Lake, Xichang, SC 27°5TN 102°16'E 1,511 8 January 2008 IF Ye C.Y., pers.comm. Hangzhou Bay, ZJ 30°18'N 120°45'E 0 January February 2008 7 Jiang eta/. 2011 Mianyang, SC 31°28'N 104°45'E 447 29 February 2008 2 Li B., pers. comm. Lianyungang, JS 35°04'N 1 19°21'E 0 28 February 2009 2 F Lu etal. 2010* Dayi, Chengdu, SC 30°36'N 103°10'E 1,150 10 February 2010 IF HeY., pers.comm. (Notes: BJ= Beijing, ,FJ= Fujian, GD= Guangdong, GS= Gansu, HB= Hebei, HU= Heilongjiang, HN= Henan, HuN= Hunan, IM= Inner Mongolia, JS= Jiangsu, LN= Liaoning, QH= Qinghai, SC= Sichuan, SZ= Shenzhen City, XJ= Xinjiang, ZJ= Zhejiang; M= male, F= female, juv= juvenile; CBR 2006, 2007= China Bird Report 2006, 2007; *= the first record of corresponding provinces, **= specimen perserved at I0Z; '?'= lack of data.) Table 2. Records of Red-breasted Goose Branta ruficollis in China Site Location Elevation (m) Date Number Source Dongting Lake, HuN 29°18'N 112°59'E 15 15 January 1960 1 juv Cheng I960*; No. 38021 ** Dongting Lake, HuN 29°18'N 112°59'E 15 January1960 1 No. 38022** Poyang Lake, JX 29°10'N 116°00'E 12 1992 1 Shi 2000* Poyang Lake, JX 29°10'N 116°00'E 12 2004 1 Li etal. 2009 Dongting Lake Wetland, HuN 29°18'N 1 1 2°59'E 15 7 7 Zhong etal. 2007 Dachahu, Poyang Lake, JX 29°09'N 116°04'E 9 2005 i Tuefr;/. 2009 Hanchihu, Poyang Lake, JX 29°04'N 1 16°25'E 11 29 December 2006 i ditto Fanghu, Poyang Lake, JX 29°49'N 116°3VE 12 29 December 2006 4 ditto Baiqiang Reservoir, Mengzhou, HN 35°00'N 112°46'E 122 28 January 2007 1 Niu etal. 2008* Baiqiang Reservoir, Mengzhou, HN 35°00'N 112°46'E 122 1 January 2008 1 A ditto Wolong Lake, Shenyang, LN 42°44'N 123°17'E 88 16—17 October 2008 7 Liefn/. 2010* Dongting Lake Wetland, HuN 29°31'N 112°47'E 26 8 October 2008 1 A Wang X. pers. comm. Shengjin Lake Reserve, AH 30°19'N117°00'E 9 15 November 2008 1 CaoL pers.comm.* Dongting Lake Wetland, HuN 29°29'N 112°48'E 23 24 February 2010 1 A Wang X. pers. comm. Shengjin Lake Reserve, AH 31°21'N 117°01'E 10 2 December 2010 1 A WangX. pers.comm. Yazihe Reserve, Guanghan, SC 31°00'N 104°12'E 483 3 January 2011 1 A QiuJ.&P. Hu* (Notes: AH= Anhui, HN=Henan, HuN= Hunan, JX= Jiangxi, LN= Liaoning, SC= Sichuan; A= adult, juv= juvenile; *= the first record of corresponding provinces, **= specimen perserved at I0Z; 7 = lack of data.) Forktail 28 (2012) SHORT NOTES 141 (1960). We believe that the newly discovered specimen was also collected by Guan and his colleagues during the same field trip in January 1960. Combining with these historical records and recent sightings, we speculate that this species might not be a vagrant but rare winter vistor to the lower Yangtze River wetlands. The scarcity of records of this species is probably largely due to lack of observation in this vast area in the past. Undoubtedly, more surveys are needed in future to clarify the status of this endangered species in China. The avifauna of Sichuan is well known for its rich component of endemics, with 37 of the 71 species, 52%, endemic to China (Zheng 2005, Xu 2008). By contrast waterbirds in Sichuan have been neglected. Few papers focus on this group of birds in Sichuan (e.g. Deng etal. 1983, Cui etal. 1992, Liu etal. 2004). It is remarkable that since 2005 no fewer than three birds new to Sichuan have turned up at this tiny Yazihe Provincial Reserve, Guanghan, which is just 16 km long and covers only 481 ha, the third one being Yellow-billed Loon Gavia adamsii on 14 December 2005 (Que & Ran 2006), a species which like Long-tailed Duck is usually found on the sea. It seems likely that the main reason for this interesting development is that birdwatching is becoming increasingly popular in China. Since 2000, nearly 20 new bird records to China has been added to this country's checklist, more than half of them contributed solely or mainly by birdwatchers (e.g. Holt in CBR 2004 2005, Luo etal. 2007, Li etal. 2008, Wu etal. 2010, Chang etal. 2010). Naturally for difficult-to-identify and more threatened species unusual records require solid documentation involving photographs or detailed descriptions. Acknowledgements We thank Sergei Kharitonov from the Bird Ringing Centre of Russia (Moscow), Jesper Hornskov and Adrian Walker, who revised and helped improvement of earlier drafts; Nigel Collar from BirdLife International for help with the manuscript; Fie Peng at Institute of Zoology, China Academy of Science, Beijing, for helping us access to the specimen collection; Dr Cao Lei and Wang Xin from University of Science and Technology of China, Dr Huang Xiao-feng from Jiangxi Academy of Forestry, Zhao Yi-ding from Southwest Jiaotong University and Song Ye for kindly sharing their observations of the Red-breasted Goose in Anhui, Hunan, Jiangxi and Yazihe Reserve, Sichuan; and Sha Jian-binfrom Northeast Forestry University, Huang Qinfrom Zhejiang Museum of Natural History, He Yi and Li Bin from Chengdu Bird Watching Society, Xiong Yan and Li Zhen-zhong from Henan Bird Watching Society, Que Pin-jia from Beijing Normal University, fortheir patient replies to our inquiry of locations of the Long-tailed Duck and Red-breasted Goose recorded in Heilongjiang, Henan, Tianjin and Sichuan province. 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Han Xiao-dong, Wu Jing-cai & Zhao Zheng-jie (1994) A preliminary observation of wintering Long-tailed Ducks around Dawangjia Island, Changhai, Liaoning. Chinese J. Zool. 29 (4): 22-24. (In Chinese.) Holt, P. (2008) Eight new ornithological records from Xinjiang China. Arid Land Geography 3 1 : 243-248. Jiang Ke-yi, Wu Ming & Shao Xue-xin (2011) Community composition, seasonal dynamics and interspecific correlation of waterbirds in the Qiantangjiang River estuary and Hangzhou Bay. Zoological Research 32: 631-640. La Touche, J. D. D. (1925-1934) A handbook of the birds of eastern China, 1 and 2. London: Taylor and Francis. Li Fei (2009) [Migratory waterbirds survey in Dunhuang, Gansu province.] Master Dissertation, Beijing Forestry University, Beijing. (In Chinese.) Li Gui-yuan (1995) The colour handbook to the bird of Sichuan. Beijing: Chinese Forestry Publishing House. (In Chinese.) Li Hai-tao, Chen Liang, Ho Chi-kong & Liu Yang (2008) A new bird record in China: Red-breasted Flycatcher (Ficedula parva). Zoological Research 29:325-327. Li Lai-xing (1996) A checklist of birds of Qinghai-Tibet Plateau wetlands and conservation. J. Qinghai Environment 1: 19-26. (In Chinese.) Li Xin, Wu Jian-ping, Zhu Cheng-wei, Zhao Yang & Li Yan-jie (2010) Migration of waterfowls in Wolong Nature Reserve. Chinese J. Wildlife 3: 135-138. (In Chinese.) Li Yan (2008) [Study on wetland bird diversity and migration law in Dalai Lake Natural Reserve.] Masters Dissertation, Northeast Normal University, Changchun. (In Chinese.) Li, Z. W. D„ Bloem, A., Delany, S., Martakis, G. & Quintero, J. O. (2009) Status of waterbirds in Asia: results of the Asian Water bird Census: 1987-2007. Kuala Lumpur: Wetlands International. Liu Hao, Shi Hong-yan & Zhang Li-quan (2004) Analysis of water birds diversity in Mianyang, Sichuan province, China. Chinese J. Zool. 4: 85- 89. (In Chinese.) 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E-mail: sunyh@ioz.ac.cn ZHANG Jun, Guanghan Bureau of Forestry, Guanghan, China, Postcode 61 8300 QIU Jing, Chengdu Bird Watching Society, Chengdu, China, Postcode 610041 WEI Qian, China Bird Watch, Beijing, China, Postcode 100026 DONG Lei, College of Art and Communication, Southwest Jiaotong University, Chengdu, China, Postcode 610031 Western Hoolock Hoolock hoolock preying on chicks of Greater Racket-tailed Drongo Dicrurus paradiseus in Lawachara National Park, Bangladesh DELIP K. DAS On 3 May 2011 at 1 2h 1 8—1 2h33, Indranil Kishor and I observed a group of Western Hoolocks Hoolock hoolock being mobbed by two Greater Racket-tailed Drongos Dicrurus paradiseus at Lawachara National Park, District Moulavibazar, Bangladesh (24°20'N 91°47'E). The gibbon group consisted of one adult female with infant, one adult male and one unsexed black-coloured juvenile. I soon noticed that the female Hoolock was holding a half-grown drongo chick in her hand, and that another chick was present in a nest next to her. She was busy eating the flesh of the chick, unconcerned by the strong mobbing of the drongos. Afterfinishing the first chick she took the second chick from the nest. The chick Plate 1. Female Western Hoolock about to take chick from nest of Greater Racket-tailed Drongo Dicrurus paradiseus. was calling and waving its legs continuously to escape. The female Hoolock inspected the chick for some time, holding it upside down in front of her face, and shaking it a couple of times whilst looking around her, causing the victim's parents to become frantic in their mobbing. She shifted to another branch and again held the chick upside down while it opened its gape to its widest extent. After a while, she bit off the chick's head, ate it, and then started eating the body. The whole event, from picking the live chick from nest to biting off its head, took c.3 minutes; Plates 1-4 illustrate the sequence. While this was happening, I noticed that the adult male gibbon, c.6- 8 m distant in another tree, was consuming a third chick, which it Plate 2. Greater Racket-tailed Drongo mobbing the female Hoolock. Forktail 28 (2012) SHORT NOTES 143 Plate 3. Female Hoolock inspecting the chick before eating it. must have taken from the nest before moving away and allowing the female access to the other two. The usual clutch-size of Greater Racket-tailed Drongo is three, sometimes four (Rocamora & Yeatman-Berthelot 2009), so it would appear that the gibbons predated the entire brood of the pair in this instance, although the juvenile evidently got nothing: it sat silently observing the adult male at a distance of 2 m. Tight scheduling precluded observations from continuing long enough to document post-predation behaviour. A reference search on hoolocks (Tilson 1979, Gittins &Tilson 1984, Mukherjee 1986, Choudhury 1991, Ahsan 1992, Alfred 1992, Feeroz & Islam 1992, Islam & Feeroz 1992, Feeroz et al. 1994, Bujarbarua & Das 2001, Kakati 2004) revealed no reports of these gibbons predating bird chicks. However, Western Black Crested Gibbons Nomascus concolor have been reported predating nestlings and eggs of birds in China (Fan & Jiang 2008), and there is a case of predation by White-handed Gibbon Hylobates lar of a hen Gallus (Carpenter 1940, Newkirk 1973). Islam & Feeroz (1992) reported that several species of bird (magpies Cissa, drongos Dicrurus and laughingthrushes Garrulax ) sometimes chase Western Hoolocks when they are in food trees, with drongos being more aggressive, continuing confrontations until the gibbons leave; but actual predation has not apparently been observed in the wild before. However, given how often the gibbons are high in trees, thereby hindering precise observations of food items, and the frequency with which mobbing occurs, such predation may be commoner than the lack of records might suggest. The feeding ecology of Western Hoolock is well studied. It is primarily vegetarian, consuming, in some habitats, over 100 species of plants although in some places apparently many fewer (Alfred 1992, Islam & Feeroz 1992, Ahsan 2001). It also consumes some prey items, predominantly invertebrates but also birds' eggs (Mukherjee 1986, Alfred 1992, Ahsan 2001). In their diet, figs dominate and fruits are very important; however, lianas, flowers, shoots, petioles, exudates, nectar and other plant foods, including Plate 4. Female Hoolock eating the chick's headless body. lichens, are also consumed (Islam & Feeroz 1992, Ahsan 2001, Bujarbarua & Das 2001, Kakati 2004). Acknowledgements I thank Indranil Kishor, Munir and Shamim Reza Rubel for company during the field trip and also Shimanto Dipu and CNRS for logistics during field work. I specially thank Raju Kasambe, Project Manager, IBA Programme, Bombay Natural History Society (BNHS), India, and Asad Rahmani, Director, BNHS, India, for commenting on and revising the first draft of the manuscript. References Ahsan, M. F. (1992) Feeding ecology of the primates of Bangladesh. Pp.79- 86 in B. Thierry, J. R. Anderson, J. J. Roeder & N. Herrenschmidt, eds. Current primatology, I: ecology and evolution. Strasbourg: Universite Louis Pasteur. Ahsan, M. F. (2001 ) Socio-ecology of the hoolock gibbon (Hylobates hoolock) in two forests of Bangladesh. Pp. 286-299 in Brookfield Zoo, ed. The apes: challenges for the 2 1st century. Conference proceedings, May 10-13, 2000. Brookfield, Illinois: Brookfield Zoo. Alfred, J. R. B. (1992) The hoolock gibbon: Hylobates hoolock. Prim. Rep. 34: 65-69. Bujarbarua, P. & Das, J. (2001 ) Hoolock gibbon (Hylobates hoolock) feeding on lichens. J. Bombay Nat. Hist. Soc. 98: 432. Carpenter, C. R. (1940) A field study in Siam of the behaviour and social relations of the gibbon, Hylobates lar. Comp. Psychol. Monogr. 1 6: 1 -21 2. Choudhury, A. (1 991 ) Ecology of the hoolock gibbon ( Hylobates hoolock), a lesser ape in the tropical forests of north-eastern India. J. Trop. Ecol. 7: 147-153. Fan, P. F. & Jiang, X. L. (2008) Predation on giant flying squirrels ( Petaurista philippensis) by Black-crested gibbons (Nomascus concolor jingdongensis) at Mt. Wuliang, Yunnan, China. Primates 50: 45-49. Feeroz, M. M. & Islam, M. A. (1992) Ecology and behaviour of Hoolock gibbons of Bangladesh. Dhaka, Bangladesh: Multidisciplinary Action Research Centre. 144 SHORT NOTES Forktail 28 (2012) Feeroz, M. M., Islam, M. A. & Kabir, M. M. (1994) Food and feeding behaviour of Hoolock Gibbon ( Hylobates hoolock), Capped Langur (Presbytis pileata) and Pigtailed Macaque (Macaca nemestrina ) of Lawachara. Bangladesh J. Zoo!. 22: 1 23-1 32. Gittins, S. P. & Tilson, R. L. (1984) Notes on the ecology and behaviour of the Hoolock gibbon. Pp. 258-266 in H. Preuschoft, D. J. Chivers, W. Y. Brockelman & N. Creel, eds. The lesser apes: evolutionary and behavioural biology. Edinburgh: Edinburgh University Press. Islam, M. A. & Feeroz, M. M. (1 992) Ecology of Hoolock gibbon of Bangladesh. Primates 33: 45 1 -464. Kakati, K. (2004) Impact of forest fragmentation on the Hoolock gibbon in Assam, India. Ph.D thesis. Cambridge: University of Cambridge. Mukherjee, R. P. (1986) The ecology of the Hoolock gibbon, Hylobates hoolock, in Tripura, India. Pp.1 15-123 in J. G.EIse&P. C. Lee, eds. Primate ecology and conservation. Cambridge: Cambridge University Press. Newkirk, J. B. (1973) A possible case of predation in the gibbon. Primates 14: 301-304. Rocamora, G. J.& Yeatman-Berthelot, D. (2009) Family Dicruridae (drongos). Pp.1 72-226 in J. del Hoyo, A. Elliott & D. A. Christie, eds. Handbook of the birds of the world, 14. Barcelona: Lynx Edicions. Tilson, R. L. (1979) Behaviour of Hoolock gibbon (Hylobates hoolock) during different seasons in Assam, India. J. Bombay Nat. Hist. Soc. 76: 1 -1 6. Delip K. DAS, ICCR Scholar, Room No. 1 106, International Student House(ISH), University of Mumbai, C Road, Churchgate, Mumbai 400020, Maharastra, India; and c/o Ranjit Kumar Das, l ////.+ Post.: Tezkhali, Thana: Bancharampur, Dist.: B. Baria, Bangladesh. Email: bisharga 1 095@gmail.com Notes on the life-history and taxonomy of Muscicapa dauurica umbrosa, an overlooked Bornean canopy bird FRANK E. RHEINDT & JAMES A. EATON Introduction Erwin's (1982) experiments into tropical canopy insect diversity suggested that the rainforest canopy may harbour the majority of the planet's terrestrial animal diversity, most of it undescribed. While birds are relatively well known (e.g. Mayr 1 946), avian species of the tropical rainforest canopy feature among the least known, and a disproportionate number of new avian taxa have involved high- canopy forms such as Neotropical Herpsilochmus antwrens (e.g. Davis & O'Neill 1 986, Whitney & Alvarez 1 998, Whitney etal. 2000) and Old World Muscicapa flycatchers (e.g. Wells 1982, King etal. 1999). At Danum Valley (Sabah, Malaysia), one of the ornithologically best- explored sites in the Sunda Islands, Edwards etal. (2009) recently photographed a strikingly distinct new Dicaeum flowerpecker. Here we concentrate on a Bornean flycatcher taxon, Muscicapa dauurica umbrosa, endemic to the high canopy of lowland rainforest. Recently described (Wells 1982), little is known about this form. We present some of the first life-history data including a documentation of its nest, and offer comments on its taxonomy. Observations FER visited Lambir Hills National Park in Sarawak (Malaysia; 4°11'52"N 1 14°02'34"E; 50 m a.s.l.), not far from the border with Sabah, from 2-1 2 June 2010. On most days during this period, he visited the canopy tower near the visitor centre of the national park in the early morning (approximately 06h30-07h30). On the first of these visits, he discovered a nest attended by two individuals of Muscicapa dauurica umbrosa (for identification see below). The height of the nest above ground was estimated at 38 m, using the tower for reference. The nest was about 20 m from the tower platform across an 'aerial clearing'. It constituted an open- cup structure of twigs and mosses with an outer diameter of c.25 cm built in the crevice of a loose branch hanging perpendicularly from a vertical main branch of the canopy crown of an emergent rainforest tree (Plate 1 ). Two birds regularly attended this nest. They were never seen at the nest simultaneously, but occasionally sat together on a nearby branch. During the whole period, no pulli or juveniles were perceived. The nest was unattended for up to 4 minutes at a time, after which one of the two adults was usually seen flying to the nest and spending 1-4 minutes in and around it. Both adults were moderately streaked on flanks and throat, and were rather rich rufous-brown on their upperparts, with rump perhaps more intensely rufous (Plates 2, 3). No eye-ring was perceived, although Plate 2 erroneously suggests one. The basal half of the lower mandible was rich orange. Discussion Breeding phenology The only previous information on the life history of M. dauurica umbrosa comes from Wells & Francis (1984) who collected a pair of adults with enlarged testes and brood-patch, respectively, on 27 July 1983 near Sepilok (Sabah, Malaysia), and subsequently collected a pairtending afullygrownjuvenileon 8 July 1984atthe same locality. Their data coincide well with ours to suggest that the annual breeding period of M. d. umbrosa (at least June-July) encompasses the height of the northern summer, when no northern migrants of M. dauurica would be present in Borneo. As Wells (1982) pointed out when discussing the biogeography of the widespread Siberian breeder M. d. dauurica (or M. latirostris cinereoalba, following the nomenclature of the time), the presence of a resident form in a small part (north-eastern Borneo) of the wintering range of the former is unusual for Oriental birds, and nothing is known about the ecological interactions between the two. The timing of breeding of umbrosa in June and July may well be due solely to the regional climate, which is slightly drier from April to October than at other times, if it is not simply evading northern migrants during the rearing of the young. Coloration and identification of this taxon Little is known about M. d. umbrosa in life. The species is rarely observed by field ornithologists, probably owing to its canopy- inhabiting lifestyle and vocal and behavioural inconspicuousness, as in other Muscicapa flycatchers. During a total of eight months of birdwatching activity in Sabah and Sarawak, JAE has observed M. d. umbrosa on only two occasions, both in the Danum Valley conservation area (Sabah; 5°1'12"N 1 17°44'48"E) in June 2007 and May 2009. Encounter rates for other field ornithologists with experience in Borneo are similarly low (R. O. Hutchinson verbally). Although there are field observations from Brunei (Mann 1987), the few specimens available are all from within Sabah, namely the type from Tawau (Wells 1 982), one adult from Lahad Datu and four adults and one juvenile from Sepilok Forest (Wells & Francis 1984). There is thus a need to clarify the field identification of this bird. The pair seen nesting at Lambir appeared moderately streaky on flanks and throat, not unlike M. [d.] williamsoni, and were Forktail 28 (2012) SHORT NOTES 145 .. Plate 1 . The nest (with one individual of M. dauurica umbrosa inside), Lambir National Park (Sarawak), June 2010. (F. E. Rheindt) Plate 2. Front view of an adult M. dauurica umbrosa from across an aerial clearing at c.20 m distance, Lambir National Park (Sarawak), June 2010. (F. E. Rheindt) Plate 3. Back view of an adult M. dauurica umbrosa, Lambir National Park (Sarawak), June 201 0. (F. E. Rheindt) matched in this trait by both individuals seen by JAE at Danum Valley. Only some of the specimens listed above display this level of streakiness. But Wells & Francis (1984) mention this could well be a variable character akin to the geographically proximate Ashy¬ breasted Flycatcher M. randi from the Philippines. Our observations from Lambir and Sabah coincide with specimens with respect to the lack of a perceptible eye-ring, which sets this taxon apart from northern migrating nominotypical individuals. The most salient features of the pair observed nesting at Lambir was their rich dark rufous-brown back coloration. The description of this feature by Wells (1982) and Wells & Francis (1984) is somewhat equivocal. Wells (1982) described umbrosa as dull grey- brown above, darkerthan the nominotypical. Two years later. Wells & Francis (1984) described the follow-up specimens as umber- brown above and identical in that character to the type. Umber is a relatively rich earthen-brown hue, but D. R. Wells {in litt. 2012) confirms that none of the existing specimens of umbrosa appears quite as rufescent as the bird in Plate 3. However, Wells etal. (1 986) showed that another Sundaic taxon, williamsoni, displays striking differences between earlier (unworn) rufescent and later (worn) drab-brown plumage stages during the breeding season. The taxon umbrosa may well exhibit a similar plumage cycle. This is supported by the fact that all five unequivocally adult umbrosa specimens were collected 4-8 weeks after the Lambir observations, in a period from 8-27 July, at a time when their plumage moult may have advanced and the rufescent coloration may have worn off. Wells et al. (1986) noticed similar changes from bright-rufescent to drab- brown back coloration in williamsoni over the course of mid- to late June. Therefore, the plumage of umbrosa may resemble williamsoni more closely than has hitherto been recognised. The alternative explanation — that the Lambir observations refer to the taxon williamsoni — seems improbable The latter has only been found breeding in the Thai-Malay peninsula, no further south than northernmost Malaysia. Although it has been recorded wintering further south on the Malay peninsula, Sumatra and Siberut (Wells et al. 1986), there is only one claimed specimen record from extreme westernmost Sarawak (Wells etal. 1986) as well as unconfirmed sightings from Brunei (Mann 1 989) which may well refer to umbrosa. We have not inspected the Sarawak specimen (in the Natural History Museum, Tring, UK) and do not know whether umbrosa was considered as a possibility by its identifier. But even if it does constitute a migrant record of williamsoni on Borneo, we note that it is geographically removed from all records of umbrosa, and that there has never been a suggestion of breeding activity of williamsoni anywhere on Borneo. The Lambir Hills are almost 1 ,500 km from the nearest breeding ground of williamsoni in northern peninsular Malaysia, and our observations at the nest confirm their identity as breeders. Taxonomic treatment of Malay-Sundaic forms The Asian Brown Flycatcher Muscicapa dauurica (formerly called M. latirostris ) is a common inhabitant of boreal, temperate and subtropical forests in East and South Asia. Siberian nominotypical populations winter widely in South-East Asia and constitute one of the region's commonest wintering songbirds in wooded habitats. This winter abundance in the Thai-Malay peninsula and Borneo has promoted the conflict in taxonomic treatments of individuals collected in this region. Investigating plumages of Thai-Malay peninsular birds, Deignan (1957) noted that most museum specimens are northern winter visitors. However, he described a new species, M. williamsoni, based on a small number of individuals that seem to breed somewhere on the peninsula. Muscicapa williamsoni was diagnosed from northern M. d. dauurica on the basis of several plumage characters, but especially its distinct rufous rather than grey-brown upperparts (particularly rump). Earlier, M. d. siamensis from the mountains of 146 SHORT NOTES Forktail 28 (2012) Vietnam and northern Thailand had been described as a subspecies on grounds of upperparts that are more rufous than those of northern Asian nominotypical birds, but Deignan (1957) insisted williamsoni is not a mere extension of this southward trend to a more rufous coloration: the northernmost members of williamsoni are much more rufescent than the southernmost members of siamensis, although the two approach each other geographically somewhere in south-eastern Burma. Wells (1977) argued against Deignan's (1957) classification of M. williamsoni as a species, pointing out the existence of intermediate birds between williamsoni and nominotypical dauurica, e.g. in the Kuala Lumpur region in winter. Wells et al. (1986) then clarified that williamsoni breeds in Burma at least to 1 7°N and south to theThai-Malay border area, and winters as given above, from the Malay Peninsula south to Sumatra and Siberut Island. It was in this context that Wells (1982) described M. d. umbrosa from the lowland rainforests of Sabah (Malaysia) as an additional subspecies that does not engage in continent-wide long¬ distance migration. He considered umbrosa rather similar to M. rand/ from the Philippines in plumage. However, as noted above, the current adult specimens may be worn birds, and umbrosa may have a similar plumage cycle to williamsoni (Wells et al. 1 986). The distribution gap between williamsoni and umbrosa narrowed during the last ice age (as recently as 1 7,000 years ago) and gene- flow could have been regular. Thus umbrosa should be suspected to be more closely related to williamsoni than to Philippine randi, which has never had a land connection to Borneo during the existence of this species complex. Taxonomic treatment of the Malayan breeding form williamsoni has been contentious: some authorities treat it as a species (e.g. King et al. 1975, Jeyarajasingam & Pearson 1999, Robson 2000), others as a subspecies (e.g. Mayr etal. 1 986, MacKinnon & Phillipps 1993, Clement 2006). The form umbrosa, in contrast, has been treated as a subspecies of M. dauurica by all authorities. Based on the morphological traits indicated by Wells (1982), Wells & Francis (1984) and herein, such as wing formula, lack of or only indistinct eye-ring, and rich brown back coloration when breeding, umbrosa and williamsoni are comparatively different from northern M. d. dauurica. In fact, certain traits, such as wing formula (Wells 1982), link them with other resident South-East Asian taxa (e.g. M. randi and Sumba Brown Flycatcher M. segregata) that have recently been considered to constitute good species (e.g. Clement 2006). The summer timing of breeding in both umbrosa and williamsoni may well happen to act as an isolating mechanism, preventing hybridisation with northern migrants. Given these considerations, we argue that the current practice of affording some South-East Asian taxa species status (e.g. M. randi) while lumping others into M. dauurica (e.g. M. dauurica umbrosa) is mistaken. Instead, we hold that, depending on the species concept employed, there are three potential treatments: (1 ) either all Asian forms mentioned should be lumped into M. dauurica, or (2) williamsoni and umbrosa should be split out as one species (M. williamsoni), or (3) both williamsoni and umbrosa should be given individual species status. We advocate treatment (2) pending further studies into the differences between the two taxa. The study of vocalisations — useful in other songbirds — may not be a fruitful avenue to test the correctness of this treatment, as Muscicapa flycatchers are infrequent and unsophisticated songsters, but molecular data may help establish how much gene-flow occurs or has occurred between the two forms. Acknowledgements We thank Kate Eldridge for help locating literature references, and Rob Hutchinson for discussions about umbrosa. David Edwards and David Wells provided helpful comments. FER is indebted to Dr Campbell Webb for granting him a teaching fellowship with the Harvard University field course that led to the discovery of the nest, and to Scott Edwards for providing the photographic equipment. Birdtour Asia Ltd. sponsored most of JAE's field travel in the region. References Clement, P. (2006) Genus Muscicapa. Pp.1 10-120 in J. del Hoyo, A. Elliott & D. A. Christie, eds. Handbook of the birds of the world, 1 1. Barcelona: Lynx Edicions. Davis, T. J. & O'Neill, J. P. (1986) A new species of antwren (Formicariidae: Herpsilochmus ) from Peru, with comments on the systematics of other members of the genus. Wilson Bull. 98: 337-352. Deignan, H.G. (1 957) A new flycatcher from southeastern Asia, with remarks on Muscicapa latirostris Raffles. Ibis 99: 340-344. Edwards, D. P., Webster, R. E. & Rowlett, R. A. (2009) 'Spectacled Flowerpecker': a species new to science discovered in Borneo? BirdingASIA 1 2: 38-41 . Erwin, T. L. (1982) Tropical forests: their richness in Coleoptera and other species. Coleopterist's Bull. 36: 74-75. Jeyarajasingan, A. & Pearson, A. (1999) A field guide to the birds of West Malaysia and Singapore. Oxford: Oxford University Press. King, B„ Woodcock, M. & Dickinson, E. C. (1975) Birds of South-East Asia. London: Collins. King, B., Rostron, R, Luijendijk, T., Bouwman, R. & Quispel, C. (1999) An undescribed Muscicapa flycatcher on Sulawesi, Indonesia. Forktail 15: 104. MacKinnon, J. & Phillipps, K. (1993) The birds of Borneo, Sumatra, Java, and Bali. Oxford: Oxford University Press. Mann, C. F. (1987) Notable bird observations from Brunei, Borneo. Forktail 3:51-56. Mann, C. F. (1989) More notable bird observations from Brunei, Borneo. Forktail 5: 1 7-22. Mayr, E. (1946) The number of species of birds. Auk 63: 64-69. Mayr, E., Traylor, M. A. & Watson, G. E. (1986) Check-list of the birds of the world, 1 1 . Cambridge, Mass.: Museum of Comparative Zoology. Robson, C. (2000) A guide to the birds of South-East Asia. London: New Holland. Wells, D. R. (1977) Muscicapa williamsoni Deignan: a reappraisal. Bull. Brit. Orn. Club 97: 83-97. Wells, D. R. (1982) Notes on some representatives of the Brown Flycatcher Muscicapa latirostris Raffles in Southeast Asia. Bull. Brit. Orn. Club 102: 148-153. Wells, D. R. & Francis, C. M. (1984) Further evidence of a resident Brown Flycatcher Muscicapa latirostris in Borneo. Bull. Brit. Orn. Club 104: 1 25- 127. Wells, D. R., Round, P. D. & Scharringa, J. (1986) New information on the 'Brown-streaked' Flycatcher Muscicapa latirostris williamsoni. Forktail 1 : 15-20. Whitney, B. M. & Alvarez, J. (1998) A new Herpsilochmus antwren (Aves: Thamnophilidae) from northern Amazonian Peru and adjacent Ecuador: the role of edaphic heterogeneity of terra firme forest. Auk 1 15: 559-576. Whitney, B. M„ Pacheco, J. F„ Buzzetti, D. R. C. & Parrini, R. (2000) Systematic revision and biogeography of the Herpsilochmus pileatus complex, with description of a new species from northeastern Brazil. Auk 1 1 7: 869- 891. Frank E. RHEINDT, Harvard University, Department of Organismic and Evolutionary Biology, 26 Oxford St, Cambridge, MA 02138, U.S.A. Email: frankrheindt@yahoo.com.au James A. EATON, 17 Keats Avenue, Littleover, Derby, DE23 4EE, U.K. Email: jameseaton@birdtourasia.com Forktail 28 (2012) SHORT NOTES 147 Apparent inter- and intraspecific brood-parasitism in a nest of Tufted Duck Aythya fuligula YANG LIU & SERGEY PYZHJANOV The cuckoos and their passerine hosts represent a classic example of an interspecific brood-parasitism system (Davies 2000). Meanwhile, intraspecific brood-parasitism, where conspecific females lay their eggs in the nest of other females, has been documented in a range of taxa that includes waterbirds, pheasants and passerines (Yom-Tov 2001). Duck species probably more commonly exhibit facultative (inter- and intraspecific) brood- parasitism than other avian groups (Geffen & Yom-Tov 2001). However, a brood parasitised with both inter- and intraspecific eggs is rarely reported in breeding duck colonies. During our fieldwork at Lake Baikal, Russia, on 5 July 2008, we recorded what we believe to have been such a nest on an anonymous island near Kurma (53°1 0'N 1 06°58'E), by the central- western shore of Lake Baikal. A female Tufted Duck Aythya fuligula that was incubating the eggs was flushed when we approached the nest. The nest was located in low dense scrub intermixed with dry and fresh grass. Inside the nest were 1 6 eggs, including 1 1 pale olive or olive-grey eggs, four creamy-white eggs and one apparently larger creamy-white egg (Plate 1 ). The eleven pale olive or olive-grey eggs, forming the majority of this brood, fit the general description of eggs of Tufted Duck (Cramp & Simmons 1 977). We also found several broods of Tufted Ducks nearby Lake Baikal and thistype of egg was common within the colonies we visited (Plate 2). The four creamy-white eggs were of the same size as the eggs of Tufted Ducks. We assumed that these eggs also belonged to T ufted Ducks but represented a colour variation for the following two reasons. First, we have seen thistype of egg in clutches of Tufted Ducks. Second, it is unlikely that this type of egg belongs to other sympatric breeding ducks at Lake Baikal based on egg size and the characteristics of the nesting site. The only egg similar in shape is that of Common Pochard Aythya ferina; however, this species rarely breeds on the islands in the Kurma region, but nests in the marshes at the Selenga Delta at the south-eastern shore of Lake Baikal. Because it is impossible that the same female can lay eggs of two colours in the same clutch (Kilner 2006), we assumed that another female laid these four creamy-white eggs. The outlier among the 16 eggs was one creamy-white egg, c.25% larger in size. Although we did not take the measurements of this egg, given its exceptionally larger size we can exclude the Plate 1 . The clutch of a Tufted Duck Aythya fuligula with (A) a parasitic egg from White-winged Scoter Melanitta deglandi, (B) eggs laid by another female Tufted Duck found near Kurma, Russia, 5 July 2008. possibility of any Anas and Aythya ducks in the region (unless a double yolk was involved — an extremely remote possibility: R. M. Kilner in litt. 2012). The only two candidate species that fit the size of this egg are White-winged Scoter Melanitta deglandi and Red-breasted Merganser Mergus serrator (Cramp & Simmons 1977, Baicich & Harrison 1997). However, the latter very rarely builds nests on the ground in vegetation (but a ground nest has been found in the south of the Selenga delta: I. Fefelov in litt.), as it usually nests in cavities under cliffs (Cramp & Simmons 1977); indeed, we located several such nests on rocky islands near Kurma. In contrast, this egg seems more likely to have been laid by a female White-winged Scoter, since this species has the same nesting preferences as Tufted Duck, which favours bushy sites (Cramp & Simmons 1977, Kear 2005). White-winged Scoters were observed in close proximity, although we were unable to locate a nest of this species; but subsequently, staff at the Wildfowl and Wetlands Trust, Slimbridge, UK, were able to review Plate 1 and confirm that this large egg indeed belongs to a scoter (B. Hughes in litt. 2012). Confirmation of inter- or intraspecific brood-parasitism involves several methods, including morphological identification, biochemical and genetic diagnosis, and abnormal laying events, large clutch-size or late hatching events (Yom-Tov 1980, 2001). Applying these methods for determining parasitic eggs in breeding colonies may be challenging due to difficulties of egg-collecting in the field. Intraspecific parasitic eggs are easier to identify than those of conspecifics because of larger egg variation between species. In our case, it was not possible to use biochemical and genetic methods to identify these eggs unambiguously. Nonetheless, our method of combining information of local breeding duck species, egg morphology and nesting preference supports the assumption of this usual nest being a case of inter- and intraspecific brood-parasitism. Inter- and intraspecific brood-parasitism has prompted several hypotheses (Sayler 1992, Beauchamp 1998). However, brood- parasitism in the Anatidae can be influenced by several phylogenetic and environmental factors (reviewed in Kear 2005), highlighting the importance of documenting more cases in the field. To our knowledge, this is the first case of a female T ufted Duck incubating a parasitic egg from White-winged Scoter. Plate 2. Nest of Tufted Duck Aythya fuligula found near Kurma, Russia, 3 July 2008. (Both photographs by the authors.) 148 SHORT NOTES Forktail 28 (2012) Acknowledgements We thank Nicole A. Schneider, Dr. Igor Fefelov and Prof. Wei Liang for valuable comments on a previous version of the manuscript. YL is grateful to the financial support for the field trip to Lake Baikal from the Swiss Veterinary Office (BVET). R. M. Kilner and B. Hughes kindly reviewed the manuscript. Nigel Collar generously edited the manuscript into its final version. References Baicich, P.J.& Harrison, C. J.O. (1997) A guide to the nests, eggs, and nestlings of North American birds. San Diego: Academic Press. Beauchamp, G. (1998) The relationship between intra-and interspecific brood amalgamation in waterfowl. Condor 100: 153-162. Cramp, S. & Simmons, K. E. L„ eds. (1977) The birds of the Western Paiearctic, 1. Oxford: Oxford University Press. Davies, N. B. (2000) Cuckoos, cowbirds and other cheats. London: T. & A. D. Poyser. Geffen, E. & Yom-Tov, Y. (2001 ) Factors affecting the rates of intraspecific nest parasitism among Anseriformes and Galliformes. Anim. Behav. 62: 1027-1038. Kear, J. (2005) Ducks, geese and swans. Oxford: Oxford University Press. Kilner, R. M. (2006) The evolution of egg colour and patterning in birds. Biol Rev. 22:383-406. Sayler, R. D. (1 992) Ecology and evolution of brood parasitism in waterfowl. Pp. 290-322 in B. D. J. Batt, A. D. Afton, M. G. Anderson, C. D. Ankney, D. H. Johnson & J. A. Kadlec, eds. Ecology and management of breeding waterfowl. Minneapolis: University of Minnesota Press. Yom-Tov, Y. (1980) Intraspecific nest parasitism in birds. Biol. Rev. 55: 93- 108. Yom-Tov, Y. (2001) An updated list and some comments on the occurrence of intraspecific nest parasitism in birds. Ibis 143: 133-143. Yang LIU, Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, 3012, Bern, Switzerland. Present address: State Key Laboratory of Biocontrol and School of Life Sciences, Sun Yat-Sen University, Guangzhou 51 0275, China. Email: yang.liu@iee.unibe.ch Sergey PYZHJANOV, Irkutsk State Pedagogical University, Nizhnjaja Naberezhnaja str. 6, RU-664011 Irkutsk- II, Irkutsk, Russia. Email: pyjyanov@igpu.ru Annual survival rate and mean life-span of Lemon-bellied White-eyes Zosterops chloris flavissimus on Kaledupa island, Wakatobi, south-east Sulawesi, Indonesia DAVID J. KELLY & NICOLA M. MARPLES White-eyes (Zosteropidae) are known to be the most rapidly speciating family of birds on the planet (Moyle etal. 2009). One of the reasons that white-eyes manage to adapt and exploit new habitats so well is because of their relatively short generation time (and greater Darwinian fitness); some Zosterops species may breed within six months of hatching (Moyle et at. 2009). However, while the breeding age of white-eyes appears to be relatively well known, there is less information on the longevity of these species. One might anticipate that such small species are destined to have rather short life-spans (Hulbert etal. 2007). Ourteam makes regulartripstosouth-eastSulawesi, Indonesia, as part of a long-running island biogeography project on local bird species, in cooperation with Operation Wallacea Limited. As part of this work, we mist-net and colour-ring birds in the area. Recently (August 2010), we retraced the footsteps of a previous expedition (August 2007) with great precision. This allowed us the opportunity to assess site-fidelity and longevity of a number of regularly trapped small passerine species, notably: Lemon-bellied White-eye Zosterops chloris, Olive-backed Sunbird Cinnyris jugularis, Grey¬ sided Flowerpecker Dicaeum celebicum and Island Monarch Monarcha cinerascens. We visited three sites (Bakau, Air Nounou, Latafe) on the island of Kaledupa, Wakatobi, south-east Sulawesi in both 2007 (using yellow colour rings) and 2010 (using mauve colour rings) and mist-netted with similar equipment in similar habitats. As many of the local shrubs grow at remarkable speeds, it was not always possible to identify previous net-ride locations. However, local guides and GPS co-ordinates confirmed the accuracy of our site selection. During our 2010 visit we trapped four birds bearing the yellow rings we had used during our 2007 visit. All -of these birds were Lemon-bellied White-eyes. We are confident that the Lemon-bellied White-eyes trapped in 2007 were adult birds. We have occasionally trapped white-eyes in juvenile plumage on the Wakatobi islands (5/548 white-eyes trapped), but have rarely trapped beyond early September. The breeding season for Lemon-bellied White-eyes on the nearby islands of Muna and Buton is between September and October (van Balen 2008). It therefore seems likely that the breeding season for the Wakatobi island white-eyes is similar. If this is the case, then the retrapped birds, when they were trapped in 2007, must have been at least one calendar year old. So, when the birds were retrapped in 2010, they must have been at least four years old. We used these ages to calculate a minimum adult survival rate for the Kaledupa birds. We trapped a total of 48 Lemon-bellied White-eyes in 2007 and retrapped four of those birds in 2010. This gives a minimum percentage of 8.3% of birds surviving into their fourth calendar year on Kaledupa. In order to allow 8.3% of the adult population to survive intotheirfourth calendaryear, the minimum annual survival rate of the local population must be 43.6%. This value is in excess of the annual survival rates recorded for African Yellow White-eye Z. senegalensis (34%) near Jos, Nigeria (McGregor et a!. 2007) and SilvereyesZ. lateralis (24-26%) in central Victoria, Australia (Burton 1 996). Longevity may be derived from annual survival rate, where mean life-span = -1/ln(annual survival rate) (Seber 1 982). Applying this formula, we get a mean life-span for the Lemon-bellied White- eyes of Kaledupa of 2.2 years (1.2 years from survival rate + 1 year at initial capture). As the survival rate value is a minimum, the calculated life-span is a minimum value too. However, this minimum value for the Kaledupa birds is greater than the mean life-spans of the African Yellow White-eyes near Jos (1 .92 years = 0.92 years from survival rate + 1 year at initial capture) and the Silvereyes in central Victoria (9.8-10.2 months) (Burton 1996). It is likely that the value cited by Burton (1996) includes birds born during the year of capture. Annual survival rates of adults are usually higher than those of fledglings (Freed & Cann 2009). The Capricorn White-eye of Heron Island Z. lateralis chlorocephalus has an annual adult mortality of 38.5% (Brook & Kikkawa 1 998). This is equivalent to an annual adult survival of 61 .5% (a value in excess of the Kaledupa birds) and translates to an average life span of 3.1 years (2.1 years from survival rate + 1 year at initial capture). The Lemon-bellied White-eyes, when retrapped in 2010, appeared to be in breeding pairs (one male and one female Forktail 28 (2012) SHORT NOTES 149 bird caught in the same net at the same time). We caught two 'breeding' pairs, one at each of two different sites (Bakau and Latafe). If we consider the retrapping rates at the two sites independently (Bakau-1 2.5%, Latafe-20%), this gives us annual survival rates of 50% and 58.5% respectively and mean life¬ span estimates of 2.4 years (1 .4 years from survival rate + 1 year at initial capture) and 2.9 years (1.9 years from survival rate + 1 year at initial capture) respectively. These values approach those of the Heron Island Silvereyes (3.1 years). The Heron Island white-eyes are known to show density dependence in their breeding success (McCallum et al. 2000), but it is unclear whether the local population density has any effects on annual survival (Kikkawa 1980). We did not attract birds to the nets with recorded calls or songs on our trips to Kaledupa, so it is unlikely that we caught all of the local population on either of those visits. It is unclear whether catching a larger sample would have increased or decreased our longevity estimate. The presence of 'breeding' pairs at two of the netting sites suggests a high degree of site fidelity, at least by some individuals. Other data demonstrate that there is little or no movement of the Wakatobi Lemon-bellied White-eyes between islands (Kelly et at. unpubl. data), supporting the idea that the Lemon-bellied White-eyes of the Wakatobi are generally sedentary in nature. While we mist-netted on other islands across the Wakatobi archipelago during our 2010 field season (Wangi-Wangi, Hoga, Tomia and Binongko), those other islands had only been visited previously in 2005, not 2007. Furthermore, we did not make the same effort on those other islands to revisit our former netting sites. The only birds we retrapped in 2010, from previous expeditions, were those on Kaledupa. Therefore, it is unclear if the Lemon-bellied White-eyes of Kaledupa are especially long-lived in comparison to the populations of the species on the other Wakatobi islands. Irrespective of this, it does appear that the Lemon-bellied White- eyes of Kaledupa are longer-lived than mainland populations of African Yellow White-eye and Silvereye. While the current dataset is rather small, there appears to be a tendency for populations of Zosterops species to live longer on oceanic islands than on the mainland. We will endeavour to collect more data on the longevity of the Wakatobi bird populations to allow further investigation of these findings. Acknowledgements We are very grateful to Operation Wallacea Limited and their staff for providing logistical support throughout our island biogeographic studies and travels, as well as the people of the Wakatobi islands for hosting our visits. We would like to thank the project students and general volunteers who assisted us during our many netting sessions across the Wakatobi islands. We are also grateful to two anonymous referees for their comments on a draft of this paper. References van Balen, B. (2008) Family Zosteropidae (white-eyes). Pp.402-485 in J. del Hoyo, A. Elliott & D. A. Christie, eds. Handbook of the birds of the world, 13. Barcelona: Lynx Edicions. Brook, B. W. & Kikkawa, J. (1998) Examining threats faced by island birds: a population viability analysis on the Capricorn silvereye using long-term data. J. Appl. Ecol. 35: 491-503. Burton, T. C. (1996) Changes in the abundance of silvereyes in a central Victorian vineyard during the grape-ripening period. Corella 20: 61-66. Freed, L. A.&Cann, R. L. (2009) Negative effects of an introduced bird species on growth and survival in a native bird community. Current Biol. 1 9: 1 736- 1740. Hulbert, A. J., Pamplona, R., Buffenstein, R. & Buttemer, W. A. (2007) Life and death: metabolic rate, membrane composition, and life span of animals. Physiology Reviews 87: 1 1 75-1213. Kikkawa, J. (1980) Winter survival in relation to dominance classes among silvereyes Zosterops lateralis chlorocephala of Heron Island, Great Barrier Reef. Ibis 1 22: 437-446. McCallum, H., Kikkawa, J. & Catterall, C. (2000) Density dependence in an island population of silvereyes. Ecology Letters 3: 95-100. McGregor, R„ Whittingham, M. J. & Cresswell, W. (2007) Survival rates of tropical birds in Nigeria, West Africa. Ibis 1 49: 61 5-618. Moyle, R. G., Filardi, C. E., Smith, C. E. & Diamond, J. (2009) Explosive Pleistocene diversification and hemispheric expansion of a 'great speciator'. Proc. Natn. Acad. Sci. 1 06: 1 863-1868. Seber, G. A. F. (1982) The estimation of animal abundance and related parameters. London: Griffin. David J. KELLY and Nicola M. MARPLES, Trinity College Dublin, Department of Zoology, School of Natural Sciences, Dublin 2, Ireland; and Trinity Centre for Biodiversity Research , Trinity College Dublin, College Green, Dublin 2, Ireland. Email: djkelly@tcd.ie;nmarples@tcd.ie Migrating dragonflies: famine relief for resident Peregrine Falcons Falco peregrinus on islands CHANG-YONG CHOI & HYUN-YOUNG NAM The diet of the Peregrine Falcon Falco peregrinus has been well documented around the world. Peregrines are powerful predators which feed mainly on birds, and more than 1,000 avian species ranging from 1 0 to 3,000 g in weight have so far been recorded as prey (Ferguson-Lees & Christie 2001 ). However, there are also many reports of occasional consumption of insects (e.g. Pruett-Jones et al. 1980, Ritchie 1 982, White & Brimm 1990, Oro& Telia 1995, White etal. 2002) as well as reptiles (Oro & Telia 1995) and mammals (e.g. bats and rodents: Byre 1990, Bradley & Oliphant 1991). Although insects are an uncommon food for Peregrines, such prey are diverse from small ones like the Plecoptera (stoneflies: Sumner & Davis 2008) to large ones, which include some Hemiptera (cicadas: Pruett- Jones et al. 1980, Ellis etal. 2007), Orthoptera (grasshoppers and crickets: Pruett-Jones etal. 1 980, White & Brimm 1 990, White etal. 2002) and Odonata (dragonflies and damselflies: White etal. 2002). Insects may be more important in Peregrine diets than is commonly believed (Snyder & Wiley 1976, Ellis etal. 2007). This article reports two adult Peregrines hunting migratory dragonflies, and discusses the implications of dragonflies being a food source for falcons on remote islands. Hongdo, the study area, is a small island in the Republic of Korea located c.120 km south-west of the Korean Peninsula and 430 km from mainland China at 34°41 N 125°12'E, and is a key stopover site for migratory birds that cross the Yellow Sea. More than 327 bird species (about 63% of the total recorded in Korea) have been recorded on this island, but only ten, including a pair of Peregrines, are resident (NPRI 2009). The first observation of an adult Peregrine foraging on dragonflies in flight was on 27 August 2009; it took three dragonflies during 8 minutes of observation. Over the next few days, the foraging activities of two adults hunting dragonflies were occasionally but repeatedly observed, including at least 20 more dragonfly captures (Figure 1). Most such foraging attempts were made in foggy conditions with still air, apparently irrespective of 150 SHORT NOTES Forktail 28 (2012) Figure 1. A Peregrine Falcon Falco peregrinus feeding on a dragonfly (Lesser Emperor Anax parthenope) on Hongdo, Republic of Korea. time of day. No such foraging was seen after 30 August, even though the survey continued right through the year. On 29 November 2009, close examination of the pair's plucking perches at the top of some rocky cliffs revealed the remains of a dragonfly wing amidst the feathers and carcasses of diverse avian species. Newly fledged Peregrines often pursue flying insects, to improve their hunting skills rather than for any energetic reward (Dekker 1 999, Razafimanjato et al. 2009); however, the chasing of dragonflies by these adults on Hongdo was clearly unnecessary for such learning, and indicates that these Peregrines were targeting dragonflies as a food source. In the study area, based on field collection and observations, the dominant migratory dragonflies in August were Globe Skimmers Pantala flavescens. Scarlet Skimmers Crocothemis servilia, Lesser Emperors Anax parthenope and a species of meadowhawk or darter Sympetrum kunckeli were also recorded but more rarely. Since Hongdo lacks wetlands, none of the dragonfly species breeds there, but huge (more than tens of thousands) swarms of them routinely pass over the study site during their southward migration from July to September. Although it was impossible to identify to species all dragonflies taken, the two species confirmed as prey were the largest, Lesser Emperor, and the most abundant, Globe Skimmer. Like many dragonflies that cover distances of hundreds orthousands of kilometres, both species are intercontinental long¬ distance migrants with worldwide distributions, forming big Figure 2. Monthly changes in the number of bird species and abundance counted every day in 2007 on Hongdo. Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec swarms during ocean crossings (Corbet 2004, Anderson 2009, Borisov 2009). As residents of Hongdo, we conducted daily bird counts throughout the year and managed a bird-ringing programme to monitor bird migration. There are no breeding seabird colonies nearby, so the Hongdo Peregrines rely mainly on the diverse transient birds in spring (March to May) and autumn (September to November) and on the abundant wintering seabirds from November to January. The lowest numbers of bird species and individuals have been recorded in summer, particularly from June to August, between the two migratory seasons (Figure 2), and this pattern of occurrence results in a reduced availability of birds in the diets of the two Peregrines when large numbers of migratory dragonflies arrive at Hongdo whilst crossing the Yellow Sea. The wet weights of the Lesser Emperors and Globe Skimmers at the study site were only 0.95±0.26 g (n=6) and 0.31 ±0.04 g (n=16), respectively. However, migratory dragonflies in swarms, particularly Globe Skimmers, have been suggested to be a potential food source for migrating raptors, particularly Amur Falcons Falco amurensis, which cross the Indian Ocean without any other identified prey (Anderson 2009). The Hongdo Peregrines' behaviour suggests that dragonflies migrating in large numbers may also be famine relief food for sedentary raptors on remote islands. Prey abundance is an important factor in prey selection of Peregrines (Bradley &Oliphant 1 991 ), and they may take advantage of the easily obtainable prey occurring in large concentrations (Byre 1990), even though the prey is not part of their usual diet. The proportion of dragonflies in the overall diet of Peregrines is probably low in both frequency and biomass, and the energetic reward from hunting dragonflies is unclear; nevertheless, this report implies that migratory and swarming insects may be important in Peregrine diets under certain circumstances such as low preferred- prey abundance and limited visibility. Acknowledgements We thank all researchers and staff at the Migratory Birds Center of the Korea National Park Service who are involved in the migratory bird monitoring programme with us on Hongdo. References Anderson, R. C. (2009) Do dragonflies migrate across the western Indian Ocean? J. Trop. Ecol. 25: 347-358. Borisov, S. N. (2009) Study of dragonfly (Odonata) migrations in the western Tien Shan Mountains using ornithological traps. Entomol. Review 89: 1025-1029. Bradley, M.&Oliphant, L.W. (1 991 ) The diet of Peregrine Falcons in Rankin Inlet, Northwest Territories: an unusually high proportion of mammalian prey. Condor 93: 193-197. Byre, V. J. (1990) A group of young Peregrine Falcons preying on migrating bats. Wilson Bull. 102: 728-730. Forktail 28 (2012) SHORT NOTES 151 Corbet, P. S. (2004) Dragonflies: behaviour and ecology of Odonata. Colchester, U.K.: Harley Books. Dekker, D. (1999) Bolt from the blue: wild Peregrines on the hunt. Surrey, U.K.: Hancock House. Ellis, D. H., LaRue, C. T., Fackler, J. K. & Nelson, R. W. (2007) Insects predominate in Peregrine Falcon predation attempts in Arizona. Western Birds 38:261-267. Ferguson-Lees, J. & Christie, D. A. (2001) Raptors of the world. London: Christopher Helm. [NPRI] National Park Research Institute (2009) Annual report on migratory bird researches. Namwon, Korea: National Park Research Institute. Oro, D. & Telia, J. L. (1995) A comparison of two methods for studying the diet of the Peregrine Falcon. J. Raptor Res. 29: 207-210. Pruett-Jones, S. G., White, C. M. & Devine, W. R. (1980) Breeding of the Peregrine Falcon in Victoria, Australia. Emu 80: 253-269. Razafimanjato, G., Rene de Roland, L.-A., Rabearivony, J. & Thorstrom, R. (2009) Nesting biology and food habits of the Peregrine Falcon Falco peregrinus radama in the south-west and central plateau of Madagascar. Ostrich 78: 7-12. Ritchie, R. J. (1 982) Porcupine quill and beetles in Peregrine castings, Yukon River, Alaska. Raptor Res. 16: 59-60. Snyder, F. N. & Wiley, J. W. (1976) Sexual size dimorphism in hawks and owls of North America. Ornithol. Monogr. 20: 1-96. Sumner, J. & Davis, K. (2008) Observations of adult Peregrine Falcons capturing stoneflies. Western Birds 39: 220-224. White, C. M. & Brimm, D. J. (1990) Insect hawking by a Peregrine Falcon {Falco peregrinus) in Fiji. Notornis 37: 140. White, C. M., Clum, N. J., Cade, T. J. & Hunt, W. G. (2002) Peregrine Falcon (Falco peregrinus). In A. Poole and F.Gill, eds. The birds of North America. 660. Washington DC: Academy of Natural Sciences Philadelphia, and American Ornithologists' Union. Chang-Yong CHOI, Migratory Birds Center, National Park Research Institute, Korea National Park Service, Jin-ri, Heuksan-myeon, Shi nan- gun, Jeonnam Province 535-91 7, the Republic of Korea. Email: subbuteo@hanmail.net Hyun-Young NAM, Migratory Birds Center, National Park Research Institute, Korea National Park Service, Jin-ri, Heuksan-myeon, Shi nan- gun, Jeonnam Province 535-91 7, the Republic of Korea. Email: stern0223@lycos.co.kr Breeding of the Japanese Murrelet Synthliboramphus wumizusume in South Korea DONG-WON KIM, CHANG-WAN KANG, HWA-JUNG KIM, YOUNG-SOO KWON & JIN-YOUNG PARK The global population of Japanese Murrelet Synthliboramphus wumizusume is believed to number only 4,000-10,000 birds, and owing to a rapid population decline the species has been designated as Vulnerable on the IUCN Red List (Carter et al. 2002, Bird Life International 201 1,IUCN 2011). Almost all of the population breeds on uninhabited rocky islands in Japan, mainly in Kyushu and on the Izu Islands, between mid-February and early May (Bi rd Life International 2001, Carter et al. 2002). There is also Figure 1. Map of South Korea and the locations of (A) Daegugul Island, (B) Dok Island, and (C) Jeju Island. evidence of breeding in Russia: a dead juvenile was found in Boysman Bay in July 1984 (BirdLife International 2001). In South Korea, breeding was first recorded at Daegugul Island (Daeguguldo), Shinan county, Jeollanam province, off the south¬ west coast of South Korea (Figure 1): three breeding pairs were found here for the first time on 1 0 May 1 983 (Kyunghyang Shinmun 1983, Won 1992). The Korean Government designated Daegugul Island as National Monument no. 341 in 1 984 for the protection of breeding seabirds, and designated the species itself as no. 450 in March 2005. Subsequent records in both breeding and non-breeding seasons were restricted to the south coast of South Korea (Park 2002, Oh 2004) until the discovery of a second breeding site: Dok Island (Dokdo), Ulleung county, Gyeongsangbuk province. Dok Island is located in the East Sea (Sea of Japan), c.220 km from mainland South Korea, and is composed of two main islands (Dong Island and Seo Island) and dozens of small islets (Figure 1 ). On 28 May 2005, an adult and a chick were found dead on Dong Island (37°14'21"N 1 31 °52'07"E) and Seo Island (37°14'35"N 1 31 °51'53"E), respectively, by YSK. The dead adult was lying on the shore and the dead chick was floating on the sea with its head pecked by an unknown predator (Figure 2a; Kwon & Yoo 2005). In 2009, a fledgling and two adults were filmed leaving the island at night by the Seoul Broadcast System in a programme entitled Dokdo, Saengmyeong-ui Ddang [Dok Island, the Land of Life] (see http://www.pandora.tv/ video.ptv?c1=08&c2=0175&ch_userid=loveasia&prgid=39061826 and http://www.pandora.tv/video.ptv7d =08&c2=01 75&ch_userid =loveasia&prgid=39061 822}. Subsequently, a dead adult was found on Dong Island on 15 July 2010 by JYP. These records suggest that Japanese Murrelet breeds on the island, although active nests have not yet been found. Here, we report a third breeding area of this species in South Korea: Jeju Island in Jeju Special Self-Governing Province (hereafter 'Jeju province'), which is the southernmost island in South Korea (Fig. 1).The possibility of Japanese Murrelet breeding on Jeju Island was anticipated owing to regular observation of adults during the breeding season at sea between Gapa and Mara Islands south-west 152 SHORT NOTES Forktail 28 (2012) Figure 2. Japanese Murrelet: (a) a dead chick found on Dok Island on 28 May 2005; (b) a chick on Jeju Island on 29 April 2011; and (c) an adult and two juveniles on Jeju Island on 25 May 2011. of Jeju Island: one individual in May 2006, five in March and 14 in May 2007 (Kim 2008); ten in April and seven in May 2008 (Kim etal. 2010); one in April and five in May 2009 (CWK); one in April and two in May 2010 (CWK); and five in April and three in May 201 1 (DWK). Breeding was suspected when two Synthllboramphus chicks were observed for a few seconds by HJK and CWK on the coast at 33°11'51"N 126°17'43''E from a boat heading to Mara Island, but no photographs were obtained. No adults were seen nearby, so it was not possible to determine if the chicks were Japanese or Ancient Murrelet 5. antiquus. The latter breeds in South Korea, but it is not known from Jeju Island during the breeding season (Kim et al. 201 1 ). Finally, breeding of Japanese Murrelet on Jeju Island was confirmed on 29 April 201 1 when DWK identified a live Japanese Murrelet chick on the coast of Seogwipo city at 33°14'21"N 1 26°36'32"E (Figure 2b). The identification was based on the dark upperparts contrasting with the white underparts and the dark cap contrasting with white cheeks and throat (Ancient Murrelet chicks have dark cheeks with some white behind the eye: Harrison 1 983). The chick appeared to be a few days old; it was very small and covered with down. It was observed from 10h40 to 1 1 h40 swimming along the shore, occasionally attempting but failing to jump up onto rocks. Murrelet chicks typically stay with adults at sea after fledging (del Hoyo et al. 1 996), but no adults were seen. The chick may have become separated from the adults during fledging, or the latter may have been predated (e.g. by Peregrine Falcon Falco peregrinus). The chick was not observed the next day, despite searching the southern coast of Jeju Island. Subsequently, on 25 May 201 1, one adult and two juveniles were seen and photographed by CWK at sea between Gapa and Mara Islands at 33°8'1 3"N 1 26°1 5'E (Figure 2c). The juveniles appeared older than the chick observed in April, based on their size and head pattern. These observations confirm breeding around Jeju Island, but it is still unknown how many of the islands in Jeju province support breeders, or how large the population is. Similarly, surveys are required on Dok Island. On Daegugul Island, surveys in April 2006 by HJK recorded a mixed flock of Japanese and Ancient Murrelet numbering c.200 individuals, but in unknown proportions (Kim 2006). Subsequent surveys on Daegugul Island have been mainly focused on Swinhoe's Storm-petrel Oceanodroma monorhis (Lee etal. 20 10). Therefore, surveys of all three known sites are required as a prerequisite for conservation and management. Forktail 28 (2012) SHORT NOTES 153 Acknowledgements We acknowledge the support of the National Institute of Environmental Research and the National Institute of Biological Resources of South Korea. We thank Sa-Ra Oh, Seong-Sik An and fellows of Jeju Wildlife Research Center for their assistance in the fields. We are also very grateful to Yun-Kyoung Lee and Hang-Soo Cho for their helpful advice on the first draft of this paper, and also to Jin-Ho Song and Kwan-Mok Kim who made this manuscript better. Finally, we thank two anonymous reviewers for comments. References BirdLife International (2001) Threatened birds of Asia: the BirdLife International Red Data Book. Cambridge, U.K.: BirdLife International. BirdLife International (2011) Species factsheet: Synthliboramphus wumizusume, Available at: http://www.birdlife.org/datazone/ speciesfactsheet.php?id=3314. Carter, H. R., Ono, K., Fries, J. N., Hasegawa, H., Ueta, M., Higuchi, H., Moyer, J.T., Ochikubo Chan, L. K., de Forest, L. N„ Hasegawa, M. & van Vliet, G. B. (2002) Status and conservation of the Japanese Murrelet (Synthliboramphus wumizusume) in the Izu Islands, Japan .J. Yamashina Inst.Orn. 33:61-87. del Hoyo, J., Elliott, A. & Sargatal, J. (1996) Handbookof the birds of the world, 3. Barcelona: Lynx Edicions. Flarrison, P. (1983) Seabirds: an identification guide. Revised edition. Australia: Croom Flelm. IUCN (2011) IUCN Red List of threatened species. Version 2011.1. . Downloaded on 20 June 201 1 . Kim, Eun-Mi, Park, Chan-Ryul & Kang, Chang-Wan (2010) The status on the legally protected birds of Korea in Jeju Island for the last three years. Korean J. Orn. 17: 259-273. (In Korean with English abstract.) Kim, Hwa-Jung (2006) Cheonyeonginyeommul Je Sambaeksasipilho Guguldo Haejoryu (Bbulsoe-ori, Badajebi, Seumsae) Beonsikji Monitoring Bogoseo (2006 nyeon) [Monitoring for the seabirds' breeding (the Japanese Murrelet, the Swinhoe's Storm Petrel, and the Streaked Shearwater) in Gugul Island, National Monument no.341 (2006)]. Pp.5-22 in The 2006 Cheonyeonginyeommul Monitoring [Monitoring of National Monument]. Cultural Heritage Administration of Korea. (In Korean.) Kim, Wan-Byung (2008) Avifauna of Mara Island, Korea. J. Korean Nature 1:143-148. Kim, Wan-Byung, Kim, Young-Ho & Oh, Hong-Shik (201 1) A study about checklist research of the birds of Jeju Island. Korean J. Orn. 18: 93-1 13. (In Korean with English abstract.) Kwon, Young-Soo & Yoo, Jeong-Chil (2005) Breeding record of the Crested Murrelet ( Synthliboramphus wumizusume ) at Dokdo Island. Korean J. Orn. 12: 83-86. (In Korean with English abstract.) Kyunghyang Shinmun (1983) Jeonnam Gugulseomseo Huigwijo Bbulsoe-ori Balgyeon [The observation of rare bird, the Japanese Murrelet, at Guguldo Island]. 3 June 1983. . Downloaded on 27 July 2011. (In Korean.) Lee, Kyung-Gyu, Ko, Kyung-Nam, Jegal, Gil-Myung & Park, Chun-An (2010) A survey report on avifauna, shorebirds and seabirds of Shinan, 1004 Islands. Shinan County. South Korea. (In Korean.) Oh, Hong-Shik (2004) A study on the management of migratory bird sanctuary and the status of waterbird migration on Jeju Island. Korean J. Orn. 11:11 -32. (In Korean with English abstract.) Park, Jin-Young (2002) Current status and distribution of birds in Korea. Ph.D. thesis, Kyung-Hee University, Seoul, South Korea. (In Korean.) Won, Pyong-Oh (1992) Yeorumcheolsae Doraeji, Beonsikji mit Haejoryu Beonsikji Haksuljosa Bogoseo [Survey report of the breeding sites for seabirds and other summer visitors in Korea], Korea Institute of Ornithology, Kyung-Hee University. (In Korean.) Dong-Won KIM, Nature Conservation Research Division, National Institute of Environmental Research, Incheon 404-708, South Korea. Email: foowl@korea.kr Chang-Wan KANG, The Korea Association For Bird Protection Jeju, Seogwipo 697-340, South Korea. Email: jejubirds@hanmail.net Hwa-Jung KIM, National Institute of Biological Resources, Incheon 404-708, South Korea. Email: hwajung@korea.kr Young-Soo KWON, National Park Research Institute, Korea National Park Service, Namwon 590-811, South Korea. Email: auk 1 005@hanmail.net Jin-Young PARK, Nature Conservation Research Division, National Institute of Environmental Research, Incheon 404-708, South Korea. Email: birdkorea@korea.kr An intraspecific adult killing in female Japanese Great Tits Parus major minor NORIMASA 5UGITA,TOSHITAKA N. SUZUKI, CRAIG A. BARNETT & KEISUKE UEDA Introduction Intraspecific killing has been documented in a wide variety of avian taxa. Adults sometimes kill eggs and chicks of conspecifics when birds are competing for nesting sites (Bel les-lsles & Pieman 1986, Stanback & Koenig 1992, Inoue et al. 2010) or when birds are expanding their territories (Lee et al. 201 1). Parents may also practise infanticide in cases of intraspecific brood parasitism when they identify parasitic chicks (Shizuka & Lyon 2010). In species that practise obligate brood reduction, the death of the victim is normally caused by a sustained assault by an older nestling and its evolution is concentrated in four orders: Gruiformes, Pelecaniformes, Accipitriformes and Sphenisciformes (Mock & Parker 1997, Simmons 2002). Fighting among adults is also common, but there are few instances where birds have been observed to fight to the death, especially in small passerines. Direct observations of intraspecific killing between adults suggest that it can occur to provide food (i.e. cannibalism, Anderson 2004) or in response to nest-site competition (Flux & Flux 1992). However, the scarcity of reports means that there is a need for more observations of fighting between adults which result in death of one individual. This would lead to greater understanding of the conditions under which such events occur. In this paper, we report a case of intraspecific killing in Japanese Great Tits Parus major minor, providing a direct observation in which an adult female attacked and killed another adult female. Observations Our observation was made on a street beside a building at Rikkyo University, Toshima, Tokyo, Japan (35°44'N 139°42'E). Although the university campus is located in an urban area it has some open grounds with tall trees. Great Tits inhabit the campus year-round and nest in tree cavities from April to July. We saw a Great Tit lying on the ground flapping its wings at 1 1 h 1 0 (Japan Standard Time) on 16 March 2010. Immediately, another Great Tit approached, swooped on, pushed and shoved the lying tit. The aggressor stuck its beak into the eyes of the other bird and plucked feathers from its back and abdomen. The 154 SHORT NOTES Forktail 28 (2012) Figure 1. Video frames of a female Great Tit (A) attacking another conspecific female (B). (a) The aggressor flew towards the victim, (b) pressed it and pecked its abdomen, (c) stabbed its eye and (d) continued to attack the victim. aggressor then returned to a perch near the victim and gave warning calls for a few minutes. It then flew back down to the victim and continued its attack. We observed six assaults with physical contacts over approximately 20 minutes. We recorded the last assault with a HDR-XR500 video camera (Sony Corporation, Tokyo, Japan) (Figure 1 ; clip available on website of Movie Archives of Animal Behavior, data number: momol 20606pm01 b, URL: http://zoo2.zool. kyoto- u.ac.jp/ethol/mov/12/1206/momo120606pm01.MP4). Both birds were adult females since female Great Tits have narrower and more obscure black mid-line stripes from the bib to vent than do males. (a) Figure 2. Marks of attacks by a female Great Tit on the dead body of a conspecific female, (a) Some feathers on the back were pulled out. (b) An eye was injured. We waited until the aggressor had stopped its attacks and left the vicinity. We confirmed the death of the victim and collected its body. We dissected the dead bird for further inspection of its injuries and body condition. The dead tit had developed ovarian follicles (1 .3 x 1 .3 mm) and weighed 1 2.2 g, which is lighter than would be expected for an adult female (mean ± SE: 1 5.0 ± 0.8 g, N = 95: Ohori 2007). Feathers of the bird had been plucked from its neck, backand abdomen, and both its eyeballs had been ruptured (Figure 2). However, there were no wounds (such as holes in the flesh) that would suggest cannibalism. Discussion Since we did not see the beginning of the fight between the females, it is difficult to know why one of the tits was lying on the ground and killed by the other tit. However, the body mass of the victim was lighter than the average body mass of female Great Tits. Therefore, it is possible that the victim was in a poor body condition when it was fighting and less able to defend itself from attacks. It is also possible that the victim had been injured as a result of an accident (such as flying into a window) before being attacked. The attacking tit might have seen the victim lying on a street as it passed by and decided to attack. The black stripe on the breast of the Great Tit has been shown to be a releasing signal for aggressiveness in this species (Jarvi & Bakken 1984). Scriba & Goymann (2008) also showed that stuffed mounts induced more extreme stress responses from European Robins Erithacus rubecula than did caged live birds. Therefore, the victim lying on the ground with its breast- stripe showing might have served to produce the extreme response of the attacker we report above. Previous studies suggest that limited availability of desirable nest-sites causes aggressive interactions between females (e.g. Forktail 28 (2012) SHORT NOTES 155 Rosvall 2008). Such competition might occasionally result in intraspecific killing. For example, in a New Zealand population of European Starlings Sturnus vulgaris females competed with one another intensely for access to nest-boxes. This competition occasionally resulted in one female stabbing the other with their claws, which could result in fatal injuries (Flux & Flux 1992). In a Belgian population of European Starlings with higher nest-box densities (and therefore less competition for nest-sites), females were less likely to fight for access to nest-boxes (Pinxten etal. 1 989). Female European Starlings respond most aggressively towards other female starlings in the pre-laying phase of the breeding season, when competition is most intense for nest-sites (Sandell & Smith 1 996). In the long-term study of Great Tits at Wytham Woods, dead bodies of females have been found early in the breeding season with their heads pecked (A. Gosler In litt. 201 2). The attack we report here took place in early spring, when Great Tits are starting to breed and searching for suitable nest cavities. Therefore, female-female competition for nest-sites is another possible cause for the fight between the females. The sustained nature of the attack is interesting as the attacking female continued to attack the prone bird for over 20 minutes. Cases of physical aggression in small passerines (including Great Tits) are normally much shorter than the length of time that we observed this fight (CAB pers. obs.). This is because fighting, whilst sometimes necessary to defend resources or offspring, is costly even for victors. There are direct energetic costs associated with fighting and as well as risks of injury (e.g. Haley 1994). Additionally, individuals may be subject to increased predation risk whilst they are engaged in fighting because they may not be so vigilant for predators and/or more conspicuous (e.g. Jacobsson et al. 1995). Finally, there may be opportunity costs in fighting such as lost foraging opportunities (e.g. Neat et al. 1 998). Therefore, it would seem that this prolonged attack may not have been in the best interest of even the attacking female and might have been induced as a result of the behaviour and posture of the dying victim. Our observations provide the first evidence of intraspecific killing between female Great Tits. There are some reports of intraspecific adult killing in other species (e.g. Lombardo 1 986, Flux & Flux 1992, Anderson 2004). However, most observations are unlikely to be published. Therefore, we would encourage the publication of further observations of intraspecific killing in order to better understand the conditions that lead to escalation of fighting and to the death of one individual. Acknowledgements We are very grateful to Yumi Ito for providing information about the fighting of Great Tits. We also thank Hiromi Kamigaichi for dissecting the dead tit body and Sachiko Endo and Masayoshi Kamioki for providing comments on the manuscript. Norimasa Sugita and Toshitaka N. Suzuki contributed equally to this work. References Anderson, E. M. (2004) Intraspecific predation among Northwestern Crows. Wilson Bull. 116: 180-181. Bel les-lsles, J.-C. & Pieman, J. (1986) House Wren nest-destroying behavior. Condor 88: 1 90-193. Flux, J. E. C. & Flux, M. M. (1992) Nature red in claw: how and why starlings kill each other. Notornis 39: 293-300. Haley, M. P. (1994) Resource-holding power asymmetries, the prior residence effect, and reproductive payoffs in male northern elephant seal fights. Behav. Ecol. Sociobiol. 34: 427-434. Inoue, Y., Yoda, K., Fujii, H„ Kuroki, H. & Niizuma, Y. (2010) Nest intrusion and infanticidal attack on nestlings in Great Cormorants Phalacrocorax carbo: why do adults attack conspecific chicks? J. Ethol. 28: 221-230. Jacobsson, S., Brick, O. & Ku 1 1 berg, C. (1995) Escalated fighting behaviour incurs increased predation risk. Anim. Behav. 49: 235-239. Jarvi,T. & Bakken, M. (1984) The function of the variation in the breast stripe of the Great Tit ( Parus major). Anim. Behav. 32: 590-596. Lee, S., Seo, K., Lee, W„ Kim, W„ Choe, J. C. & Jablonski, P. (2011) Non-parental infanticide in a dense population of the Black-billed Magpie (Pica pica). J. Ethol. 29: 401-407. Lombardo, M. P. (1986) A possible case of adult intraspecific killing in the Tree Swallow. Condor 88: 1 1 2. Mock, D. W. & Parker, G. A. (1997) The evolution of sibling rivalry. Oxford: Oxford University Press. Neat, F. C., Taylor, A. C. & Huntingford, F. A. (1998) Proximate costs of fighting in male cichlid fish: the role of injuries and energy metabolism. Anim. Behav. 55: 875-882. Ohori, S. (2007) Shijyuukara. Bird Res. News 4: 2-3. (In Japanese.) Pinxten, R„ Eens, M. & Verheyen, R. F. (1989) Polygyny in the European Starling. Behaviour 111: 234-256. Rosvall, K. A. (2008) Sexual selection on aggressiveness in females: evidence from an experimental test with Tree Swallows. Anim. Behav. 75: 1603- 1610. Sandell, M. I. & Smith H. G. (1996) Female aggression in the European Starling during the breeding season. Anim. Behav. 53: 1 3-23. Scriba, M. F. & Goymann, W. (2008) The decoy matters! Behavioural and hormonal differences in the reactions of European Robins towards stuffed and live decoys. Gen. Comp. Endochrinol. 1 55: 51 1-516. Shizuka, D. & Lyon, B. E. (2010) Coots use hatch order to learn to recognize and reject conspecific brood parasitic chicks. Nature 463: 223-226. Simmons, R. E. (2002) Siblicide provides food benefits for raptor chicks: re¬ evaluating brood manipulation studies. Anim. Behav. 64: F19-F24. Stan back, M.T. & Koenig, W. D. (1992) Cannibalism in birds. Pp. 277-298 in M. A. Elgar & B. J. Crespi, eds. Cannibalism: ecology and evolution among diverse taxa. New York: Oxford University Press. Norimasa SUGITA, Toshitaka N. SUZUKI (corresponding author), Craig A. BARNETT and Keisuke UEDA, Department of Life Science, Rikkyo University, 3-34-1 Nishi-lkebukuro, Toshima, Tokyo 171-8501, Japan. E-mail: toshi.n.suzuki@gmail.com Nesting Fork-tailed Swifts Apus pacificus in north-eastern Vietnam VLADIMIR DINETS The breeding distribution of many bird species in Tonkin, northern Vietnam, is still poorly known (Pilgrim et al. 2009). That of Fork¬ tailed Swift Apus pacificus is no exception. In the most up-to-date overview of the region's avifauna, Robson (2011) lists this species as resident in West, but not East, Tonkin, which suggests a gap of at least 600 km between breeding sites in north-western Vietnam and those in southern China. Observations reported hereshowthat such a gap does not exist. On 1 7 June 201 1 a colony of Fork-tailed Swifts was found on a small limestone islet (20°45'N 1 07°03'E) close to Cat Ba Island in Ha Long Bay, north-eastern Vietnam. The swifts were nesting in a deep crevice in a vertical rockface rising from the sea, approximately 25 m above the high-tide mark. The nests were not visible, but swifts (up to four at a time) were repeatedly observed entering and leaving the crevice. House Swifts A. affinis were nesting on a more exposed rock surface about 30 m from the crevice in larger numbers (at least ten active nests), providing a good opportunity for comparing the birds side-by-side. The Fork-tailed Swifts were larger, with longer, narrower wings, and narrower white rump-bands. Their calls also differed obviously from those of the House Swifts, being 156 SHORT NOTES Forktail 28 (2012) long screeches rather than warbling trills. The swifts were observed in the evening, and then at dawn of the following day, both times in calm overcast weather. Recently Leader (2011) presented evidence to support the splitting of Fork-tailed Swift into four species. Of these, only/), (p.) cooki is known to breed in Indochina, although the breeding range of A. (p.) kurodae is not far from north-eastern Vietnam (it includes Guangdong and probably eastern Guangxi, China). Leader stated that A. (p.) cooki habitually breeds in limestone caves. However, cave-breeding cannot be used for differentiating between the taxa, because A. (p.) kurodae and A. (p.) pacificus also sometimes breed in caves, including caves in sea-cliffs (Leader 201 1; also pers. obs. in Vityaz Cove, Ussuriland). The plumage of the Ha Long Bay birds seemed almost black with no brown tinge. This feature and the narrow rump-bands suggest that the birds were A. (p.) cooki. Green iridescence characteristic of/), (p.) cooki was not seen, probably due to lack of direct sunlight. Ha Long Bay is a World Heritage Site, and parts of it (although not the colony site) are protected in national parks. It is not a particularly popular birding destination. The area receives heavy tourist traffic, but this is focused in popular areas. The bay contains many limestone islands and so more remote parts may harbour additional nesting sites of Fork-tailed Swift. Assuming the Ha Long Bay birds are A. (p.) cooki, the known range of this taxon is extended by 300-400 km to the east, and down to sea-level. It is possible that further studies will find A. (p.) cooki to be sympatric with A. (p.) kurodae in extreme north-eastern Vietnam or Guangxi. Acknowledgements I thank Anastasiia Tsvietkova for help in the field, and Simon Mahood and Nicholas Wilkinson for help with background information. References Leader, P. J. (201 1) Taxonomy of the Pacific swift Apus pacificus Latham, 1802 complex. Bull. Brit. Orn. Club 131: 81-93. Pilgrim, J. D., Bijlmakers, P., De Bruyn, T., Doppagne, S., Mahood, S. P. & Tordoff, A. W. (2009) Updates to the distribution and status of birds in Vietnam. Forktail 25: 1 30-1 36. Robson, C. (201 1) A field guide to the birds of South-East Asia. London: New Holland. Vladimir DINETS, 1 705 Laurel Ave. #2, Knoxville, TN 37916, USA. Email: dinets@gmail.com Notes on the 2009 autumn crane migration in Muraviovka zakaznik, Amur oblast, Russian Federation ADAM C. STEIN & GALINA NOSACHENKO Introduction Direct visual observations and counts have been a useful tool in understanding population size and dynamics of many conspicuous migratory species (Dingle 1996). In particular, observations or counts that take advantage of geographical features which funnel a large number of migrants into a relatively small area can prove highly valuable. For example, Smith (1980) was able to count over 500,000 migratory raptors as they funnelled through Panama during their migration from North to South America. Throughout the world, these geographical features have been identified by researchers and counts are now conducted annually in many of these locations providing long-term trends with certain migratory species (Dingle 1996). In the Russian Far East, the large wetland complex of Muraviovka zakaznik (Russian game reserve) serves as one of these geographical features where large numbers of White-naped Cranes Grus vipio and Hooded Cranes G. monacha become concentrated during autumn migration. This 34,000 ha zakaznik is surrounded by agricultural fields in the southern portion of Amur oblast in the Russian Federation. Given that the zakaznik is located on the Russian/Chinese border, data collected here not only have the potential to provide unique insights into the health, demographics and numbers of cranes during their migration but may also aid in detecting changes in any of these which could result from theirtransition to new political and cultural hosts. Given that both White-naped and Hooded Cranes are listed as globally Vulnerable with a declining population ( Bi rd Life International 2008), information regarding their population, environmental use and where efforts should be concentrated (both in a political and life-cycle context) will be valuable for the proper management of their populations. During the northern autumn of 2009, we monitored cranes that utilised Muraviovka zakaznik as a stopover site during their migration from their breeding grounds in Russia to their wintering sites further south in China, the Korean peninsula and southern Japan. More specifically, we performed a morning census throughout the stopover period when the birds departed from their roosts and travelled to their feeding sites. This census allowed us to derive (1 ) an approximate number of individuals of each of the species migrating through the reserve, (2) the size of flocks departing from their roosts, (3) the time at which they departed from their roosts, and (4) the approximate ratio of juveniles to adults birds. In addition, we were able to view and evaluate injured birds as well as hybrid pairs. Our key observations are presented here. Methods Autumn crane activity was observed on a near-daily basis throughout the months of September and October 2009 within Muraviovka zakaznik. From 1 5 to 24 September 2009 early morning counts were conducted from 06h15 to 09h00. During this period the migratory cranes would leave their roosts within the reserve to begin feeding within the agricultural fields east of the reserve. Counting and monitoring of the cranes was conducted along a north-south farm road that bordered the reserve and the adjacent fields. To ensure an accurate and complete count, two to four observers were spaced approximately 0.5-1 km apart along the road during the monitoring period. Utility poles were used as markers, so that each observer knew the boundaries in which he/ she was to conduct his/her monitoring and to ensure that no crane was double-counted. The cranes were counted only after they passed the transect (road) travelling from west to east. At no time during the counts were cranes observed to cross the transect from east to west. In addition to counting the cranes, observers noted any birds with apparent injuries and, when the flocks flew close enough, identified juveniles from adults. Each bird within a given flock was counted, and the time noted when each flock passed over the transect (within ten-minute intervals). Temperature data were provided for the dates of our counts by the Amur Oblast Meteorological Station located c.60 km to the west in the city of Blagoveshchensk. Forktail 28 (2012) SHORT NOTES 157 Results Numbers Migrating cranes began to arrive in small numbers (<100 individuals) within the reserve as early as 9 September, and small flocks (<20 individuals) continued to migrate through the reserve as late as mid-October. The largest density of cranes occurred during the middle part of September. The peak number of individuals we observed for both crane species was on 15 September. That morning we scored 1,095 Hooded and 255 White-naped Cranes, representing c.9% and 4% of the world's populations of these species respectively (BirdLife International 2008). Numbers decreased after this date and by 24 September fewer than 500 individual cranes were present within the reserve (Figure 1). The reserve is also used by a small number of Red-crowned Cranes Grus japonensis as well as a few individual Eurasian Cranes G. grus (<5 annually). Red-crowned Cranes did not follow the daily movements out of the reserve to feed like the other species, and therefore were not included in the autumn crane census. However, there were probably no more than 1 5 individuals within the reserve during the autumn migration. o ~o > 73 C X! C . Downloaded on 05 September 2011. BirdLife International (2008) Grus vipio. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.1. . Downloaded on 05 September 201 1 . Ballard, B. M„ Thompson, J. E., Merindo, M. D., Ray, J. D., Roberson, J. A. & Tacha, T. C. (1 999) Demographics of the Gulf Coast subpopulation of mid-continent Sandhill Cranes. Proc. Ann. Conf. Southeast Assoc. Fish & Wildl. Agencies 53: 449-463. Brazil, M. (2009) Birds of East Asia: China, Taiwan, Korea, Japan, and Russia. Princeton: Princeton University Press. Dingle, H. (1996) Migration, the biology of life on the move. Oxford: Oxford University Press. Johnsgard, P. (1983) Cranes of the world. Bloomington: Indiana University Press. Smith, N. G. (1980) Hawk and vulture migrations in the Neotropics. Pp. SI- 65 in A. Keast & E. S. Morton, eds. Migrant birds in the neotropics. Washington, D.C.: Smithsonian Institution. Adam C. STEIN, Monteverde Institute, Monteverde, Puntarenas, Costa Rica. Email: acstein@mvinstitute.org Galina NOSACHENKO, Oksky Reserve, Ryazan, Ryazan Oblast, Russian Federation. Email: gnosachenko@rambler.ru Woolly-necked Stork Ciconia episcopus at Napahai wetland, Yunnan, China JAMES W. BURNHAM & ERIC M. WOOD The Woolly-necked Stork Ciconia episcopus is a wading bird species of Least Concern (IUCN 201 1 ). It ranges over tropical Africa, India, Sri Lanka and South-East Asia, and primarily feeds on fish, amphibians, reptiles and invertebrates. Although the population is stable throughout its range (IUCN 201 1), the Asian population potentially merits management attention owing to land-use pressures affecting suitable marsh and wetland habitat. Napahai wetland, covering c.660 km2, is located on the Zhongdian Plateau within China's Three Parallel Rivers World Natural Heritage region in north-western Yunnan province (27.879°N 99.638°E) at an elevation of 3,790 m. Located next to Xianggelila (Zhongdian), Napahai is important to a variety of migratory waterbirds. Large numbers of wintering Bar-headed Geese Anser indicus and Black-necked Crane Grus nigricollis led to its designation in 1985 as a national nature reserve; some 90% of the crane's central population are estimated to use Napahai as wintering habitat (Li & Yang 2005, Liu et at. 2010). The wetland cycles between summer high water, driven by monsoonal rain and snow melt from surrounding mountains, and winter low water when a mosaic of wetland and agricultural areas are available for wintering waterbird use. Typically, wintering waterbirds roost in the shallow waters of Napahai and forage in surrounding wetlands or agricultural fields that surround the perimeter of the wetland (Kong etal. 201 1). At 1 3h30 on 1 1 June 201 1 we detected a Woolly-necked Stork Ciconia episcopus at the north end of Napahai wetland. JWB has Forktail 28 (2012) SHORT NOTES 159 ongoing research at Napahai and at Poyang Lake and is familiar with Chinese waterbirds. We viewed the bird through 8.5x42 binoculars forfive minutes. It was loosely associating with a dispersed group of five Black Storks C. nigra and two Black-necked Cranes, and was foraging along the flooded edge of Napahai wetland with its head extended forward, close to the ground. It was a similar size to the Black Storks and had a black cap, solid white nape and neck, black plumage with purplish tones, large bill and white undertail-coverts. These are all standard field marks for this species. This observation came one day after the bird was documented by Han etal. (201 1 ). Colleagues at the Kunming Institute of Zoology (KIZ) were notified of the bird's presence and they, in turn, monitored the Woolly-necked Stork while it was at Napahai. Field reports indicate that the bird was present in the Napahai wetland from June to September of 201 1 and was last seen on 28 September 201 1 (Wu Heqi pers. comm.). The historic status of the Woolly-necked in China is unclear. Han et at. (201 1) suggest the 201 1 Napahai bird is the first record for China, but S. Chan (in litt. 2012) reports that a museum specimen at KIZ derives from an individual on display atthe Kunming Zoo in the 1960s. The origin of that bird is unknown, but transportation and political considerations at that time suggest that it is likely to have been caught in China. The 201 1 bird at Napahai thus seems likely to be the second for China. Robson (2008) lists the species as a 'rare to locally fairly common resident' in North Myanmar, adjacent to Yunnan, so it is perhaps surprising that there have not been more prior records. Future observers at Napahai and other wetlands in Yunnan should be aware of the potential for the presence of the Woolly-necked Stork and other species that occur in neighbouring countries but are not regularly recorded in China. Acknowledgements This sighting would not have been possible without support by the National Science Foundation under Grant No. DGE-0549369 IGERT: Training Program on Biodiversity Conservation and Sustainable Development in Southwest China atthe University of Wisconsin-Madison. Dr. Wu Heqi at KIZ was helpful in providing information regarding the stork's duration of stay at Napahai. Simba Chan kindly indicated the existence of the specimen in KIZ. References Han L.-X., Han B„ Den Z.-W., Yu H.-Z. & Zhao J.-L. (2011) Wooly-necked stork, a new bird record of China. Zoological Research 32: 575-576. (In Chinese with English abstract.) IUCN (2011) IUCN Red List of threatened species. Version 2011.2. . Downloaded on 1 3 January 201 2. Kong D.-J., Yang X.-J., Liu Q., Zhong X.-Y. & Yang J.-X. (201 1 ) Winter habitat selection by the vulnerable Black-necked Crane Grus nigricollis in Yunnan, China: implications for determining effective conservation actions. Oryx 45: 258-264. Li F. & Yang F. (2005) [Distribution and population of Black-necked Cranes on the Yunnan-Guizhou Plateau.] Pp. 29-43 in Wang Qi-Shan and Li Fengshan, eds. Crane research in China. Kunming: Yunnan Nationalities Publishing House. (In Chinese.) Liu Q, Yang J.-X., Yang X.-J., Zhao J.-L. & Yu H.-Z. (2010) Foraging habitats and utilization distributions of Black-necked Cranes wintering at the Napahai Wetland, China. J. Field Orn. 81 : 21-30. Robson, C. (2008) A field guide to the birds of South-East Asia. London: New Holland. James 1/1/. BURNHAM, Department of Forest and Wildlife Ecology, University of Wisconsin-Madison, 1630 Linden Drive, Madison, Wisconsin 53706, USA; and International Crane Foundation, E-l 1376 Shady Lane Road, P.O. Box 447, Bara boo, Wisconsin 53913, USA. Email: burnham@wisc.edu Eric M. WOOD, Department of Forest and Wildlife Ecology, University of Wisconsin-Madison, 1 630 Linden Drive, Madison, Wisconsin 53706, USA. Email: emwood@wisc.edu First records of Javan Munia Lonchura leucogastroides in Peninsular Malaysia TIM ROBINSON The Javan Munia Lonchura leucogastroides isanestrildidfinch native to the Indonesian Islands of Java, Bali and Lombok. It is also established in southern Sumatra, where it was probably introduced (MacKinnon & Phillips 1993, Wells 2009), and in Singapore where it was introduced around 1922. By the 1980s it had become the commonest estrildid finch in Singapore (Lever 1 987) but its numbers have declined in recent years (Wells 2009), and the species is now considered uncommon there. Despite the narrowness (approximately 1 km) of the strait separating Singapore from Malaysia's southern state of Johor, the only published record of this species in Peninsular Malaysia is of a single bird in a Johor Bahru park in April 2003 that was considered to be an escapee (mentioned in Wells 2009). This paper discusses recent observations of Javan Munias in Johor that suggest that the species is now a breeding resident there. At 09h00 on 21 July 201 1 , three adult and two immature Javan Munias were observed at Sungai Tiram, Johor (1°37'03"N 1 03°47'29"E), where they were seen perching on and moving in and out of tall (>1 m) grass stems. The white bellies of the birds drew my attention, as did the black throat and vent on the adult birds. Their rump and tail were dark in colour, distinguishing them from White-bellied Munia L. leucogastra, which has an olive-yellow tail fringe. These birds were seen again in the same location on 22 July 2011 at 07h50, where they were observed for over 1 0 minutes at a distance of 7 m. When disturbed, they would move a short distance and resume foraging amongst the grass stems. These birds were present at the same site on 1 1 August and 6 October 201 1 , by which time the two juvenile birds had developed adult plumage. Following discussions with the local birding community, Paul Wu, a member of the Malaysian Nature Society, provided photographs of this species that he had taken in April and May 2009 at a Temple near Kulai, some 30 km west of the Sungai Tiram site. He estimated there were three to four pairs of birds nesting in potted plants around the temple, and had photographed one bird carrying nesting material. In some Asian countries, estrildid finches are traded as prayer birds; this often involves transfer of birds from their point of capture to another location where they are released into the wild as part of religious rituals (Severinghaus & Chi 1999). In Malaysia, estrildid finches are the most commonly traded prayer birds, and are both imported from Indonesia (Shepherd 2006) and netted from native stocks for trade at local bird markets. On 7 October 201 1, during a search of the local bird market near the Sungai Tiram site, I found three captive Javan Munias for sale, together with 62 other estrildid finches. The trader stated that the birds had been trapped on the eastern reaches of Sungai Tiram, and that his shop had been selling Javan Munias irregularly for at least ten years. Following this development, between 7 and 1 6 October 201 1 , all bird markets that could be located in Johor south of 2°N were surveyed (n=10). During this survey, an additional five Javan Munias were discovered for sale 160 SHORT NOTES Forktail 28 (2012) Serial Date Location Notes 1 Apr 2003 Johor Bahru 1 bird identified in a city park of Johor Bahru, considered to be an aviary escapee or deliberately released (mentioned in Wells 2009) 2 Apr-May 2009 Kulai 3-4 pairs of wild birds observed carrying nesting material at Seng Kang Temple, Kulai. One pair photographed 3 Jul-Oct 2011 Sungai Tiram 5 wild birds, two with juvenile plumage, observed and photographed in same location regularly Jul-Oct 201 1 4 Oct 201 1 Ulu Tiram 3 captive birds for sale, trapped on eastern reaches of Sungai Tiram (8) 5 Oct 2011 Kulai 5 captive birds for sale, trapping location unknown 6 Dec 2011 PULADA Army Camp 2 wild birds observed 7 Dec 2011 Ulu Tiram 19 captive birds for sale, trapped on eastern reaches of Sungai Tiram (8) Total birds = 41 Figure 1. Records of Javan Munia in Peninsular Malaysia. at the Kulai bird market. The 2,021 captive estrildid finches found in Johor's bird markets during this survey comprised 1,070 Scaly- breasted Munias L. punctulata (52.9%), 601 Black-headed Munias L atricapilla (29.7%), 342 White-headed Munias L. maja (16.9%) and 8 Javan Munias (0.4%). I made two further observations of Javan Munias during December 201 1. One was at the PULADA Army Camp (1°37'15"N 103°49'55''E), where two birds were observed flying into and then sheltering insidean ornamental shrub.Theotherwasatthe UluTiram bird market where an additional 19 captive birds were on sale; these were also reportedly captured on the eastern reaches of Sungai Tiram.The location of these and previous observations are given in Figure 1 . Given the number of both wild and captive birds recorded, the photograph of a bird carrying nesting material, and the presence of juvenile birds in the wild, it would appear that the Javan Munia is now established as a breeding resident on the northern rural fringe of Johor Bahru. Anecdotal evidence from traders supports this, suggesting that, as predicted by Wells (2009), the species has been present in small numbers in Johor for a number of years. It is difficult to say with certainty whether Johor's Javan Munias have become established entirely through natural immigration of wild birds from Singapore, or whether the population originated from the release of prayer birds, imported from neighbouring countries such as Singapore or Indonesia. Possibly, combinations of these factors have occurred, with released prayer birds augmenting a small naturally immigrated population. Given the proximity of Johor to Singapore, and that Javan Munia has not yet been recorded in other urban areas of Malaysia where prayer birds are used, it is more probable that the Johor population has originated from Singapore rather than Indonesia. Currently, the most northern record of Javan Munia in Johor is approximately 50 km from the Singapore Central Business District, implying that this species has extended its range a similar distance in the 90 years of its presence in Singapore. Based on this precedent, it is probable that, unassisted, this species will be slow to disperse further north. My survey of bird markets in October 2011 established that Javan Munia comprised less than 1% of the captive estrildid finches sold in Johor's prayer bird trade. As there is no price difference between species of estrildid finch sold, it is unlikely that Javan Munia is specifically targeted for capture. However, the seemingly low number of individuals and localised occurrence of Javan Munia in Johor makes the species vulnerable. It is likely that trapping for the prayer bird trade is disrupting the formation of a stable population base in the SungaiTiram and Kulai local areas. Because captured Javan Munia make up only a minority of estrildid finches for sale, and are generally sold in mixed-species groups, their release may isolate individual birds, and will not necessarily facilitate the colonisation of new areas. If in the future Johor's Javan Munia population increases in abundance as Singapore's did in the 1 980s, the proportion of captive Javan Munias being traded will probably increase, and the liberation of prayer birds into suitable habitats such as gardens and temples could enable a faster dispersion of this species across the state than would be expected if natural processes only were involved. There is potential for competition to occur with other Lonchura native to Malaysia, perhaps most interestingly with White-bellied Munia, which unlike the Malaysia's other Lonchura, is not found at the Peninsula's southernmost tip, and did not experience range overlap with Javan Munia in Singapore. The status and distribution of both Javan and White- bellied Munia in Malaysia should be closely monitored in the future. Acknowledgements David Bakewell, Paul Wu and Yong Ding Li were all generous in sharing their knowledge and experiences with this species, and I am also thankful to Mike Connor, David Pauli and two anonymous referees for their helpful comments on an earlier draft of this manuscript. References Hails, C. & Jarvis, F. (1987) Birds of Singapore. Singapore: Times editions. Lever, C. (1987) Naturalised birds of the world. New York: Longman Scientific and Technical. Mackinnon, J.& Phillips, K. (1993) A field guide to the birds of Borneo, Sumatra, Java and Bali. United Kingdom: Oxford University Press. Restall, R. (1996) Munias and mannikins. Sussex: Pica Press. Shepherd, C. (2006) The bird trade in Medan, north Sumatra: an overview. Bir ding ASIA 5: 16-24. Severinghaus, L. & Chi, L. (1999) Prayer animal release in Taiwan. Biol. Conserv. 89: 301-304. Wells, D. (2009) The birds of the Thai-Malay Peninsula, 2: passerines. London: Academic Press. Tim ROBINSON, 10/165 Blarney Cres, Campbell ACT 2612 Australia. Email: tim.robinson58@gmail.com Forktail 28 (2012) SHORT NOTES 161 Brahminy Kites Haliastur indus fishing with Irrawaddy dolphins Orcaella brevirostris in the Mekong River GERARD E. RYAN Although the Brahminy Kite Haliastur indus is a widespread and often relatively common species in coastal areas and near rivers (Bird Life International 2009), relatively little attention has been paid to the species's feeding ecology (Iqbal etal. 2009). Brahminy Kites are reported as catholic feeders with a diet offish and opportunistic take of other small vertebrates, and often carrion (e.g. Irby 1861, Strange 2002, Robson 2005, Davidson 2009). Few studies, however, report specific details of the species's diet, and much understanding of the species's foraging strategy appears to be based on diffuse casual observation by many observers throughout its range. Fishing in association with dolphins is well known for diving seabirds (e.g. Evans 1982, Ballance et al. 1997). Generally this involves birds taking advantage of groups of dolphins herding large schools offish within diving range, nearthe surface. In rivers, such association between birds and dolphins appears to be much rarer. There are, to the best of my knowledge, no records of birds fishing with freshwater dolphins, or of accipitrids fishing with dolphins. Here I report on incidental observations of Brahminy Kites fishing in association with Irrawaddy dolphins Orcaella brevirostris in the Mekong River, in north-eastern Cambodia. Methods Observations of kites feeding around dolphins were made opportunistically on surveys for Irrawaddy dolphins on the Mekong River from 2009 to 201 2. Dolphin surveys covered the area between Kratie town to the Lao PDR border and back, a direct linear distance of 1 90 km north-south (Ryan etal. 2011). Within the study area the river morphology consists of complex braided channel mosaic with a wide variety of habitats ranging from shallow rocky rapids to deep pools that provide important habitat for fish and dolphins (Poulsen etal. 2002). High levels of avian and biological diversity are reported in the area, which includes a Ramsar site in the northern end; Try & Chambers (2006) and Bezuijen et al. (2008) provide extensive overviews of the northern and southern halves of the study area, in which Brahminy Kites are commonly observed. Surveys consisted of slow travel in a traditional-style wooden boat, traversing the river for signs of dolphins (details in Ryan etal. 201 1).The observations reported are from seven surveys between March 2009 and February 2012, each taking around ten days. Irrawaddy dolphins can move throughout most of the main channel, but congregate around deep pools in the dry season. Consequently, a large proportion of observation time is spent around deep pool areas. Attempts were made to take careful notes of the birds' behaviour at such times, but my priority activity of photographing dolphins limited the extent of my attention. Results Three observations of Brahminy Kites fishing in association with Irrawaddy dolphins were made: in April 2009, March 2010 and February 201 2. All of these observations were of a single bird, and occurred at Kampi pool, the southernmost end of the dolphin's range, around 1 4 km north of Kratie town, in Kratie province. In 2009 the kite spent around 20-30 minutes following a group of 4-6 feeding dolphins. The bird circled slowly around the area where the dolphins fed and swooped down to the water surface at the same time as, and very near to where, the dolphins surfaced. At least four such dives were observed, two of which broke the water surface but neither of which was successful in catching fish. The bird eventually moved on for no apparent reason, while the dolphins continued to fish in the same area. The 2010 observations were similar although less detail was recorded. The kite appeared to fish around a small group of dolphins, following them around the local area and swooping at least three times near surfacing dolphins. Again, no successful swoops were observed and the encounter is thought to have lasted around 10-15 minutes. In 201 2 the kite was observed to follow a group of socialising and feeding dolphins for around 50 minutes. The mammals were in a deep area, and large numbers of small fish, believed to be a common cyprinid of the genus Henicorhynchus, known as trey riel in Cambodian, intermittently broke the surface. The bird was observed to swoop abortively, without touching the surface, at least six times, and broke the water surface at least four times over this period, two of which were very near surfacing dolphins. The bird disappeared around the time the dolphins moved to feed in another area, suggesting the fish being targeted by both species had moved on. Discussion These observations are potentially the first record of a Brahminy Kite using dolphins to find prey, and at the least a rare report of an accipitrid fishing with dolphins. The scarcity of such observations, compared to the hundreds of hours of observation I made of dolphins throughout this period, suggests that the mode of feeding is not a common one. Indeed, that all observations occurred at a similar time of year and at the same location suggests it may even be the same individual observed each time. In fact, given the concentrations of both dolphins and fish in deep pools throughout the dry season (Poulsen et al. 2002), piscivorous birds would sometimes occur in close proximity to Irrawaddy dolphins in the Mekong simply by chance. For example, in January 2012 an Osprey Pandion haliaetus was also observed slowly circling a group of around six dolphins at Anlung Cheuteal, a deep pool on the border of Cambodia and Lao People's Democratic Republic. The bird appeared to follow the dolphins for around five minutes without swooping, before continuing across the pool searching. Observations of other avian species feeding independently of, but near to, dolphins in areas of high prey density have been recorded elsewhere (Acevedo 1 991 ). This is probably the case regarding the Osprey observation. However, that the Brahminy Kites were observed to begin to dive just before dolphins appeared at the surface strongly suggests an association in theirforaging strategy, probably related to dolphins chasing fish toward the surface. This pattern fits similar observations where White-fronted Terns Sterna striata were observed to follow single dolphins and feed around them (Brager 1998). As Brahminy Kites are reported to use a wide variety of food resources, it is unsurprising that they may take this opportunity when it arises, and thus this behaviour is likely to be some form of facultative commensalism. It would be of interest to understand better such interactions between birds and riverine dolphins, including the extent and frequency of feeding associations, and the species that may be involved. Better knowledge of these unusual events could also be used as an ecotourism drawcard to support the conservation of the species in question. Acknowledgements These observations were recorded during dolphin surveys conducted with the generous support of private donors through WWF-Switzerland and WWF-Germany, and the permission of the Cambodian Fisheries Administration. GERalso wishes to thank the Cambodian Mekong Dolphin 162 SHORT NOTES Forktail 28 (2012) Conservation Project field team, as well as two anonymous reviewers for their helpful comments. References Acevedo, A. (1991) Behaviour and movements of bottlenose dolphins, Tursiops truncatus in the entrance to the Ensenada De La Paz, Mexico. Aquatic Mammals 17: 137-147. Ballance, L. T„ Pitman, R. L. & Reilly, S. B. (1997) Seabird community structure along a productivity gradient: importance of competition and energetic constraint. Ecology 78: 1 502-1518. Bezuijen M. R., Timmins, R. & Seng, T., eds. (2008) Biological surveys of the Mekong River between Kratie and Stung Treng Towns, northeast Cambodia, 2006-2007. WWF Greater Mekong - Cambodia Country Programme, Cambodia Fisheries Administration and Cambodia Forestry Administration, Phnom Penh. Bird Life International (2009) Haliastur indus. In: IUCN 201 1 . IUCN Red List of Threatened Species. Version 2011.1. . Downloaded on 1 7 September 2011. Brager, S. (1998) Feeding associations between White-fronted Terns and Hector's dolphins in New Zealand. Condor 100: 560-562. Davidson, P. (2009) A photographic guide to the birds of Vietnam, Cambodia and Laos. London: New Holland. Evans, P. G. H. (1982) Associations between seabirds and cetaceans: a review. Mammal Review 1 2(4): 1 87-206. Iqbal, M., Mulyono, H.,Takari, F. & Anwar, K. (2009) Aerial feeding on a large prey item by a Brahminy Kite Haliastur indus. Australian Field Orn. 26: 33-35. Irby, H. L. (1861) Notes on birds observed on Oudh and Kumaon. Ibis 3: 217-251. Poulsen, A., Ouch Poeu, Sintavong Viravong, Ubolratana Suntornratana & Nguyen Thanh Tung (2002). Deep pools as dry season fish habitats in the Mekong Basin. MRC Technical Paper No. 4, Mekong River Commission, Phnom Penh. 22 pp. ISSN: 1683-1489. Strange, M. (2002) A photographic guide to the birds of Southeast Asia: including the Philippines and Borneo. Periplus Editions, Hong Kong. Robson, C. (2005) The New Holland field guide to the birds of South-East Asia. London: New Holland. Ryan, G. E., Dove, V., Trujillo, F. & Doherty, P. F. (201 1) Irrawaddy dolphin demography in the Mekong River: an application of mark-resight models. Ecosphere 2: art58 Try, T. & Chambers, M. (2006) Situation analysis: Stung Treng Province, Cambodia. Mekong Wetlands Biodiversity Conservation and Sustainable Use Programme, Vientiane, Lao PDR. 93 pp. Gerard E. RYAN, WWF-Cambodia, #21, Street 322, Beoung Keng Kang I, Khan Chamcar Morn, Phnom Penh, Cambodia. Email: gryan@wwf.panda.org New information on the distribution of White-fronted Microhierax latifrons and Black-thighed Falconets M. fringillarius in Kalimantan, Indonesia MOHAMMAD IRHAM, E. MEIJAARD & S. (BAS) VAN BALEN The White-fronted Falconet Microhierax latifrons is well-known as the near-endemic falconet in Sabah (Ferguson-Lees & Christie 2001 , Sheldon et al. 2001, Mann 2008). Although it is a forest-edge species, it is listed as Near Threatened owing to its restricted range and reliance on lowland forests coupled with the likely decline and degradation of these habitats (BirdLife International 201 2). To the west and south of its range, itscongenerthe Black-thighed Falconet M. fringillarius occupies the remaining, larger part of the island of Borneo. During a biodiversity survey in East Kalimantan (Indonesian territory), on 1 June 201 1 , at 08h1 5 to08h20, Ml saw two individuals of male White-fronted Falconet in the Simenggaris area of Nunukan Regency, East Kalimantan Province, Indonesia (4°27'N 117°16'E; Figure 1 ). This site is located c.20 km south by the main road to the border between Indonesia and Sabah (Malaysia). The falconets had white forecrowns extending to above the rear end of their eyes, plain black upperparts, underparts white from throat to breast, tawny-coloured belly, black flanks and undertail (Plate 1). They were easily distinguished from Black-thighed Falconet by an unbarred blackish undertail, no curved white line behind the eyes and a tawny rather than rufous abdomen. When first spotted, the falconets were perched on the top of a c.20 m tall snag along a logging road through secondary forest. One bird was eating a cicada while the other watched for flying prey insects (Plate Figure 1. Localities of records of White-fronted Falconet Microhierax latifrons (dots) and Black-thighed Falconet M. fringillarius (triangle). Forktail 28 (2012) SHORT NOTES 163 Plate 1 . Two male White-fronted Falconets Microhierax latifrons, 1 June 201 1 , Nunukan, East Kalimantan (M. Irham). 1). Several hunting manoeuvres were observed in which insects were caught in the air with a loop flight and consumed in c.2 minutes after returning to the initial posts. The present record confirms an earlier tentative one of the species in Kalimantan: on 24 June 2005, EM and M. Leighton saw a group of five falconets c.40 km south from the Sabah border, north of the Sebuku River, in Sebakis (c.4°05'N 1 1 7° 1 0'E), the northernmost forest block within PT Adindo's concession area (Nunukan district, East Kalimantan Province; Figure 1). The birds were hunting for insects towards dusk from the top of a tall, leafless tree in logged-over forest. Although the birds were quite well seen (at 50 m using 7x40 binoculars), and a large forehead-patch, black eye-stripe and yellowish rather than rusty wash on the lower breast identified them as White-fronted Falconets, this record was treated as tentative because of the poor light conditions. The distribution of the White-fronted Falconet was traditionally defined as nearly exclusively Sabah territory, south to the Lawas River (west coast, Sarawak) and Darvel Bay (east coast) (Smythies 1999, Mann 2008). Previous records of the falconet nearest to the present area include Kinabatangan, Tiger Estate and Tawau (Thompson 1 966, Sheldon etal. 2001 ; Figure 1 ), at 50-80 km north¬ east to east of Simenggaris. The records above redefine the ranges of both species of falconet on Borneo, where a record of a Black-thighed Falconet seen on 15 November 1997 by SvB (O'Brien 1997) near Paking village (c.3°30'N 116°30'E, Malinau district. East Kalimantan; Figure 1), is the northernmost for the species in its eastern range. This suggests that the two species may occur sympatrically in the transition zone between their respective ranges. In the area between the rivers Sebuku and Sesayap no falconets have to date been recorded and it is therefore unclear which of the two rivers may act as an effective natural boundary, or even what biological or physical mechanism separates the two species. The ranges of the two Bornean falconet species reflect those of White-crowned Shama Copsychus stricklandii and Black-and-crimson Pitta Pitta ussheri, which are also endemic to lowlands in the north-east part of the island and have sister taxa on the rest of Borneo (Moyle et at. 2005). We note, however, that for the former species Collar (2004) suggested that based on extensive hybridisation and lack of consistent morphological differences the two Copsychus species should be lumped (see also Lim etal. 2010). Additional records of Microhierax from the area of potential range overlap and study of museum specimens are needed to clarify whether hybrids occur, and what possible mechanisms separate the two species. Acknowledgements The biodiversity survey in 201 1 was fully funded by DIPA 201 1 Research Center for Biology and Ml would like to thank PT. Adimitra Lestari for accommodating the field survey in Simenggaris. SvB extends his thanks to the Wildlife Conservation Society-lndonesia Programme (Bogor), and Center for International Forestry Research (Bogor) for providing the opportunity to visit the Tubu uplands. References Anonymous (2005) High conservation value forest report for PT Adindo Hutan Lestari, East Kalimantan. The Nature Conservancy. BirdLife International (2012) Species factsheet: Microhierax latifrons. Downloaded from http://www.birdlife.org on 30/01/2012. Collar, N. J. (2004) Species limits in some Indonesian thrushes. Forktail 20: 71-87. Ferguson-Lees, J. & Christie, D. A. (2001) Raptors of the world. London: Christopher Helm. Lim, H. C., Zou, F„ Taylor, S. S., Marks, B. D., Moyle, R. G., Voelker, G. & Sheldon, F. H. (2010) Phylogeny of magpie-robins and shamas (Aves: Turdidae: Copsychus and Trichixos ): implications for island biogeography in Southeast Asia. J. Biogeogr. 37: 1 894-1906. Mann, C. F. (2008) The birds of Borneo: an annotated check-list. London: British Ornithologists' Union (Check-list No. 23). Moyle, R. G., Schilthuizen, M., Rahman, M. A. & Sheldon, F. H. (2005) Molecular phylogenetic analysis of the white-crowned forktail Enicurus leschenaulti in Borneo. J. Avian Biol. 36: 96-1 01 . O'Brien, T. G., ed. (1997) Bulungan Biodiversity Survey, 5 November 1997 to 5 December 1997. Bogor: WCS-Indonesia Programme & CIFOR. Sheldon, F. H., Moyle, R. G. & Kennard, J. (2001) Ornithology of Sabah: history, gazetteer, annotated checklist, and bibliography. Orn. Monogr. 52: 1-285. Smythies, B. E. (1999) The birds of Borneo. 4th edn (Davison, G.W.H (ed.)) Kota Kinabalu, Sabah, Malaysia: Natural Hstory Publications and Sabah Society & the Malayan Nature Society. Thompson, M. C. (1966) Birds from North Borneo. Univ. Kansas Pub!. Mus. Nat. Hist. 1 7: 377-433. Mohammad IRHAM, Museum Zoologicum Bogoriense, Research Center for Biology-lndonesian Instititute of Sciences, Jl. Raya Jakarta-Bogor KM. 46 Cibinong Indonesia 16911. Email: irham.mzb@gmail.com Erik MEIJAARD, People and Nature Consulting International, Ciputat, Jakarta, 15412, Indonesia; and School of Biological Sciences, University of Queensland, Brisbane, QLD 4072, Australia. Email: emeijaard@gmail.com S. (Bas) van BALEN, Basilornis Consults, Roompotstraat 44, 6826 EP Arnhem, The Netherlands. Email: bvanbalen001@hotmail.com 164 SHORT NOTES Forktail 28 (2012) Breeding ecology of the Northern Boobook Ninoxjaponica totogo in central Taiwan WEN-LOUNG UN, SI-MIN UN & HUI-YUN TSENG Introduction The Brown Hawk Owl complex Ninox scutulata, also known as the Brown Boobook (King 2002), is widely distributed in East Asia, from south-eastern Siberia, North Korea, Japan, Taiwan and the Philippines to the Moluccas and the Lesser Sundas (del Hoyo etal. 1999, Konig etal. 1999). Such a wide range has produced phenotypic variation between populations, including in size, morphs and migration status. Recent authors have recognised 1 1 (del Hoyo etal. 1999) or 13 (Konig etal. 1999) subspecies, but King (2002) analysed sonagrams of territorial songsfrom 1 1 subspecies and separated the entire complex into three distinct species: 'Brown Boobook' N. scutulata, 'Chocolate Boobook' N. randl and 'Northern Boobook' N. japonica. Accordingly, the taxonomic designation of this owl in the Far East is N. japonica. This species consists of two subspecies: N.j. japonica, a summer visitor in northern regions such as Japan and Korea, and N.j. totogo, a resident of the southern regions. King (2002) also indicated the extreme paucity of research on the entire species complex. The type locality of N.j. totogo is Lanyu, a tiny offshore islet located 50 km south-east of Taiwan. Although a breeding population of N.j. totogo on this islet has been known for many years (Severinghaus 2007), the resident status of N. japonica in Taiwan has been the topic of long and lasting debate. The species in Taiwan was originally treated as a migrant or transient (Hachisuka & Udagawa 1951, Wang etal. 1991), whereas later literature considered it a resident because it was found to occur throughout the year (Mees 1970, Dickinson 2003). Recent literature has proposed the occurrence of both resident and migratory populations in Taiwan (Brazil 2009, Severinghaus etal. 2010). However, breeding records of the species in Taiwan were not published until 2010, when Tseng & Lin (2010) and Severinghaus et al. (2010) separately reported partial information from their occasional observations on the breeding behaviour of this owl. Nevertheless, breeding by this owl in East Asia has never been documented in detail. In this study, we provide: (1 ) robust evidence for the occurrence of a breeding population in Taiwan (N.j. totogo ); (2) a first quantitative report on the breeding ecology of this species; (3) basic morphometries of these breeding individuals; and (4) a comparison of ecological and morphological differences between the summer visitors (N. j. japonica) and residents (N.j. totogo) in Far East Asia. Methods Study area and nest-site search The study area is located in central-western Taiwan, within latitudes 23°55'N and 24°03'N and longitudes 120°44'E and 120°53'E. The majority of the landscape in this region is subtropical hardwood forest at 70-700 m elevation, with a mosaic of villages and cultivated lands. The Northern Boobook primarily inhabits densely vegetated woodland, and its tree-cavity nest-sites are very difficult to locate. From 1999 to 2009, every suspected or confirmed pair was intensively followed in the period 1 January to 30 April, a time when the pairs are very active and the frequency of breeding calls and copulation could be recorded by directly approaching the pairs using spotlights. The nest holes were found by tracing the routes of male and female owls and by locating the source of calls. The route-tracing method was based on the observation that if one individual (usually the female) disappeared from sight, this could indicate preparation for egg-laying involving staying near or in the nest-site. The nest-site could then be found by tracing the routes — the direction in which the owls flew after sunset and before sunrise — taken by male and female owls at the time of pair- formation. The call-location method was based on the observation that the owls began calling at sunset, usually initially near the nest- site. We located the first calls of the owls and approached their probable location. This method provided excellent opportunities to find nests. Landscape structure of the nesting site and breeding phenology After the nest was found, we identified the tree to species and measured its diameter at breast height (DBH). A circle with a radius of 500 m, which was considered suitable for the territorial extent of a small owl (e.g. Pande etal. 2007, Santhanakrishnan etal. 2011), was plotted around the nest-site. The landscape composition (as a percentage of the total area) within this region was estimated from an aerial photograph. The following categories of landscapes were identified: primary forest (also termed original forest) (PF), secondary forest (SF), artificial forest (MF), orchards (OC), betel nut palms (BP), trees along streets (ST), roads (RD), buildings (BD) and urban parks (UP). Multiple direct visits to the nests were conducted to confirm the clutch size and brood size for every breeding attempt. A video camera attached on an elongated aluminium pole with mobile monitor was used to record the subsequent stages. We marked the egg(s) to estimate the incubation period in each nest. When the nestlings were approaching fledging, we visited the nest-site every day to confirm fledging date and number of successfully fledged nestlings. A nest that produced at least one fledgling was recorded as successful. We observed and recorded the activities of young and parents every 3-7 days until they dispersed from the nesting area. Morphometries Morphological measurements of the adults were taken during the incubation or early nestling periods. Adults were directly caught by hand or hand-net when inside the nest holes. The length of bill (from tip to cere), head (from bill-tip to end of skull), wing chord, tail and tarsus were measured to the nearest 0.1 mm with digital calipers, and the body mass to 0.1 g with an electronic scale. The individuals were banded with an aluminium ring and a coloured ring for individual identification. Males and females were identified by observing their mating, guarding and incubating behaviours (males do not incubate). All measurements were compared between males and females with a nonparametric Mann-Whitney U test. Results Nest-sites and nest-trees Ten nests of the Northern Boobook were recorded during the study period, with no nesting site being re-used. Eight were located in a relatively natural environment, at altitudes of 350-700 m.The other two nests were found in suburbs, both at 70 m elevation. The landscape structure around the rural nests was dominated by secondary forest (range 35-95%, mean 75%), followed by artificial forest (range 5-25%, mean 13%). In contrast, the suburban nests were surrounded by a landscape including 57-58% of man-made structures (including roads and buildings) (Table 1). Six species of nest-tree were identified (Table 1). Forktail 28 (2012) SHORT NOTES 165 Table 1. Data on ten nests of Taiwanese Northern Boobooks: year, location, elevation, nest-tree species, nest-tree diameter at breast height (DBH), and landscape composition. For landscape composition abbreviations see Methods. No. Year Location Elevation(m) Nest tree DBH (cm) Landscape composition (radius 500 m ) (%) PF SF MF BP 0C ST UP BD RD 1 1999 Dakeng 380 Machiluszuihoensis 50 95 5 2 2000 Yuchi 365 Cinnamomum camphora 48 60 35 5 3 2002 Yuchi 700 Helicia formosana 55 48 25 25 2 4 2004 Wufeng 375 Helicia formosana 70 88 12 5 2005 Wufeng 350 Cinnamomum camphora 50 80 15 5 6 2006 Wufeng 375 Machiluszuihoensis 62 80 15 5 7 2006 Wufeng 415 Aluerites fordii 50 90 10 8 2006 Wufeng 400 Aluerites fordii 45 85 15 9 2008 Nantou city 70 Eucalyptus robusta 77 20 22 33 25 10 2009 Nantou city 70 Melaleuca leucadendra 60 21 22 33 24 Behaviour prior to egg-laying Male Northern Boobooks began calling and occupying territories from January to February. The pair-bond formed from mid-February to early March. During this period the male and female were very vocal and usually moved abouttogether. Copulation was observed at this stage. The frequencies of calling and copulation showed the same pattern, which increased before and decreased after egg- laying. Both behaviours reached their peak at around 1 1 March. However, copulatory behaviour was also observed at the incubation stage (Figure 1). 10 o Copulation • Calling o* # V* •• •• • • » O 0*> °0 „ • • V r • % ® o o O 00 o o Qd*o ° • o • • StocP \ -\. — «N«»* 10 10 20 30 40 50 60 70 80 90 100 110 120 Day of year Figure 1. Frequency of calling and copulation in Taiwanese Northern Boobook during the breeding season. Data combined from the ten nests during the ten years. Breeding phenology Eggs were laid from 1 6 March through April, and the mean first egg- laying date was 28 March ± 8.8 days. Mean clutch size was 3.2 ± 0.4 (range 3-4). Mean incubation period was 26.8 ± 1 .8 days (range 25- SI ), and the eggs were incubated only by females. The nestlings hatched between 12 April and 7 May, and the hatching rate and brood size were 80.8 ± 22.9% (33-100%) and 2.6 ± 0.8 (1-4 chicks) respectively. Mean nestling period for successful nests (n=9) was 25.9± 1.1 days (range 25-28). Mean number of fledglings in all nests was 2.1 ± 1 .0, and the fledgling success rate was 86.7 ± 32.2% (Table 2). In the unsuccessful nest, four young drowned during the nestling period due to heavy rain. Most fledglings appeared in May, and the post-fledging period extended from mid-July through to late August. Morphology of adults, eggs and nestlings Ten pairs of adults were measured during the breeding period. No significant differences were found between adult males and females (Table 3). The eggs were 20.2 ±0.6x1 8.2 ± 0.4 mm, with an average mass of 20.4 ± 1 .2 g (n=13). Mean body mass of nestlings at ages 1 , 5, 1 0, 1 5 and 20 days was, respectively, 1 6.8 ± 1 .0 g (n=6), 41 .7 ± 4.9 g (n=3), 1 00.6 ± 1 1 .6 g (n=5), 1 1 8.3 ± 7.4 g (n=3) and 1 43.6 ± 1 1 .6 g (n=5) (Figure 2). The growth equation between ages 1 and 20 days was y = 6.8602x + 14.232 (r2= 0.96), where y was the body mass and x was the age in days (Figure 2). Discussion This study is one of the very few attempts to quantify some basic parameters in the breeding ecology of birds in the Ninox scutulata complex. The only previous work has been on the Japanese Table 2. First egg-laying day (number = day of year) (FD), clutch size (CS), incubation period (IP), brood size (BS), hatching rate (HR), number of fledglings (NF), fledgling success rate (FSR), and nestling period (NP) for the ten Ninox nests in central Taiwan. The incubation period was recorded from one or two eggs. In all, 1 7 eggs were used to estimate the mean incubation period. No. FD CS IP (days) BS HR (%) NF FSR (%) NP(days) 1 85 (26 Mar) 3 25 3 100 3 100 25 2 88 (29 Mar) 4 27,31 3 75 3 100 26 3 95 (5 Apr) 3 25,26 2 67 2 100 25 4 94 (4 Apr) 3 27,28 3 100 3 100 28 5 80(21 Mar) 3 26 3 100 2 67 27 6 75 (16 Mar) 4 28,31 4 100 0 0 - 7 80 (21 Mar) 3 25 1 33 1 100 25 8 77 (18 Mar) 3 25,26 3 100 3 100 26 9 95 (5 Apr) 3 26,27 2 67 2 100 26 10 100 (10 Apr) 3 26,27 2 67 2 100 25 Mean±SE 86.9±8.8 3.2±0.4 26.8±1 .8 2.6±0.8 80.8+22.9 2.1±1.0 86.7±32.2 25.9±1.1 166 SHORT NOTES Forktail 28 (2012) Table 3. Morphological data for breeding adult Taiwanese Northern Boobooks. M-W = Mann-Whitney. Male(m Morphological character mean =10) SE Female(n=10) mean SE M-W U test P Bill (mm) 13.1 1.0 13.6 1.4 41.5 0.253 All head (mm) 51.1 3.4 49.8 2.9 39 0.207 Wing chord (mm) 217.2 6.5 214.0 20.1 47.5 0.412 Tail (mm) 119.4 10.1 118.8 5.8 40.5 0.229 Tarsus (mm) 33.9 2.6 33.3 2.8 44.5 0.468 Body mass (g) 167.5 22.4 168.2 27.5 49 0.470 Wing/tail 1.83 0.16 1.79 0.13 39 0.207 180 160 „ 140 - 120 on I 100 S 80 o cn 60 40 20 0 y = 6.8602x+ 14.232 r* = 0.9608 0 5 10 15 20 25 Age (day) Figure 2. Mean body mass of nestling Taiwanese Northern Boobooks at 1 , 5, 1 0, 1 5 and 20 days of age. population of N.j.japonica, for which breeding data were briefly presented by Masuda (1974) and Oba (1996). A notable difference between the Taiwanese and Japanese populations is the marked difference in the timing of their breeding seasons. Whereas in Taiwan territory occupation and pair-formation begins in mid- February and mean first-egg date is 28 March, breeders in Japan do not arrive until April (Brazil & Yabuuchi 1991, Oba 1996), from which we deduce that the mean first egg-laying date cannot be earlier than 30 April. Moreover, the approximate length of the entire breeding season in Japan is only four months, a much shorter period than the seven months we have observed in Taiwan. These differences raise the issue of the degree of morphological differentiation between migratory and resident birds in South-East Asia. A comparison of our morphometric data on N.s. totogo with the measurements of N. s.japonica in King (2002) shows that the Taiwanese breeders have a slightly shorter wing chord (21 4.0 mm for males and 21 7.2 mm for females vs 220.8 mm for N. s.japonica) and slightly longer tail (1 1 9.4 mm for males and 118.8 mm for females vs 115.8 mm for N. s. japonica), a result consistent with the tendency of migratory birds to have a higher wing/tail ratio to facilitate their over-water dispersal (Averill 1 920, Yong & Moore 1994). This suggests that the migratory and resident birds in Taiwan and South-East Asia might have developed a degree of local adaptation and genetic differentiation. The species diversity of Ninox owls in the Far East might therefore still be underestimated, and this needs to be tested with genetic markers in future. Acknowledgements We thank Ju-Ying Wu for his substantial assistance with the field work. We are also grateful to Dr. Ying Wang, Shou-Hsien Li and Lucia Liu Severinghaus, who provided valuable suggestions during this research. References Averill, C. K. (1920) Migration and physical proportions: a preliminary study. Auk 37: 572-579. Brazil, M. (2009) Birds of East Asia: China, Taiwan, Korea, Japan, and Russia. New Jersey: Princeton University Press. Brazil, M. A. & Yabuuchi, M. (1991) The birds of Japan. Washington: Smithsonian Institution Press. del Hoyo, J., Elliott, A. & Elliott, A. eds. (1999) Handbook of the birds of the world, 5. Barcelona: Lynx Edicions. Dickinson, E. C. (2003) The Howard and Moore complete checklist of the birds of the world. New Jersey: Princeton University Press. Hachisuka, M. & Udagawa, T. (1951) Contributions to the ornithology of Formosa, Part II. Q. J. Taiwan Museum 4: 1-1 79. Konig, C., Weick, F. & Becking, J. H. (1999) Owls: a guide to the owls of the world. Robertsbridge, East Sussex: Pica Press. Konig, C., Weick, F. & Becking, J. H. (2009) Owls of the world. New Haven: Yale University Press. King, B. (2002) Species limits in the Brown Boobok Ninox scutulata complex. Bull. Brit. Orn. Club 1 22: 250-256. Masuda, A. (1974) ['Brooding and rearing of Japanese Brown Hawk-Owl, Ninox scutulata japonica.'] Res. Rep. Kochi Univ. Nat. Sci. 22(12): 1 93— 200. (In Japanese.) Mees, G. F. (1970) Notes on some birds from the island of Formosa (Taiwan). Zoo/. Meded. 44: 285-304. Oba, T. (1996) Vocal repertoire of the Japanese Brown Hawk Owl Ninox scutulata japonica: with notes on its natural history. Nat. Hist. Res. 2: 1- 64. Pande, S„ Pawashe, A., Mahajan, M. N., Joglekar, C. & Mahabal, A. (2007) Effect of food and habitat on breeding success in Spotted Owlets ( Athene brama) nesting in villages and rural landscapes in India. J. Raptor Res. 41: 26-34. Santhanakrishnan, R., Ali, A. M. S. & Anbarasan, U. (201 1) Breeding biology of the Spotted Owlet ( Athene brama Temminck, 1821) in Madurai district, Tamil Nadu, India. Internatn. J. Env. Sci. 1 : 1682-1691. Severinghaus, L. L. (2007) Cavity dynamics and breeding success of the Lanyu Scops Owl. J. Orn. 148 (suppl. 2): S407-S416. Severinghaus, L. L., Ting, T. S., Fang, W. H„ Tsai, M. C. & Yen, C. W. (2010) [‘The avifauna of Taiwan, 2.'] Taiwan: Forestry Bureau, Council of Agriculture. (In Chinese.) Tseng, Y.-S. & Lin, W.-L. (2010) [‘Owls of Taiwan.'] Taichung County: Wild Bird Rescue Institute of Taichung. (In Chinese.) Wang, J.-S., Wu, S.-H., Huang, K.-Y., Yang, X.-Y., Tsai, T.-H., Tsai, M.-C. & Xiao, Q.-L. (1991) [A field guide to the wild birds of Taiwan.'] Taipei: Taiwan Wild Bird Information Centre and Wild Bird Society of Japan. (In Chinese.) Yong, W. & Moore, F. R. (1 994) Flight morphology, energetic condition, and the stopover biology of migrating thrushes. Auk 111: 683-692. Wen-Loung UN, Wild Bird Rescue Institute of Taichung, National Taiwan Normal University, Taiwan. Email: ketupaf lavipes@yahoo.com. tw Si-Min UN, National Taiwan Normal University, Taiwan. Email: fish@ntnu.edu.tw Hui-Yun TSENG, National Museum of Natural Science, Tunghai University, Taiwan. Email: hytsengl 21 6@gmail.com Forktail 28 (2012) SHORT NOTES 167 The vocalisations of Red-collared Woodpecker Picus rabieri SIMON P. MAHOOD & JAMES A. EATON Introduction Red-collared Woodpecker Picus rabieri is a poorly known species restricted to Lao PDR, Vietnam from North and East Tonkin south to Central Annam, and extreme north-east Cambodia, with one old record from Yunnan province, China (BirdLife International 201 1, F. Goes pers. comm.). It occurs in evergreen and semi-evergreen forest and locally in tall deciduous forest below 700 m elevation, ascending locally to 1 ,000 m (BirdLife International 2001 ). It is rarely encountered and considered uncommon throughout the Vietnamese portion of its range, although it is locally abundant in Laos (Dersu 2008, Duckworth 2008, Thewlis etal. 1998, Duckworth et at. 1999, 2010, Timmins 2009, Suford in press). Where it is most abundant in Lao PDR, forests are generally semi-evergreen in character on level or at most gently sloping ground up to 300- 400 m elevation (J. W. Duckworth pers. comm. 2012). It is probably not as common in Vietnam, perhaps owing to an almost complete loss of forest on gentle slopes and of forest below 400 m elevation on all except the steepest slopes; where forest at optimal altitudes for the species remains, it is usually on limestone karst. The species's abundance drops off rapidly above 400 m and it is unclear if its apparent preference for forest with a moderately dry substrate (semi¬ evergreen forest and forest on limestone karst) represents an actual preference or simply a lack of other forest types at favoured elevations (J. W. Duckworth pers. comm. 2012). It is classified as Near Threatened by BirdLife International (201 1 ), owing to deforestation, particularly in the lowlands and more gentle terrain; as well as being the prime habitat of this woodpecker, these same areas are also favoured for agriculture, including plantation forestry. Red-collared Woodpecker vocalisations have not been documented in the published literature. For instance, Robson (201 1 ) does not list any calls but reports that the species 'drums in fast rolls'. A number of online trip reports and birding websites mention a short, explosive keck, often repeated in quick succession, and superficially similar to contact calls of similar-sized sympatric woodpeckers, e.g. Laced Woodpecker Picus vittatus. Similarly, Suford (in press) noted that it was vocally similarto congeners. Here we document the keck call and two previously undescibed vocalisations. Vocalisations Call 7. — Although not documented in the published literature, the keck call is relatively well known by birders. Examples of this vocalisation can be found on xeno-canto.org (XC69142-6, previously published on AVoCet as AV5442-6); a sonagram is shown in Figure 1. It is a single abrupt but full and rich note 0.08-0.09 seconds in length. The fundamental frequency is at approximately 2.8 kHz and there are strong harmonics with the next strongest frequency at approximately 1 .4 kHz, and successively weaker ones at approximately 4.0, 5.6 and 7 kHz. It is repeated at 1.5-second intervals. Call 2. — This call is remarkably similar to the territorial vocalisation of Blue-naped Pitta Pitta nipalensis, a species that overlaps in distribution with the woodpecker. This call was elucidated in forest around Ban Nahin, close to the well-publicised Bare-faced Bulbul Pycnonotus hualon site, although it was assumed that a Blue-naped Pitta made it. Birders at this site first heard it in 2009, but despite much effort the bird was not seen (de Win 2009). JAE and other subsequent would-be observers of Blue-naped Pitta at Ban Nahin forest have had similar experiences. At 07h00 on 21 December 2010, SPM was attempting to locate Red-collared Woodpecker in Ban Nahin by speculatively whistling the keck call. He heard a keck given close by in response, so he whistled an imitation of the call back to the bird. Immediately afterwards he heard a Blue-naped Pitta type vocalisation which emanated from the same place. The bird was out of sight because of the density of the vegetation and the angle of the slope, but the sound seemed to comefrom the ground in an area of bamboo and saplings within broadleaved evergreen forest. SPM whistled an imitation of this sound and a Red-collared Woodpecker flew from the position under observation up to a liana c.1 m above the ground. It perched there for less than ten seconds, allowing an identification to be made, and then flew away into the forest. The keck call now emanated from the direction in which the woodpecker had fled. SPM continued to imitate the 'Blue-naped Pitta' call, and the woodpecker responded by flying towards him and perched about 20 m up in a tree close by, where it was partially obscured by leaves. Although throughout this time it did not vocalise, it made the 'Blue-naped Pitta' vocalisation twice more when not in view . Knowing of SPM's experience, during a Birdtour Asia tour visit in January 201 1 JAE, along with seven other birders, heard the 'Blue- naped Pitta' call three times from within 500 m of the site where SPM had seen the Red-collared Woodpecker. Unfortunately there was no sign of any bird, but the call appeared to come from a small patch of bamboo inside primary evergreen forest. Despite waiting 30 minutes, with playback of both the keck and 'Blue-naped Pitta' calls, there was no response. The following day, in exactly the same spot, JAE speculatively played the 'Blue-naped Pitta' vocalisation, receiving an immediate response. When the bird called a second time, JAE was able to obtain a single sound recording (Figure 2). Unfortunately, despite spending over two hours in the area, and investigating the area of bamboo, nothing was seen. In mid-201 1 Dave Farrow drew SPM's attention to a recording of a drumming Red-collared Woodpecker made by Craig Robson in Ke Go Nature Reserve, Ha Tinh province, Vietnam, at <500 m elevation. At the end of the recording a Blue-naped Pitta type vocalisation can be heard. Although it cannot be confirmed that the vocalisation was made by a Red-collared Woodpecker, it is extremely unlikely that Blue-naped Pitta occurs at Ke Go Nature Reserve owing to the location and to the presence of Blue-rumped Pitta P. soror there. Although Blue-naped and Blue-rumped Pittas occur sympatrically in northern Vietnam (e.g. atTam Dao National Park), at these sites they separate out attitudinally, with Blue- rumped Pitta exclusively at lower elevations. Outside of the range of Blue-rumped Pitta, Blue-naped Pitta occurs at much lower elevations; for example it occurs at less than 1 00 m elevation at a P'— jf £ fr~ Figure 1. The keck call of Red- collared Woodpecker (JAE, March 2012, Phong Nha Ke Bang National Park, Vietnam). . . . i - 1 - - - 1 - - - 1 - - - 1 - - - 1 - - - 1 - - - 1 - - - r . — - - r- sO 1 23456789 10 168 SHORT NOTES Forktail 28 (2012) -20- : kus0 1 2 3 4 5 6 7 8 9 10 Figure 2. The 'Blue-naped Pitta' vocalisation of Red-collared Woodpecker (JAE, January 201 1, Ban Nahin, Lao PDR). ku 20- 0 000-- -20- s0 Figure 3. The territorial vocalisation of Blue-naped Pitta (JAE, March 2007, Tam Dao National Park, Vietnam). 20 0 000 -20 Figure 4. The 'Blue-rumped Pitta' vocalisation of Red-collared Woodpecker (JAE, March 201 2, Phong Nha Ke Bang National Park, Vietnam). site close to Kaziranga National Park, Assam, India (JAE pers. obs.). In Indochina, semi-evergreen forest on karst supports a number of moderate- and high-altitude species at surprisingly low elevations (e.g. Green-backed Tit Parus monticolus and Green Cochoa Cochoa viridis, both of which have been regularly recorded at <400 m elevation at Ban Nahin), so it would not be unreasonable to assume that Blue-naped Pitta might also occur at similar elevations in karst at sites from which Blue-rumped Pitta is absent. It is not surprising that birders suspected the Ban Nahin call of being made by Blue-naped Pitta (compare Figures 2 and 3). Like one of the vocalisations of that species it consists of a loud, mellow, throaty, upslurred whistle. However, it is possibly somewhat more abrupt than that of Blue-naped Pitta, although both are c.0.5 seconds in length. The woodpecker's vocalisation is perhaps not as modulated, with the first half of the strophe 1-1 .8 kHz in pitch rising to just over 2 kHz at the end (perhaps slightly lower than the maximum frequency achieved by the pitta), and without dipping slightly in frequency in the first quarter as noticeably as in the pitta. The call is given singly or in a series. Call 3. — This third vocalisation (Figure 4) is somewhat intermediate between a Blue-rumped Pitta and a macaque species Macaca. It was elucidated by JAE in Phong Nha Ke Bang National Park, Vietnam, during a Birdtour Asia tour in March 2012. He was attempting to find Red-collared Woodpecker using the keck and 'Blue-naped Pitta' calls when he heard a novel sound. He recorded it and played it back, and almost immediately a Red-collared Woodpecker flew in, repeating the call. It consists of a short, modulated, slightly nasal, panicked-sounding strophe given singly or in a series. It is 0.2 seconds in duration and begins at 1-2 kHz, rising in pitch to 3.5 kHz before quickly dropping back to 1 .5 kHz. The highest frequency in the strophe is located two-thirds of the way through: the rise in pitch is almost twice as long as the drop at the end. It is possible that this sound was previously overlooked as a variant Blue-rumped Pitta vocalisation. Discussion Bird vocalisations have evolved to travel efficiently through the environment in which the birds that make them live. There is therefore often convergence in structure and pitch between distantly related bird species that occupy a similar habitat niche. Although superficially Red-collared Woodpecker and Blue-naped Pitta are species with very different habits, they are quite similar. Blue-naped Pitta obtains all or almost all of its food from the forest floor. Although data are scant and few detailed observations of the woodpecker have been published, most encounters with it are of birds seen foraging on the forest floor, or flushed from the forest floor, or in woody understorey tangles. A typical behaviour when encountered is to flush from the forest floor to a low perch on a liana before flying off out of sight (J. W. Duckworth pers. comm. 2012). It is not known if the woodpecker seen by SPM in Ban Nahin forest was vocalising from the ground, as it was out of sight and might have been perched on a low stem or branch. Blue-naped Pitta usually vocalises from the ground or, when agitated, from a perch up to 3 m of the ground. The two species therefore spend most of their time in the same part of the forest stratum and presumably often vocalise from the same elevation above the ground. Interestingly, both species apparently vocalise relatively infrequently or at least vocalise infrequently when they are in view or have been detected by an observer. In Indochina, Blue-naped Pitta and Red-collared Woodpecker rarely occur at the same location. Blue-naped Pitta is decidedly scarce in Vietnam, with confirmed records from only one location, Tam Dao, where Red-collared Woodpecker has not been recorded and native forest has been lost at lower elevations (Tordoff 2002). It has also been reported from Pu Mat National Park (SFNC 1998), although this record is considered unconfirmed by Tordoff (2002). Its status in Laos is unclear, although it has been recorded at only a small number of locations, including Nam Kading NBCA (down to 500 m elevation), Nam Xam NBCA and Nam Et NBCA. Red-collared Woodpecker has been recorded at a number of sites within the range of Blue-naped Pitta in Lao PDR, including Nam Xam NBCA where it was recorded at 1 ,000 m (an exceptionally high elevation, although there is no reason to question the veracity of the record) and Nam Kading NBCA. In northern Vietnam it has been recorded at only one site (Ban Bung IBA) where Blue-naped Pitta could conceivably occur (Hill & Kemp 1996) but it has been recorded in Pu Mat National Park (Tordoff 2002). There is very little remaining lowland forest in areas where both Red-collared Woodpecker and Blue-naped Pitta might co-occur in northern Indochina, but even the small numberof records available suggests that their geographical distributions might overlap. At sites where they co-occur Blue-naped Pitta usually occurs at higher elevations than Red-collared Woodpecker, which occupies the lowlands and indeed is scarce outside gentle terrain. Heard-only records of Blue-naped Pitta from within the range of Red-collared Woodpecker (and vice versa), especially those from lower elevations, should be treated as unconfirmed owing to the potential for previously unforeseen confusion with that species. Although it is possible that knowledge of a fuller range of the vocalisations of Red-collared Woodpecker will lead to more records of this poorly known bird, it may be that the paucity of lowland forest on level ground or gentle slopes and the relative infrequency of observer effort within such habitat will mean that this species continues to be rarely recorded, especially by birdwatchers. Forktail 28(2012) SHORT NOTES 169 Acknowledgements We thank Will Duckworth for his comprehensive comments on an early draft of this manuscript and Pamela Rasmussen for making the sonagrams. Dave Farrow provided us with the Craig Robson recording of the Blue-naped Pitta type vocalisation of Red-collared Woodpecker, whilst Craig Robson and Pamela Rasmussen both provided helpful comments on the manuscript. References Bird Life International (2001 ) Threatened birds of Asia: the BirdLife International Red Data Book. Cambridge, UK: BirdLife International. BirdLife International (2011) Species factsheet: Picus rabieri. Downloaded from http://www.birdlife.org on 30/07/201 1. Robson, C. R . (201 I) A field guide to the birds of South-East Asia. London: New Holland. Dersu (2008) C880: Wildlife program phase 1. Baseline inventory. Wildlife and habitat studies of the Nakai plateau. Vientiane: Dersu and Associates for Nam Theun 2 Power Company. Duckworth, J. W. (2008) A reconnaissance wildlife survey of the BCI pilot villages in the BCI Xe Pian-Dong Hua Sao corridor. Vientiane: WWF Lao office. Duckworth, J. W., Salter, R. E. & Khounboline, K., compilers (1999) Wildlife in Lao PDR: 1 999 status report. Vientiane: IUCN/WCS/CPAWM. Duckworth, J. W., Hallam, C. D., Phimmachak, S., Sivongxay, N., Stuart, B. L. & Vongsa, O. (2010) A conservation reconnaissance survey of north-east Vilabouli district, Savannakhet province, Lao PDR. Vientiane: WCS report to Lane Xang Minerals Ltd. Hill, M. & Kemp, N. (1 996) Biological survey of Na Hang Nature Reserve, Tuyen Quang province, Vietnam: part 1, Ban Bung sector. London: Society for Environmental Exploration. SFNC (1998) Status and distribution of the bird fauna of the Pu Mat Nature Reserve. SFNC Project, Vinh, Vietnam. [Suford] Sustainable Forestry for Rural Development project (in press) Preliminary biodiversity assessment and management recommendations of Suford-AF Production Forest Areas. Vientiane, Lao PDR: Suford project, Department of Forestry. Thewlis, R. M., Timmins, R. J., Evans, T. D. & Duckworth, J. W. (1998) The conservation status of birds in Laos: a review of key species. Bird Conservation International 8 (supplement): 1-159. Timmins, R. J. (2009) Biodiversity significance and management of the Phou Theung forest area, Xekong Province, Lao PDR. Vientiane: WWF. Tordoff, A. W., ed. (2002) Directory of Important Bird Areas in Vietnam. Hanoi: BirdLife International in Indochina and the Institute of Ecology and Biological Resources. de Win, S. (2009) http://www.birding2asia.com/W2W/Laos/NaHin.html Simon P. MAHOOD, Wildlife Conservation Society Cambodia Programme, House 21 , Street 21 , SangkatTonie Bassac, Phnom Penh, Cambodia. Email: s.mahood@wcscambodia.org James A. EATON, A-3A-5, Casa Indah, Kota Damansara, Selangor, Malaysia. E-mail: jameseaton@birdtourasia.com Diet of the Speckled Boobook Ninox punctulata in north Sulawesi, Indonesia JAMES A. FITZSIMONS, ERIK MEIJAARD, IWAN HUNOWU, DEWI PRAWIRADILAGA, JANELLE L. THOMAS & JOHNY S.TASIRIN Introduction The ecology of most of Sulawesi's owl species is poorly known (Bishop 1 989, Debus 2002, Fitzsimons 2010). The Speckled Boobook Ninox punctulata is one of four Ninox species that are endemic to Sulawesi and its satellite islands, although the recent discovery by Madika et al. (2011) could bring up this number to five. It occurs throughout the island and inhabits forests and disturbed lowland habitats (White & Bruce 1 986, Coates & Bishop 1 997). Konig et al. (2008: 469) stated it to be 'widespread and common within its restricted range'. Despite being one of the commoner Ninox species on Sulawesi, little is known about its diet. Marks ef al. (1999: 236) suggested that there is 'almost no information on diet'. Konig et al. (2008: 469) described its food as 'presumably mainly insects' and that 'the biology and ecology.. ..of this species needs study'. Coates & Bishop (1997: 363) noted it 'has been recorded foraging along narrow streams within primary forest'. Rozendaal & Dekker (1 989) reported a bird killing a Swift Fruit Bat Thoopterus nigrescens' in a net over a river in dense primary forest'. Here we describe components of the diet of the Speckled Boobook based on prey remains and pellets collected at a sheltered roost site in north Sulawesi, with a view to increasing ecological understanding of this species, and possibly helping predict its sensitivity to ecological disturbance. Methods and study area Pellets of the Speckled Boobook were located by JAF and JLT on 19 July 2009 in an abandoned, dilapidated but sheltered dwelling at the headquarters of Bogini Nani Wartabone National Park atToraut, north Sulawesi (0°34'N 1 23°54'E; 220 m asl).The site borders the extensive rainforest that makes up the 287,000 ha national park, a river and an area of agricultural land in the Dumoga Valley (for a description of rainforest at Toraut see Whitmore & Sidiyasa 1986). This dwelling was a known roost site for Speckled Boobook (e.g. Farrow 2008), and an individual of this species was heard by JAF, JLT and park rangers within metres of the dwelling on the night of 18 July 2009. Both Sulawesi Masked Owl Tyto rosenbergii and Sulawesi Scops Owl Otus manadensis also use habitat in this area, and both were heard on the night of 1 8 July 2009. Sulawesi Masked Owl is known to roost in one of the large rainforest trees near the park headquarters (Fitzsimons 2010). Based on various forms of evidence (known roost site, presence of Speckled Boobook feathers, pellet size and composition), we determined that the pellets were those of Ninox punctulata and not the other species. Approximately ten separate piles of pellet material (each pile most likely consisting of multiple pellets) in varying states of decomposition were located throughout the dwelling, with only one pellet seemingly intact. Ants were removing insect material from these pellet remains at the time and moth pupae were also causing disintegration of the material. Prey remains were analysed by EM, in consultation with Sulawesi small mammal expert Guy Musser, by visually comparing the lower mandibles within the sample. We compared these to descriptions in the taxonomic literature for small mammals in Sulawesi (e.g. Musser 1972, 1981a, b, 1982, 1991, Ruedi 1995). Tooth cusp patterning was also examined for distinguishing features. The minimum number of prey individuals was recorded (by counting skulls and matching left and right jawbones) and weights sourced from the published literature. Although the remains of invertebrates were in a degraded state at the time of analysis and could not be systematically or accurately assessed, we identified key distinguishing features such as beetle wings. 170 SHORT NOTES Forktail 28 (2012) Results It was not possible to quantify the number or type of invertebrate material in the pellets, owing to their state of decomposition. However, beetles (Coleoptera) were identified as present. Based on the obvious presence of a diastema and tooth shape on the mandible, all mammal specimens were all identified as rodents. No remains of insectivorous mammals were found. Fifteen of the mandibles were identified as from Pacific Rat Rattus exulans and seven from Black Rat Rattus rattus complex (previously, individuals from this complex in Sulawesi were considered a separate species, Tanezumi Rat Rattus tanezumi: Musser & Carleton 2005, G. Musser pers. comm. 2012). Measurements on the maxillary toothrow (LM13 in Musser 1 979) were only possible on a few maxillary fragments. These were inconclusive as to species identity, and it is possible that other small rodent species were among the vertebrate remnants. Discussion Both the Pacific Rat and Black Rat are introduced fauna in Sulawesi, and are common in disturbed areas. These species also inhabit degraded forests, but are absent in primary forests where tracks are not present (G. Musser pers. comm. 2012). In contrast, the entire (native) murid fauna of Sulawesi originally evolved in forested habitats (Musser 1 987), although some native species can occupy secondary growth and scrub but usually only if good forest is nearby (G. Musser pers. comm. 2012). Interestingly, Durden (1986a,b) found the native Musschenbroek's Spiny Rat Maxomys musschenbroekii to be the most commonly trapped rodent in forests near our study site, while other native species (Yellow-tailed Rat Rattus xanthurus, Hoffmann's Rat R. hoffmanni, Sulawesi Giant Rat Paruromys dominator and Hellwald's Spiny Rat Maxomys hellwaldii) were also present (Durden & Watts 1 988). In the immediate vicinity of the park headquarters where the pellets were collected, the range of habitats — scrub, forest with tracks, agricultural fields — is likely to contain both introduced Rattus species. This situation suggests that Speckled Boobooks in this locality are using some or all of these habitats for hunting. Both Pacific and Black Rats inhabit the ground, but are also good climbers. Thus it is not possible to infer whether Speckled Boobooks have a preference for hunting ground-dwelling or arboreal prey. As indicated above, the only previously documented prey of the Speckled Boobook is Swift Fruit Bat, a species which, although physically larger than the rodents in our study, weighs 67-99 g (Bergmans & Rozendaal 1 988) — thus falling between the average weights of rodents from our study (40 g for Rattus exulans and 280 g for R. rattus: Dickman & Watts 2008, Watts & Aplin 2008). Considering the diversity of insectivorous mammals in Sulawesi (namely shrews: Ruedi 1995), and the similar sizes and general behaviour of these mammals to rodents, it is interesting that they were not present in the Speckled Boobook's diet, although admittedly our sample size is small. When first describing Cinnabar Boobook Ninox ios, which she suggested may take soft-bodied insects in flight, Rasmussen (1999: 462) described the Speckled Boobook (along with some Melanesian taxa) as'strikingly different in plumage and morphology [from other Sulawesi Ninox species], with short tails, very heavy tarsi, and Athene-like plumage pattern and toe bristles; in fact some had been placed in that genus (among others) in the past'. There is limited information on the weight of Speckled Boobooks: Konig etal. (2008: 415) stated 'about 200 g', and Marks ef al. (1999: 236) stated 'one male [weighed] 151 g'. Considering the average weight of Black Rats is 280 g, this suggests that the Speckled Boobook is able to take prey almost twice its own weight. The comparatively heavier Southern Boobook N. novaeseelandiae boobook in south-eastern Australia (males 1 94-360 g, females 1 70-298 g: Konig et al. 2008), while known to take Black Rats (and occasionally birds up to rosella Platycercus size), prefers smaller mammals such as House Mouse Mus musculus and insects (e.g. Trost et al. 2008). As the insect remains within the Speckled Boobook pellets were being removed by ants and being eaten by moths when we collected them, we cannot comment on the importance of vertebrates vs invertebrates in the diet of this owl species. Problems in determining the proportions of vertebrates and invertebrates in diets are also evident in the comparatively better-studied Ninox species, the Southern Boobook (e.g. Rose 1996, McNabb 2002, Fitzsimons & Rose 2007, Trost et al. 2008, Olsen 201 1); survey methods and condition of pellet material contribute to significant differences. Our findings should not be taken as representative of the total diet of Speckled Boobooks in north Sulawesi, but potentially representative of vertebrates in their diet. The old building in which the Speckled Boobook in our study roosted has been used regularly fora number of years (see Farrow 2008, De Win 2010; 2009 photos and videos on Internet Bird Collection — http://ibc.lynxeds.com/species/speckled-hawk-owl- ninox-punctulata). Another small Ninox, the Southern Boobook, is also known to use such buildings (e.g. Fitzsimons & Rose 2007). Our findings suggest that the Speckled Boobook feeds on both small mammals (rodents) and insects. In our study area, the species appears to select prey species that predominate in open agricultural and scrub areas or forest edges, rather than primary forests, suggesting considerable ecological flexibility. However, more research is required to obtain a better understanding of this species's diet, hunting preferences, and habitat. The increased number of ornithologists, as well as birdwatching tours to Sulawesi, and Wallacea more generally, which often promote themselves on locating endemic owl species, provides an opportunity to increase our knowledge of the basic ecology of little-studied owls in this region. Acknowledgements Thanks to Idjong Datunsolang, Hendrik Rumaer and Arifin Ali of Bogani Nani Wartabone National Park for showing us the roost site and granting permission to collect the pellets. Guy Musser kindly provided his expert opinion on the identity of the prey remains and information on the habits of Sulawesi rodents more generally. Stephen Debus and Nigel Collar provided constructive comments on this paper. References Bergmans, W. & Rozendaal, F. G. (1988) Notes on collections of fruit bats from Sulawesi and some off-lying islands (Mammalia, Megachiroptera). Zoologische Verhandelingen 248: 1-74. Bishop, K. D. (1989) Little known Tyto owls of Wallacea. Kukila 4: 37-43. Coates, B. J. & Bishop K. D. (1997) A guide to the birds of Wallacea: Sulawesi, the Moluccas and Lesser Sunda Islands, Indonesia. Alderley, Queensland, Australia: Dove Publications. Debus, S. (2002) Distribution, taxonomy, status, and major threatening processes of owls in the Australasian region. Pp. 355-363 in I. Newton, R. Kavanagh, J. Olsen and I. Taylor, eds. Ecology & conservation of owls. Collingwood: CSIRO Publishing. De Win, S. (2010) B2A Sulawesi & Halmahera 2010 tour report. 03-21 August. Available http://www.birding2asia.com/tours/reports/Sulawesi- HalmaheraAugust10_1 .html Dickman, C. R. & Watts, C. H. S. (2008) Black Rat Rattus rattus. Pp. 707-709 in S. Van Dyck and R. Strahan, eds. The mammals of Australia. Third Edition. Sydney: Reed New Holland. Durden, L. A. (1986a) Rats and ectoparasites on Project Wallace. Antenna 10:29-30. Durden, L. A. (1986b) The reinfestation of Forest Rats ( Maxomys musschenbroekii ) by epifaunistic arthropods in Sulawesi, Indonesia. J. Trop. Ecol. 2: 283-286. Durden, L. A. & Watts, C. H. S. (1988) A collection of ticks (Ixodidae) from Sulawesi Utara, Indonesia. Biotropia 2: 32-37. Forktail 28 (2012) SHORT NOTES 171 Farrow, D. (2008) Sulawesi & Halmahera 14 September-5 October 2008Tour Report. Available: http://www.birdquest.co.uk/pdfs/report/ INDONESIA%20(SULAWESI)%20REP%2008.pdf Fitzsimons, J. A. (2010) Notes on the roost sites of the Sulawesi Masked Owl Tyto rosenbergii. Forktail 26: 142-145. Fitzsimons, J. A. & Rose, A. B. (2007) The diet of a Southern Boobook Ninox novaeseelandiae in Box-lronbark country, central Victoria. Austral. Zool. 34: 85-88. Konig, C., Weick, F. & Becking, J. (2008) Owls of the world. Second edition. London: Christopher Helm. Madika, B., Putra, D. D., Harris, J. B. C., Yong, D. L., Mallo, F. N., Rahman, A., Prawiradilaga, D. M. & Rasmussen, P. C. (201 1) An undescribed Ninox hawk owl from the highlands of Central Sulawesi, Indonesia? Bull. Brit. Ornithol. Club 131:21-29. Marks, J. S., Cannings R. J.& Mikkola, H. (1 999) Family Strigidae (typical owls). Pp 76-151 in J. del Hoyo, A. Elliott and J. Sargatal, eds. Handbookofthe birds of the world, 5. Barcelona: Lynx Edicions. McNabb, E. G. (2002) Notes on the diet and observations of the Southern Boobook (Ninox novaeseelandiae) in southern Victoria. Pp. 1 92-1 98 in I. Newton, R. Kavanagh, J. Olsen and I. Taylor, eds. Ecology & conservation of owls. Collingwood: CSIRO Publishing. Musser, G. G. (1 972) The species of Hapalomys (Rodentia, Muridae). Amer. Mus. Novitates 2503: 1-27. Musser, G.G. (1979) Results of the Archbold Expeditions. No 102. The species of Chiropodomys, arboreal mice of Indochina and the Malay Archipelago. Bull. Amer. Mus. Natur. Hist. 1 62: 379-445. Musser, G. G. (1981a) Results of the Archbold expeditions. No. 105. Notes on the systematics of Indo-Malayan murid rodents, and descriptions of new genera and species from Ceylon, Sulawesi, and the Philippines. Bull. Amer. Mus. Natur. Hist. 168: 229-334. Musser, G. G. (1 981 b) The giant rat of Flores and its relatives east of Borneo and Bali. Bull. Amer. Mus. Natur. Hist. 1 69: 67-1 76. Musser, G.G. (1982) Results of the Archbold Expeditions. No. 110. Crunomys and the small-bodied shrew rats native to the Philippine Islands and Sulawesi (Celebes). Bull. Amer. Mus. Natur. Hist. 1 74: 1-95. Musser, G. G. (1 987) The mammals of Sulawesi. Pp. 73-93 in Whitmore, T. C., ed. Biogeographical evolution of the Malay Archipelago. Oxford: Oxford University Press. Musser, G. G. (1991) Sulawesi rodents: descriptions of new species of Bunomys and Maxomys (Muridae, Murinae). Amer. Mus. Novitates 3001 : 1-44. Musser, G. G.& Carleton, M. D. (2005) Superfamily Muroidea. Pp. 894-1531 in D. E. Wilson and D. M. Reeder, eds. Mammal species of the world: a taxonomic and geographic reference. Baltimore: Johns Hopkins University Press. Olsen, J. (2011) Australian High Country owls. Melbourne: CSIRO Publishing. Rasmussen, P. C. (1 999) A new species of Hawk-owl Ninox from north Sulawesi, Indonesia. Wilson Bull. 1 1 1: 457-630. Rose, A. B. (1996) Notes on the diet of the Southern Boobook Ninox novaeseelandiae in New South Wales. Aust. Bird Watcher 1 6: 339-343. Rozendaal, F. G.& Dekker R. W. R. J. (1989) An annotated checklist of the birds of Dumoga-Bone National Park, North Sulawesi. Kukila 4: 85-109. Ruedi, M. (1 995) Taxonomic revision of shrews of the genus Crocidura from the Sunda shelf and Sulawesi with description of two new species. Zool. J. Linnean Soc. 115:211 -265. Trost, S„ Olsen, J., Rose, A. B. & Debus, S. J. S. (2008) Winter diet of Southern Boobooks Ninox novaeseelandiae in Canberra 1997-2005. Corella 32: 66-70. Watts, C. H. S. & Aplin, K. P. (2008) Pacific Rat Rattus exulans. Pp. 704-705 in S.Van Dyck and R. Strahan, eds. The mammals of Australia.Jbird Edition. Sydney: Reed New Holland. White, C. M. N. & Bruce, M. D. (1986) The birds ofWallacea (Sulawesi, the Moluccas and Lesser Sunda Islands, Indonesia): an annotated check-list. London: British Ornithologists' Union (Check-list no. 7). Whitmore, T. C. & Sidiyasa, K. (1 986) Composition and structure of a lowland rain forest atToraut, northern Sulawesi. Kew Bull. 41: 747-756. James A. FITZSIMONS, The Nature Conservancy, Suite 3-04, 60 Leicester Street, Carlton VIC 3053, Australia ; and School of Life and Environmental Sciences, Deakin University, 221 Burwood Highway, Burwood VIC 3125, Australia. Email: jfitzsimons@tnc.org Erik MEIJAARD, People & Nature Consulting International, Vila Lumbung House No. 6, Jalan Raya Petitenget 1000X, Kerobokan, Badung 8036 1, Bali, Indonesia. Email: emeijaard@gmail.com Iwan HUNOWU, Wildlife Conservation Society, Indonesia Program - Sulawesi, Jl. Sam Ratulagi Number 4 1, PO Box 1580, Manado, 95000, Indonesia. Email: ihunowu@wcs.org Dewi PRAWIRADILAGA, Division of Zoology, Research Centre for Biology-LIPI, Jl. Raya Bogor Km 46, Cibinong Science Centre, Bogor 1691 1, Indonesia. Email: dmprawiradilaga@gmail.com Janelle L. THOMAS, BirdLife Australia, Suite 2-05, 60 Leicester Street, Carlton VIC 3053, Australia. Email: janelle.thomas@birdlife.org.au JohnyS. TASIRIN, Wildlife Conservation Society, Indonesia Program - Sulawesi, Jl. Sam Ratulagi Number 41, PO Box 1 580, Manado, 95000, Indonesia. Email: jtasirin@wcs.org Mobbing to death of a Japanese Long-eared Bat Plecotus sacrimontis by two species of tit TOSHITAKAN. SUZUKI Introduction Mobbing is a widespread antipredator behaviour that occurs when individuals approach and cooperatively drive off a potential predator (Wilson 1975, Curio 1978). Birds use mobbing against a variety of predators (e.g., hawks, owls and snakes). This behaviour includes distinctive calls that attract additional mobbers from the same and different species (Curio 1 978, Hurd 1 996). Some species of bird have evolved the ability to adjust their mobbing response according to the predator species (Griesser 2009, Suzuki 2011, 2012). On the other hand, there are several reports of birds occasionally mistaking harmless animals as targets for mobbing. For example, tits have been documented mobbing Common Cuckoo Cuculus canorus mounts at winter feeders because the plumage coloration and patterns of cuckoos mimic those of Sparrowhawks Accipiter nisus (Davies & Welbergen 2008). Nighthawks (Caprimulgidae) are similar to owls of the family Strigidae in their cryptic plumage, and occasionally induce mobbing by forest birds (Marks etal. 201 1). I describe an instance in which a Willow Tit Poecile montanus and a Great Tit Parus major minor simultaneously mobbed a flying Japanese Long-eared Bat Plecotus sacrimontis, a small nocturnal mammal that measures approximately 50 mm in size, including the head and body (Ohdachi etal. 2009). This bat preys exclusively on arthropods (Ohdachi et al. 2009) and poses no threat to birds, although some other bat species in other geographic regions have been reported to prey on birds (reviewed in Ibanez et al. 2001). 172 SHORT NOTES Forktail 28 (2012) There are several reports of predation by birds on bats (Lee & Kuo 2001, Chacon-Madrigal & Barrantes 2004, Hernandez et ai 2007, Estok et ai 2010), but mobbing of flying bats by small passerines has been rarely reported (e.g. Tugendhat 1 966). Observations A Japanese Long-eared Bat was observed on 25 April 2008, at 1 1 h45 (Japan Standard Time), flying about 4 m above a stream in a mixed deciduous-coniferous forest in Karuizawa, Nagano, Japan (36°22'N 138°36'E). I then saw a Willow Tit and a Great Tit fly towards the bat. The Willow Tit flew at an angle as if it was trying to intercept the bat's flight path and came within 1 m of the bat, repeatedly producing mobbing calls (, zi-taa calls: Haftorn 2000). The Great Tit also flew at the bat in this manner and came within 2 m of the bat. The two birds sometimes perched on trees beside the stream but immediately resumed flying towards, approaching closely, and mobbing the bat. The tits chased the bat for at least 20 m, although they did not make direct attacks on it. Although it was unclear if the mobbing had started before I started my observation, I observed the mobbing for more than 20 seconds. During the mobbing, the bat seemed to be exhausted since it flew up and down with a quite low speed and its flying height gradually decreased. Finally, the bat fell into the stream. After this, both tits stopped mobbing and left the vicinity. The bat moved its wings for c.10 seconds but could not escape from the water. I went down the stream and confirmed that the bat was dead. No signs of injury or attack were observed on the bat's body. Discussion Mobbing of bats by small passerines has rarely been reported. Another account described an observation of Barn Swallows Hirundo rustica chasing and mobbing a flying Pipistrelle Bat Pipistrellus pipistrellus during the day in England (Tugendhat 1 966). My report provides the first observation in which Willow and Great Tits simultaneously mobbed a Japanese Long-eared Bat, which may have contributed to its death. I do not know how the two tits started to mob the bat. Japanese Long-eared Bats are nocturnal mammals and normally roost in cavities during the day (Ohdachi etal. 2009), so diurnal birds are unlikely to encounter bats. However, Great Tits are secondary cavity-nesters and from March to May often enter tree-cavities to assess potential nesting sites (pers. obs.). Since the incident in question took place in the last week of April, it is possible that the bat was disturbed in its roost by the Great Tit, which then proceeded to mob it. It remains unclear why the Willow and Great Tits mobbed the bat. It seems unlikely they mistook it for a potential predator, since bats are morphologically very different from small-bird predators such as shrikes, owls and hawks. Moreover, it is unlikely that this was an attempted predation, although several previous reports have shown that raptors, owls, crows and motmots consume flying bats (Lee & Kuo 2001, Chacon-Madrigal & Barrantes 2004, Hernandez etal. 2007). In Hungary, Great Tits have been observed to kill and prey on bats, but only when the bats were inactive and hibernating in caves (Estok etal. 2010). Further observations of bat mobbing by birds might provide insight into predator recognition and the feeding ecology of birds, as well as the evolution of nocturnal behaviour in bats (Rydell & Speakman 1 995). References Chacon-Madrigal, E. & Barrantes, G. (2004) Blue-crowned Motmot ( Momotus momota) predation on a Long-tongued Bat (Glossophaginae). Wilson Bull. 116: 108-110. Curio, E. (1978) The adaptive significance of avian mobbing. Zeitschr. Tierpsych. 48: 1 75-183. Davies, N. B. & Welbergen, J. A. (2008) Cuckoo-hawk mimicry? An experimental test. Proc. Roy. Soc. 6 275:181 7-1822. Estok, P., Zsebok, S. & Siemers, B. M. (2010) Great Tits search for, capture, kill and eat hibernating bats. Biol. Lett. 6: 59-62. Griesser, M. (2009) Mobbing calls signal predator category in a kin group¬ living bird species. Proc. Roy. Soc. B 276: 2887-2892. Haftorn, 5. (2000) Contexts and possible functions of alarm calling in the Willow Tit, Parus montanus ; the principle of 'better safe than sorry'. Behaviour 1 37: 437-449. Hernandez, D. L„ Mell, J. J. & Eaton, M. D. (2007) Aerial predation of a bat by an American Crow. Wilson J. Orn. 1 19: 763-764. Hurd, C. R. (1996) Interspecific attraction to the mobbing calls of Black- capped Chickadees (Parus atricapillus). Behav. Ecol. Sociobiol. 38: 287- 292. Ibanez, C„ Juste, J., Garcia-Mudarra, J. L. & Agirre-Mendi, P. T. (2001) Bat predation on nocturnally migrating birds. Proc. Natn. Acad. Sci. 98: 9700-9702. Lee, Y.-E. & Kuo, Y.-M. (2001) Predation on Mexican Free-tailed Bats by Peregrine Falcons and Red-tailed Hawks. J. Rapt. Res. 35: 1 1 5-1 23. Marks, J. S., Crabtree, C. S., Benz, D. A. & Kenne, M. C. (2011) Mobbing of Common Nighthawks as cases of mistaken identity. Wilson J. Orn. 1 23: 183-185. Ohdachi, S. D., Ishibashi, Y., Iwasa, M. A. & Saitoh, T. (2009) The wild mammals of Japan. Kyoto: Shoukadoh Book Sellers. Rydell, J.& Speakman, J. R. (1995) Evolution of nocturnality in bats: potential competitors and predators during their early history. Biol. J. Linn. Soc. 54: 183-191. Suzuki, T. N. (2011) Parental alarm calls warn nestlings about different predatory threats. Curr. Biol. 21 : R1 5-R16. Suzuki, T. N. (2012) Referential mobbing calls elicit different predator¬ searching behaviours in Japanese Great Tits. Anim. Behav. 84: 53-57. Tugendhat, M. (1966) Swallows mobbing Pipistrelle Bats. Brit. Birds 59:435. Wilson, E. O. (1975) Sociobiology. Cambridge: Belknap Press. Toshitaka N. SUZUKI, Department of Life Science, Rikkyo University, 3-34-1 Nishi-lkebukuro, Toshima, Tokyo 171-8501, Japan. Email: toshi.n.suzuki@gmail.com Forktail 28 (2012) Guidelines for contributors Forktail publishes original papers in the English language treating any aspect of the ornithology (e.g. distribution, biology, conservation, identification) of the region bounded by the Indus River to the west, the Russian Far East, Korean Peninsula, Japan, and Lydekker's Line (i.e. the eastern boundary of Wallacea) to the east, the Chagos Archipelago, Lesser Sundas, Christmas Island and Cocos (Keeling) Islandsto the south (see map in Oriental Bird Club Bull. 31:7). Submissions are considered on the understanding that they are being offered solely for publication by the Oriental Bird Club, which will retain copyright. All submissions are reviewed by referees and those accepted are normally published in order of receipt. 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Rare and endangered birds of the Far Eosf.Vladivostok: Far East Science Center, Academy of Sciences of the USSR. (In Russian.) Sien Yao-hua, Kuan Kuan-Hsun & Zheng Zuo-xin (1964) ['An avifaunal survey of the Chinghai province.'] Acta Zoo!. Sinica 16: 690-709. (In Chinese.) Smythies, B. E. (1 981 ) The birds of Borneo. Third edition. Kota Kinabalu and Kuala Lumpur: The Sabah Society and the Malayan Nature Society. Somadikarta, S. (1986) Collocalia linchi Horsfield & Moore -a revision. Bull. Brit. Orn. Club 1 06: 32-40. White, C. M. N. & Bruce, M. D. (1 986) The birds of Wallacea (Sulawesi, the Moluccas and Lesser Sunda Islands, Indonesia): an annotated check-list. London: British Ornithologists' Union (Check-list no. 7). 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