A Taxonomic Review of Hipposideros halophyllus , with Additional Information on H. ater and H....

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions,research libraries, and research funders in the common goal of maximizing access to critical research.

A Taxonomic Review of Hipposideros halophyllus, with Additional Informationon H. ater and H. cineraceus (Chiroptera: Hipposideridae) from Thailand andMyanmarAuthor(s): Bounsavane Douangboubpha, Sara Bumrungsri, Pipat Soisook, Susan W. Murray,Sébastien J. Puechmaille, Chutamas Satasook, Si Si Hla Bu, David L. Harrison and Paul J. J. BatesSource: Acta Chiropterologica, 12(1):29-50. 2010.Published By: Museum and Institute of Zoology, Polish Academy of SciencesDOI: http://dx.doi.org/10.3161/150811010X504572URL: http://www.bioone.org/doi/full/10.3161/150811010X504572

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological,and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and bookspublished by nonprofit societies, associations, museums, institutions, and presses.

Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance ofBioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.

Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercialinquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

http://dx.doi.org/10.3161/150811010X504572

http://www.bioone.org/doi/full/10.3161/150811010X504572

http://www.bioone.org

http://www.bioone.org/page/terms_of_use

INTRODUCTION

As recently noted by Guillén-Servent andFrancis (2006), the taxonomy of bats in the genusHipposideros has been complicated by the morpho-logical similarity of many species and the lack of adequate field collections in many regions of theOld World tropics. This is particularly the case with respect to the H. bicolor group, where despite a series of papers since Tate (1941) and Hill (1963) describing several new species (Khajuria, 1970; Hilland Yenbutra, 1984; Kock and Bhat, 1994; Fran-cis et al., 1999; Guillén-Servent and Francis, 2006;Bates et al., 2007) and a number of additional pub-lications which have contributed to an understand-ing of the taxonomy and phylogeny of this group,for example Jenkins and Hill (1981) and Bogda no -wicz and Owen (1998), there remains considerabletaxonomic confusion. This in turn has led to a lackof authoritative information on species diagnosis,distributions, ecology, and population status.

In this study, three species of small hipposideridfrom the bicolor subgroup of the bicolor group (sen-su Corbet and Hill, 1992) were studied, primarily inThailand but with additional information fromMyanmar and northern Malaysia. The species areHipposideros halophyllus Hill and Yenbutra, 1984,which is essentially restricted to Thailand but withone additional record from northern Malaysia (Fran -cis, 2008), and two geographically widespread taxa,H. cineraceus Blyth, 1853 and H. ater Temp leton,1848. The study was based on a thorough review ofthe existing literature and extensive recent fieldwork in Thailand (2006–2009), and some previousfield work in Myanmar (2000–2003, 2006) andnorthern Malaysia (2002–2004).

MATERIALS AND METHODS

Study Areas

New material for this study was collected from 28 localitiesin 17 provinces of Thailand, four localities in three states in

Acta Chiropterologica, 12(1): 29–50, 2010PL ISSN 1508-1109 © Museum and Institute of Zoology PAS

doi: 10.3161/150811010X504572

A taxonomic review of Hipposideros halophyllus, with additional information on

H. ater and H. cineraceus (Chiroptera: Hipposideridae) from Thailand and Myanmar

BOUNSAVANE DOUANGBOUBPHA1, 2, 8, SARA BUMRUNGSRI2, PIPAT SOISOOK3, SUSAN W. MURRAY4, SÉBASTIEN J. PUECHMAILLE5, CHUTAMAS SATASOOK2, 3, SI SI HLA BU6, DAVID L. HARRISON7, and PAUL J. J. BATES7

1Faculty of Environmental Sciences, National University of Laos, Dong Dok Campus, Xaythany District, Vientiane Capital, Lao PDR

2Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai 90112, Thailand 3Princess Maha Chakri Sirindhorn Natural History Museum, Faculty of Science, Prince of Songkla University,

Hat Yai 90112, Thailand4Department of Biology, Boston University, 5 Cummington Street, Boston, MA 02215, USA

5School of Biological and Environmental Sciences, University College Dublin, Belfield, Dublin 4, Ireland6Department of Zoology, Mandalay University, Mandalay, Myanmar

7Harrison Institute, Centre for Systematics and Biodiversity Research, Bowerwood House, St. Botolph’s Road, Sevenoaks, Kent, TN13 3AQ, Great Britain

8Corresponding author: E-mail: [email protected]

Based on recent field surveys in Thailand, Myanmar, and northern peninsular Malaysia, this paper reviews the taxonomy,morphometric and acoustic characters, distribution and ecology of the little known, globally endangered species Hipposideroshalophyllus. It lists nine new localities records, including the first from northern and peninsular Thailand, which represent a substantial increase in the species’ known range; it confirms the record from northern peninsular Malaysia. In addition, it providesfurther information on two other small species of the Hipposideros bicolor group, H. ater and H. cineraceus.

Key words: Hipposideros halophyllus, H. ater, H. cineraceus, distribution, ecology, echolocation, baculum, Southeast Asia

Myanmar, and one locality in peninsular Malaysia. In order toprovide information for further research, brief descriptions ofthese sites are given below.

ThailandChiang Mai Province: (1) Pha Dang Cave, Srilanna Nation -

al Park, Chiang Dao District, 19°20.769’N, 99°01.416’E, 480 ma.s.l. [loc. H1, Fig. 1; loc. C9, Fig. 2], surveyed on 22–23 Octo -ber, 2006. This limestone outcrop, which has several caverns in-side, is surrounded by hill evergreen forest and ricefields. Sevenspecies were collected in the cave and its vicinity: Hipposideroshalophyllus, H. cineraceus, H. larvatus, Taphozous longima-nus, Rhinolophus coelophyllus, R. malayanus, and R. pusillus.(2) Khimee Cave, Chiang Dao Wildlife Sanctuary, KhongDistrict, 19°21.266’N, 98°43.837’E, 718 m a.s.l. [loc. C8, Fig.2], was surveyed in April 2006. This large limestone cavern issurrounded by mixed deciduous and some dry evergreen forestand was the roosting site of H. cineraceus.

Phitsanulok Province: Tham Pha Cave, Tha Phol Non-hunt-ing area, Neon Maprang District, 16°30’N, 100°40’E [loc. H2,Fig. 1] was surveyed on 11 January, 2000. This small group oflimestone outcrops is situated in a flat area and surrounded bysugar cane plantations and fruit orchards. The vegetation on theoutcrops is primarily shrubby woodland. During the survey thearea was very dry. Seven species of bat were collected fromcave surveys and harp trapping: Cynopterus sphinx, Hipposi -deros pomona, Rhinolophus thomasi, R. coelophyllus, R. micro -globosus, Megaderma spasma, and Tadarida plicata.

Uthai Thani Province: (1) Lup Lae Cave, Ban Rai District,15°03.077’N, 99°28.879’E, 200 m a.s.l., [loc. H4, Fig. 1; loc.C13, Fig. 2], was surveyed on 23 March, 2007. It is situated ina limestone outcrop, which is covered with mixed deciduousand bamboo forest and surrounded by tamarind and mangoplantations, and corn fields. Eight species of bat were collectedwithin the cave and its vicinity: Hipposideros halophyllus,H. cineraceus, H. larvatus, H. pomona, Megaderma spasma,Rhinolophus malayanus, R. coelophyllus, and R. yunannensis.(2) Khi Nok Cave, Huai Kha Khaeng Wildlife Sanctuary15°37.222’N, 99°18.210’E, 380 m a.s.l. [loc. H3, Fig. 1; loc.C11, Fig. 2], was surveyed on 19 February 2008. A harp trapwas set at the entrance of the cave, which was situated in a lime-stone outcrop surrounded by bamboo forest. Six species werecaptured: R. malayanus, H. pomona, H. cineraceus, H. halo-phyllus, H. larvatus, and H. armiger. (3) Rathana Kiri Cave,Khao Kwang Thong District, 15°22.282’N, 99°40.984’E, 88 ma.s.l. [loc. C12, Fig. 2], was surveyed on 1 May, 2007. A harptrap was set at the entrance of the cave which is situated in a limestone outcrop. Seven species were captured: H. cinera -ceus, H. pomona, H. larvatus, R. coelophyllus, R. thomasi,R. mala ya nus, and Myotis siligorensis.

Lop Buri Province: (1) Khao Samor Khon, Ta Wung Dis -trict, 14°54.548’N, 100°30.342’E, 3 m a.s.l. [loc. H7, Fig. 1],

was surveyed on 24 January, 2007. A harp trap, which was setacross a small trail in jamun (Syzygium cumini) forest, collectedH. halophyllus, R. coelophyllus, R. mala yanus, and R. pusillus.Subsequently, on 21 March, 2007, hand nets were used in thenearby Ob Cave, 14°54.525’N, 100°29.491’E, 38 m a.s.l. [alsoloc. H7, Fig. 1]. Situated in a lime stone outcrop, it is coveredwith mixed deciduous and dipterocrap forests and surroundedby ricefields. Two species were collected: H. halophyllus and M. si ligorensis. (2) Khao Don Deung, Ban Mi District,15°08.888’N, 100°36.819’E, 40 m a.s.l. [loc. H6, Fig. 1; loc.C14, Fig. 2], was surveyed on 22 March, 2007. It is a limestoneoutcrop covered by mixed deciduous and dipterocarp forestsand surrounded by teak plantations, sun flower and corn fields.Eight species were collected H. halophyllus, H. cineraceus, H. larvatus, H. pomona, Rhinolophus coelophyllus, R. mala ya -nus, R. microglobosus, and R. pusillus.

Sara Buri Province: Ton Chan Cave, Phaphouthabat Dis -trict, 14°43.193’N, 100°47.684’E, 33 m a.s.l, [loc. H8, Fig. 1],was surveyed on 21 March, 2007. The cave, which is in a lime-stone outcrop, includes a Chinese temple and is surrounded byvillages and mixed deciduous forest. Only H. halophyllus wasfound.

Sa Kaeo Province: Khao Singto, Meuong District,13°59.417’N, 102°00.465’E, 12 m a.s.l. [loc. H9, Fig. 1; loc.C17, Fig. 2], was surveyed on 20 March, 2007. The limestoneoutcrop is covered by mixed deciduous forest and surroundedby a eucalyptus plantation. Seven species were collected fromwithin the cave and its vicinity: H. halophyllus, H. ci ne ra -ceus, H. galeritus, H. larvatus, H. pomona, M. spasma, andR. shameli.

Kanchana Buri Province: Ma Duea Fun Caves, 14°01.822’N,99°23.887’E, 60 m a.s.l. [loc. H10, Fig. 1] and Ma Duea,14°01.673’N, 99°24.095’E, 64 m a.s.l. [loc. H11, Fig. 1; loc.C19, Fig. 2], Sri Mongkhon District, were surveyed on 18 De -cember, 2007. This limestone outcrop, which is covered withmixed deciduous and bamboo forests, is adjacent to the KwaeNoi River. Five species were collected: H. halophyllus, H. cine -raceus, H. pomona, Aselliscus stoliczkanus, and R. malayanus.

Ratcha Buri Province: Khao Bin Cave, Chom Bung Dis -trict, 13°35.527’N, 99°40.012’E, 61 m a.s.l. [loc. C20, Fig. 2],was surveyed on 9 February, 2007 and 24 March, 2007. Thislimestone outcrop, which is covered with mixed deciduous andbamboo forests, is a tourist attraction. Five species were collect-ed: H. cineraceus, H. pomona, A. sto licz kanus, R. malayanus,and R. thomasi.

Phetcha Buri Province: Khao Yoi Cave, Khao Yoi District,13°14.014’N, 99°49.708’E, 53 m a.s.l. [loc. H14, Fig. 1; loc.C21, Fig. 2], was surveyed on 11 February, 2007. It is situatedin a limestone outcrop and frequented by tourists. This outcropis surrounded by a mixed deciduous forest, villages, ricefieldsand a temple. Eight species were collected in the vicinity of the cave: H. halophyllus, H. cineraceus, H. larvatus, H. pomona,R. coelophyllus, R. mala yanus, R. pusillus, and M. siligorensis.

30 B. Douangboubpha, S. Bumrungsri, P. Soisook, S. W. Murray, S. J. Puechmaille, et al.

FIG. 1. Distribution map of H. halophyllus — black triangles represent localities visited by one or more authors during the currentstudy; open triangles are literature records. H1: Pha Dang Cave, Chiang Mai Province; H2: Tham Pha Cave, Phitsanulok Province;H3: Khi Nok Cave, Uthai Thani Province; H4: Lup Lae Cave, Uthai Thani Province; H5: Vi Moke Cave, Lop Buri Province; H6:Khao Don Deung, Lop Buri Province; H7: Khao Samor Khon, Lop Buri Province; H8: Ton Chan Cave, Sara Buri Province; H9:Khao Singto, Sa Kaeo Province; H10: Ma Duea Fun Cave, Kanchana Buri Province; H11: Ma Duea Cave, Kanchana Buri Province;H12: Khao Chong Phran, Ratchaburi Province; H13: Khao Bin Cave, Ratchaburi Province; H14: Khao Yoi Cave, Phetcha BuriProvince; H15: Koh Mai Pai, Phang Nga Province; H16: Khaun Khang Peatswamp, Nakon Srithammarat Province; H17: Bukit

Jerneh and Tumang Lembing Caves, Perlis State

�

A taxonomic review of Hipposideros halophyllus from Thailand and Myanmar 31

Chumphon Province: (1) Khao Kram Cave, Patiew District,10°55.131’N, 99°22.440’E, 67 m a.s.l. [loc. C22, Fig. 2], wassurveyed on 10 October 2006. The cave, which includes a tem-ple, is surrounded by a rubber plantation. Four species were collected: H. cineraceus, H. larvatus, H. pomona, and R. affinis.(2) Silawan Cave, Patiew District, 10°41.461’N, 99°14.476’E,68 m a.s.l. [loc. C23, Fig. 2], was surveyed on 11 January, 2007.Situated in a limestone outcrop, it is surrounded by disturbedforest, oil palm, rubber and orchard plantations. Seven specieswere collected in the area: H. cineraceus, H. larvatus, H. gale-ritus, R. lepidus, R. malayanus, M. siligorensis, and Tylo nyc terispachypus.

Ranong Province: Pra Kayang Cave, Kraburi District,10°19.569’N, 98°45.923’E, 3 m a.s.l. [loc. C24, Fig. 2], wassurveyed on 13 January, 2007. It is situated in a limestone out-crop, which is covered with deciduous forest and surrounded bynipa palm (Nipa fruticans) and mangrove forest. Six specieswere collected in the vicinity of the cave: H. cineraceus, H. ga-leritus, H. lekaguli, R. affinis, Myotis horsfieldi, and Miniopte -rus medius.

Phang Nga Province: (1) Ao Mai Ngam, Surin Island Na -tion al Park, 9°25.873’N, 97°51.975’E, 20 m a.s.l. [loc. C25,Fig. 2], was surveyed on 2 February, 2006. A specimen of H. cineraceus was collected on a trail in seasonal evergreen for-est. (2) Koh Mai Pai, Muang District, 8°21.364’N, 98°32.429’E,20 m a.s.l. [loc. H15, Fig. 1; loc. C26, Fig. 2], was surveyed on27 June, 2008. Harp traps were set in moist evergreen forestnear a limestone outcrop, which is surrounded by mangrove for-est. Eight species were captured at this site: H. halo phyl lus,H. cineraceus, H. bicolor, H. galeritus, H. larvatus, R. co elo -phyllus, R. malayanus, and R. stheno.

Trang Province: Khao Chong Waterfall, Khao BanthatWildlife Sanctuary, 7°32.894’N, 99°47.196’E, 81 m a.s.l. [loc.C27, Fig. 2], was surveyed on 20 January, 2007. Seven specieswere collected in secondary forest: H. cineraceus, H. bicolor,H. larvatus, R. affinis, R. lepidus, Keri voula hardwickii, and Cy -no pterus brachyotis.

Songkla Province: (1) Khao Rak Kiat Cave, Rattaphum Dis -trict, 6°38.767’N, 99°37.383’E, 100 m a.s.l. [loc. C33, Fig. 2],was surveyed on 16 December, 2006. Situated in a limestoneoutcrop, it is surrounded by disturbed forest and a rubber plan-tation. Six species were collected in the cave and its vicinity: H. cineraceus, H. larvatus, R. affinis, R. le pidus, R. stheno, andM. medius. (2) Wildlife Educa tion Centre, Ton Nga Chang Wild -life Sanctuary, Rat taphum District, 6°56.739’N, 100°14.467’E,107 m a.s.l. [loc. C31, Fig. 2], was surveyed on 29 November,

2006. Harp traps were set in lowland tropical rain forest. Tenspecies were collected: H. cineraceus, H. bicolor, H. diadema,Co elops frithii, R. acuminatus, R. affinis, R. trifoliatus, Keri vou -la hardwickii, Murina cyclotis, and Megaerops ecaudatus. (3)Khao Nouy Cave, Rattaphum District, 6°59.537’N, 100°08.470’E,130 m a.s.l. [loc. C32, Fig. 2], was surveyed on 7 October, 2006and 7 February, 2007. Situated in a limestone outcrop, it is sur-rounded by disturbed forest, plantations, and human habitation.Three species were collected in the cave and its vicinity: H. ci -ne raceus, H. bicolor, and H. armiger. (4) Outa phao Watershed,Rattaphum District, 6°47.777’N, 100°14.092’E, 197 m a.s.l.[loc. C30, Fig. 2], was surveyed on 25 February, 2007. Harp trapswere set in a rubber plantation and on a trail in disturbed forestnear a stream. Six species were collected: H. cineraceus, H. bi-color, H. larvatus, R. affinis, K. hardwickii, and M. cyclotis.

Nakon Srithammarat Province: Khaun Khang Peatswamp,Thalenoi Non-Hunting Area, Cha Aod District, 7°53.780’N,100°6.347’E, [loc. H16, Fig. 1], was surveyed on 31 January2009. Harp traps were set on trails adjacent to pools in man-grove forest near to a river and about 500 meters from a smalllimestone outcrop. The area is surrounded by rubber plantationsand villages. Four species of bats were captured: R. affinis,R. lepidus, R. stheno, and H. halophyllus.

Satun Province: (1) Talow Wao-Talow Oulang Road, Taru -tao Islands National Park [loc. C36, Fig. 2] was surveyed on 7 March, 2007. The harp traps were set at two sites on the roadfrom Talow Wao to Talow Oulang. The first, under a ca nopy oftrees, surrounded by old-growth dry evergreen forest (lowland),was between km 1–2 (6°36.501’N, 99°40.435’E, 73 m a.s.l.).The second in lowland evergreen forest, close to a limestoneoutcrop was at km 6 (6°39.500’N, 99°40.833’E, 100 m a.s.l.).Five species were found: H. cineraceus, H. bicolor, R. coelo-phyllus, R. lepidus, and K. hardwickii. (2) Crocodile (Jorrake)Cave, Tarutao Island National Park, 6°41.946’N, 99°39.096’E,65 m a.s.l. [also loc. C36, Fig. 2], was surveyed on 8 March,2007. A river runs through the limestone cave, which is coveredin lowland dry evergreen forest and surrounded by mangroveforest and sea. One species was collected: Hipposideros ciner-aceus. (3) Ao Son, Tarutao Islands National Park, 6°38.767’N,99°37.383’E, 5 m a.s.l [also loc. C36, Fig. 2]. A specimen of H. cineraceus was collected on 3 March, 2003 in moist ever-green forest close to the beach.

Narathiwat Province: Sirinthorn Waterfall, Bala Forest,Hala-Bala Wildlife Sanctuary, 5°48.017’N, 101°50.000’E, 300 m a.s.l. [loc. C37, Fig. 2]; a specimen of H. cineraceus wascollected on 5 July, 2003 in lowland evergreen forest.


FIG. 2. Distribution map of H. ater (A: circles) and H. cineraceus (C: triangles) — black symbols represent localities visited by oneor more authors during the current study; open symbols are literature records. A1: Kan Thar Yar Beach, Rakhine State; A2: Myeik,Tanintharyi Division; A3: Tharabwin Village, Tanintharyi Division. C1: Nam Tamai Valley, Kachin State; C2: Mogok, MandalayDivision; C3: Gokteik, Shan State; C4: Mingun, Mandalay Division; C5: nr Sagaing, Sagaing Division; C6: Saddan Sin Cave, MonState; C7: Nagamauk II, Mon State; C8: Khimee Cave, Chiang Mai Province; C9: Pha Dang Cave, Chiang Mai Province; C10: BanJagae and Lum Khao Ngu GSs., Tak Province; C11: Khi Nok Cave, Uthai Thani Province; C12: Rathana Kiri Cave, Uthai ThaniProvince; C13: Lup Lea Cave, Uthai Thani Province; C14: Khao Don Deung, Lop Buri Province; C15: Khao Yai NP., NakhonRatchasima Province; C16: Phu Nam Tok Tub Kwang (Tap Kwang), Sara Buri Province; C17: Khao Singto Cave, Sa Khao Province;C18: Wat Khao Wong Kot, Chantha Buri Province; C19: Ma Duea Cave, Kanchana Buri Province; C20: Khao Bin Cave, Ratcha BuriProvince; C21: Khao Yoi Cave, Phetcha Buri Province; C22: Khao Kram Cave, Chumphon Province; C23: Silawan Cave, ChumphonProvince; C24: Pra Kayang Cave, Ranong Province; C25: Ao Mai Ngam, Phang Nga Province; C26: Koh Mai Pai, Phang NgaProvince; C27: Khao Chong Waterfall, Trang Province; C28: Noy Cave, Songkla Province; C29: Syson Cave, Songkla Province;C30: Outaphao Watershed, Songkla Province; C31: Wildlife Education Centre, Songkla Province; C32: Khao Nouy Cave, SongklaProvince; C33: Khao Rak Kiat Cave, Songkla Province; C34: Ka Cave, Songkla Province; C35: Ban Wang Bla Chan, Satun

Province; C36: Tarutao NP., Satun Province; C37: Sirinthorn Waterfall, Narathiwat Province

�

MyanmarRakhine State: Kan-Thar-Yar, Gwa Township, 17°44’N,

94°32’E, 1 m a.s.l. [loc. A1, Fig. 2]; a single specimen of H. aterwas collected on 15 November, 2000 whilst flying around thebathroom of a wooden chalet next to a sandy beach. Numerouspalm trees were growing in the vicinity. The area also includeddisturbed lowland forest, other human habitation and orchards.

Mon State: (1) Saddan Sin Cave, 16 km north-east of Maw -la myine, 16°19’N, 97°42’E, [loc. C6, Fig. 2]. A single specimenof H. cineraceus was collected on 2 November 2002 in the cave,which is situated in a large isolated limestone outcrop setamongst extensive paddy fields and many toddy palms. (2) Na -gamauk II, 16°18.992’N, 97°42.352’E, 29 m a.s.l. [loc. C7, Fig.2] was surveyed on 5 and 7 December 2006. Mist-nets were set-up at the entrance of the cave where eight species were cap -tured: A. stoliczkanus, Cra se o nycteris thonglongyai, Pipi strel -lus sp., H. cineraceus, H. po mo na, M. spasma, R. malaya nus,and Taphozous melanopogon.

Tanintharyi Division: Tharabwin Village, 12°18’N, 99°04’E[loc. A3, Fig. 2], a single specimen of H. ater was collected on13 November, 2003 in the roof of a village house. Tharabwinvillage is surrounded by agricultural land and some remnants ofdisturbed lowland forest.

MalaysiaPerlis State: Bukit Jerneh Cave and Tumang Lembing

Cave, c.o. 6°32’N, 100°16‘E [loc. H17, Fig. 1]. The caves arelocated in adjacent limestone hills (Bukit Jerneh and BukitTumang Lembing, respectively) approximately 25 km from the border with Thailand and are part of the Chuping formation,a group of individual limestone outcrops running north to southin Central Perlis State (Price, 2001). The hills are relativelyclose to Perlis State Park, which is contiguous with Thale BanNational Park in Thailand and includes the Nakawan Range oflimestone hills. The natural vegetation of both parks compriseevergreen rain forest and mixed deciduous forest. However,Butkit Jerneh cave and Tumang Lembing cave are surroundedby mixed agricultural land including fruit and rubber planta-tions, and rice fields. Tumang Lembing cave has three main entrances, each associated with a large chamber, and water waspresent in one of the chambers. Gua Bukit Jerneh consists of a river flowing through Jerneh Hill, with two main entrances.Spe cies collected at the caves included: H. armiger, H. halo-phyllus, H. bicolor, H. larvatus, H. pomona, H. sp. (most likelyH. cineraceus), R. malayanus, R. affinis, M. spasma, T. melano-pogon, Myotis ridleyi, M. ater, and Minio pterus sp. A colony offruit bats was observed in both caves but no individuals werecollected.

Measurements

In this study, 86 voucher specimens were measured. Theyare held in the Princess Maha Chakri Sirindhorn Natural HistoryMuseum, Prince of Songkla University, Thailand (PSUZC), theChiang Dao Wildlife Research Station, Thailand (CD), theHarrison Institute, UK (HZM and MM [field numbers]) and theNatural History Museum, London, UK (BMNH).

Measurements were taken with a digital caliper and whererelevant followed Bates and Harrison (1997). They included:FA: forearm length – from the extremity of the elbow to the extremity of the carpus with the wings folded; TL: tail length —from the tip of the tail to its base adjacent to the anus; TIB: tibia length — from the knee joint to the extremity of the heel

behind the os calcis; HF: foot — from the extremity of the heelbehind the os calcis to the extremity of the longest digit, not including the hairs or claws; EAR L: ear length — from the low-er border of the external auditory meatus to the tip of the pinna;EAR W: ear width – greatest width across the pinna; 3MT, 4MT,5MT: the distance from the extremity of the carpus to the distalextremity of the third, fourth and fifth metacarpals, respective-ly; 3D1P, 3D2P, 4D1P, 4D2P — length of the first and secondphalanges of the third and fourth digits, respectively;3D1P/3MTx100 — % length of first phalanx of third digit rela-tive to its metacarpal; BL: greatest length of baculum; GTL:greatest length of skull — the greatest antero-posterior diameterof the skull, from the most projecting point at each extremity regardless of what structure forms these points; CCL: condylo-canine length — from the exoccipital condyle to the anterioralveolus of the canine; CBL: condylo-basal length — from theexoccipital condyle to the alveolus of the anterior incisor; MW:mastoid width — the greatest distance across the mastoid re-gion; ZB: zygomatic breadth — the greatest width of the skullacross the zygomata; BB: breadth of the braincase — width ofthe braincase at the posterior roots of the zygomatic arches; PC:postorbital constriction — the narrowest width across the con-striction posterior to the orbits; ML: mandible length — fromthe most posterior part of the condyle to the most anterior partof the mandible, including the lower incisors; C1–C1: anteriorpalatal width — taken across the outer borders of the upper ca-nine; M3–M3: posterior palatal width — taken across the outerborders of the third upper molar; C–M3: upper toothrow length– from the front of the upper canine to the back of the crown ofthe third upper molar; C–M3: lower toothrow length — from thefront of the lower canine to the back of the crown of the thirdlower molar; RW: rostral chambers width — greatest widthacross the rostral chambers. MASS: recorded in grams (g).

Sound Analysis

The echolocation calls were detected with a PetterssonUltrasound Detector D 240X, which was set at 10× time expan-sion rate and 1.7 seconds max storage time. Calls were digitisedand recorded on to a MP3 Recorder (Multi-codec Jukebox) oran iHP-120 Recorder (Multi-codec Jukebox). The recordingswere converted from MP3 to Wave File using the softwareGold Wave. The sound was analyzed using the software Bat SoundPro v3.1 (Pettersson Elektronik AB). For each bat, ten pulseswere chosen over the period of the recorded sound, to avoidpossible periodical changes. The spectrogram was analyzed us-ing Power spectrum, FFT (Fast Fourier Transforms) size 1024and a Hanning window. The frequency of maximum energy(FMAXE) corresponds to the frequency of the constant frequen-cy segment of the second harmonic. The sound was measured in each spectrum, with the score for each specimen based on the mean of the ten measurements. Owing to constraints withthe detector, in H. halophyllus, the frequency was sometimes estimated by measuring the first harmonic and doubling it, andsometimes by measuring the second harmonic directly.

Morphometric Analysis

To determine whether there was sexual dimorphism withinthe taxa and significant differences in metric characters betweentaxa, a series of T-tests were run at a confidence limit of 95%.Principal Components Analysis (PCA) performed on the corre-lation matrix was used for multivariate comparisons.



RESULTS

Morphometrics

External measurements were available for 81spec imens from the study area and all descriptionsbelow are based on this material, unless other-wise stated. Significant variation in size between the sexes was found in five external measurementsfor H. halophyllus and two external measure-ments for H. cineraceus (Table 1). These measure-ments were mostly concerned with aspects of thewing, especially metacarpal length. External meas-urements were available for only two specimens ofH. ater from the study area, both were females.Males and females of H. halophyllus and H. cine-ra ceus were treated separately in all subsequent analyses.

Hipposideros cineraceus is significantly smallerthan H. halophyllus in all external measurements(Table 2) except for ear length (EAR L) and width (EAR W) and the first phalanx of the third digit(3D1P), in which it exceeds H. halophyllus in size.Hippo si deros ater appears to be a larger taxon based on external measurements but the sample size (n = 2) is too small to make any meaningful con-clusions. A multivariate analysis based on eight ex ternal characters for female specimens onlyshows that H. ater, H. halophyllus and H. cinera -ceus form three discrete groupings (Fig. 3 andTable 4).

Based on material from the study area, it is ap-parent that the shape of the noseleaf is diagnostic(Figs. 4 and 5). In all specimens, H. halophyllushas a highly characteristic kidney-shaped internarialseptum, a narrow anterior leaf with deep emargi-nation on its anterior border, and is without supple -men tary leaflets. In H. cineraceus, the internarialseptum is small, parallel-sided or slightly triangularin shape, the anterior leaf is without a deep emar-gination and there are no supplementary leaflets. InH. ater from Myanmar, the internarial septum isslightly inflated and triangular in shape, the anteriorleaf is without an emargination and there is one pairof rudimentary supplementary leaflets.

Penis and baculum shape also appear to be diag-nostic in helping to discriminate between the threetaxa (Figs. 6 and 7). In H. halophyllus, the penis isthin with a pointed tip and the baculum is minute,less than 0.5 mm in length, with a straight shaft,slightly expanded base and a rounded tip. In H. cine -raceus, the penis is short and thin with a bluntlyrounded tip; the baculum is 1.8–2.5 mm in length,

with a narrow shaft, rounded base and characteristi-cally bifid tip. No male specimens of H. ater fromMyanmar with intact genitalia were available forthis study. However, in specimens of H. ater fromIndia, the penis is relatively large with a rounded tipand the baculum has a simple rounded tip and a nar-row shaft.

Eighty-three skulls were measured including fiveH. ater from the study area. No significant differ-ence in skull size was found between male and fe-male specimens of either H. halophyllus or H. cine -ra ceus (Table 3). Insufficient data were available forH. ater to test for sexual dimorphism. In contrast tothe external body measurements, the skulls of H. halophyllus averaged the smallest in all measure-ments except for mastoid width (MW), zygomaticbreadth (ZB), and anterior palatal width, takenacross the upper canines (C1–C1), where they areequal to or slightly exceed H. cineraceus (Tables 2and 3). The postorbital constriction (PC) is particu-larly reduced in H. halophyllus in comparison to theother two taxa. H. ater appears to have the longest and broadest skulls overall. However, toothrowlengths only slightly exceed those of H. cineraceus.A multivariate comparison based on nine cranialcharacters for female specimens only shows a mi-nor overlap between H. halophyllus and H. cinera -ceus, with H. ater forming a discrete group (Fig. 8and Table 5).

The rostrum of H. halophyllus is characteristical-ly downward sloping when viewed laterally, where-as it is more robust in H. cineraceus and H. ater(Fig. 9). The dorsal process on the posterior part ofeach zygoma is variably developed. In some speci-mens of H. halophyllus it is scarcely evident, where-as in others it is tall and almost semi-circular in out line. In H. cineraceus, the zygomata are usuallywith out processes, if present they are low and com parable in size to the smallest of those in H. ha -lophyllus. In H. ater it varies from well developed in some specimens to scarcely evident in others.

Echolocation

Echolocation calls of H. halophyllus and H. cine -raceus were recorded at 19 localities in Thailand(nine localities for H. halophyllus and 16 locali-ties for H. cineraceus). The hand held frequency(FMAXE) of H. halophyllus averaged 177.6 kHz formales and 181.6 kHz for females in Thailand(Tables 1 and 2, Fig. 10). The mean call frequencyfor H. halophyllus in Malaysia was slightly higher,185.1 kHz for 16 individuals, and ranged from 181.0

36 B. Douangboubpha, S. Bumrungsri, P. Soisook, S. W. Murray, S. J. Puechmaille, et al.T

AB

LE

1. E

xter

nal

mea

sure

men

ts (

in m

m),

ech

oloc

atio

n ca

ll (

in k

Hz)

and

bod

y m

ass

(in

g) o

f th

ree

spec

ies

of H

ippo

side

ros.

FM

AX

E—

fre

quen

cy o

f m

axim

um e

nerg

y; F

A —

for

earm

leng

th; T

L —

tail

; TIB

— ti

bia;

HF

— f

oot;

EA

R L

— e

ar le

ngth

; EA

R W

— e

ar w

idth

; 3M

T, 4

MT,

5M

T —

thir

d, f

ourt

h, f

ifth

met

acar

pal l

engt

hs; 3

D1P

, 3D

2P, 4

D1P

, 4D

2P —

fir

st a

ndse

cond

pha

lang

es o

f th

e th

ird

and

four

th d

igit

s; 3

D1P

/3M

Tx1

00 —

% l

engt

h of

fir

st p

hala

nx o

f th

ird

digi

t re

lati

ve t

o th

e th

ird

met

acar

pal;

BL

— b

acul

um l

engt

h; M

AS

S —

bod

y m

ass.

Mea

n, r

ange

and

sta

ndar

d de

viat

ion;

sam

ple

size

s di

ffer

ing

from

thos

e re

port

ed u

nder

nar

e gi

ven

in p

aren

thes

es.T

-tes

t com

pari

son

for

sexu

al d

imor

phis

m (

sex.

dim

.; ns

= n

ot s

igni

fica

nt)

nS

exF

MA

XE

FAT

LT

IBH

FE

AR

LE

AR

W3M

T4M

T

Hip

posi

dero

s ha

loph

yllu

s (T

hail

and)

15♂

♂17

7.6,

9.0

37

.6, 0

.725

.5, 1

.817

.6, 0

.65.

8, 0

.413

.3, 1

.010

.7, 0

.630

.0, 0

.830

.9, 0

.615

6.4–

187.

9 (1

4)36

.4–3

8.7

22.4

–28.

7 (1

4)16

.4–1

8.5

4.9–

6.4

11.1

–15.

49.

9–11

.628

.7–3

1.3

29.5

–31.

513

♀♀

181.

6, 6

.938

.2, 0

.7

26.7

, 1.6

17.7

, 0.8

5.

8, 0

.513

.1, 1

.1

11.2

, 0.5

31.0

, 0.8

31.7

, 0.5

160.

6–18

6.2

(12)

37.2

–39.

223

.8–2

8.6

16.4

–18.

64.

5–6.

311

.4–1

5.1

10.3

–12.

4 (1

2)29

.5–3

2.2

(12)

30.7

–32.

5 (1

2)S

ex. d

im.

nsns

nsns

nsns

P=

0.0

31P

= 0

.018

P=

0.0

01

Hip

posi

dero

s ci

nera

ceus

(Tha

ilan

d an

d M

yanm

ar)

25♂

♂14

9.0,

5.3

34.6

, 0.9

24.2

, 2.1

15.1

, 0.5

5.3,

0.5

16.1

, 1.5

12.2

, 0.9

25.3

, 0.9

27.9

, 0.8

134.

8–15

4.1

(21)

32.4

–36.

519

.0–2

7.8

14.5

–16.

24.

4–5.

913

.4–1

8.3

10.1

–13.

423

.7–2

7.6

26.4

–29.

726

♀♀

150.

1, 3

.635

.0, 0

.9

24.8

, 2.2

15.6

, 1.1

5.

2, 0

.416

.1, 1

.4

12.5

, 0.8

25.8

, 0.8

28.4

, 0.8

141.

4–15

4.1

(17)

33.3

–37.

2 20

.4–2

8.6

14.4

–19.

9 4.

3–5.

813

.4–2

0.2

10.4

–13.

624

.5–2

7.1

26.4

–29.

7S

ex. d

im.

nsns

nsns

nsns

nsns

P=

0.0

23

Hip

posi

dero

s at

er (M

yanm

ar)

2♀

♀–

39.5

, 40.

324

.7, 2

8.7

16.2

, 17.

86.

5, 7

.016

.6, 1

6.7

12.3

, 12.

431

.0, 3

3.4

33.1

, 34.

8

–187.6 kHz. The frequencies of H. cine raceus weresignificantly lower, 149.1 kHz for males and 150.1for females (Tables 1 and 2, Fig. 11). Unfortunately,no echolocation data are available for H. ater.

SYSTEMATIC DESCRIPTIONS

The results indicate that the three small speciesof H. bicolor group in Thailand and Myanmar areclearly distinct. However, the taxonomic status of H. ater (sensu Hill, 1963) in Myan mar requires fur-ther study. It differs from H. halophyllus and H. ci -ne raceus, but also appears to differ in a number offeatures from H. a. ater described from Sri Lanka.The more important discriminating characters ofeach of the three species are discussed below, to-gether with information on the distribution, taxo-nomic status and ecology.

Hipposideros halophyllus Hill and Yenbutra, 1984Thailand Roundleaf Bat

Hipposideros halophyllus Hill and Yenbutra, 1984; Khao SamorKhon (=Khao Sa Moa Khon), Tha Woong (=Ta Woong),Lop Buri, Thailand

External and bacular charactersHipposideros halophyllus has a forearm length of

36.4–39.2 mm (Table 1). The ears are rounded witha broad, ill-defined tip (Fig. 4). The noseleaf is smalland without supplementary leaflets (Fig. 5). The in-ternarial septum is expanded into a small kidney-shaped, disc-like structure, with a width of 0.8–1.2mm. The anterior leaf is narrow, slightly elongatedand with a narrow but deep emargination in the an-terior border. There is a well-defined frontal sac be-hind the posterior leaf in both sexes. The fourthmetacarpal is about equal in length to the third butexceeds the fifth. The first phalanx of the third digitis short, between 46.5–49.6% the length of the rel-atively long third metacarpal. The penis is thin with a pointed tip (Fig. 6). The baculum is minute, < 0.5 mm in length, with a straight shaft, slightly expanded base and a rounded tip (Fig. 7).

Cranial and dental charactersThe skull is small and elongate, with a condylo-

canine length of 12.6–13.0 mm. The mastoid widthis 7.6–8.0 mm, which exceeds zygomatic breadth(Table 3). When viewed in lateral profile, the anteri-or part of the rostrum slopes gently downwards (Fig.9). The posterior lateral compartments of the ros-trum are well inflated; their posterior borders are almost straight and are clearly defined from the nar-row postorbital constriction. The sagittal crest is

well-developed anteriorly. Each zygoma has a dor-sal process on its posterior part which is variable insize, being scarcely evident in some specimens buttall and almost semi-circular in outline in others.The length of the upper toothrow (C–M3) is 4.6–5.0mm. The first upper premolar (P2) is very small,sometimes lying within the toothrow and sometimesextruded, and the upper canine (C1) is not in contactwith the second upper premolar (P4). The crownarea of the anterior, lower premolar (P2) is aboutequal to that of the second (P4), and about one half tothree quarters of its height.

EcholocationHipposideros halophyllus uses constant frequen-

cy echolocation calls. The frequency of maximumenergy (FMAXE) averaged 177.6 kHz (156.4–187.9kHz) for males and 181.6 kHz (160.6–186.2 kHz)for females based on calls by BD using a PetterssonD240X (Table 1). However, in KanchanaburiProvince, Thailand a frequency of 188.8–191.2 kHzwas recorded by SJP using a Pettersson D1000X,and in peninsular Malaysia a frequency range of181.0–187.6 kHz was recorded by SM using a Pet -tersson D960. Previously, S. Waengsothorn, J. Na -bhitabhata, K. Muangkhum, S. Charavuthanukul,and T. Moonchan (in litt.) had recorded calls of173–191 kHz for individuals from Lop BuriProvince. Individual variation seems to be relativelyhigh (Fig. 10). It exceeds variation between the sex-es and/or geographical variation and may possiblyreflect constraints with BD’s detector (see Materialsand Methods).

Taxonomic notesThere appears to be little geographical variation

and all populations are referred here to the nominaterace, H. h. halophyllus.

Conservation status and distributionHipposideros halophyllus was originally includ-

ed as ‘Lower Risk, near threatened’ in Hutson et al.(2001) and Simmons (2005). However, more recent-ly it was awarded ‘Endangered’ status in Boitani etal. (2006) and in IUCN (2009). Population size wasestimated at between 1,000–1,400 individuals andconsidered to be decreasing (Waengsothorn et al.,2006a, 2006b; IUCN, 2009).

Hipposideros halophyllus is known from Thai -land and Malaysia (Francis, 2008, based on S. Mur -ray, personal communication — Fig. 1). The currentstudy has extended its known range by approx-imately 435 km to north and includes the first


nS

ex5M

T3D

1P3D

2P4D

1P4D

2P3D

1P/3

MT

x100

BL

MA

SS

Hip

posi

dero

s ha

loph

yllu

s(T

hail

and)

15♂

♂27

.2, 0

.614

.5, 0

.514

.4, 0

.610

.1, 0

.47.

3, 0

.348

.3, 0

.90.

4, 0

.04.

1, 0

.226

.0–2

8.1

13.8

–15.

313

.2–1

5.3

9.5–

10.9

6.6–

8.0

46.5

–49.

60.

4–0.

4 (3

)4.

0–4.

8 (1

4)13

♀♀

27.8

, 0.4

15.0

, 0.3

14.8

, 0.3

10.4

, 0.3

(12

)7.

3, 0

.348

.3, 0

.8–

4.8,

0.8

27.1

–28.

5 (1

2)14

.6–1

5.6

(12)

14.2

–15.

3 (1

2)10

.0–1

0.9

6.7–

7.7

(12)

46.7

–49.

5 (1

2)–

4.0–

7.0

(12)

Sex

. dim

. P

= 0

.012

P=

0.0

06ns

nsns

nsP

= 0

.006

Hip

posi

dero

s ci

nera

ceus

(Tha

ilan

d an

d M

yanm

ar)

25♂

♂26

.7, 0

.915

.0, 0

.613

.5, 0

.89.

2, 0

.77.

1, 0

.559

.3, 1

.82.

0, 0

.33.

5, 0

.625

.3–2

8.9

14.1

–16.

612

.2–1

5.2

8.2–

15.2

6.3–

8.9

53.8

–62.

51.

8–2.

5 (7

)3.

0–5.

526

♀♀

27.0

, 0.7

15.4

, 0.5

13.9

, 1.3

9.3,

0.4

6.8,

0.5

59.6

, 1.8

–3.

9, 0

.525

.8–2

8.4

14.5

–16.

712

.1–1

9.1

8.5–

10.0

5.6–

7.9

57.0

–64.

1–

2.7–

5.0

Sex

.dim

.ns

P=

0.0

08ns

nsns

nsns

Hip

posi

dero

s at

er(M

yanm

ar)

2♀

♀31

.2, 3

3.2

15.9

, 16.

815

.3, 1

5.5

10.0

, 10.

37.

9, 8

.747

.7, 5

4.1

–4.

5 (1

)

TA

BL

E1.

Ext

ende

d

records from peninsular Thailand and provides fur-ther information on the record from Malaysia.

Thailand: Chiang Mai Province: Pha DangCave, Srilanna National Park, Chiang Dao District,(19°20.769’N, 99°01.416’E, 480 m a.s.l.) [loc. H1,Fig. 1] (PSUZC collection); Phitsanulok Province:Tham Pha Cave, Tha Phol Non-hunting area, NeonMaprang District (16°30’N, 100°40’E) [loc. H2,Fig. 1] (unpublished data of SB); Uthai ThaniProvince: Khi Nok Cave, Huai Kha Khang WildlifeSanctuary, Lan Sak (approx. 15°27’N, 99°17’E)[loc. H3, Fig. 1] (Yenbutra and Felten, 1986; Robin -son et al., 1995; and PSUZC collection); Lup LaeCave, Ban Rai District, (15°03.077’N, 99°28.879’E,200 m a.s.l.) [loc. H4, Fig. 1] (PSUZC collection);Lop Buri Province: Vi Moke Cave, Ban Mi District(approx. 15°08’N, 100°37’E) [loc. H5, Fig. 1] (Wa -thanakul, 1995); Khao Don Deung, Ban Mi District,(15°08.888’N, 100°36.819’E, 40 m a.s.l.) [loc. H6,Fig. 1] (PSUZC collection); Khao Samor Khon(Khao Sa Moa Khon; Khao Samokhon), Tawung

(Tha Woong) District, (14°54.548’N, 100°30.342’E,3 m a.s.l.) [loc. H7, Fig. 1] (PSUZC collection; Hilland Yen butra, 1984; Yenbutra and Felten, 1986; S. Waeng sothorn, J. Nabhitabhata, K. Muangkhum,S. Cha ravuthanukul, and T. Moonchan, in litt.); SaraBuri Province: Ton Chan Cave, PhaphoutthabatDistrict, (14°43.193’N, 100°47.684’E, 33 m a.s.l.)[loc. H8, Fig. 1] (PSUZC collection); Sa Kaeo Prov -ince: Khao Singto, Meung District, (13°59.417’N,102°00.465’E, 12 m a.s.l.) [loc. H9, Fig. 1] (PrachinBuri Province in Yenbutra and Felten, 1986; Waeng -sothorn et al., 2006a, 2006b; and PSUZC collec-tion); Kanchana Buri Province: Ma Duea Fun Cave(14°01.822’N, 99°23.887’E, 60 m a.s.l.) [loc. H10,Fig. 1] (SJP, unpublished data; Mahidol Univer-sity Collection); Ma Duea Cave (14°01.673’N,99°24.095’E, 64 m a.s.l.) [loc. H11, Fig. 1] (SJP, un-published data); Ratchaburi Province: Khao ChongPhran (approx. 13°40’N, 99°46’E) [loc. H12, Fig. 1](Hillman, 1999); Tham Khao Bin (= Khao BinCave) (13°35.527’N, 99°40.012’E, 61 m a.s.l.) [loc.

TABLE 2. Echolocation call (FMAXE in kHz), external and cranio-dental measurements (in mm) and body mass (in g) of males and females of H. halophyllus and H. cineraceus, presented as 0 ± SD. P = significance value, based on T-tests; ns = not significant

Measurements♂♂ ♀♀

H. halophyllus H. cineraceusP-value

H. halophyllus H. cineraceusP-value

Acoustic and external measurementsFMAXE 177.6 ± 9.0 149.0 ± 5.3 < 0.001 181.6 ± 6.9 150.1 ± 3.6 < 0.001FA 37.6 ± 0.7 34.6 ± 0.9 < 0.001 38.2 ± 0.7 35.0 ± 0.9 < 0.001TL 25.5 ± 1.8 24.2 ± 2.1 < 0.05 26.7 ± 1.6 24.8 ± 2.2 < 0.05TIB 17.6 ± 0.6 15.1 ± 0.5 < 0.001 17.7 ± 0.8 15.6 ± 1.1 < 0.001HF 5.8 ± 0.4 5.3 ± 0.5 < 0.001 5.8 ± 0.5 5.2 ± 0.4 < 0.001EAR L 13.3 ± 1.0 16.1 ± 1.5 < 0.001 13.1 ± 1.1 16.1 ± 1.4 < 0.001EAR W 10.7 ± 0.6 12.2 ± 0.9 < 0.001 11.2 ± 0.5 12.5 ± 0.8 < 0.0013MT 30.0 ± 0.8 25.3 ± 0.9 < 0.001 31.0 ± 0.8 25.8 ± 0.8 < 0.0014MT 30.9 ± 0.6 27.9 ± 0.8 < 0.001 31.7 ± 0.5 28.4 ± 0.8 < 0.0015MT 27.2 ± 0.6 26.7 ± 0.9 < 0.05 27.8 ± 0.4 27.0 ± 0.7 < 0.0013D1P 14.5 ± 0.5 15.0 ± 0.6 < 0.01 15.0 ± 0.3 15.4 ± 0.5 < 0.013D2P 14.4 ± 0.6 13.5 ± 0.8 < 0.001 14.8 ± 0.3 13.9 ± 1.3 < 0.054D1P 10.1 ± 0.4 9.2 ± 0.7 < 0.001 10.4 ± 0.3 9.3 ± 0.4 < 0.0014D2P 7.3 ± 0.3 7.1 ± 0.5 ns 7.3 ± 0.3 6.8 ± 0.5 < 0.0013D1P/3METx100 48.3 ± 0.9 59.3 ± 1.8 < 0.001 48.3 ± 0.8 59.6 ± 1.8 < 0.001MASS 4.1 ± 0.2 3.5 ± 0.6 < 0.001 4.8 ± 0.8 3.9 ± 0.5 < 0.001

Cranial and dental measurementsGTL 15.1 ± 0.1 15.5 ± 0.2 < 0.001 15.1 ± 0.2 15.6 ± 0.2 < 0.001CCL 12.9 ± 0.1 13.1 ± 0.2 < 0.001 12.8 ± 0.1 13.2 ± 0.2 < 0.001CBL 13.1 ± 0.1 13.4 ± 0.2 < 0.001 13.1 ± 0.2 13.5 ± 0.2 < 0.001MW 7.8 ± 0.1 7.8 ± 0.1 ns 7.8 ± 0.0 7.8 ± 0.1 nsZB 7.4 ± 0.1 7.3 ± 0.2 ns 7.3 ± 0.2 7.3 ± 0.2 nsBB 7.0 ± 0.2 7.2 ± 0.2 < 0.001 6.9 ± 0.2 7.3 ± 0.2 < 0.001PC 1.9 ± 0.1 2.6 ± 0.2 < 0.001 1.9 ± 0.1 2.5 ± 0.2 < 0.001ML 8.9 ± 0.1 9.0 ± 0.2 < 0.05 8.9 ± 0.1 9.1 ± 0.2 < 0.001C1–C1 2.8 ± 0.1 2.6 ± 0.2 < 0.01 2.8 ± 0.1 2.7 ± 0.2 < 0.05M3–M3 4.8 ± 0.1 5.0 ± 0.1 < 0.001 4.9 ± 0.1 5.0 ± 0.1 < 0.01C–M3 4.9 ± 0.1 5.0 ± 0.1 < 0.001 4.8 ± 0.1 5.0 ± 0.1 < 0.001C–M3 5.0 ± 0.2 5.2 ± 0.2 < 0.01 5.0 ± 0.2 5.3 ± 0.1 < 0.001RW 3.6 ± 0.1 3.9 ± 0.2 < 0.001 3.6 ± 0.1 4.0 ± 0.1 < 0.001



FIG. 3. PCA between first and second principal components based on eight external metric characters for female specimens only. Black circles represent H. halophyllus, black squares are H. ater and black triangles are H. cineraceus

H13, Fig. 1] (Hill and Yenbutra, 1984); PetchaburiProvince: Khao Yoi Cave, Khao Yoi District,(13°14.014’N, 99°49.708’E, 53 m a.s.l.) [loc. H14,Fig. 1] (PSUZC collection; Hill and Yenbutra,1984); Phang Nga Province: Koh Mai Pai, MuangDistrict, (8°21.364’N, 98°32.429’E, 20 m a.s.l.)[loc. H15, Fig. 1] (PSUZC Collection); NakonSritham marat Prov ince: Khaun Khang Peat swamp,Tha lenoi Non-Hunting Area, Cha Aod Dis trict(7°53.780’N, 100°6.347’E) [loc. H16, Fig. 1](PSUZC Collection).

Malaysia: Perlis State: Bukit Jerneh Cave(06°32.934’N 100°16.152’); Tumang LembingCave (06°33.234’N 100°16.268’E ) [loc. H17, Fig.1] (Francis, 2008).

Ecology and behaviour During this study, H. halophyllus was collected

at 13 localities in Thailand with an altitudinal rangeof sea level to 480 m a.s.l., with eight of the sites being below 55 m a.s.l. (for further details seeMaterials and Methods). In each case, it was collect-ed either within a limestone cave or in its vicinity. It is not known whether H. halophyllus is exclusive-ly dependent on limestone or whether this simply reflects a bias of collecting effort. The surround-ing vegetation included a range of forest types (hillevergreen, mixed deciduous, dipterocarp, bambooand mangrove). In Malaysia, it was also found in an area of limestone. Despite the previous report of Waengsothorn et al. (2006b), in both Thailand

and Malaysia, H. halophyllus appears to be toler-ant of highly disturbed habitats that include fruit and rubber plantations, orchards, rice paddy and cornfields. At six of the sites, it co-existed with H. cine -ra ceus (see Materials and Methods); other leaf-nosed bats found with it at various locations includ-ed H. armiger, H. bicolor, H. galeritus, H. larvatus,and H. pomona.

Hipposideros cineraceus Blyth, 1853Least Leaf-nosed Bat

Hipposideros cineraceus Blyth, 1853; Near Pind Dadan Khan,Salt range, Punjab

External and bacular charactersHipposideros cineraceus has a forearm length of

32.4–37.2 mm (Table 1). The ears are relativelylarger than those of H. halophyllus (Fig. 4). Thenoseleaf is small and lacks supplementary lateralleaflets. The internarial septum is small, parallel-sided or slightly triangular in shape, not kidney-shaped as in H. halophyllus and without a deepemargination on the anterior border of the anteriorleaf as found in this latter species (Figs. 4 and 5). A well developed frontal sac is sometimes present.The fourth metacarpal exceeds the fifth and third inlength. Compared to H. halophyllus, the first pha-lanx of the third digit appears relatively long sincethe third metacarpal is short (3D1P/3MTx100 = 53.8–64.1%). The tibia is short. The penis is short and thin with a bluntly rounded tip whereas

-8 -6 -4 -2 0 2 4 6PC1

PC

2

3

2

1

0

-1

-2

-3

-4

40 B. Douangboubpha, S. Bumrungsri, P. Soisook, S. W. Murray, S. J. Puechmaille, et al.T

AB

LE

3. C

rani

al m

easu

rem

ents

(in

mm

) of

thr

ee s

peci

es o

f H

ippo

side

ros.

GT

L —

gre

ates

t sk

ull

leng

th,

CC

L —

con

dylo

-can

ine

leng

th,

CB

L —

con

dylo

-bas

al l

engt

h, M

W —

mas

toid

wid

th, Z

B —

zyg

omat

ic b

read

th, B

B —

bre

adth

of

brai

ncas

e, P

C —

pos

torb

ital

con

stri

ctio

n, M

L —

man

dibl

e le

ngth

, C1 –

C1

— a

nter

ior

pala

tal

wid

th, M

3 –M

3—

pos

teri

or p

alat

al w

idth

,C

–M3

— u

pper

too

thro

w l

engt

h, C

–M3

— l

ower

too

thro

w, R

W —

ros

tral

wid

th. M

ean,

ran

ge a

nd S

D a

re i

nclu

ded.

T-t

est

com

pari

son

for

sexu

al d

imor

phis

m (

ns =

not

sig

nifi

cant

)

nS

exG

TL

CC

LC

BL

MW

ZB

BB

PC

ML

C1 –

C1

M3 –

M3

C–M

3C

–M3

RW

Hip

posi

dero

s ha

loph

yllu

s (T

hail

and)

15♂

♂15

.1, 0

.112

.9, 0

.113

.1, 0

.17.

8, 0

.17.

4, 0

.17.

0, 0

.21.

9, 0

.18.

9, 0

.12.

8, 0

.14.

8, 0

.14.

9, 0

.15.

0, 0

.23.

6, 0

.114

.8–1

5.2

12.6

–13.

012

.8–1

3.3

7.6–

8.0

7.2–

7.5

6.7–

7.3

1.8–

2.1

8.6–

9.1

2.7–

3.0

4.8–

5.0

4.8–

5.0

4.5–

5.5

3.5–

3.7

12♀

♀15

.1, 0

.2 (

11)

12.8

, 0.1

13

.1, 0

.27.

8, 0

.0

7.3,

0.2

6.

9, 0

.2

1.9,

0.1

8.

9, 0

.12.

8, 0

.14.

9, 0

.1

4.8,

0.1

5.

0, 0

.23.

6, 0

.1

14.8

–15.

312

.7–1

3.0

12.7

–13.

47.

8–7.

97.

1–7.

76.

6–7.

41.

8–2.

18.

7–9.

22.

6–2.

94.

7–5.

04.

6–5.

04.

6–5.

23.

5–3.

7S

ex. d

im.

nsns

nsns

nsns

nsns

nsns

nsns

ns

Hip

posi

dero

s ci

nera

ceus

(Tha

ilan

d an

d M

yanm

ar)

25♂

♂15

.5, 0

.3 (

24)

13.1

, 0.2

13.4

, 0.2

(24

)7.

8, 0

.17.

3, 0

.27.

2, 0

.22.

6, 0

.29.

0, 0

.22.

6, 0

.25.

0, 0

.1

5.0,

0.1

5.

2, 0

.23.

9, 0

.2

14.8

–16.

112

.9–1

3.6

12.9

–13.

87.

6–8.

17.

0–7.

76.

7–7.

62.

2–2.

98.

8–9.

4 (2

4)2.

3–2.

84.

7–5.

2 4.

8–5.

34.

7–5.

5 (2

4)3.

7–4.

226

♀♀

15.6

, 0.2

13.2

, 0.2

13

.5, 0

.27.

8, 0

.1

7.3,

0.2

7.

3, 0

.2

2.5,

0.2

9.

1, 0

.22.

7, 0

.25.

0, 0

.15.

0, 0

.15.

3, 0

.14.

0, 0

.115

.2–1

6.1

12.8

–13.

712

.9–1

4.1

7.5–

8.1

6.9–

7.6

6.9–

7.6

2.3–

2.9

8.9–

9.6

2.2–

2.9

(24)

4.8–

5.2

4.7–

5.2

5.0–

5.6

3.7–

4.1

Sex

. dim

.ns

nsns

nsns

nsns

nsns

nsns

nsns

Hip

posi

dero

s at

er(M

yanm

ar)

1♂

**

**

**

2.3

9.6

3.0

5.2

5.1

5.3

4.0

4♀

♀15

.8, 0

.213

.7, 0

.113

.8, 0

.18.

3, 0

.17.

6, 0

.17.

6, 0

.22.

6, 0

.19.

4, 0

.22.

9, 0

.15.

3, 0

.15.

1, 0

.05.

3, 0

.14.

2, 0

.115

.6–1

6.2

13.5

–13.

713

.8–1

4.0

8.2–

8.4

7.5–

7.8

7.4–

7.9

2.4–

2.6

9.2–

9.7

2.8–

3.0

5.2–

5.4

5.1–

5.2

5.2–

5.4

4.1–

4.3 in H. ha lophyllus the tip is pointed (Fig. 6). The bac-

ulum is 1.8–2.5 mm in length with a narrow shaft,round ed base, and a characteristically bifid tip (Fig.7). Those from Thailand correspond both in size andmorphology to material from Vietnam and Malaysiaillustrated in Topál (1975) and Bates and Harrison(1997), respectively. The baculum exceeds in sizeand differs in shape to those of H. halophyllus.

Cranial and dental charactersThe skull is small and elongated, with a condylo-

canine length of 12.8–13.7 mm and a mastoid widthof 7.5–8.1 mm, which exceeds zygomatic breadth(Table 3). When viewed in lateral profile, the ros-trum is horizontal, not gently sloping downwards asin H. halophyllus (Fig. 9). The junction between thepostorbital constriction and the rounded posteriorborder of the rostrum is less clearly defined than inH. halophyllus. Each zygoma is usually without a dorsal process on posterior part; if present, it is lowand comparable in size to the smallest processes ofH. halophyllus. Upper toothrow length is 4.7–5.3mm. As in H. halophyllus, the first upper premolar(P2) is very small and either included in or extrudedfrom the toothrow. The canine (C1) may or may notbe in contact with the second upper premolar (P4).The crown of the first lower premolar (P2) is fromtwo-thirds to equal that of the second (P4), and aboutone half to three quarters its height.

EcholocationHipposideros cineraceus uses constant frequen-

cy echolocation calls. In Thailand, the frequency ofmaximum energy (FMAXE) averaged 149.0 kHz(134.8–154.1 kHz) in males and 150.1 kHz (141.4–154.1 kHz) in females (BD based on a Pet tersson D240X). Pre viously, Robinson (1996) reported 135.0 kHz, whereas SJP, using a PetterssonD1000X, recorded 154.2–156.3 kHz from Kancha -naburi and Uthai Prov inces and 154.0–158.1 kHzfrom Naga mauk, Mon State, 16°19’N, 97°42’E,Myan mar. In both sexes, the calls average lowerthan those of H. halophyllus. Individual variation infrequency appears to exceed sexual or geographicalvariation (Fig. 11).

Taxonomic notesFollowing Corbet and Hill (1992), specimens

from throughout Myanmar and Thailand are refer-able to the nominate race H. c. cineraceus.

Conservation status and distributionHipposideros cineraceus was included as ‘Lower

Risk, least concern’ in Hutson et al. (2001),


FIG. 4. Face and noseleaf of two species of Hipposideros. A — H. halophyllus ♂, SB061023.2, Pha Dang Cave, Chiang MaiProvince, Thailand; B — H. halophyllus, ♀, PSUZC-MM 2007.41, Khao Samor Khon, Lop Buri Province, Thailand. C — H. cineraceus, ♂, PSUZC-MM 2006.125, Ton Nga Chang Wildlife Sanctuary, Songkla Province, Thailand; D — H. cineraceus,♂, PSUZC-MM 2006.69, Khao Kram Cave, Chumphon Province, Thailand; E and F — H. ater, HZM.5.35011, ♀, Kan-Thar-Yar

beach, Rakhine State, Myanmar. (Not to scale)

A B

C D

E F


Simmons (2005), Boitani et al. (2006), and IUCN(2009).

Hipposideros cineraceus is known from north-eastern Pakistan, northern India, Myanmar, Thai -land, Lao PDR, Vietnam, Cambodia, Malaysia, Su -matra, Krakatau Island, Borneo, and probably Phil -ippines (Corbet and Hill, 1992; Matveev, 2005;Simmons, 2005). Its distribution in Myanmar andThailand is mapped in Fig. 2.

Myanmar: Kachin State: Nam Tamai Valley (c.o.27°50’N, 97°45’E) [loc. C1, Fig. 2] (BMNH collec-tion); Mandalay Division: Mogok (22°55’N,96°30’E) [loc. C2, Fig. 2] (BMNH collection);Mingun (22°03’N, 96°01’E) [loc. C4, Fig. 2](BMNH collection); Shan State: Gokteik (22°21’N,96°55’E) [loc. C3, Fig. 2] (BMNH collection); Sa -ga ing Division: nr Sagaing (21°52’N, 95.59’E) [loc.

FIG. 5. Noseleaves of three species of Hipposideros. A: H. halophyllus, ♂, PSUZC-MM 2006.71, Pha Dang Cave, Chiang MaiProvince, Thailand; B: H. cineraceus, ♂, PSUZC-MM 2006.126, Ton Nga Chang Wildlife Sanctuary, Songkla Province, Thailand;

C: H. ater, ♀, HZM.10.35983, Tharabwin Village, Tanintharyi Division, Myanmar. Scale = 1 mm

C5, Fig. 2] (BMNH collection); Mon State: SaddanSin Cave, 16 km NE of Mawlamyine (16°19’N,97°42’E) [loc. C6, Fig. 2] (HZM collection); Naga -mauk II (16°18.992’N, 97°42.352’E, 29 m a.s.l.)[loc. C7, Fig. 2] (SJP survey). Thailand: Chiang MaiProvince: Khimee Cave, Chiang Dao Wildlife Sanc -tuary, Khong District, (19°21.266’N, 98°43.837’E,718 m a.s.l.) [loc. C8, Fig. 2] (Chiang Dao Wild-life Research Station collection); Pha Dang Cave,Sri lan na National Park, Chiang Dao District(19°20.769’N, 99°01.416’E, 480 m a.s.l.) [loc. C9,Fig. 2] (PSUZC collection); Tak Province: BanJagae Guard Station and Lum Khao Ngu GuardStation, Thung Yai Naresuan Wildlife Sanctuary (ca.15°41.310’N, 98°54.070’E, 718 m a.s.l.) [loc. C10,Fig. 2] (Robinson et al. 1995); Uthai Thani Prov -ince: Khi Nok Cave, Huai Kha Khaeng WildlifeSanctuary (15°37.222’N, 99°18.210’E, 380 m a.s.l.)[loc. C11, Fig. 2] (PSUZC Collection); Rathana KiriCave, Khao Kwang Thong District (15°22.282’N,99°40.984’E, 88 m a.s.l.) [loc. C12, Fig. 2]; Lup LeaCave, Ban Rai District, (15°03.077’N, 99°28.879’E,200 m a.s.l.) [loc. C13, Fig. 2] (PSUZC collection);Lop Buri Province: Khao Don Deung, Ban Mi Dis -trict, (15°08.888’N, 100°36.819’E, 40 m a.s.l.) [loc.C14, Fig. 2] (PSUZC collection); Nakhon Ra tcha simaProv ince: Khao Yai National Park, Pak Chong Dis -trict (ca. 14°32’N, 101°24’E) [loc. C15, Fig. 2] (Yen-butra and Felten, 1986); Sara Buri Prov ince: PhuNam Tok Tub Kwang (Tap Kwang), Kaeng KhoiDistrict (ca. 14°35’N, 100°08’E) [loc. C16, Fig. 2](Yenbutra and Felten, 1986); Sa Kaeo Prov ince:Khao Singto Cave, Meung District, (13°59.417’N,

CharactersEigenvector

1 2 3

FA -0.41 0.11 -0.17TIB -0.35 0.23 0.313MT -0.41 0.20 -0.154MT -0.42 0.08 -0.225MT -0.34 -0.34 -0.543D1P -0.08 -0.83 0.034D1P -0.37 0.09 0.224D2P -0.32 -0.28 0.68

Eigenvalue 5.38 1.30 0.54% of total variation explained 67.3 83.5 90.3

TABLE 4. Eigenvectors and eigenvalues of PCA of eight externalmeasurements of 49 female specimens of H. halophyllus,H. cineraceus and H. ater. See also Fig. 3

A B C


102°00.465’E, 12 m a.s.l.) [loc. C17, Fig. 2] (Waeng -sothorn et al., 2006b and PSUZC col lection); ChanthaBuri Province: Wat Khao Wong Kot, Tha Mai District(ca. 12°55’N, 101°58’E) [loc. C18, Fig. 2] (Yenbutraand Felten, 1986); Kanchana Buri Province: MaDuea Cave, Sri Mongkhon District (14°01.673’N,99°24.095’E, 64 m a.s.l.) [loc. C19, Fig. 2] (PSUZCcollection); Ratcha Buri Province: Khao Bin Cave,Chom Bung District, (13°35.527’N, 99°40.012’E,61 m a.s.l.) [loc. C20, Fig. 2] (PSUZC collection);Petcha Buri Province: Khao Yoi Cave, Khao YoiDistrict, (13°14.014’N, 99°49.708’E, 53 m a.s.l.)[loc. C21, Fig. 2] (Yen butra and Felten, 1986 andPSUZC collection); Chum phon Province: Khao KramCave, Patiew District, (10°55.131’N, 99°22.440’E,67 m a.s.l.) [loc. C22, Fig. 2] (PSUZC collec-tion); Si lawan Cave, Patiew District, (10°41.461’N,99°14.476’E, 68 m a.s.l.) [loc. C23, Fig. 2] (PSUZCcollection); Ranong Prov ince: Pra Kayang Cave,Kraburi Dis trict, (10°19.569’N, 98°45.923’E, 3 ma.s.l.) [loc. C24, Fig. 2] (PSUZC collection); PhangNga Prov ince: Ao Mai Ngam Natural Trail, SurinIslands Nat ional Park, (9°25.873’N, 97°51.975’E,20 m) [loc. C25, Fig. 2] (PSUZC collection); KohMai Pai, Muang District; (8°21.364’N, 98°32.429’E,20 m a.s.l.) [loc. C26, Fig. 2]; Trang Province: KhaoChong Waterfall, Khao Bant hat Wildlife Sanctuary,(7°32.894’N, 99°47.196’E, 81 m a.s.l.) [loc. C27,Fig. 2] (PSUZC collection); Song kla Province: NoyCave (ca. 7°4.300’N, 100°15.100’E, 40 m a.s.l.) [loc.C28, Fig. 2] (Bum rung sri, 1997); Syson Cave (ca.7°4.500’N, 100°10.010’E, 60 m a.s.l.) [loc. C29, Fig.2] (Bum rungsri, 1997); Outaphao Watershed, Rat ta -phum Dis trict, (6°47.777’N, 100°14.092’E, 197 ma.s.l.) [loc. C30, Fig. 2] (PSUZC collection); Wild lifeEdu cation Centre, Ton Nga Chang Wildlife Sanc-tuary, (6°56.739’N, 100°14.467’E, 107 m a.s.l.) [loc.C31, Fig. 2] (PSUZC collection); Khao Nouy Cave,Rat ta phum District, (6°59.537’N, 100°08.470’E,130 m a.s.l.) [loc. C32, Fig. 2] (Bum rung sri, 1997and PSUZC collection); Khao Rak Kiat Cave, Rat ta phum District, (7°4.254’N, 100°15.098’E, 100 m a.s.l.) [loc. C33, Fig. 2] (Bum rungsri, 1997and PSUZC collection); Ka Cave (ca. 6°36.400’N,

100°51.300’E, 60 m a.s.l.) [loc. C34, Fig. 2] (Bum -rung sri, 1997); Satun Province: Ban Wang BlaChan, Muang Satun District (ca. 6°45’N, 100°10’E)[loc. C35, Fig. 2] (Yenbutra and Felten, 1986); AoSon, Tarutao Islands National Park, (6°38.767’N,99°37.383’E, 5 m a.s.l.) [loc. C36, Fig. 2] (PSUZCcol lection); Jorrake (Crocodile) Cave, Tarutao Is -lands National Park (6°41.946’N, 99°39.096’E, 65 m a.s.l.) [also loc. C36, Fig. 2] (PSUZC collec-tion); km 1–2 road from Talow Wao to TalowOulang, Taru tao Islands Nat ional Park (6°36.501’N,99°40.435’E, 73 m a.s.l.) [also loc. C36, Fig. 2](PSUZC collection); Nara thi wat Province: Si rin -thorn Water fall, Bala forest, Hala-Bala WildlifeSanc tuary (5°48.017’N, 101°50.000’E, 300 m a.s.l.)[loc. C37, Fig. 2] (PSUZC collection).

Ecology and behaviour Hipposideros cineraceus was collected at 20 lo -

cations in Thailand with an altitudinal range of 3 to 718 m a.s.l.; 15 of the sites were between 40 and300 m a.s.l. (for further details see Materials andMethods). Although most of the locations were inlimestone areas, a minority (such as Surin IslandNational Park in Phang Nga Province and BalaForest in Na ra thi wat Province) were found in forestwith no obvious lime stone outcrops. As with H. ha -lophyllus, the spe cies was found in a range of foresttypes (hill and lowland evergreen, mixed deciduous,dipterocarp, mangrove and bamboo) and is tolerantof highly disturbed habitats that include plantations,orchards, rice paddies, corn fields and human habi-tation. It co-exists with a wide range of other batspecies. Preg nant females were found in Februaryand March in Khao Nouy Cave, Outaphao Water -shed and Tarutao Islands National Park.

Hipposideros ater Templeton, 1848Dusky Leaf-nosed Bat

Hipposideros ater Templeton, 1848; Colombo, Sri Lanka

External and bacular charactersHipposideros ater has a forearm length of 39.5

and 40.3 mm, all measurements based on Myanmar

FIG. 6. Lateral view of penis of three species of Hipposideros. A: H. halophyllus, PSUZC-MM 2006.71, Pha Dang Cave, Chiang MaiProvince; B: H. cineraceus, PSUZC-MM 2006.126, Ton Nga Chang Wildlife Sanctuary, Songkla Province; C: H. ater, MM3,

Thiruneer Malai, 26 km from Central Madras, Tamil Nadu, India. Scale = 2 mm

A B C


material unless otherwise stated (Table 1). The earsare slightly larger than those of H. cineraceus andconsiderably larger than H. halophyllus (Fig. 4). Thenoseleaf is small and both specimens from Myanmarexamined for this character have one pair of rudi-mentary supplementary lateral leaflets. These leaf -lets were not seen in any specimens of H. halo-phyllus or H. cineraceus (Fig. 5) (or specimens of H. ater from India and Sri Lanka). The internarialseptum is slightly inflated with a greatest width of0.8 mm; it exceeds that of H. cineraceus but is with-out the kidney-shaped disc of H. halophyllus. Thereis a small gland behind the posterior leaf. The fourthmetacarpal averages long er than the third and fifth.Since both Myanmar specimens examined in detailare females, no penis or baculum were observed.

FIG. 7. Dorsal (left) and lateral (right) views of the bacula ofthree species of Hipposideros. A: H. halophyllus, PSUZC-MM2007.203, Ob cave, Khao Smorkhon, Lop Buri Province,Thailand; B: H. cineraceus, PSUZC-MM 2007.178, Crocodile(Jorrake) Cave, Tarutao Islands National Park, Satun Province,Thailand; C: H. ater, HZM.4.35004, 16 km west of Tirunelveli,

southern India. Scale = 1 mm

CharactersEigenvector

1 2 3

CCL -0.40 0.18 -0.12MW -0.30 -0.44 0.25ZB -0.28 -0.44 0.29PC -0.32 0.29 0.22ML -0.42 -0.05 -0.11C1–C1 -0.15 -0.51 -0.74M3–M3 -0.36 -0.11 0.38C–M3 -0.37 0.32 -0.27C–M3 -0.34 0.34 -0.17

Eigenvalue 4.75 1.94 0.57% of total variation explained 52.8 74.3 80.6

TABLE 5. Eigenvectors and eigenvalues of PCA of nine cranial and dental measurements of 51 female specimens of H. halophyllus, H. cineraceus and H. ater. See also Fig. 8

However, in specimens from India, the penis is larg-er than those of H. halophyllus and H. cineraceus,with a rounded tip (Fig. 6). The baculum (for furtherexamples from India see Topál, 1975 and Bates andHarrison, 1997) has a simple rounded tip, unlike the bifid tip of H. cine raceus, and a narrow shaft; at 1.7 mm, it is over four times the length of that of H. halophyllus but has a comparable shape (Fig. 7).

Cranial and dental charactersThe skull has a condylo-canine length of

13.5–13.7 mm and mastoid width of 8.2–8.4 mm,which exceeds that of H. halophyllus and H. ciner-aceus (Table 3). As in the other two species, mastoidwidth exceeds zygomatic breadth. The rostrum, in lateral profile, is horizontal, similar to that of H. cineraceus and differing from the gently slopinganterior profile of H. halophyllus (Fig. 9). The sagit-tal crest is low. Each zygoma has a variably devel-oped jugal projection on its posterior part; althoughpresent in some specimens, in others it is scarcelyevident. Upper toothrow length (C–M3) is 5.1–5.2mm. The first upper premolar (P2) is very small andextruded from the toothrow; the second upper pre-molar (P4) is sometimes in contact with the uppercanine (C1). The crown of the first lower premolar(P2) is about two-thirds of the second (P4), and abouthalf of its height.

EcholocationNo data are available from the study region. In

India, H. ater has a resting frequency of 163.1–169.5 kHz (Jones et al., 1994). In Sabah, a fre-quency of 139.5 kHz was recorded (Francis and Ha -bersetzer, 1998). In Australia, Heller and Hel versen

A

B

C


(1989) recorded a frequency of 154.0 kHz and R. B.Coles (in litt.) recorded a frequency of 154–164 kHz.

Taxonomic notesSpecimens of H. ater from Myanmar differ in

a number of characters from those seen from south-ern India and Sri Lanka (the type locality). Spec -imens from the Indian Subcontinent held in theHarrison Institute and The Natural History Museum,London (n = 3♂♂ and 2♀♀) have smaller wings(FA = 37.5 mm, 36.4–38.2), shorter metacarpals(3MET = 28.5 mm, 27.2–30.1 mm; 4MET = 30.1mm, 29.1–31.2 mm; 5MET = 28.7 mm, 27.3–30.1mm) and are without a supplementary lateral leaflet adjacent to the anterior noseleaf. However, the In dian skulls average larger (CCL = 13.8 mm, 13.5– 14.2 mm; ZB = 8.2 mm, 7.9–8.3 mm; C–M3 = 5.2 mm, 4.7–5.4 mm) and the sagittal crest appearsto be more developed anteriorly. The taxonomic significance of these differences is not currentlyknown but it is probable that the Myanmar popula-tion belong to a discrete species.

Corbet and Hill (1992) referred specimens fromMyanmar to H. ater saevus Andersen, 1918. How -ever, with a condylo-canine length of 15.1 mm (An -dersen, 1918) saevus, which was named from KaiIsland, Indonesia, appears to be considerably largerand would not appear to be related to the geograph-ically distant H. ater. Externally, the specimens fromMyanmar are similar in size to the geographically

isolated H. a. nicobarulae Miller, 1902 from theNicobar Islands. However the skulls of nicobarulae(CCL = 14.3–15.0, n = 6, from Bates and Harrison,1997) are considerably larger.

Conservation status and distributionHipposideros ater was included as ‘Lower Risk,

least concern’ in Hutson et al. (2001), Simmons(2005), Boitani et al. (2006), and IUCN (2009).

According to Simmons (2005), H. ater rangesfrom Sri Lanka and India to western Malay sia, Indo -nesia, Philippines, New Guinea to Austral ia. It wasnot included in Lekagul and McNeely (1977) forThailand. Subsequently, Yenbutra and Felten (1986)included a single record from Ratchaburi Province.This specimen (BMNH.78. 2344) from Khao BinCave, with a condylo-canine length of 12.7 mm anda mastoid width of 7.7 mm is actually referable to H. halophyllus (see Hill and Yenbutra, 1984).Corbet and Hill (1992) mapped the species’ range asincluding much of western and peninsular Thailandbut did not include any justification. In the recentstudy, no specimens of H. ater were collected in themany exhaustive surveys undertaken throughout thecountry and its presence in Thailand needs confir-mation (see also Fig. 2).

Myanmar: Rakhine State: Kan Thar Yar Beach,Gwa Township, (17°43’N, 94°31’E), [loc. A1, Fig.2] (Pearch et al. 2003); Tanintharyi Division: Myeik(12°26’N, 98°36’E) [loc. A2, Fig. 2], (= Mar gui in

FIG. 8. PCA between first and second principal components based on nine cranial and dental metric characters for female specimens only. Black circles represent H. halophyllus, black squares are H. ater and black triangles are H. cineraceus

-6 -5 -4 -3 -2 -1 0 1 2 3 4 5PC1

PC

2

4

3

2

1

0

-1

-2

-3

-4


Lindsay, 1926); Tharabwin Village (12°18’N,99°04’E), [loc. A3, Fig. 2] (HZM collection).

Ecology and behaviourIn Myanmar, both specimens collected during re-

cent surveys were found roosting in human habita-tion. The first was observed just before dusk flyingaround the bathroom of one of a row of woodenchalets next to the sandy beach at Kan Thar Yar,Rak hine State. Numerous palm trees were grow-ing between a beach and the chalets, behind whichran a little-used road with dense vegetation beyond(Pearch et al. 2003). The second was found in theroof of a village house in Tharabwin, TanintharyiDivision.

DISCUSSION

From a conservation viewpoint, the studyshowed that H. halophyllus is much more wide-spread than previously thought. In a recent review

published in Boitani et al. (2006) the potential rangewas restricted to a small crescent-shaped area of ap-proximately 12,500 km2 around Bangkok and thepopulation was thought to number between 1,000and 1,400 (Waengsothorn et al., 2006a, 2006b; S. Waengsothorn, J. Nabhitabhata, K. Muang khum, S. Charavuthanukul, and T. Moon chan, in litt.). Thespecies was given a Red List Category of Endan -gered with a criteria of B2ab(iii); C1 (IUCN, 2009).This was based on an inferred population decline ofat least 10% in ten years; few known localities andabsence from localities where it was previouslyknown. It was suggested that more surveys wereneeded to resolve its distribution, which was provi-sionally given as two small isolated areas to thenorth-east and south-west of Bangkok.

The current study identified nine new localitiesin Thailand (Fig. 1) including Pha Dang Cave inChiang Mai Province, which extends the knownnorthern range by 435 km and Koh Mai Pai in PhangNga Province and Khaun Khang Peatswamp in

FIG. 9. The skulls of three species of Hipposideros. A: H. halophyllus, ♂, PSUZC-MM 2007.38, Khao Yoi Cave, Phetcha BuriProvince, Thailand; B: H. cineraceus, ♂, PSUZC-MM 2006.125, Ton Nga Chang Wildlife Sanctuary, Songkla Province, Thailand;

C: H. ater, ♀, HZM.10.35983, Tharabwin Village, Tanintharyi Division, Myanmar. Scale = 5 mm

A

B

C


FIG. 10. Echolocation calls of H. halophyllus from 11 localities in Thailand. Frequency of call (FMAXE in kHz) plotted against latitude (degrees North) of locality. Black squares represent males and open squares are females

Nakon Srithammarat Province in peninsular Thai -land, which extends its southern range in Thailandby over 700 km. It also confirms the first recordfrom Malaysia, previously reported in Francis(2008). In addition, it notes the continuing presenceof H. halophyllus at four previously known loca-tions (Khao Samor Khon [type locality], Khao Sing -to, Khao Yoi Cave, and Khi Nok Cave). How ever, atone site, Khao Bin Cave, from which a specimenhad been collected in September, 1969 (Hill andYen butra, 1984), only H. cineraceus was found dur-ing the present study. Two other sites (Vi Moke

Cave, and Khao Chong Phran) from which H. ha-lophyllus had been recorded previously were notvisited and consequently the current status of thepopulations cannot be determined. Specific threatsto the species were observed at three sites. In KhaoSamor Khon, bats were collected by local hunterswhilst in Ton Chan and KhaoYoi there was distur-bance from tourists visiting Buddhist temples with-in the caves.

Currently, H. halophyllus is only known fromtwo localities in peninsular Thailand, and one site in peninsular Malaysia, and appears to be less

FIG. 11. Echolocation calls of H. cineraceus from 21 localities in Thailand and Myanmar. Frequency of call (FMAXE in kHz) plottedagainst latitude (degrees North) of locality. Black squares represent males and open squares are females

Fre

quency (

kH

z)

Latitude (°N)

195

190

185

180

175

170

165

160

155

6 8 10 12 14 16 18 20

Fre

quency (

kH

z)

Latitude (°N)

160

155

150

145

140

135

130

4 6 8 10 12 14 16 18 20


abundant in this area than H. cineraceus, which waslocated at 12 sites. In central and northern Thailand,it shared seven of the 10 H. cineraceus roosts. It hasnot been found in Myanmar.

In contrast, the taxon currently referred to H. ateris entirely restricted to Myanmar and was not foundin any part of Thailand. Its distribution suggests thatit favours areas with a high annual rainfall, in excessof 2,500 mm annually (for details of Myanmar cli-mate, see Bender, 1983). It also suggests that it mayprove to be more widespread in both the seaboardareas of western Myanmar and possibly peninsularThailand. In Boitani et al. (2006), it was suggestedthat H. ater is likely to be a species complex and thecurrent study would lend some support to this view.Despite small sample sizes, specimens fromMyanmar appear to differ in a number of morpho-metric characters from material from Sri Lanka(type locality of H. ater) and also southern India.Further studies, including molecular analysis wouldbe of considerable interest to help resolve the rela-tionships of this taxon with those from the IndianSubcontinent, including the Nicobars, Malaysia andelsewhere in Southeast Asia. It should be noted thatdespite the apparent taxonomic differences betweenthe populations in Sri Lanka and Myanmar, theirecology and behaviour have much in common. Bothwere found in wet lowland regions and both roost inthe lofts of houses and old dwellings (Bates andHarrison, 1997 and this study). This is unlike H. ci -ne raceus and H. halophyllus which appear to prefermore natural roosting sites such as caves.

H. cineraceus is the most widely distributed ofthe three taxa in the study area, where its range extends from the north of Myanmar to the south ofthe Thai peninsula. Francis (personal communica-tion) in Boitani et al. (2006) suggested that there‘appears to be more than one species in peninsularMalaysia’. However within the study area, there appeared to be no evidence of cryptic species. Atpresent, H. cineraceus appears to be widespread,relatively abundant and with no significant threats toits overall population status within Thailand andMyanmar.

ACKNOWLEDGEMENTS

In Lao PDR, we would like to thank the late AssociateProfessor Dr. Bounnam Phathumthong of Department of Biol -ogy, Faculty of Science, the National University of Laos, whowas BD’s advisor, and supported this project in Lao PDR. We would also like to thank his successor, Associate ProfessorBoun thob Praxaysombath as well as the staff of the Faculty of Environmental Science, the National University of Laos for their encouragement. In Thailand, thanks are due to the

students of the Bat Research Unit, Department of Biology,Prince of Song kla University, including P. Kingsada, A. Dejta -radol, P. Phommexay, S. Karnpun, T. Jongsiri, T. Srithongchuay,A. Thaisauriyan, U. Pimsai, H. Jansod, K. Nualcharoen, M. Yo -kubol, P. Piyapan, and B. Pongsee for their help. We would alsolike to thank the staff of the Princess Maha Chakri Sirin dhornNatural History Museum, Prince of Songkla University. We aremost grateful to the staff of the Chiang Dao Wildlife ResearchStation, Thailand for the loan of specimens and help in the field.In Myanmar, we would like to thank Professor Daw Tin Nwe(retired), Dr Khin Maung Swe, Dr Khin Mie Mie and all the batteam of Yangon University for their help in organising and par-ticipating in a range of field surveys throughout their countryand Dr Iain Mackie of Aberdeen University, who co-led a num-ber of the expeditions. We are most grateful to Dr Zubaid Akbarof the Universiti Kebangsaan Malaysia and to the EconomicPlanning Unit of the Malaysia Prime Minister’s Department forgranting SM permission to conduct wildlife research inMalaysia. In the United States, SM would like to thank DrThomas H. Kunz of Boston University and Dr Tigga Kingston(formerly of Boston University, now of Texas Tech University)for their help and supervision. In Ireland, SJP would like tothank the Science Foundation Ireland (RFP GEN0056) for theirfinancial support and in Thailand, Dr Meriadeg Ar Gouilh andthe IRD UR 178, Mahidol University for their logistic support.In the UK, we thank Paula Jenkins, Louise Tomsett, RobertoPortela Miguez, and all the staff of the Mammal Section of theNatural History Museum, London for their help in accessing thecollection. At the Harrison Institute, we would like to thankMalcolm Pearch for his help during the senior authors study vis-it to the UK, and all the ‘Friends of the Harrison Institute’ whoparticipated in the 2000 Myanmar expedition to RakhineDivision. Finally, we would like to thank the Graduate School,Prince of Songkla University and the British Government fortheir financial support of the project through the DarwinInitiative (Nos: 14011, 18002); the British Council through theirsupport through the PMI2 programme; and the Foreign andCommon wealth Office through their SEAGP Funding. Withoutall of the above and the collaboration of many others, this proj-ect would not have been possible.

LITERATURE CITED

ANDERSEN, K. 1918. Diagnoses of new bats of the familiesRhinolophidae and Megadermatidae. Annals and Magazineof Natural History, 2: 374–384.

BATES, P. J. J., and D. L. HARRISON. 1997. Bats of the IndianSub continent. Harrison Zoological Museum Publication,Sevenoaks, xvi + 258 pp.

BATES, P. J. J., S. J. ROSSITER, A. SUYANTO, and T. KINGS-TON. 2007. A new species of Hipposideros (Chiroptera: Hippo sideridae) from Sulawesi. Acta Chiropterologica, 9:13–26.

BENDER, F. 1983. The geology of Burma. Gebrüder Borntraeger,Berlin, 293 pp.

BLYTH, E. 1853. Report of the curator, Zoological Department.Journal of the Asiatic Society of Bengal, 22: 408–417.

BOGDANOWICZ, W., and R. D. OWEN. 1998. In the Minotaur’slabyrinth: phylogeny of the bat family Hipposideridae. Pp.27–42, in Bat biology and conservation (T. H. KUNZ and P. A. RACEY, eds.). Smithsonian Institution Press, Washing -ton, D.C., xiv + 365 pp.

BOITANI, L., G. CATULLO, I. MARZETTI, M. MASI, M. RULLI, and

S. SAVINI. 2006. The Southeast Asian mammal databank. A tool for conservation and monitoring of mammal diversi-ty in Southeast Asia. www.ieaitaly.org/samd.

BUMRUNGSRI, S. 1997. Roost selection of cave dwelling bats inSongkla and Satun Provinces. M.Sc. Thesis, Kasetsart Uni -ver sity, Bangkok, 79 pp. [in Thai].

CORBET, G. B., and J. E. HILL. 1992. The mammals of the Indo -malayan region: a systematic review. Natural History Mu -seum Publications, Oxford, 488 pp.

FRANCIS, C. M. 2008. A field guide to the mammals of Thailandand South-East Asia. Asia Books Co., Bangkok, 392 pp.

FRANCIS, C. M., and J. HABERSETZER. 1998. Interspecific and in-traspecific variation in echolocation call frequency and mor-phology of horseshoe bats, Rhinolophus and Hipposideros.Pp. 169–179, in Bat biology and conservation (T. H. KUNZ

and P. A. RACEY, eds.). Smithsonian Institution Press, Wash -ington, D.C., xvi + 365 pp.

FRANCIS, C. M., D. KOCK, and J. HABERSETZER. 1999. Siblingspecies of Hipposideros ridleyi (Mammalia, Chiropte ra, Hip -po sideridae). Senckenbergiana Biologica, 79: 255–270.

GUILLÉN-SERVENT, A. G., and C. M. FRANCIS. 2006. A new spe -cies of bat of the Hipposideros bicolor group (Chiroptera:Hip posideridae) from central Laos, with evidence of con-vergent evolution with Sundaic taxa. Acta Chiropterologica,8: 39–61.

HELLER, K.-G., and O. VON HELVERSEN. 1989. Resource parti-tioning of sonar frequency bands in rhinolophoid bats.Oeco logia, 80: 178–186.

HILL, J. E. 1963. A revision of the genus Hipposideros. Bulletinof the British Museum of Natural History (Zoology), 2(1):1–129.

HILL, J. E., and S. YENBUTRA. 1984. A new species of the Hippo -sideros bicolor group (Chiroptera: Hipposideridae) fromThailand. Bulletin of the British Museum of Natural History(Zoology), 47(1): 77–82.

HILLMAN, A. 1999. A study of wrinkled-lipped bats (Tadaridaplicata) at Khao Chong Phran, Non-hunting area, Ratcha -buri Province. Journal of Thai Forestry Research, 1: 72–83.[in Thai].

HUTSON, A. M., S. P. MICKLEBURGH, and P. A. RACEY (comp.).2001. Global status survey and conservation action plan:Micro chiropteran bats. IUCN/SSC Chiroptera SpecialistGroup, IUCN, Gland, 258 pp.

IUCN. 2009. The IUCN Red List of Threatened Species.www.iucnredlist.org.

JENKINS, P. D., and J. E. HILL. 1981. The status of Hipposiderosgaleritus Cantor, 1846 and Hipposideros cervinus (Gould,1854) (Chiroptera: Hipposideridae). Bulletin of the BritishMuseum of Natural History (Zoology), 41(5): 279–294.

JONES, G., K. SRIPATHI, and D. A. WATERS. 1994. Individual var -iation in the echolocation calls of three sympatric Indianhipposiderid bats, and an experimental attempt to jam batecholocation. Folia Zoologica, 43: 347–362.

KHAJURIA, H. 1970. A new leaf-nosed bat from central India.Mammalia, 34: 622–627.

KOCK, D., and H. R. BHAT. 1994. Hipposideros hypophyllus n. sp.of the H. bicolor-group from peninsular India (Mammal ia:Chiroptera: Hipposideridae). Senckenbergiana Biologica,73(1–2): 25–31.

LEKAGUL, B., and J. A. MCNEELY. 1977. Mammals of Thailand.Association for the Conservation of Wildlife, Bangkok,758 pp.

LINDSAY, H. M. 1926. Report No. 39: Mergui Archipelago.Bom bay Natural History Society’s Mammal Survey ofIndia, Burma and Ceylon. Journal Bombay Natural HistorySociety, 30: 42–48.

MATVEEV, V. A. 2005. Checklist of Cambodian bats (Chiro pte -ra), with new records and remarks on taxonomy. RussianJournal of Theriology, 4: 43–62.

MILLER, G. S. 1902. The mammals of the Andaman and NicobarIslands. Proceedings of the United States National Museum,24(1296): 751–795.

PEARCH, M. J., KHIN MIE MIE, P. J. J. BATES, TIN NWE, KHIN MAUNG SWE, and SI SI HLA BU. 2003. First record of bats (Chiroptera) from Rakhine State, Myanmar (Burma). Natural History Bulletin of Siam Society, 51:241–259.

PRICE, L. 2001. Caves and karst of peninsular Malaysia. GuaPublications, Kuala Lumpur, 98 pp.

ROBINSON, M. F. 1996. A relationship between echolocationcalls and noseleaf widths in bats of the genera Rhinolophusand Hipposideros. Journal of Zoology (Lon don), 239:389–393.

ROBINSON, M. F., A. L. SMITH, and S. BUMRUNGSRI. 1995. Smallmammals of Thung Yai Naresuan and Huai Kha KhaengWildlife Sanctuaries in Western Thailand. Natural HistoryBulletin of Siam Society, 43: 27–54.

SIMMONS, N. B. 2005. Order Chiroptera. Pp. 312–529, in Mam -mal species of the World: a taxonomic and geographic ref -erence (D. E. WILSON and D. M. REEDER, eds.). The JohnsHopkins University Press, Baltimore, 2142 pp.

TATE, G. H. H. 1941. A review of the genus Hipposideros withspecial reference to Indo-Australian species. Bulletin of theAmerican Museum of Natural History, 78: 353–393.

TEMPLETON, W. E. 1848. In Report to the curator (E. BLYTH).Journal of the Asiatic Society of Bengal, 17: 247–255.

TOPÁL, G. 1975. Bacula of some Old World leaf-nosed bats.Verte brata Hungarica, 16: 21–53.

WAENGOSOTHORN, S., J. NABHITABHATA, T. MOONCHAN, and K. MUANGKHAM. 2006a. Intensive survey of the current sta-tus of Thai endemic mammals and their habitats outsidePro tected Areas in central Thailand. BRT 2006 ResearchRe ports, 285–297. [in Thai].

WAENGOSOTHORN, S., J. NABHITABHATA, T. MOONCHAN, and K. MUANGKHAM. 2006b. The current distribution and popu-lation structures of Thailand roundleaf bat (Hipposideroshalo phyllus), a Thai endemic mammal. Journal of Wildlifein Thailand, 13(1): 43–60. [in Thai].

WATHANAKUL, W. 1995. Identification of bats of Thailand.Department of Biology, Faculty of Science and Technology,Petchaburi Rajabhat University, Petchaburi, Thailand, 855pp. [in Thai].

YENBUTRA, S., and H. FELTEN. 1986. Bats species and their dis-tribution in Thailand according to the collections in TISTRand SMF. Pp. 9–45, in Contributions to the knowledge ofthe bats of Thailand (H. FELTEN, ed.). Courier Forschungs -institut Senckenberg, 87: 1–112.


Received 14 April 2009, accepted 25 January 2010

Hipposideros halophyllus — PSUZC-MM 2006.70–71,77–79, 82–83, 3♂4♀, Pha Dang Cave, Srilanna National Park,Chiang Mai Province (19°20.769’N, 99°01.416’E, 480 m a.s.l.);PSUZC-MM2008.81–83, ♂2♀, Khi Nok Cave, Huai KhaKhaeng Wildlife Sanctuary, Uthai Thani Province(15°37.222’N, 99°18.210’E, 350 m); PSUZC-MM 2007.204,♀, Lup Lae Cave, Ban Rai District, Uthai Thani Province(15°03.077’N, 99°28.879’E, 200 m a.s.l.); BM(NH). 78.2344,♂, Khao Bin Cave, Chom Bung District, Ratcha Buri Province(13˚35.527’N, 99˚40.012’E, 61 m a.s.l.); PSUZC-MM2007.201–202, ♂♀, Khao Don Deung, Ban Mi District, LopBuri Province (15°08.888’N, 100°36.819’E, 40 m a.s.l.);PSUZC-MM 2007.40–41, 240–241, 2♂2♀, Khao Samor Khon,Tawung District, Lop Buri Province (14°54.548’N,100°30.342’E, 3 m a.s.l.); PSUZC-MM 2007.203, ♂, Ob Cave,Khao Smorkhon, Tawung District, Lop Buri Province(14°54.525’N, 100°29.491’E, 38 m a.s.l.); PSUZC-MM2007.195–200, 3♂3♀, Khao Singto, Meung District, Sa KaeoProvince (13°59.417’N, 102°00.465’E, 12 m a.s.l.); PSUZC-MM 2007.24, 35–39, BM(NH).78.2346, 6♂♀, Khao Yoi Cave,Khao Yoi District, Petcha Buri Province (13°14.014’N,99°49.708’E, 53 m a.s.l.); PSUZC-MM 2008.96–97, 2♂, KohMai Pai, Muang District, Phang Nga Province (8°21.364’N,98°32.429’E, 20 m a.s.l.); SB090131.1, 11, 2♀, Khaun KhangPeatswamp, Thalenoi Non-Hunting Area, Cha Aod District,Nakon Srithammarat Province (7°53.780’N, 100°6.347’E).

Hipposideros cineraceus — BM(NH).79.11.21.160 (type ofmicropus), ?, Dehra Dun, near Simla, north-western India;HZM.3.34873, ♀, Saddan Sin Cave, 16 km north-eastMawlamyine, Mon State, Myanmar (16°19’N, 97°42’E); CD-B-0041, ♀, Khimee Cave, Chiang Dao Wildlife Sanctuary,Khong District, Chiang Mai Province (19°21.266’N,98°43.837’E, 718 m a.s.l.); PSUZC-MM 2006.72–76, 80–81,5♂2♀, Pha Dang Cave, Srilanna National Park, Chiang MaiProvince (19°20.769’N, 99°01.416’E, 480 m a.s.l.); PSUZC-MM2008.84, ♂, Khi Nok Cave, Huai Kha Khaeng WildlifeSanctuary, Uthai Thani Province (15°37.222’N, 99°18.210’E,350 m); PSUZC-MM 2007.192, ♂, Lup Lea Cave, Ban RaiDistrict, Uthai Thani Province (15°03.077’N, 99°28.879’E, 200m a.s.l.); PSUZC-MM 2005.39, ♀, East Thung Yai WildlifeSanctuary (15°41’31”N, 98°54’07”E, 900 m a.s.l.); PSUZC-MM 2007.22–23, 191, 193, 4♀, Khao Bin Cave, Chom BungDistrict, Ratcha Buri Province (13°35.527’N, 99°40.012’E, 61m a.s.l.); PSUZC-MM 2007.185–190, 3♂3♀, Khao DonDeung, Ban Mi District, Lop Buri Province (15°08.888’N,

100°36.819’E, 40 m a.s.l.); PSUZC-MM 2007.184, ♀, KhaoSingto Cave, Meung District, Sa Kaeo Province (13°59.417’N,102°00.465’E, 12 m a.s.l.); PSUZC-MM 2007.21, ♂, Khao YoiCave, Khao Yoi District, Petcha Buri Province (13°14.014’N,99°49.708’E, 53 m a.s.l.); PSUZC-MM 2006.69, 84–86, 88–89,3♂3♀, Khao Kram, Patiew District, Chumphon Province(10°55.131’N, 99°22.440’E, 67 m a.s.l.); PSUZC-MM 2007.9,♂, Silawan Cave, Patiew District, Chumphon Province(10°41.461’N, 99°14.476’E, 68 m a.s.l.); PSUZC-MM 2007.10,♂, Pra Kayang Cave, Kra Buri District, Ranong Province(10°19.569’N, 98°45.923’E, 3 m a.s.l.); PSUZC-MM 2006.18,♀, Ao Mai Ngam, Surin Islands National Park, Phang NgaProvince (9°25.873’N, 97°51.975’E, 20 m a.s.l.); PSUZC-MM2007.8, Khao Chong Waterfall, Khao Banthat WildlifeSanctuary, Trang Province (7°32.894’N, 99°47.196’E, 81 ma.s.l.); PSUZC-MM 2005.62, ♂, Ao Son, Tarutao IslandsNational Park, Satun Province (6°38.767’N, 99°37.383’E, 5 ma.s.l.); PSUZC-MM 2007.177–179, 181, 4♂, Crocodile Cave,Tarutao Islands National Park, Satun Province (6°41.946’N,99°39.096’E, 65 m a.s.l.); PSUZC-MM 2007.180, 182, 2♀, km1–2 road from Talow Wao to Talow Oulang, Tarutao IslandsNational Park (6°36.501’N, 99°40.435’E, 73 m a.s.l.); PSUZC-MM 2005.63, 2006.10, 144, ♂2♀, Khao Rak Kiat Cave,Rattaphum District, Songkhla Province (7°4.254’N,100°15.098’E, 100 m a.s.l.); PSUZC-MM 2006.125–126, 2♂,Wildlife Education Centre, Songkhla Province (9°56.739’N,100°14.467’E, 107 m a.s.l.); PSUZC-MM 2007.171–175, ♂4♀,Khao Nouy Cave, Rattaphum District, Songkhla Province(6°59.537’N, 100°08.470’E, 130 m a.s.l.); PSUZC-MM2007.194, ♀, Outaphao Watershed, Songkhla Province(6°47.777’N, 100°14.092’E, 197 m a.s.l.); PSUZC-MM2005.76, ♂, Sirinthorn Waterfall, Bala Forest, Hala-Bala Wild -life Sanctuary, Narathiwat Province (5°48.017’N, 101°50.000’E,300 m a.s.l.).

Hipposideros ater — HZM.2.28189, 3.28190, Field No.MM2-3, 2♂2♀, Thiruneer Malai, 26 km from central Madras,Tamil Nadu, India (13.05N, 80.00E); HZM.4.35004, ♂,Sankanthira Mandapam, northern Cheranmadevi, 16 km west-ern Tirunelveli, southern India (8.45N, 77.43E); HZM.10.35983, ♀, Tharabwin Village, Tanintharyi Division,Myanmar (12°18’N, 99°04’E); HZM.5.35011, ♀, Kan Thar YarBeach, Gwa Township, Rakhine State, Myanmar (17°43’N,94°31’E); BM(NH).23.11.11.2, ♂, Mergui, (TanintharyiDivision), Myanmar; BM(NH).21.1.17.52–53, 2♀, TenasserimVillage (Tanintharyi Division) Myanmar.

APPENDIX

List of specimens


A Taxonomic Review of Hipposideros halophyllus , with Additional Information on H. ater and H....

Documents

Transcript of A Taxonomic Review of Hipposideros halophyllus , with Additional Information on H. ater and H....