PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):271–302. 2004. The mammals of Palawan Island, Philippines

Jacob A. Esselstyn, Peter Widmann, and Lawrence R. Heaney

(JAE) Palawan Council for Sustainable Development, P.O. Box 45, Puerto Princesa City, Palawan, Philippines (present address: Natural History Museum, 1345 Jayhawk Blvd., Lawrence, KS 66045, U.S.A.) (PW) Katala Foundation, P.O. Box 390, Puerto Princesa City, Palawan, Philippines; (LRH) Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605 U.S.A.

Abstract.—The mammal fauna of Palawan Island, Philippines is here doc- umented to include 58 native species plus four non-native species, with native species in the families Soricidae (2 species), Tupaiidae (1), Pteropodidae (6), Emballonuridae (2), Megadermatidae (1), Rhinolophidae (8), Vespertilionidae (15), Molossidae (2), Cercopithecidae (1), Manidae (1), Sciuridae (4), Muridae (6), Hystricidae (1), Felidae (1), Mustelidae (2), Herpestidae (1), Viverridae (3), and Suidae (1). Eight of these species, all microchiropteran bats, are here reported from Palawan Island for the first time (Rhinolophus arcuatus, R. ma- crotis, Miniopterus australis, M. schreibersi, and M. tristis), and three (Rhin- olophus cf. borneensis, R. creaghi, and Murina cf. tubinaris) are also the first reports from the Philippine Islands. One species previously reported from Pa- lawan (Hipposideros bicolor)isremoved from the list of species based on re- identificaiton as H. ater, and one subspecies (Rhinolophus anderseni aequalis Allen 1922) is placed as a junior synonym of R. acuminatus. Thirteen species (22% of the total, and 54% of the 24 native non-flying species) are endemic to the Palawan faunal region; 12 of these are non-flying species most closely related to species on the Sunda Shelf of Southeast Asia, and only one, the only bat among them (Acerodon leucotis), is most closely related to a species en- demic to the oceanic portion of the Philippines. Of the 28 insectivorous bats, 18 species are somewhat to highly widespread in Indo-Australia, 2 are shared only with the Sunda Shelf and Indochina, 1 with the Sunda Shelf alone, 3 occur on the Sunda Shelf and the oceanic Philippines, 1 occurs in Palawan, Sulawesi, and the oceanic Philippines, 2 occur only on Palawan and in the oceanic Philippines, and 1 occurs on Borneo, Sulawesi, and throughout the Philippines. Though the insectivorous bats tend to be widely distributed, these data, particularly the distributions of the non-volant species, strongly reinforce the perception of Palawan Island (and associated smaller islands) as a biogeo- graphic unit of the Sunda Shelf, with only limited similarity to other portions of the Philippine Islands.

The Philippine archipelago is remarkable world (Heaney et al. 1998, Heaney & Re- for the large number of indigenous land galado 1998, Mittermeier et al. 1997). mammal species (ca. 175), and especially These species, especially the endemics, are for the number of endemic species (ca. not distributed homogeneously over the 112). Given its relatively small land area, country; rather, there is a large number of the Philippines has perhaps the greatest discrete biogeographic units, and these cor- concentration of endemic mammals in the respond to the limits of the islands that ex- 272 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON isted during periods of low sea level during ney 1980), but the fauna is still poorly the late Pleistocene (Heaney 1986, 1991a, known in many respects. Little information 1991b, 2000). With a single exception, cur- has been available for most species on ecol- rent geological evidence indicates that none ogy and distribution, including habitat re- of these ‘‘Pleistocene islands’’ has had dry- quirements, and only a few studies have land connections to the Asian mainland or considered phylogenetic relationships (e.g., to other areas. Rather, each arose as a de shrews: Heaney & Ruedi 1994; bats: Mus- novo oceanic island, some from a combi- ser et al. 1982; squirrels: Heaney 1979; mu- nation of oceanic crust and volcanic mate- rid rodents: Musser 1979, Musser & New- rials, and some as uplifted areas of conti- comb 1983, Musser & Heaney 1992; pan- nental rock that had been submerged for golin: Feiler 1998; leopard cats: Groves long periods, and all of these have remained 1997). In particular, microchiropteran bats isolated by sea channels (Hall 1998, 2002; have been only superficially documented. Heaney 1985, 1986, 1991a). The sole ex- The limitations of available data have thus ception is the Palawan faunal region, which limited understanding of the Southeast generally has been considered to be a por- Asian fauna, and the Philippine fauna in tion of the Sunda Shelf, both geologically particular, from both biogeographic and and biogeographically, with many species ecological perspectives, and hence limited shared with Borneo (Dickerson 1928, Ev- conservation planning in a nation that is of- erett 1889, Heaney 1986). Although Pala- ten cited as one of the most in need of ef- wan was initially also a de novo oceanic fective conservation action (Mittermeier et island, its biogeographic affinity to the Sun- al. 1999, Ong et al. 2002, Wildlife Conser- da Shelf has been thought to be due to the vation Society of the Philippines 1997). presence of a shallow shelf between Borneo Twoofus(Esselstyn and Widmann) re- and Palawan with an intervening depth of cently conducted extensive surveys of the ca. 145 m (Heaney 1986, 1991a). Previous- mammals of Palawan Island, focusing on ly, evidence indicated that sea levels the 15 sites described below. Esselstyn dropped to about 165 m below present lev- worked from December 1999 to November els during the penultimate glacial episode 2000 at Sites 1–11, emphasizing (though (Gascoyne et al. 1979), which would have not exclusively) insectivorous bats, which resulted in a dry-land connection of Pala- are the mammals most poorly known in the wan to Borneo at about 165,000 BP (Hea- Philippines (Heaney et al. 1998, Heaney & ney 1985, 1986, 1991a). However, recent Mallari 2002), and Widmann conducted evidence suggests that sea level dropped studies of bats, rodents, and larger mam- only to about 135 m (Rohling et al. 1998) mals from 1997 to 2002 at Sites 12–15; or perhaps as little as 115 m below present Heaney visited briefly in April 2000. In this levels (Siddall et al. 2003, Voris 2000) dur- paper, we report information collected dur- ing that glacial episode, leaving open the ing these studies, emphasizing new data on question of when Palawan was connected bats, and we include additional unpublished to Borneo, or if the gap simply became very records of mammals. We summarize infor- narrow. mation on all additional species that were The mammals of Palawan Island, the not taken during this study but have been largest part of the faunal region at 11,785 documented on the island, and re-examined km2, and the associated smaller islands have some key specimens from prior studies. We been documented over the course of more include descriptions of the habitats where than a century (Allen 1910, Allen 1922, Ev- we conducted our surveys because of a gen- erett 1889, Heaney et al. 1998, Hoogstraal eral paucity of such information, and use all 1951, Kuntz 1969, Reis & Garong 2001, available information to evaluate conser- Sanborn 1952, Taylor 1934, Timm & Bir- vation status of the mammals. We include VOLUME 117, NUMBER 3 273 measurements of the skulls of selected spe- tion was frequently changed. Mist nets were cies of insectivorous bats that have been es- continuously monitored during peak activ- pecially poorly known. ity periods from 1800 to 2000 h (at Sites 12 and 13, until 2400 h) and then checked Methods again in the early morning. The harp trap was checked periodically between 1800 and At Sites 1–11, small non-volant mam- 2100 h and again in the early morning. In mals were captured using locally made live forested areas, we searched for bats in po- (cage) traps and Victor (snap) rat traps. Ap- tential roosting locations (e.g., hollow trees, proximately 90% of live traps used mea- rock formations, new banana leaves, etc.). sured 11 ϫ 11 ϫ 24 cm, and 10% measured Most bats were released at the site of cap- 13 ϫ 16 ϫ 13 cm. Trap lines consisting of ture. For many of the bats, we report the approximately 70 traps (50 live traps and proportion of adult females that were preg- 20 snap traps) were placed in areas of tra- nant on certain dates. These were palpated versable terrain. Individual traps were externally to determine the presence or ab- placed in locations of likely capture (e.g., sence of an embryo before being released. near holes, along fallen logs, near root but- Forearm and cranial measurements were tresses, etc.) along the line spaced at 5–15 taken by L. R. Heaney and D. S. Balete at m intervals. Most traps were set at ground the Field Museum of Natural History level, but in forest habitats we placed 5– (FMNH). All voucher specimens from Es- 15% of the traps in elevated locations up to selstyn, which are cited below as specimens 2mabove ground level on fallen logs, hor- examined, were preserved in fluid (with izontal vines, etc. All but two trap lines skulls later removed and cleaned) and cat- were set for three nights; the exceptions at aloged at the FMNH; half of the vouchers Sites 3 and 7 were set for five and two have been deposited at the National Muse- nights, respectively. At Sites 12–14, a mix- um of the Philippines (NMP). Additionally, ture of live traps were used (see Site de- data on several previously unreported spec- scriptions). Most trap lines were baited with imens housed in the University of Michigan fresh grilled coconut coated with peanut Museum of Zoology (UMMZ) and United butter. Other trap lines were baited with live States National Museum of Natural History earthworms or bananas. All traps were (USNM) are included here. checked in the early morning and late af- ternoon. Baits were changed at least once Site Descriptions daily, usually in the afternoon, and as nec- essary in the early morning. Most live an- See Fig. 1 for approximate locations of imals were released at the site of capture. study sites. The province of Palawan in- We captured bats using a harp trap (ca. 2 cludes the main island called Palawan and ϫ 2m,4bank), mist nets (2 ϫ 6m,16 many smaller, nearby islands. The island is mm mesh), butterfly nets, and hand capture politically divided into 13 municipalities, at Sites 1–11; only nets were used at Sites one of which is Puerto Princesa City; the 12–15. Harp trap and net locations were se- municipalities and the city are subdivided lected to be locations of likely capture (e.g., into barangays, and barangays into sitios. natural canopy breaks, over streams and Site 1 (10Њ03Ј00ЉN, 119Њ00Ј44ЉE) was in trails, around fruiting trees, and near poten- lowland primary forest located along the tial roosting locations). The harp trap and Tarabanan River in northern Puerto Prin- mist nets were usually set in a location for cesa Municipality, at elevations ranging be- three nights, and occasionally for only one tween ca. 100 and 200 m. Slope in the area or two nights. During surveys of caves, we was generally moderate, rising from the riv- primarily used the harp trap, and its loca- er to the ridge-tops. Forest in the area was 274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Map of Palawan Island, showing the locations of the primary towns (solid circles) and research sites (solid triangles), and the position of Palawan in the Philippines (inset). nearly undisturbed, which is uncommon at lection of minor forest products and hunting this low elevation. We were aware of only of Sus barbatus and Macaca fascicularis. one small human-made clearing (ca. 0.5 ha) Canopy ranged from 20 to 30 m in height in the area; other disturbances included col- and was multi-layered. Canopy trees ranged VOLUME 117, NUMBER 3 275 in diameter from 40–80 cm and had light Ten harp-nights yielded 1 Hipposideros buttress development. Leaf litter was thin. diadema, 16 Rhinolophus arcuatus, 10 R. We surveyed small volant and non-volant creaghi, 2 R. virgo, and 2 Kerivoula hard- mammals for 13 days in January 2000. wickii. Four trap lines were run for three nights, Site 3 (10Њ07Ј28ЉN, 118Њ59Ј36ЉE) was in yielding 3 Tupaia palawanensis, 21 Max- primary montane and mossy forest located omys panglima, and 1 juvenile Viverra tar- near the peak of Cleopatra’s Needle (max- galunga in 792 trap-nights. Three of the imum elevation 1603 m), at elevations lines were baited with coconut and peanut ranging from 1300 to 1600 m. Slope was butter, while one line was baited with live moderate to extreme. Other than a trail to earthworms. Forty-eight net-nights pro- the peak occasionally traveled by tourists, duced 1 Cynopterus brachyotis and 12 there was little human-caused disturbance harp-nights produced 3 Rhinolophus acu- in the area. No above-ground sources of minatus, 3 R. arcuatus, and 1 R. creaghi, water were found near the site, but mist was plus other means produced 11 Megaderma frequently present: during our stay of two spasma and 1 R. acuminatus. weeks during dry season, the area was al- Site 2 (9Њ42Ј14ЉN, 118Њ32Ј01ЉE) was in most continuously shrouded by a dense fog. lowland primary forest located mid-way up Vegetation type was montane forest up to Mt. Salakot, between 300 and 700 m ele- ca. 1500 m. At this elevation, the vegetation vation. Slope was rolling to moderately began a transition to mossy forest. In mon- steep. Several small streams dissected the tane forest, the canopy reached a height of area. The only major human-caused distur- approximately 10 m. Trees, rocks, fallen bance in the area was an unused helicopter logs, and other stable surfaces were covered landing pad and an abandoned road; the with a thin layer of moss; epiphytic ferns road had a dense regrowth of ferns and and orchids were abundant. Many trees had small trees. Sus barbatus was hunted, and an adventitious root system, but maintained some almaciga trees (Agathis sp.) near the straight boles. The canopy was more open upper reaches of the site had fallen due to here than at lower elevations. Above 1500 over-collection of resin. Agoho trees (Ca- m, trees were shorter (2–4 m in height) and suarina sp.) gradually became more com- took on a shrub form above 1550 m. Moss mon as elevation increased. Canopy ranged growth was heavier at this elevation, and from 15 to 30 m in height and was multi- vegetation became extremely dense at the layered. Canopy trees ranged in diameter upper reaches. Pitcher plants (Nepenthes) from 35–60 cm with the largest emergents began to appear at about 1500 m and were reaching 80–90 cm. Buttress systems were abundant by 1550 m. We surveyed small only slightly developed and stilt root sys- volant and non-volant mammals at this site tems were present above 600 m, but rare. for 12 days during February and March Leaf litter was slightly deeper than at lower 2000. Three trap lines yielded 1 Tupaia pa- elevations. We surveyed small volant and lawanensis, 33 Maxomys panglima, and 3 non-volant mammals at this site for 20 days Rattus tiomanicus in 740 trap-nights. Two during March and July 2000. Six trap lines lines were run for three nights each and one (three coconut and peanut butter-baited line was run for five nights; all three lines lines, one banana-baited line, and two were baited with coconut and peanut butter. earthworm-baited lines) yielded 16 Tupaia Forty-eight net-nights produced 2 Rhinolo- palawanensis, 55 Maxomys panglima, and phus arcuatus, and 12 harp-nights produced 1 Sundamys muelleri from 1272 trap nights. no captures; 2 Pipistrellus javanicus were Fifty-six net-nights yielded 1 Cynopterus captured by hand. brachyotis, 1 Hipposideros diadema, 8 Site 4 (9Њ33Ј45ЉN, 118Њ27Ј54ЉE) is a cave Rhinolophus arcuatus, and 1 R. creaghi. known locally as ‘‘Ma-ngit’’. It is located 276 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON along the Iraan River near Sitio Pamolkoan, 2000. A total of 575 bats belonging to 10 Barake, Aborlan, at ca. 430 m elevation. species (18 Cynopterus brachyotis, 1 Me- The cave is in a small valley, restricted by gaderma spasma, 100 Hipposideros ater, mountains to the east and west. A small, 239 H. diadema, 14 Rhinolophus arcuatus, seasonal stream flows through the cave. At 33 R. creaghi, 10 R. macrotis, 86 R. virgo, least six entrances to the cave were evident, 15 Miniopterus australis, and 59 M. schrei- and many small tunnels connected medium bersi) were captured. to small caverns, which ranged from well- Site 6 (9Њ28Ј25ЉN, 118Њ30Ј21ЉE) was a lit to completely dark. Very little distur- mostly abandoned agricultural area located bance was evident in or around the cave in Barake, Aborlan Municipality, at eleva- due to its isolated location (ca. 10 hour hike tion ranging from 40–80 m. A small stream from the nearest road). The cave was sur- flowed through the area and topography rounded by a large expanse of primary low- was flat to rolling. Vegetation was a mosaic land forest, but some agricultural areas of grassland (primarily Imperata cylindri- were present within ca. 5 km. Disturbances ca) with sparse trees (mostly Vitex sp.), to local vegetation included collection of cashew plantations, dense brush, and very rattan (Calamus spp.). We surveyed bats in small (Ͻ1 ha) areas of secondary growth. Ma-ngit Cave for six days during December Frequent fires appeared to maintain this 1999. We captured 819 bats belonging to area as a grassland. We surveyed small vo- seven species (9 Eonycteris spelaea, 43 lant and non-volant mammals for four days Hipposideros diadema, 367 Rhinolophus during June 2000. A single trap-line baited creaghi, 4 R. virgo, 9 Miniopterus australis, with live earthworms yielded 17 Rattus ex- 386 M. schreibersi, and 1 M. tristis) inside ulans in 186 trap-nights. We captured 7 Cy- the cave. nopterus brachyotis in 6 net nights, and 1 Site 5 (10Њ05Ј00ЉN, 118Њ51Ј06ЉE) is a Kerivoula whiteheadi in 3 harp-nights. complex area of limestone karst containing Site 7 (9Њ29Ј15ЉN, 118Њ29Ј24ЉE) was in a probably greater than 100 caves in Baran- narrow band of secondary forest in Barake, gays Tagabinet and Cabayugan, Puerto Aborlan Municipality, located between dis- Princesa; elevation is ca. 50 m. Caves in the turbed habitat at lower elevation (Site 6) area ranged from tiny cracks too small to and primary and good secondary forest at enter, to large complexes of multiple cav- higher elevation. Elevation ranged from erns with multiple entrances. Local terrain 80–140 m; slope was rolling to moderately was generally flat except for the sometimes- steep, and two small streams dissected the massive limestone outcrops, which form area. Canopy height varied from 5–20 m. high-rising cliffs throughout the area. Caves Woody vines and lianas were common and probably are present all over these complex vegetation was quite dense in areas. Wild formations, but only the very few found bananas (Musa spp.) were abundant, but near ground level were accessible. We cap- patchy in distribution; leaf litter depth was tured bats in and around five different highly variable. We surveyed this site for caves/cave complexes. These represented small volant and non-volant mammals for some of the most accessible caves in the four days during June 2000. A single trap area. Disturbance at the caves was moder- line baited with live earthworms yielding ate, with vandalism and guano excavation 120 trap-nights produced 1 Rattus exulans. evident at most caves. Most of the caves Six net-nights produced 27 Cynopterus were surrounded immediately by agricul- brachyotis and 1 Macroglossus minimus, tural development. Both primary and sec- and 3 harp-nights yielded 1 Hipposideros ondary lowland forests were present in the diadema and 1 Kerivoula pellucida. surrounding hills. We surveyed bats at these Site 8 (9Њ59Ј47ЉN, 118Њ56Ј43ЉE) is a caves for 14 days between March and May large cave complex located in a limestone VOLUME 117, NUMBER 3 277 karst formation on top of the first ridge up Minor damage had been done by both trea- from San Rafael, Puerto Princess, at an el- sure hunters and guano collectors. The sur- evation of ca. 250 m. The cave complex is rounding vegetation was dominated by ag- known locally as ‘‘Taraw’’. The cave sys- ricultural areas with some strips and patches tem appeared to be quite large; we were of residual forest. Large expanses of sec- unable to explore much of it due to a lack ondary and logged-over forest were found of climbing equipment and expertise. Some nearby in the vicinity of Lake Manguao. We caverns exceeded 20 m in height, while oth- captured 115 bats belonging to 4 species ers were quite small. We found no perma- (48 Eonycteris spelaea, 1 Rousettus am- nent water in or around the cave, but evi- plexicaudatus, 64 Hipposideros diadema, dence of storm flow was present. Evidence and 2 Miniopterus tristis) using a harp trap of vandalism and guano collection was pre- and butterfly net at one of the entrances to sent. A mixture of habitats surrounded the the cave during three days in October 2000. cave complex: between the cave and the Site 11 (10Њ46Ј34ЉN, 119Њ31Ј52ЉE) was community of San Rafael, the vegetation located around the perimeter of Lake Man- was dominated by brushland and agricul- guao, in Barangays Poblacion and Bantu- tural developments, while on the other side lan, Taytay Municipality. The area was of the cave secondary and primary forest dominated by secondary and logged-over dominated, with mixed areas of slash-and- forest, with disturbance from slash and burn burn fields. We surveyed bats at this cave agriculture being found throughout the area. for five days during July 2000. We captured The area retained ca. 60% forest cover. 2775 bats representing 10 species (7 Hip- Slope was generally moderate to steep, and posideros ater, 43 H. diadema, 90 Rhino- elevation ranged from ca. 40–250 m. We lophus acuminatus, 240 R. arcuatus, 239 R. trapped for small volant and non-volant creaghi, 25 R. macrotis, 151 R. virgo, 1257 mammals in forest, agricultural habitats, Miniopterus australis, 711 M. schreibersi, and two small caves near the lake for 14 4 immature M. sp., and 8 Myotis macrotar- days during October and November 2000. sus). We totalled 471 trap-nights in three lines Site 9 (9Њ39Ј40ЉN, 118Њ27Ј48ЉE) is a baited with coconut and peanut butter, and small cave located in Sitio Labtay, Napsan, captured 3 Tupaia palawanensis, 23 Max- Puerto Princesa City. The cave, which con- omys panglima, and 1 Rattus exulans. For- sisted of a single chamber 1–2 m wide, 3– ty-two net-nights yielded 53 Cynopterus 6mhigh, and ca. 30 m long, was in a nar- brachyotis, 2 Macroglossus minimus, and 1 row canyon along the Panagurian River at Rousettus amplexicaudatus, and 13 harp- ca. 280 m. Vegetation in the area consisted nights produced 1 Megaderma spasma, 1 of good-quality secondary forest, second- Rhinolophus acuminatus, 2 Kerivoula hard- growth forest, and agricultural develop- wickii, and 5 Tylonycteris pachypus. Addi- ments. We trapped bats with a harp trap, tionally, we captured 4 Megaderma spasma mist nets, and a butterfly net in the cave and 4 R. acuminatus by hand. and surrounding forest for five days during Site 12 (9Њ27Ј48ЉN, 118Њ32Ј16ЉE) was lo- August 2000. We captured 73 bats (4 Cy- cated at the ‘‘rainforestation’’ site in Sitio nopterus brachyotis, 68 Hipposideros dia- Kandis, Aborlan Municipality, in forest/ dema, and 1 Rhinolophus arcuatus). grassland mosaic at ca. 40 m above sea lev- Site 10 (10Њ44Ј00Љ, 119Њ34Ј23Љ)isacave el, about seven km away from the next located near sea level in Sitio Sader, Ban- good secondary forest in the foothills of the tulan, Taytay Municipality. The cave con- Victoria Range. Charcoal making, logging, sisted of a single chamber (ca. 3–8 m wide, rattan collection, grazing and burning were 3–7 m high, and 40 m long) with a large common until 1994 when such activities (Ͼ2.5 m diameter) entrance at each end. were made illegal. The terrain was flat to 278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rolling, dissected by two creeks. The site Island, Narra Municipality, a small (8.3 consisted of 5.5 ha Imperata cylindrica km2) shallow coral island, 1.8 km offshore grassland interspersed with single shrubs in the Sulu Sea. Approximately two-thirds and trees, predominantly Antidesma ghae- of the island was covered with mangrove sembilla, which forms the fire climax in and one-third with coastal forest over lime- more open situations, with Vitex pubescens, stone. About five percent of the latter had Guioa pleuropteris, Tarenna stenantha, Fa- been converted into coconut plantation. Se- graea fragrans, Lantana camara, and Mus- lective logging was done until the early saenda philippica in protected areas not af- 1990s, resulting in the complete loss of ma- fected by fire in the last ten to fifteen years. ture Intsia bijuga. The mangrove consisted About 4.5 ha consisted of regenerating for- of nine species of the genera Rhizophora, est, close to two seasonal creeks, dominated Sonneratia, Avicennia, Bruguiera, Aegicer- by Garcinia benthami, G. parviflora, Can- as, and Ceriops. Canopy height was vari- arium asperum, Polyscias nodosa, and Bar- able, usually between 8 and 15 m. Emer- ringtonia curranii. Undergrowth was mod- gent trees (e.g., Garuga floribunda and erate to very dense. Canopy height was on Pterocymbium taluto) ranged up to 42 m. average eight meters, with some taller Leaf litter layer was not closed, except un- emergents such as Nephelium sp. and Dip- der very dry conditions. Barren coral rocks terocarpus gracilis. The most conspicuous and crevices were ubiquitous. Buttresses vine was Gnetum latifolium. Macrophytic were a common feature of all emergent for- epiphytes were virtually absent. Leaf litter est trees. Under open conditions, an herbal layer was usually not closed, except in very layer consisting of Impatiens sp. was pre- dry years. Surveys were conducted regular- sent. Vines were abundant, including climb- ly from 1997–2000 with 10 medium-sized ing bamboo Dinochloa sp., often forming Sherman live traps, 10 commercial live rat dense tangles. Macrophytic orchids were traps, and 55 wire mesh traps (measure- present, but relatively scarce. Traps and nets ments equal to medium Shermans). Baits were set along a trail within the coastal for- were roasted coconut with peanut butter, est. Traps were baited with roasted coconut but mostly fruits available in the area. A with peanut butter, and a few with crickets. total of 3514 trap-nights yielded 169 small Most traps caught hermit crabs. Nets were mammals (28 Tupaia palawanensis, 8 Sun- set in the understory, which was very open dasciurus juvencus, 11 Rattus exulans, 16 from January to April 2002 and only pro- Rattus tiomanicus, 104 Maxomys panglima, vided very few fruits due to an extended and 2 Sundamys muelleri). Mist nets (2 ϫ dry spell. Trapping totaled 104 trap-nights, 6m)were set along trails in forest, in grass- and produced 3 Rattus tanezumi and 2 R. land, gaps in the shrub cover, and rarely in tiomanicus. Netting totaled 28 net nights, the canopy (ca. 8 m high). Capture sites and yielded 2 Cynopterus brachyotis, 5 Ma- were often near or in fruiting shrubs or croglossus minimus, and 5 Megaderma trees, since the main focus of the study was spasma; all bats were released, and the rats on frugivores. From 1997 to 2000, a total preserved as vouchers. of 482 net-nights yielded 1257 bats (1 Ac- Site 14 (9Њ17ЈN, 118Њ27ЈE) was in fresh- erodon leucotis, 829 Cynopterus brachy- water swamp forest in Narra Municipality, otis, 394 Macroglossus minimus, 5 Eonyc- in about 5 ha of remnant forest along Tar- teris spelaea, 4 Rousettus amplexicaudatus, itien River. The habitat was dominated by 18 Megaderma spasma, 1 Hipposideros two woody species, Nauclea orientalis and diadema, 4 Scotophilus kuhlii, and 1 Mu- Pandanus sp., at lower elevations, which rina cf. tubinaris); all but the M. cf. tubi- are flooded for at least six months. The herb naris were released. layer was not extensive and was dominated Site 13 (09Њ13ЈN, 118Њ26ЈE) was on Rasa by Acrostichum sp. The higher portions VOLUME 117, NUMBER 3 279 were dominated by pioneering species of reported a single humerus of a shrew, sub- early to medium successional stages, such stantially smaller in size than C. palawa- as Trema orientalis, Vitex pubescens, and nensis, from undated sediments in a small Commersonia bartramia. Even during ex- rock-shelter cave near Tabon Cave on Lip- treme dry spells like that in the first half of uun Point, near Malunut Bay, in Quezon 2002, there were isolated open water bodies Municipality (near the location of the town left, which connected to several creeks dur- of Quezon as shown in Fig. 1). They de- ing the rainy season. The swamp forest is scribed the specimen as being similar in bordered by ricefields and grassland. Twen- size to C. monticola from Borneo. We ten- ty trap-nights yielded 1 Rattus exulans and tatively include it in our tallies of native 1 R. tiomanicus. Twelve net-nights yielded species of Palawan, but we recommend that 27 Cynopterus brachyotis, 5 Macroglossus it be sought by trapping with small snap- minimus, and 1 Megaderma spasma; all traps baited with live earthworms and pit- bats were released and the rats preserved as fall traps. vouchers. Suncus murinus.—This introduced com- Site 15 (10Њ12ЈN, 118Њ55ЈE) was along mensal is abundant in urban and agricultur- the ‘‘jungle trail’’ near the Central Park Sta- al areas (Rabor 1986); in forest, it is rarely tion in Puerto Princesa (formerly St. Paul) present, but occasionally is common (Hea- Subterranean River National Park ney et al. 1989, Heaney & Tabaranza 1997). (PPSRNP). Primary lowland forest on steep It is found throughout Asia and Indo-Aus- slopes ascended from sea level to about 40 tralia, including the Philippines (Heaney et m. Five net-nights on 15 September 1996 al. 1998). We observed this species fre- yielded 1 Cynopterus brachyotis, 2 Rhino- quently in houses in Puerto Princesa City lophus arcuatus, 3 Rhinolophus virgo, and and the State Polytechnic College of Pala- 2 Rhinolophus sp.; all were released. Ad- wan in Aborlan Municipality. ditionally, all three authors made visual ob- servations at various times. Order Scandentia Family Tupaiidae—Tree Shrews Accounts of Species Order Insectivora Tupaia palawanensis.—This common Family Soricidae—Shrews species is endemic to the Palawan faunal region (Wilson 1993); it is related to T. glis, Crocidura palawanensis.—We never en- which is widespread on the Sunda Shelf countered this poorly known species. It is (Corbet & Hill 1992). It is widespread on endemic to the Palawan faunal region and Palawan (Taylor 1934), and is usually com- has been taken in old-growth rain forest and mon in secondary and primary lowland for- shrubby second growth (Heaney & Ruedi est, though local densities may be highly 1994); the holotype came from ‘‘deep forest variable between apparently similar habitats near the sea at . . . Brooke’s Point’’ (Taylor (Dans 1993, Hoogstraal 1951, Sanborn 1934), a second from near sea level in Ba- 1952). It is rare in montane forest, and com- buyan, Puerto Princesa (Hoogstraal 1951, mon but patchy in agricultural areas. We Sanborn 1952), and a third from 3600– captured and/or observed this species in co- 4350 ft (ca. 1100–1300 m) on Mt. Mantal- conut and cashew plantations, brushy areas ingajan (USNM); two additional specimens with a few small trees (Sites 6, 11, and 12), are from Balabac (Heaney & Ruedi 1994). secondary and logged-over forest (Sites 7 IUCN (2002) lists this species as Vulnera- and 11), and primary forest (Sites 1, 2, 3, ble, but current definitions suggest that Data and 15) from near sea level to 1400 m. Deficient would be more appropriate. IUCN (2002) lists this species as Vulnera- Crocidura sp.—Reis & Garong (2001) ble, but we concur with Heaney et al. 280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(1998) that the species should be delisted and in coastal forest on Rasa Island (Site due to the variety of habitats used and its 13). We also captured this species in pri- apparent abundance. Specimens examined: mary forest in a tree fall gap (Site 1), over 2: Site 1 (1), Site 2 (1). a stream (Site 2), and at a place with no visible disturbance (Site 15). We found Order Chiroptera them roosting in various-sized groups in Family Pteropodidae—Fruit Bats three caves at Sites 5 (there appeared be less than 50 in each of two caves) and 9 We follow Ingle & Heaney (1992) and (ca. 300 individuals); although this species Heaney et al. (1998) in regarding reports of occasionally roosts in caves on Borneo Haplonycteris fischeri (Kock 1969) and (Payne et al. 1985), there are no previous Ptenochirus minor (Yoshiyuki 1979) from records of such roosts in the Philippines. Palawan as erroneous, probably originating On several occasions we captured them car- in mislabeled specimens. Pteropus hypo- rying whole green figs (Ficus sp.) during melanus is known from Cuyo Island, at the flight; two of these individuals were return- northeast edge of the Palawan faunal region ing to a cave at Site 5 between 1900– (Heaney et al. 1998), as well as in the oce- 2000 h. anic Philippines and on islets around Bor- Out of 20 adult females caught at Site 14 neo, and should be sought on Palawan. on 20 March 2002, 15 were pregnant and 5 Acerodon leucotis.—This poorly known were carrying a single suckling young. On species is endemic to the Palawan faunal 1 and 2 April 2000, we captured three adult region. Hoogstraal (1951) found the species females at Site 5; all were carrying a single in an area with patches of ‘‘much disturbed suckling infant during flight. On 17 May remnants of original forest and dense sec- 2000 we captured eight adult females at ond growth forest’’ on Busuanga Island, Site 5; one was pregnant, one was carrying and two specimens were taken at Santiago, a suckling infant during flight, and two Iwahig (in Puerto Princesa) on Palawan were emaciated and may have recently (Sanborn 1952). A specimen from Bat Is- weaned their young. On this date, we also land (Barangay Tagburos, in Honda Bay), captured a male with enlarged mammaries taken by P. O. Glass in 1978, is housed in (see Francis et al. 1994). On 30 October the UMMZ. Heaney sighted large numbers 2000, among 25 adult females, none were of medium-sized, pale-furred flying foxes at pregnant but one was lactating. Specimens Site 15 in April 2000, in the clearing of the examined: 4, Site 1 (1), Site 5 (3). old park headquarters near the center of the Eonycteris spelaea.—This widespread park (‘‘Central Park’’), that were probably Southeast Asian species is common in ag- this species. Widmann captured one at Site ricultural areas in the Philippines, where all 12 at a height of 5 m, and saw others feed- known roosts are in caves (Heaney et al. ing in the canopy at ca.8mheight. The 1998, Rickart et al. 1993). Sanborn (1952) IUCN (2002) lists this species as Vulnera- reported a large series from a cave above ble; we regard it as Data Deficient. Tanabog, Palawan. We netted five individ- Cynopterus brachyotis.—Found through- uals at Site 12, but most of our records out Southeast Asia; in the Philippines, it is came from caves in lowland forest. We common to abundant in secondary forest found this species roosting in caves at Sites and agricultural areas, and rare in primary 4 and 10; at Site 4, the roosting population forest (Heaney et al. 1998); Sanborn (1952) appeared to exceed 2000. At Site 10, there reported many from Palawan. We netted was an extremely large population (proba- this species frequently in secondary forest bly Ͼ50,000) of small pteropodids roosting and agricultural areas at Sites 6, 7, 11, and inside the cave. We captured 49 pteropodids 12, in freshwater swamp forest at Site 14, at the entrance to the cave, 48 of which VOLUME 117, NUMBER 3 281 were E. spelaea and one was a Rousettus hunting and perhaps to electrocution on amplexicaudatus. On 19 December 1999, power lines. all three adult females we captured at Site Rousettus amplexicaudatus.—Within the 4 were pregnant. On 21 October 2000 at Philippines, this widespread Southeast Site 10, we captured 11 adult female E. spe- Asian species is commonly found in agri- laea, four of which were carrying an infant cultural habitats up to 500 m and rarely in during flight and five of which were preg- primary lowland forest (Heaney et al. nant. This species is heavily hunted in some 1998). All known roosting sites are in caves areas of the Philippines (Rickart et al. 1993, (Heaney et al. 1989, 1991, 1998, 1999; Hei- Utzurrum 1992), but we observed no evi- deman & Heaney 1989; Rickart et al. dence of that being the case on Palawan. 1993). According to Payne et al. (1985), R. Specimens examined: 5, Site 4 (3), Site 10 amplexicaudatus often roosts in association (2). with Eonycteris spelaea. We netted one in- Macroglossus minimus.—In the Philip- dividual from a cave at Site 10 containing pines, this widespread Australasian species a large population of E. spelaea, one in an is common in secondary forest and agri- agricultural clearing in Site 11, and four in cultural areas and uncommon in primary forest-grassland mosaic at Site 12; all of forest up to more than 2000 m (Heaney et these sites are in heavily disturbed areas be- al. 1998, 1999). We captured this species in low 60 m. Specimens examined: 2, Site 10 secondary lowland forest (Sites 7 and 12) (1), Site 11 (1). and agricultural clearings (Site 11), usually near wild or domestic banana plants (Musa Family Emballonuridae—Sheath-tailed spp.), and in freshwater swamp forest in Bats Narra Municipality and Rasa Island (Sites 13 and 14). Specimens examined: 3, Site 7 There are no known records of Saccolai- (1), Site 11 (2). mus saccolaimus from Palawan, but its Pteropus vampyrus.—In the Philippines, widespread distribution from India to New this widespread Southeast Asian species oc- Guinea, including the oceanic Philippines curs in primary lowland forest and adjacent (Heaney et al. 1998), suggests that it may agricultural areas (Heaney et al. 1998; Ra- be present and should be sought. bor 1955, 1986; Rickart et al. 1993; San- Emballonura alecto.—The Philippine born 1953; Taylor 1934). Widmann esti- sheath-tailed bat is known from Borneo, the mated 400 individuals on Malinau Island, Philippines, and Sulawesi (Heaney et al. Aborlan Municipality in 1998, 570 on Rasa 1998); we never encountered this species Island on 12 November 1999, and a small on Palawan, but Taylor (1934:200) captured colony (ca. 40 individuals) at Lagan on Du- five individuals ‘‘under an overhanging maran Island on 27 October 2001, based on rock along Iwahig River, near the base of departure counts. Flying foxes commonly Thumb Peak’’. sighted in Puerto Princesa City around Taphozous melanopogon.—The bearded mango and guyabano (ϭ sour sop) trees are tomb bat is widespread in southern Asia probably this species. In 1998, we found (Heaney et al. 1998). In the Philippines, it two individuals of this species that ap- is common in urban areas and lowland ar- peared to have been electrocuted on power eas with limestone caves and rare in forest lines, one at the Provincial Agriculture Cen- (Rickart et al. 1993, Sanborn 1952). There ter in Irawan, Puerto Princesa, and the other is a previous record from the vicinity of at the State Polytechnic College, Aborlan. Puerto Princesa (Allen 1922), and A. C. Al- We believe this species to be common over- cala collected 6 specimens from Sitio Mal- all, but under moderate pressure due to abusog, Tinitian, Roxas Municipality in 282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1984 which are deposited in the UMMZ. three caves in disturbed lowland forest at We never encountered this species. 50 to 250 m elevation at Sites 5 (17% of 575 captures) and 8 (Ͻ1% of captures). Family Megadermatidae—False Vampire During March to April 2000, none of the and Ghost Bats 26 females we captured at Site 5 were preg- nant or lactating, but on 19 and 20 May Megaderma spasma.—This widespread 2000, 25 of 30 adult females were pregnant. southern Asian species is common in pri- We have re-examined a specimen from Pa- mary lowland forest and disturbed forest in lawan in the UMMZ identified by Allen the Philippines (Heaney et al. 1991, 1998, (1922) as H. bicolor, and a series from the 1999; Rickart et al. 1993). We captured this Tigoplan River, Palawan in FMNH reported species from sea level to ca. 500 m in sec- by Sanborn (1952), and now consider them ondary forest (Site 7), primary forest (Sites to be H. ater; thus, we now know of no 1 and 2), in a bamboo thicket (Site 11), and records of H. bicolor from Palawan. Cranial in or near caves (Sites 5 and 11). It was the measurements of 5 individuals (Table 1) are most common insectivorous bat netted in smaller than those of H. bicolor (Heaney et forest-grassland-mosaic (Site 12), in swamp al. 1999, Ingle & Heaney 1992) but match forest (Site 14), and in coastal forest (Site those of H. ater (Ingle & Heaney 1992, 13). At Site 1, we found this species roost- Rickart et al. 1993). Specimens examined: ing in small groups (Ͻ10) in four hollow 5, Site 5 (4), Site 8 (1). trees distributed throughout the area. At Hipposideros diadema.—The diadem Site 11 we found ca. 12 individuals roosting roundleaf bat is widespread from Myanmar in a small cave (ca. 0.5–3 m wide, 0.3–1.5 to the Solomon Islands, with many previous m high, and 10 m long) along with Rhino- records from Palawan (Allen 1922, Heaney lophus acuminatus. We also found two in- et al. 1998). In the Philippines, it is com- dividuals roosting in a small cave (also Site mon in disturbed forest, agricultural areas 11) that had been severely disturbed by (Ingle 1992, Rickart et al. 1993), and pri- treasure hunters three years earlier. Cranial mary forest (Heaney et al. 1998, Rickart et measurements of three individuals (Table 1) al. 1993). Reis & Garong (2001) reported are slightly smaller than those of specimens specimens from sediments in a rock-shelter from Leyte and Biliran (Rickart et al. 1993) near Tabon Cave, Quezon Municipality dat- and southern Luzon (Heaney et al. 1999). ed to 11,130 BP. We captured this species Specimens examined: 3, Site 1 (3). from sea level to 600 m in disturbed grass- land-forest mosaic (Sites 6 and 12), second- Family Rhinolophidae—Horseshoe and ary forest (Site 7), primary forest (Site 2), Roundleaf Bats and at nearly all caves we visited (Sites 4, Several poorly known but apparently 5, 8, 9, and 10), and we observed large widespread species in this family occur on numbers (probably thousands) in the un- the Sunda Shelf and in the oceanic Philip- derground river cave at PPSRNP. All of the pines and should be sought on Palawan; roosts we identified held groups of H. dia- these include Hipposideros cervinus and H. dema numbering greater than 200. Thirteen lekaguli (Balete et al. 1995, Heaney et al. adult females captured in December 1999 1998, Ingle & Heaney 1992). included none that were pregnant or lactat- Hipposideros ater.—Occurs from India ing. At Site 5 in March to April 2000, none to Australia (Heaney et al. 1998). Known of the 54 adult females were pregnant or from lowland and montane forest and caves lactating, but between 15 and 20 May 2000 (Heaney et al. 1991, 1998; Payne et al. 26 of 43 were pregnant and one was car- 1985, Rickart et al. 1993). We found this rying a suckling infant during flight. One of species to be uncommon to abundant in 21, one of 12, and none of 13 adult females VOLUME 117, NUMBER 3 283 40.5 41.5 45.5 47.7 54.4 52.4 46.5 46.0 46.2 47.7 55.7 54.3 39.7 40.3 length Forearm (39.2–41.3) (40.8–41.9) (45.1–45.8) (46.9–48.5) (51.5–54.6) (45.2–47.7) (45.6–46.5) (46.6–48.1) (55.4–56.0) (39.4–39.9) (40.1–40.4) — — 5.12 6.85 6.74 6.31 7.14 4.87 5.21 length Palatal 5.7 (2) 5.08 (2) 6.37 (1) 6.87 (2) 6.56 (1) (4.85–5.31) (4.43–5.63) (5.64–5.76) (6.76–6.98) (6.51–6.97) (6.05–6.48) (4.83–4.94) (4.95–5.47) 3.57 3.46 3.39 3.58 5.34 4.83 4.12 4.15 4.77 4.76 3.77 3.79 3.28 3.43 Palatal breadth (3.5–3.66) (3.3–3.56) (3.5–4.04) (3.36–3.43) (3.41–3.75) (4.55–5.06) (4.03–4.21) (4.08–4.27) (4.66–4.95) (3.22–3.33) (3.41–3.45) 4.71 4.72 4.87 4.93 6.62 6.45 5.52 5.48 7.07 7.06 6.15 6.01 3.94 4.05 toothrow Molariform (4.9–4.95) (4.0–4.1) (4.59–4.85) (4.67–4.76) (4.84–4.91) (6.37–6.52) (5.46–5.57) (5.29–5.63) (6.95–7.18) (5.78–6.40) (3.72–4.21) molar 6.16 6.27 6.51 6.63 8.96 8.45 7.54 7.14 8.79 9.0 7.65 7.59 4.8 4.86 to last Canine (7.5–7.75) (6.04–6.23) (6.15–6.36) (6.49–6.53) (6.48–6.78) (8.39–8.51) (7.52–7.56) (6.95–7.47) (8.67–8.91) (4.68–4.93) (4.75–4.97) 4.84 4.87 4.20 4.37 6.7 6.25 7.37 7.57 5.08 5.21 6.23 6.17 4.46 4.78 length Orbital (4.7–4.94) (7.3–7.43) (4.3–4.56) (4.84–4.89) (4.33–4.42) (4.17–4.22) (5.87–6.59) (5.01–5.41) (5.02–5.18) (6.05–6.21) (4.61–4.94) width 8.55 8.48 9.37 9.48 9.06 9.52 9.81 9.68 8.28 8.31 11.27 11.32 Mastoid 11.12 10.62 (8.41–8.72) (8.45–8.51) (9.32–9.44) (9.39–9.56) (9.51–9.52) (8.64–9.36) (9.63–9.78) (8.26–8.30) (8.23–8.39) (10.51–10.78) (11.26–11.28) 8.87 8.72 8.63 8.66 9.79 9.53 7.83 7.82 width 11.77 11.03 10.46 9.86 6.19 7.58 6.00 4.51 — 46.39 14.54 14.17 10.89 10.70 Zygomatic (8.78–8.95) (8.68–8.78) (8.62–8.64) (8.61–8.70) (9.72–9.86) (9.41–9.62) (10.5–10.92) (7.74–7.90) (7.76–7.88) (10.83–11.27) (14.52–14.55) — — — length 16.16 17.68 16.86 8.64 8.57 4.88 6.65 5.07 4.13 5.84 42.8 22.26 18.78 18.68 20.35 14.29 14.54 16.13 (2) 17.3 (2) 20.87 (2) 19.44 (1) (20.7–21.03) (18.3–19.26) Condyloincisive (15.99–16.26) (15.72–16.48) (17.21–17.38) (17.28–18.08) (18.41–18.91) (14.21–14.37) (14.31–14.77) 3 3 3 2 1 4 2 1 2 3 1 4 3 2 f f f f1 f f f f m m m m m1 m m m Species Sex n Table 1.—Means and ranges of cranial measurements of adults of Megadermatidae and Rhinolophidae from Palawan Island, Philippines. (holotype) macrotis neensis seni aequilis cuatus natus Rhinolophus virgo Rhinolophus Rhinolophus bor- Rhinolophus creaghi Megaderma spasma Rhinolophus ander- Rhinolophus ar- Rhinolophus acumi- Hipposideros ater 284 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from July, August, and October were preg- seni aequalis (Allen 1922) was this species. nant (Sites 8, 9, and 10, respectively). Spec- Heaney compared this specimen, which is imens examined: 5, Site 4 (2), Site 5 (2), the holotype and only known specimen, to Site 9 (1). series of all species currently known from Rhinolophus acuminatus.—This poorly Palawan. It was unambiguously identified known species occurs from Thailand to as R. acuminatus;asnoted by Medway Lombok and Palawan, but not elsewhere in (1977:32), a dorsal connecting process with the Philippines (Heaney et al. 1998; speci- a prominent triangular point (as shown by mens from Negros reported by Csorba et al. Medway 1977 fig. 6a and by Ingle and Hea- (2003) as this species were mislabelled). It ney 1992 fig. 13a) is present, the base of is found in lowland forest on Borneo the sella is not expanded into a cup, the (Payne et al. 1985) and in secondary low- median groove of the horseshoe is not land dipterocarp forest on Banggi (Md. Nor broadened, and papillae are not present. 1995). We captured this species in caves The ears (18 mm) are less than half of the from ca. 60 to 250 m in caves (Sites 8 and length of head plus body, and the forearm 11), a bamboo thicket (Site 11), and pri- is 46.4 mm. The skull (measurements in Ta- mary forest (Site 1). At Site 8, we captured ble 1) is virtually identical to those in the 90 individuals out of 2775 captures. At Site series from Sites 1 & 8, including overall 11 we found ca. 20 individuals roosting in size and shape, nasal swellings, braincase a small cave (ca. 0.5–3 m wide, 0.3–1.5 m breadth and inflation, toothrows, palatal high and 10 m long) with ca. 12 Megad- bridge, foramina in the roof of the posterior erma spasma. At Site 1, we took two in- portion of the nasal passage, and bullae. Be- dividuals over a small stream just below a cause R. acuminatus was named by Peters rock outcrop containing many fissures suit- in 1871 (from Gadok, Java), we therefore able for roosting bats. We also captured a recognize R. anderseni aequalis as its ju- single individual in our temporary living nior synonym. We note that Cabrera (1909) quarters at Site 1 after we observed for sev- described Rhinolophus anderseni ‘‘proba- eral days a bat feeding inside our semi-en- bly from Luzon’’. Aside from the holotype closed tent for several minutes daily be- of R. anderseni aequalis, no specimens tween 0430 and 0600 h. Of 15 adult fe- have subsequently been referred to this spe- males taken in July 2000 at Site 8, one was cies. Csorba et al. (2003) tentatively as- pregnant and one was lactating. Cranial signed R. anderseni Cabrera as a junior measurements (Table 1) match those pre- synonym of R. arcuatus on the basis of the viously available (Ingle & Heaney 1992) original description plus new drawings of that are based on series from Balabac and the noseleaf and measurements of the skull, Busuanga reported by Kuntz (1969) and but without examining the holotype. We housed in the USNM. We also refer two provisionally accept this, but point out the specimens collected by A. C. Alcala on 13 need for direct examination and compari- July 1984, at Malabusog, Roxas Munici- sons. IUCN (2002) lists R. acuminatus as pality housed at UMMZ (162885 and Data Deficient. Specimens examined: 8, 162886) to this species, and include them Site 1 (4), Site 8 (1), Malabusog (2), and in Table 1. the holotype of R. anderseni aequalis. Sanborn (1952) reported a single speci- Rhinolophus arcuatus.—Widespread men from Palawan (housed in FMNH) that from Sumatra to New Guinea (Heaney et he regarded as the first record from Pala- al. 1998). Specimens from the Philippines wan. However, we have determined that a currently identified as R. arcuatus may con- single specimen (UMMZ 53112) collected sist of two or more species (Heaney et al. in the late 1800s by the Beal/Steere Expe- 1991, 1999; Ingle & Heaney 1992; Rickart dition and subsequently named R. ander- et al. 1993). Individuals referred to this VOLUME 117, NUMBER 3 285

‘‘species’’ have been found in agricultural area of mixed agricultural/second growth areas, secondary forest, and primary low- forest within 1 km of mature forest. Spec- land, montane, and mossy forest (Heaney et imen examined: 1, from Sabang. al. 1991, 1999; Ingle 1992; Rickart et al. Rhinolophus creaghi.—This species was 1993). We regularly captured this species at previously known from Borneo and Madura elevations from sea level to 1400 m in low- Islands, where it often roosts in caves (Cor- land primary forest (Sites 1 and 2), mon- bet & Hill 1992, Csorba et al. 2003, Koop- tane forest (Site 3), and caves (Sites 5 and man 1993, Medway 1977, Payne et al. 8). We also captured a single individual in 1985). This is the first record of this species the understory of mature but disturbed for- from Palawan Island and the Philippines. est near a cave at Site 9, and we tentatively On Palawan, we found it to be common in identified two individuals from Site 15 primary lowland forest from near sea level (which were released) as belonging to this to at least 700 m. We captured one individ- species. Of 8 adult females taken between ual at Site 1 and 11 individuals at Site 2. It 15 and 20 May 2000 at Site 5, 5 were preg- roosts in caves, often in large numbers; we nant and one was lactating. Of 22 adult fe- captured 368 (45% of captures) at Site 4, males taken in July 2000 at Site 2, one was 239 (9% of captures) at Site 8, and 33 (6% lactating, and of 189 captured in July at Site of captures) at Site 5. Of 135 adult females 8, none were pregnant but 14 were lactat- captured in December 1999 at Site 4, 8 ing. These are the first specimens of this were pregnant. Of 16 captured at Site 5 in species from the Palawan faunal region. March to April, none were reproductively Cranial measurements (Table 1) closely active; of 11 at Site 2 and 151 at Site 8 match those of specimens from Leyte (July 2000), none were pregnant but one (Rickart et al. 1993) and southern Luzon and 12 were lactating, respectively. Cranial (Heaney et al. 1999). Specimens examined: measurements (Table 1) show this to be the 5, Site 1 (2), Site 2 (1), Site 3 (2). largest member of the genus on Palawan; Rhinolophus cf. borneensis.—A single our specimens are not distinguishable from specimen taken by P. O. Glass on 31 Jan- a small series from Borneo (FMNH 47071– uary 1978 at ‘‘Sabang, Buenavista’’ (in 47075). Two previously unidentified speci- Barangay Cabayugan, near Ulugan Bay in mens from Mt. Salicod, 2300 ft. (which Puerto Princesa Municipality, ca. 10Њ05ЈN, may be the same mountain as Mt. Salakot, 118Њ49ЈE; UMMZ 161395) appears to be Site 2; P. O. Glass, in lit.), taken by P. O. this species. It was previously known from Glass in 1978 and housed in the UMMZ, Indochina, the Malay Peninsula, Java, and were taken earlier but were not reported; Borneo, as well as some smaller islands in cranial measurements from these specimens the southern South China Sea (Corbet & are included in Table 1. This species is list- Hill 1992, Csorba et al. 2003); this is the ed as Near-Threatened by IUCN (2002). first record from Palawan and from the Specimens examined: 7, Site 4 (3), Site 5 Philippines. Cranial measurements and fea- (1), Site 8 (1), Mt. Salicod (2). tures (Table 1) closely match specimens in Rhinolophus macrotis.—This poorly FMNH from Sarawak, the Natuna Islands, known species ranges from India to Su- and Sabah, and external features are similar, matra and the Philippines, where it is but because we have only a single speci- known from lowland forest with some re- men, the identification is tentative. On Bor- cords from caves (Heaney et al. 1998, Ingle neo, ‘‘the species roosts in caves, some- 1992). Our specimens represent the first re- times in colonies of several hundred indi- cord from Palawan. We captured this spe- viduals’’ (Payne et al. 1985). P. O. Glass (in cies in or near three caves in disturbed low- litt.) noted that he captured the specimen in land forest at 50–250 m at Sites 5 (10 cap- a mist net in a small banana grove in an tures) and 8 (25 captures). The species ap- 286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pears to be uncommon. At Sites 5 and 8 it shown to be distinct from Myotis muricola represented less than 5% and 1% of our and to be present on Culion Island in Pa- captures, respectively. At Site 5 on 20 May lawan faunal region), Philetor brachypte- 2000, we captured and examined two adult rus, and Pipistrellus tenuis. females; both were pregnant. Of 8 adult fe- Glischropus tylopus.—This poorly males captured in July at Site 8, none were known species is found from Myanmar to pregnant or lactating. Cranial measurements the Molucca Islands and Palawan (Heaney of 5 individuals (Table 1) fall within or near et al. 1998). In Peninsular Malaysia it roosts the range for the species in Ingle and Hea- in rock crevices, bamboo, and in new ba- ney (1992). Specimens examined: 5, Site 5 nana leaves (Payne et al. 1985). We never (4), Site 8 (1). encountered this species, but it is repre- Rhinolophus virgo.—This Philippine en- sented by a specimen in the USNM (Hol- demic is widely distributed within the Phil- lister 1913). ippines (Heaney et al. 1998). It is known Kerivoula hardwickii.—This species is from secondary forest, primary lowland widespread from India and southern China forest, reaching mossy forest on small, low- to the Lesser Sunda Islands and the Phil- lying islands, and often roosts in caves ippines (Heaney et al. 1998). It was previ- (Heaney et al. 1991, Ingle 1992, Rickart et ously known from lowland, montane, and al. 1993); Sanborn (1952) reported a series ridge-top mossy forest from 500 to 1600 m from Tanabog, Palawan. This species ap- in the Philippines (Heaney et al. 1999, peared to be rare in the cave at Site 4 (0.5% Rickart et al. 1993). Everett (1889) includ- of captures), common at Site 8 (151 cap- ed mention of this species from Palawan. A tures, about 6% of total) and abundant in previous record from UMMZ (Heaney et al. some caves at Site 5 (15% of captures). We 1998) has been re-identified as K. white- also captured this species in primary forest headi, as noted below. Payne et al. (1985) at Sites 2 and 15. Of 29 adult females cap- reported the species to ‘‘frequent the un- tured between 27 March and 4 April 2000, derstory of tall forest’’ on Borneo, and Md. none were pregnant or lactating. Of 35 fe- Nor (1995) caught one in primary lowland males captured at Site 5 between 15 and 20 dipterocarp forest on Banggi Island ‘‘in the May 2000, l6 were pregnant and 5 were axil of a leaf on a rattan vine 1 m above lactating. Of 78 females taken at Site 8 in ground’’, and netted them in the understory July 2000, 6 were pregnant and 2 were lac- of primary forest on Balambangan Island. tating. Cranial measurements (Table 1) fall We captured two adult females, one of within the range (Ingle & Heaney 1992) or which was lactating, in a bamboo thicket at very near the range (Rickart et al. 1993) of Site 11 (ca. 60 m asl), and two individuals previously available individuals. IUCN in the understory (ca. 2–4 m above the (2002) lists this species as Near-Threatened. ground) of primary lowland forest at ca. Specimens examined: 6, Site 4 (2), Site 5 650 m elevation at Site 2, all in harp-traps. (4). Specimens examined: 3, Site 2 (1), Site 11 (2). Family Vespertilionidae—Vesper and Kerivoula pellucida.—This poorly Evening Bats known species is known from the Malay Peninsula, the Sunda Shelf, Jolo, and Pa- This diverse family of bats is generally lawan (Heaney et al. 1998). Known only poorly documented, and more species from lowland forest (Payne et al. 1985). should be sought on Palawan, including Taylor (1934) reported two specimens from such widespread taxa as Harpiocephalus Palawan (no locality given) that he obtained harpia, Murina cyclotis, Myotis ater (which from a group of seven that he found flying Hill 1983 and Corbet & Hill 1992 have together in daylight: his map (Fig. 13), VOLUME 117, NUMBER 3 287 shows the locality in the vicinity of Broo- (Heaney et al. 1991, Rickart et al. 1993, ke’s Point. On 21 June 2000, using a harp- Sanborn 1952). We captured this species in trap we captured an adult female carrying varying numbers at several caves. In pri- a suckling infant in secondary lowland for- mary and disturbed lowland forest at Sites est (ca. 80 m) over a small stream at Site 4 and 5 it was scarce, represented by less 7. Cranial measurements of the adult female than 1% and less than 3% of captures, re- (Table 2) are smaller than the one specimen spectively. At Site 8 it was abundant (45% from Davao del Norte, Mindanao available of 2775 captures). Cranial measurements of to Ingle and Heaney (1992), but they are 4 individuals (Table 2) are similar to those otherwise very similar; additional speci- reported by Ingle and Heaney (1992) and mens are badly needed to examine patterns Rickart et al. (1993) from the Philippines, of variation. Specimens examined: 2, Site 7 and by Corbet & Hill (1992) from through- (2). out the species range. Specimens examined: Kerivoula whiteheadi.—This poorly 4, Site 4 (2), Site 5 (2). known species is widely distributed from Miniopterus schreibersi.—This common southern Thailand to Borneo and the Phil- species is found from Europe to the Solo- ippines (on Luzon, Mindanao, and Pala- mon Islands and is widespread in the Phil- wan; Heaney et al. 1998). In the Philip- ippines, but this is the first record from Pa- pines, it is known only from near sea level lawan (Heaney et al. 1998). It is common in disturbed forest and agricultural areas in caves throughout the lowlands in agri- (Sanborn 1952). A single specimen in the cultural areas and forest and known from UMMZ captured by P. O. Glass on 29 Sept. both lowland and montane forest (Heaney 1978 at Irawan, Puerto Princesa Municipal- et al. 1991, 1999; Rickart et al. 1993; San- ity (noted as 2 km N Irawan, at the base of born 1952). We captured this species in and Mt. Beaufort by P. O. Glass, in litt.) was around caves in disturbed and primary low- erroneously reported by Heaney et al. land forest at Sites 4, 5, and 8. At Sites 4 (1998) under both K. hardwickii and this and 8 the species was abundant, represented species. Additionally, two specimens taken by 47% and 26% of captures respectively. ‘‘under banana fronds’’ by C. A. Ross on 8 At Site 5 it was common at 10% of 575 April 1987 at Barangay Binwang, Quezon captures. Of 214 adult females captured in Municipality, are housed in the USNM. We December 1999 at Site 4, only one was captured a single individual of this species pregnant. Of 36 captured between 15 and in a harp-trap near ground level in a cogon 20 May 2000 at Site 5, 15 were pregnant grassland (Imperata cylindrica)atSite 6. and none were lactating. Of 421 captured Cranial measurements of these four individ- at Site 8 in July 2000, none were pregnant uals (Table 2) are similar to those in Ingle but 4 were lactating. Cranial measurements & Heaney (1992) of a specimen from Min- (Table 2) are similar to those reported by danao, though some variation in size is pre- Ingle and Heaney (1992) and Rickart et al. sent; more specimens are needed to assess (1993) from the Philippines, and by Corbet geographic variation. Specimens examined: and Hill (1992) from throughout the species 4, Site 6 (1), Irawan, Puerto Princesa Mu- range. IUCN (2002) lists this species as nicipality, 60 m (1), and Binwang, Quezon Near-Threatened, but its abundance in (2). heavily disturbed habitat in the Philippines Miniopterus australis.—This common (Heaney et al. 1998, Rickart et al. 1993) species is found from India to Australia; it makes this inappropriate. Specimens ex- is widespread in the Philippines, but this is amined: 6, Site 4 (4), Site 5 (1), Site 8 (1). the first record from Palawan (Heaney et al. Miniopterus tristis.—This widespread 1998). It is known to roost in caves in low- species is found from the Philippines to the land areas of agriculture or second growth Solomon Islands; this is the first record 288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 23.4 23.6 54.2 53.3 47.4 42.3 33.1 48.1 58 34.0 35.5 36.7 37.1 29.7 33.1 33.6 length Forearm (22.8–24.6) (53.3–53.3) (39.2–43.9) (36.2–37.8) (29.4–30.4) (33.4–33.7) — — 8.13 8.31 3.91 3.86 6.64 8.58 9.69 6.35 6.43 5.79 5.66 5.82 6.82 6.98 length Palatal (8.31) (3.67–4.05) (6.37–6.48) (5.56–5.79) (5.70–6.08) (6.91–7.04) 4.62 4.37 2.86 2.75 3.17 4.15 3.84 3.65 3.39 3.33 3.23 3.06 3.00 2.41 2.72 2.90 Palatal breadth (4.33–4.40) (2.71–2.81) (3.26–3.54) (2.91–3.06) (2.34–2.49) (2.84–2.96) 5.66 5.73 3.90 5.31 5.88 2.69 2.68 3.82 3.78 4.58 4.45 3.80 3.76 3.18 3.53 3.74 toothrow Molariform (5.66–5.81) (2.59–2.74) (4.32–4.56) (3.73–3.79) (3.12–3.30) (3.65–3.82) molar 5.23 7.30 8.40 7.75 7.83 3.45 3.41 4.88 4.70 6.28 6.14 5.21 5.19 5.17 5.39 5.62 to last Canine (6.0–6.25) (7.75–7.91) (3.37–3.45) (5.16–5.21) (5.07–5.22) (5.50–5.74) — 5.08 6.88 6.91 6.51 6.73 5.15 4.89 3.53 3.74 5.31 5.05 4.46 4.29 4.05 4.07 length Orbital (6.70–6.76) (3.57–3.80) (4.90–5.32) (4.16–4.40) (4.07–4.07) width 7.73 9.51 9.68 7.90 7.60 9.82 9.76 6.71 6.75 8.73 8.19 7.16 7.28 6.66 7.14 7.29 Mastoid (9.62–9.93) (6.59–6.96) (8.12–8.33) (7.23–7.33) (6.56–6.77) (7.05–7.53) width 9.11 9.31 9.01 7.98 7.75 8.88 8.42 7.21 7.39 7.56 8.48 8.70 11.8 12.25 10.72 10.76 Zygomatic (7.56–7.96) (8.30–8.57) (7.29–7.58) (7.44–7.64) (8.64–8.76) (10.74–10.78) length 14.63 17.88 19.18 18.50 18.72 10.29 10.24 13.30 13.03 15.33 14.78 13.25 13.06 12.00 13.51 8.26 7.21 3.08 5.57 3.87 2.58 6.94 31.3 13.39 13.75 incisive Condylo- (18.71–18.74) (10.10–10.34) (14.67–14.94) (12.90–13.15) (11.85–12.28) (13.63–13.87) 1 1 1 1 2 1 2 1 1 1 5 1 3 1 2 f f f f f f f4 f1 f m m m m m m m m tubinaris cf. Species Sex n Table 2.—Means and ranges of cranial measurements of adult Vespertilionidae from Palawan Island, Philippines. pus bersi Murina Myotis macrotarsus Myotis rufopictus Miniopterus tristis Pipistrellus javanicus Tylonycteris pachy- Miniopterus schrei- Miniopterus australis Kerivoula whiteheadi Kerivoula pellucida Kerivoula hardwickii VOLUME 117, NUMBER 3 289 from Palawan (Heaney et al. 1998). The fer the specimen from Palawan to this same species is known to roost in caves and for- species. Specimen examined: 1, Site 12 (1). age in disturbed forest (Rickart et al. 1993, Myotis horsfieldii.—This common spe- Sanborn 1952). We captured one specimen cies is distributed from southeastern China in a cave surrounded by old-growth forest to the Malay Peninsula, Sulawesi, and the at Site 4 and two in a cave surrounded by Philippines (Heaney et al. 1998). On Bor- disturbed areas and secondary forest at Site neo, the species ‘‘roosts in crevices or bell- 10. Miniopterus tristis appeared to be con- holes in caves, usually not far from large sistently less common than the other spe- streams or rivers’’ (Payne et al. 1985). In cies of Miniopterus. Cranial measurements the Philippines, it has been recorded in low- of 3 individuals (Table 2) are similar to land forest and agricultural areas, up to 800 those reported by Ingle and Heaney (1992) m (Heaney et al. 1998). We never encoun- and Rickart et al. (1993) from the Philip- tered this species; two specimens in the pines, and by Corbet and Hill (1992) from UMMZ taken by A. C. Alcala in Sitio Mal- throughout the species range. Specimens abusog, Tinitian, Roxas Municipality in examined: 3, Site 4 (1), Site 10 (2). 1984 were reported by Heaney et al. (1998). Murina cf. tubinaris.—A single speci- Myotis macrotarsus.—This species is men of a small tube-nosed bat (genus Mu- known from Borneo and the Philippines rina) was taken in a lowland grassland-for- (Heaney et al. 1998); it roosts in caves near est mosaic at Site 12 on 24 March 1997, sea level and forages in agricultural areas and is housed in the Staatliches Museum fur (Heaney and Utzurrum, unpubl. data). Md. Naturkunde in Stuttgart, Germany Nor (1995) caught the species over a dry (#49238). The specimen (Table 2) is very river bed and in the understory of primary similar to a series from Tonkin, Vietnam lowland forest on Balambangan Island. In (FMNH 32203–32204, 46626–46627), a cave in disturbed lowland forest at Site 8, though slightly larger. In our specimen, as this species was represented by Ͻ0.5% of in the Vietnamese series, the upper tooth- 2775 captures. We also observed small rows converge slightly, the anterior pre- numbers in the cave at PPSRNP. Cranial molars are reduced, and the canines are measurements (Table 2) of one individual short but longer than the premolars, as not- show it to be slightly larger than those re- ed by Koopman and Danforth (1989) and ported by Ingle & Heaney (1992). IUCN Corbet and Hill (1992). The length of fore- (2002) lists this species as Near-Threatened. arm (33 mm) falls at the center of the range Specimens examined: 1, Site 8 (1). given by Koopman and Danforth for M. Myotis rufopictus.—This poorly known tubinaris (28–35 mm), and at the high end Philippine endemic has been recorded from given for M. suilla (26–33 mm). Koopman primary lowland and montane forest (Hea- and Danforth (1989) considered M. florium, ney et al. 1999, Mudar & Allen 1986). We M. suilla, and M. tubinaris to be members never encountered this species; on Palawan, of a species group, and perhaps to be con- it is known from a single specimen in specific, noting that few specimens are UMMZ reported by Allen (1922). We fol- available. Corbet and Hill (1992) re-empha- low Ingle & Heaney (1992) in regarding sized the uncertainty in current taxonomy, this as one of several distinct species within but took a somewhat different view, refer- the subgenus Chrysopteron, rather than rec- ring specimens from Borneo to M. suilla, ognizing only a single species, Myotis for- rather than to M. tubinaris. While we agree mosus, within the subgenus (e.g., Corbet & entirely on the need for more specimens Hill 1992). This species is not listed by and further study, we follow Koopman & IUCN (2002); we recommend listing as Danforth (1989) on referring Bornean spec- Data Deficient. Measurements in Table 2 of imens to M. tubinaris, and provisionally re- the specimen reported by Allen (1922) were 290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON taken by Heaney. Specimen examined: 1, at Site 11. Very near the capture site we Puerto Princesa (1). observed what appeared to be more than a Pipistrellus javanicus.—This species is dozen individuals of this species foraging distributed from Korea to Java and the Phil- over a few remnant trees in a cleared area ippines (Heaney et al. 1998). Taxonomic with houses that is immediately surrounded status is uncertain; P. imbricatus has been by logged-over and secondary forest. Spec- reported from Palawan (e.g., Allen 1922, imens examined: 5, Site 11 (5). Corbet & Hill 1992, Sanborn 1952), but In- Tylonycteris robustula.—This species is gle and Heaney (1992) were unable to dis- also widespread from southern China to the tinguish more than one species of Pipis- Lesser Sunda Islands and the Philippines trellus of this size in the Philippines; de- (including records from Calauit, Luzon, and tailed study is needed. It is common in pri- Palawan); its habitat is apparently similar to mary montane forest and uncommon in that of T. pachypus (Heaney & Alcala 1986, lowland and mossy forest (Heaney et al. Heaney et al. 1998). We never encountered 1999, Ingle 1992, Sanborn 1952). We cap- this species. tured two individuals from a roost in a hol- low tree in montane forest (ca. 1300 m) at Family Molossidae—Free-Tailed Bats Site 3. The opening in the tree appeared to This family is generally poorly known in have formed where a branch had been bro- Southeast Asia, partly because they typi- ken off the tree and was quite small (ca. 1.5 cally fly high above the canopy and are ϫ 5 cm). Cranial measurements (Table 2) therefore rarely netted. At least one species fall within the range of specimens reported (Chaerophon plicata)iswidespread in the by Ingle and Heaney (1992), and are slight- region and should be sought on Palawan. ly smaller than a series from southern Lu- Cheiromeles torquatus.—This poorly zon (Heaney et al. 1999). Specimens ex- known species is found from Sumatra to amined: 2, Site 3 (2). Java, Borneo, and Palawan, but not the rest Scotophilus kuhlii.—This common spe- of the Philippines (Heaney et al. 1998). It cies is widespread from Pakistan to Taiwan roosts in large caves and hollow trees and and the Philippines (Heaney et al. 1998); it forages in open areas, over streams, and is abundant in urban and agricultural areas, above forest canopy on Borneo (Payne et and roosts in buildings and ‘‘tents’’ made al. 1985). We never encountered this spe- from modified palm leaves (Heaney et al. cies, which was documented on Palawan by 1998; Rickart et al. 1989, 1993). Hollister Sanborn (1952) based on a single specimen. (1913) and Taylor (1934) reported it from IUCN (2002) lists this species as Near- Puerto Princesa, Sanborn (1952) reported it Threatened. from Brooke’s Point, and we found it to be Mops sarasinorum.—This very poorly abundant in buildings at the Provincial Ag- known species occurs in Sulawesi and the riculture Center, Irawan, Puerto Princesa, Philippines; the Palawan record is based on and in staff houses of the State Polytechnic a single specimen in the Senckenberg Mu- College in Puerto Princesa, and in mixed seum, Frankfurt (Heaney et al. 1998). It prob- urban/agricultural areas (Site 12). ably occurs in lowland forest (Heaney et al. Tylonycteris pachypus.—This tiny bat is 1998). We never encountered this species. It widespread from India to the Philippines is listed by IUCN (2002) as Near-Threatened, (Heaney et al. 1998). In the Phillipines, it but we recommend Data Deficient. is known from bamboo stands in agricul- tural areas (Heaney & Alcala 1986); Hol- Order Primates lister (1913) reported a specimen from Family Cercopithecidae—Monkeys Puerto Princesa. We captured several indi- Macaca fascicularis.—This common viduals of this species in a bamboo thicket monkey occurs from Myanmar to Timor VOLUME 117, NUMBER 3 291 and the Philippines (Fooden 1995, Heaney (2002) as Near-Threatened; M. culionensis et al. 1998). It is known from agricultural probably deserves the same status. areas near forest, second growth, secondary forest, and primary lowland and montane Order Rodentia forest (Heaney et al. 1998, 1999; Rickart et Family Sciuridae—Squirrels al. 1993); Sanborn (1952) reported speci- mens from Iwahig, Puerto Princesa, and Hylopetes nigripes.—This large gliding Brooke’s Point. Reis & Garong (2001) re- squirrel is endemic to the Palawan faunal ported a specimen from sediments in a region; the number of museum specimens rock-shelter near Tabon Cave, Quezon Mu- (Allen 1910, Sanborn 1952) suggests that it nicipality dated to 11,130 BP. We common- is common. Reis & Garong (2001) reported ly observed this species at all of our sites two specimens from sediments in a rock- (except Site 13), in secondary and primary shelter near Tabon Cave, Quezon Munici- forest (including mangrove, swamp forest, pality dated to 11,130 BP. Taylor (1934) beach forest, and lowland forest) from sea found the species in primary and secondary level to 1000 m; at forest edge near agri- lowland forest where they nest in cavities cultural areas and houses they seem to be in large trees. We observed an individual less common and more shy. On Palawan, running up the side of a large hollow tree the species is under moderate hunting pres- in primary forest at Site 1, and we frequent- sure for meat and the local pet trade, but ly heard and twice spotlighted them in ma- appeared to have stable populations. In ture lowland forest at Site 15. We also most areas, it was quite wary of humans, heard the distinctive calls several times in but in areas such as the PPSRNP (Site 15), selectively logged but largely intact forest the species did not associate humans with near Barake, Aborlan Municipality, in the danger, and had become a regular thief of Victoria Range. According to local resi- picnic baskets. It is listed by IUCN (2002) dents, the species is common in mature for- as Near-Threatened. est and is occasionally hunted as a source of food. IUCN (2002) lists this species as Order Pholidota Near-Threatened; by current criteria, it Family Manidae—Pangolins should be listed as Data Deficient. Sundasciucus juvencus.—This tree squir- Manis culionensis.—This endemic spe- rel is endemic to central and northern Pa- cies of the Palawan faunal region, with re- lawan Island (Heaney et al. 1998). Hoogs- cords from Palawan and Culion Islands traal (1951) and Sanborn (1952) reported (Heaney et al. 1998), was formerly included this species from primary and secondary within Manis javanica (Feiler 1998). It is lowland forest. We commonly observed this known from primary and secondary low- species in primary and secondary lowland land forest, possibly localized in distribu- forest at Sites 1, 2, 7, 11, 12, and in both tion (Allen 1910, Hoogstraal 1951, Sanborn secondary forest and grassland/degraded 1952, Taylor 1934). We sighted several in forest mosaic at Site 15. We also found it lowland grassland/forest mosaic at Site 12. in a very small (Ͻ1 ha.) patch of secondary It is hunted for its skin, which is used to lowland forest surrounded by grassland and treat asthma. We have seen it for sale in agricultural areas at Site 6. We observed the Puerto Princesa and our guide at Site 11 species on Dumaran Island, but not on Mal- said that it is hunted in logged-over lowland inau or Rasa. The species is reportedly a forest in that area. The species was de- common pest in coconut plantations. It is scribed by local informants as fairly com- occasionally hunted as a source of food and mon, but hunting pressure is moderately for the local pet trade. It is listed by IUCN heavy. Manis javanica is listed by IUCN (2002) as Endangered, but this is strongly 292 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON contradicted by the available data, and we genus is apparently difficult to capture recommend de-listing. (Musser 1979); we never encountered this Sundasciurus rabori.—This poorly- species. We recommend IUCN listing as known species, described from 5 specimens Data Deficient. taken at 3600–4350 ft (ca. 1100–1300 m) Haeromys pusillus.—This species is on Mt. Mantalingajan, is endemic to Pala- known only from Borneo, Palawan, and Ca- wan Island (Heaney 1979). P. C. Gonzales lauit Islands (Musser & Carleton 1993, deposited 2 specimens at the UMMZ that Musser & Newcomb 1983). It is cited in he collected on Mt. Gorangbato in Brooke’s Heaney et al. (1998) as ‘‘Haeromys sp. A’’ Point Municipality in 1984; these are the as potentially endemic to Palawan, but we only reported specimens aside from the follow Musser (pers. comm.) in treating it original type series from Mt. Mantalingajan as conspecific with H. pusillus. We never (Heaney 1979). Although we worked in encountered this species, but Musser & some seemingly suitable habitats on Cleo- Carleton (1993) cited a specimen from Pa- patra’s Needle (Site 3), we did not specifi- lawan. A specimen of H. pusillus was taken cally seek this species, and we never en- in Sabah, Borneo, in a pit-fall trap near the countered it. The IUCN (2002) lists S. ra- edge of tall dipterocarp forest (Payne et al. bori as Vulnerable, but based on current 1985), and A. C. Alcala stated that he cap- IUCN criteria, it should be considered Data tured the specimen from Calauit (in Deficient. FMNH) by hand in a bamboo thicket (pers. Sundasciunis steerii.—This species is en- comm.). IUCN (2002) listed this species as demic to Balabac and southern Palawan Is- Vulnerable, but based on current criteria, it land (Heaney et al. 1998); Sanborn (1952) should be considered Data Deficient. reported a large series from Brooke’s Point. Maxomys panglima.—This common rat Heaney et al. (1998) listed it as common in is endemic to the Palawan faunal region; lowland forest and coconut and banana the genus is common on the Sunda Shelf, plantations. Because all of our study sites but is absent from oceanic portions of the were in central and northern Palawan, we Philippines (Musser et al. 1979). Sanborn never encountered this species. It is listed (1952) reported large series from several lo- by IUCN (2002) as Near-Threatened; since calities. We found it to be the most com- its habitat use is similar to the closely-re- monly captured small mammal in agricul- lated S. juvencus, it is probably not threat- tural/forest mosaic at Site 12 (62% of 169 ened. captures), and was common to abundant in secondary forest (Site 11), primary lowland Family Muridae—Mice (Sites 1 and 2), and montane forest (Site 3) from near sea level to at least 1550 m. We Chiropodomys calamianensis.—This captured a single juvenile in mossy forest poorly known arboreal mouse is endemic to at 1580 m at Site 3. Because we found this the Palawan faunal region; it is closely re- species to be common, although sometimes lated to species on the Sunda Shelf (Musser patchy, in all lowland and montane forested 1979). Reis & Garong (2001) reported a sites where we trapped extensively, and in specimen from sediments in a rock-shelter mixed agricultural/second growth areas at near Tabon Cave, Quezon Municipality dat- Sites 11 and 12, we consider the IUCN ed to 11,130 BP. It is known from forest (2002) listing as Near-Threatened to be un- near sea level (Taylor 1934), coconut plan- justified. Specimens examined: 5, Site 1 (3), tations, bamboo thickets, and buildings Site 3 (2). (Sanborn 1952); there are 12 specimens Mus musculus.—This introduced com- from Palawan in FMNH and NMP from the mensal has a nearly world-wide distribu- Hoogstraal expedition (Sanborn 1952). The tion, although Southeast Asian populations VOLUME 117, NUMBER 3 293 are sometimes treated as a separate species, Hoogstraal (1951) and Sanborn (1952) M. castaneus (Musser & Carleton 1993). It found them to be common on Palawan in is common in human habitations in urban some agricultural and residential areas. We and rural areas (Heaney et al. 1998). We captured three at Site 13, and three in a res- captured several in a residential area at Site idential area in Puerto Princesa; vouchers 12, and it is most likely common in such were deposited in the NMP and the collec- places throughout Palawan. tion of the Palawan Council for Sustainable Palawanomys furvus.—This poorly Development. known monotypic genus is endemic to Pa- Rattus tiomanicus.—This indigenous rat lawan Island. It has been taken from a sin- is found on the Malay Peninsula and the gle locality on Mt. Mantalingajan and prob- islands of the Sunda Shelf, including Pala- ably occurs in high mountain forest (Mus- wan (Heaney et al. 1998). Payne et al. ser & Newcomb 1983). Our survey efforts (1985) reported the species from secondary on Cleopatra’s Needle (Site 3) failed to find forest, agricultural areas and gardens, scrub, this species; perhaps it is restricted to the and grassland. We captured this species in more extensive mountain ranges of south- grassland/forest mosaic at Site 12 (9% of ern Palawan. The IUCN (2002) lists this captures), two in selectively logged forest species as Endangered; the lack of data and at Site 13, one in a ricefield at Site 14, and lack of damage to its presumed habitat three individuals from mossy forest and the (montane and mossy forest) suggest that it transition zone between mossy and montane should be listed as Data Deficient. forest at Site 3. At Site 3, two individuals Rattus exulans.—This introduced com- were taken at ca. 1580 m during the night mensal species is widespread from Bang- and one at ca. 1540 m during the day. Spec- ladesh to Easter Island (Heaney et al. 1998). imens examined: 3, Site 3 (3). The first records from Palawan were named Sundamys muelleri.—This moderately as a distinct species (luteiventris)byAllen large rat is found from southern Myanmar (1910), but it is currently treated as a junior to the Sunda Shelf, including Palawan synonym of R. exulans (Musser & Carleton (Heaney et al. 1998); the genus is absent 1993). It is common in agricultural areas from the oceanic Philippines. Sanborn (Barbehenn et al. 1973, Rabor 1986) and (1952) described a subspecies endemic to sometimes present in disturbed forest and Palawan, S. m. balabagensis, from a single rare in primary forest (Barbehenn et al. specimen taken at 3000 ft (ca. 900 m) ‘‘in 1973; Heaney et al. 1991, 1998). We found thick forest near the top of Mt. Balabag’’; this species in grassland (Site 6), agricul- two additional specimens in the USNM are tural areas (Sites 6, 11, 12, and 14), and in from Pinigisan, on the lower slopes of Mt. secondary lowland forest (Site 7). The spe- Mantalingajan at 2100–2500 ft (ca. 640– cies appears to be absent from primary 760 m). Additional specimens from the Pa- (e.g., Sites 1, 2, and 3) and logged-over for- lawan region are from Culion Island (San- est (e.g., Site 11) on Palawan. Specimens born 1952), Balabac, and Busuanga examined: 4, Site 6 (3), Site 11 (1). (USNM; Heaney et al. 1998). On Borneo, Rattus tanezumi.—This introduced com- the species occurs in forest, often near mensal, formerly included within Rattus streams (Payne et al. 1985) at elevations rattus (Musser & Carleton 1993), is wide- usually below 3500 ft (ca. 1070 m; Med- spread from Afghanistan to New Guinea way 1977). Md. Nor (1995) caught the spe- and Micronesia (Heaney et al. 1998). It is cies on Banggi, Balambangan, and Mol- often abundant in urban and agricultural ar- leangan Islands ‘‘mostly in primary forest eas and common in disturbed forest up to on low ground and near streams’’. We cap- 1800 m (Danielsen et al. 1994; Heaney et tured one individual from a riparian zone in al. 1989, 1999; Rabor 1986; Sanborn 1952). primary forest at ca. 700 m at Site 2, and 294 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON two in lowland grassland/forest mosaic at along a river trail in Barake, Aborlan Mu- Site 12. Specimen examined: 1, Site 2 (1). nicipality.

Family Hystricidae—Porcupines Family Mustelidae—Weasels, Otters, and Badgers Hystrix pumila.—The only porcupine found in the Philippines is endemic to the Amblonyx cinereus.—This otter occurs Palawan faunal region; other species occur from India to Taiwan and the Sunda Shelf widely on the Sunda Shelf and continental (Heaney et al. 1998). On Palawan, it is Asia. Reis & Garong (2001) reported 3 found in coastal rivers and bays (Hoogstraal specimens from sediments in a rock-shelter 1951, Rabor 1986, Sanborn 1952). Payne et near Tabon Cave, Quezon Municipality dat- al. (1985) and Sanborn (1952) reported the ed to 11,130 BP. It is known to occur in species feeds on crustaceans, mollusks, and secondary and primary lowland forest and fish where there is permanent water and to den in abandoned mine shafts (Hoogs- some tree cover. Rangers at PPSRNP re- traal 1951, Sanborn 1952). We observed ported to Heaney and Widmann that otters this species at dusk along the edge of sec- frequently visit along the beach and small ondary forest at Site 11, once at night at streams, and local people reported them Site 15 (where it was feeding on fruit of from the Iwahig River (Puerto Princesa), Terminalia catappa), and several times at Aborlan River (Aborlan Municipality), night in grassland/forest mosaic at Site 12. Malatgao and Taritien Rivers (Narra Mu- Local guides reported them at Site 14, at nicipality), and adjacent mangrove and the Iwahig Penal Colony in Puerto Princesa, freshwater swamp forest. We received one in Rizal Municipality, and in Dumaran Mu- report of an otter raiding a prawn pond. nicipality (on the mainland). This was re- IUCN (2002) lists this species as Near- ported as the most important game species Threatened. for the Tagbanua ethnic community in Bar- ake, Aborlan Municipality (Lacema & Wid- Mydaus marchei.—This badger is en- mann 1999); they are often dug out of their demic to the Palawan faunal region; it is subterranean dens. Not listed by IUCN related to a species that occurs on the Sunda (2002) but listing as Data Deficient or Near- Shelf. It has been documented in mixed Threatened seems justified. grassland and secondary forest (Hoogstraal 1951, Kruuk 2000, Rabor 1986, Taylor 1934), and Sanborn (1952) reported series Order Carnivora from several localities. We occasionally Family Felidae—Cats smelled its strong odor in areas of mixed Prionailurus bengalensis.—This small agriculture and secondary forest throughout cat is widespread from Siberia to Pakistan Palawan; we sighted it often in residential and Bali, with reports from the Philippines and cultivated areas, grassland, and grass- on Busuanga, Cebu, Negros, Palawan, and land/forest mosaic at Site 12, and rarely in Panay Islands only (Heaney et al. 1998, ricefields and freshwater swamp forest at Taylor 1934). The population from the Pa- Site 14. One individual living in a den on lawan faunal region was described recently campus at the State Polytechnic College as a distinct subspecies, P. b. heaneyi, by was easily followed and observed. Because Groves (1997); it is well represented by it is widespread and moderately common museum specimens (Allen 1910, Sanborn on Palawan, and is rarely hunted (Grim- 1952). Rabor (1986) reported the species wood 1976, Kruuk 2000), we agree with from agricultural areas and forest from sea Kruuk (2000) that the IUCN listing of this level to ca. 1500 m. We spotlighted one species as Vulnerable is not justified. VOLUME 117, NUMBER 3 295

Family Herpestidae—Mongooses Paradoxurus hermaphroditus.—This common species is found from Sri Lanka Herpestes brachyurus.—The only mon- to the Lesser Sunda Islands and the Phil- goose found in the Philippines is distributed ippines (Heaney et al. 1998). Recorded in from the Malaysian Peninsula to Borneo agricultural areas and forest over a wide and Palawan (Heaney et al. 1998). The Pa- elevational range (Allen 1910; Heaney et al. lawan population was named as a distinct 1991, 1999; Hoogstraal 1951; Rabor 1986); species (H. palawanus Allen 1910), but Sanborn (1952) reported large series from currently is treated as a subspecies (Corbet several localities. We often saw them feed- & Hill 1992). Allen (1910) described them ing in fruiting trees and shrubs in grassland/ based on one specimen from Iwahig; San- forest mosaic at Site 12, and we saw road- born (1952) reported one specimen from kills along the coastal highway. They are Puerto Princesa and one from Brooke’s hunted, but the large number of museum Point, and Rabor (1986) found the species specimens and sightings indicate that they most often near rivers. On Borneo, Payne traditionally have been and probably remain et al. (1985) found the species to occur in the most common carnivore on Palawan primary and secondary lowland forest, (e.g., Allen 1910, Sanborn 1952). plantations, and gardens. We never encoun- Viverra tangalunga.—This civet is found tered this species, but we received reports from the Malay Peninsula to Sulawesi and of them at Site 14. the Philippines (Heaney et al. 1998). Known from primary and secondary low- Family Viverridae—Civets land, montane, and mossy forest (Allen Arctictis binturong.—The binturong is 1910, Heaney et al. 1999, Rickart et al. known from northern Myanmar to the Sun- 1993). We captured and released a juvenile da Shelf (Heaney et al. 1998). On Borneo, of this species in a cage trap in lowland the species is arboreal and terrestrial, most- primary forest at Site 1, and we observed ly nocturnal, and occurs in old-growth and two in forest-grassland mosaic at Site 12. secondary forests, sometimes entering ag- ricultural areas near forest (Payne et al. Order Artiodactyla 1985). The Palawan population, which ini- Tragulus napu and Axis calamianensis tially was named as a distinct species (A. both occur in the Palawan faunal region, whitei Allen 1910) from four specimens, is but we found no evidence of either species still represented by few specimens (Heaney on Palawan Island. et al. 1998). Rabor (1986) reported obser- vations from primary and secondary low- Family Suidae—Pigs land forest up to 200 m. Our guide at Site 11 reported that a juvenile repeatedly en- Sus barbatus.—The bearded pig is found tered remnant trees that were fruiting in a from the Malay Peninsula to Borneo and clearing surrounded by secondary forest. At Palawan (Heaney et al. 1998). Rabor (1986) Site 2, we observed A. binturong drinking and Payne et al. (1985) reported the species water from a stream at ca. 400 m during from primary and secondary forest from sea mid-day. We spotlighted one in a fruiting level to the highest peaks; Sanborn (1952) Ficus tree at Site 15, and twice saw one in reported a series from Iwahig. Groves grassland/forest mosaic at Site 12 feeding (2001) has tentatively suggested that the on fruits of Guioa pleuropteris. Local peo- population of this species from the Palawan ple reported hunting them for food, and also faunal region, which has been recognized catching them and selling them as pets. The as a distinct subspecies, may warrant rec- IUCN (2002) listing of A. binturong whitei ognition as a distinct species, Sus ahoeno- as Vulnerable seems justified. barbus. We regularly observed this species 296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON or evidence of its occurrence in forest hab- and Pteropus) has been very poorly studied itats (including fragmented forest) from sea on Palawan and elsewhere in the Philip- level to montane forest at ca. 1500 m (Sites pines. This is the direct result of the diffi- 1, 2, 3, 4, 7, and 11). We also observed culty in capturing these species by any evidence of the species entering cultivated means other than shooting. Despite the pau- areas near forest and damaging crops. Wild city of ecological information on these spe- pigs are heavily hunted on Palawan with cies, their distribution among major island snares, low caliber rifles, and small, baited groups appears to be moderately well un- explosive devices known as ‘‘pig bombs’’. derstood because their activities and roosts The species appears to be locally common, are highly conspicuous, and no additional but is in decline due to heavy hunting pres- species have been found on Palawan in over sure (Caldecott et al. 1993, Oliver 1992). 50 years. It seems unlikely that additional The IUCN (2002) lists S. barbatus ahoe- species will be found on Palawan, with the nobarbus as Vulnerable. possible exception of P. hypomelanus. Insectivorous bats are clearly the least Discussion known of all Palawan mammals. Distribu- tions remain poorly documented, ecological Adequacy of sampling.—Small fruit bats information is scanty for most species, and on Palawan (Cynopterus, Eonycteris, Ma- the taxonomy is often uncertain. Our survey croglossus, and Rousettus) appear to have efforts and examination of previously col- been fairly completely sampled; no species lected insectivorous bats documented eight have been added in over 50 years (exclud- species (Rhinolophus arcuatus, R. cf. bor- ing the apparently erroneous reports of neensis, R. creaghi, R. macrotis, Miniopte- Haplonycteris fischeri and Ptenochirus mi- rus australis, M. schreibersi, M. tristis, and nor), despite extensive netting. It is inter- Murina cf. tubinaris)onPalawan for the esting that our mist netting in primary for- first time, and several more are noted in the est produced very few captures; for exam- text as very likely to be present. ple, in primary lowland forest at 150 m el- Our knowledge of small non-volant evation (Site 1), we captured 1 fruit bat in mammals (including Soricidae, Tupaiidae, 42 net-nights; in primary lowland forest at Sciuridae, and Muridae) on Palawan is un- ca. 500 m (Site 2), we captured 1 fruit bat even. Some lowland species (e.g., Tupaia in 56 net-nights, and in primary montane palawanensis, Sundasciurus juvencus, forest at ca. 1400 m (Site 3), we captured Maxomys panglima, Rattus tiomanicus, no fruit bats in 48 net-nights. Although our Sundamys muelleri, and the non-native mu- sample size is limited, all of these values rines) are common and well known. Very fall well below what would be typical on little is known of several other species (e.g., islands in the oceanic portion of the Phil- Crocidura palawanensis, Crocidura sp., ippines (Heaney et al. 1989, 1999), sug- Sundasciurus rabori, Chiropodomys cala- gesting that small fruit bats are not as abun- mianensis, Haeromys pusillus, and Pala- dant in primary forest on Palawan (e.g., wanomys furvus). Perhaps we failed to lo- Heideman & Heaney 1989, Heaney et al. cate these poorly known species because 1989). Indeed, all of the small fruit bats we did little trapping in trees or other places currently known from Palawan predomi- above the ground surface (C. calamianensis nately occur in disturbed habitats, in con- and H. pusillus), sampled only one site trast to the oceanic Philippines, where sev- above 1000 m (S. rabori and P. furvus), and eral endemic genera (Alionycteris, Haplon- our trapping techniques were limited, i.e., ycteris, Otopteropus, and Ptenochirus) are we did not use pitfall traps (Crocidura most common in old-growth forest. spp.). The presence of so many poorly The ecology of large fruit bats (Acerodon known species suggests that other species VOLUME 117, NUMBER 3 297 may await discovery, especially in high rids. Only one endemic species is a bat (Ac- mountain habitats and high in the canopy. erodon leucotis), and only it has its closest Musser & Newcomb (1983) suggested that relatives in the oceanic Philippines. This unknown species are yet to be discovered pattern of endemism is clearly consistent on Palawan, citing the report of Hoogstraal with the geological history of the Philip- (1951), which narrated their attempt to cap- pines and also highlights the importance of ture a ‘‘very large rat with a white tail’’ the greater vagility of bats over non-flying described to them by native Palaw’an. Oth- mammals. Of the 28 insectivorous bats, 18 er large islands in the Philippines have been species are somewhat to highly widespread shown to support diverse communities of in Indo-Australia (and some beyond), 2 are endemic small mammals which are restrict- shared only with the Sunda Shelf and In- ed to montane areas (e.g., Heaney 2001, dochina (Rhinolophus acuminatus and Heaney & Rickart 1990, Rickart 1993), and Rhinolophus cf. borneensis), 1 with the perhaps the same awaits discovery on Pa- Sunda Shelf only (Cheiromeles torquatus), lawan. 3 occur on the Sunda Shelf and the oceanic Medium to large mammals (Cercopithe- Philippines (Kerivoula pellucida, K. white- cidae, Manidae, Hystricidae, the carnivores, headi, Myotis macrotarsus), 1 occurs on and Suidae) are possibly the most thor- Palawan, Sulawesi and the oceanic Philip- oughly inventoried subset of Palawan’s pines (Mops sarasinorum), 2 occur only on mammalian fauna. Because of their large Palawan and in the oceanic Philippines size, they are easily observed compared to (Rhinolophus virgo and Myotis rufopictus), other mammals. Many of these species are and one occurs on Borneo, Sulawesi, and also commonly hunted, so obtaining speci- throughout the Philippines (Emballonura mens is often easier than for non-game spe- alecto). These data again demonstrate that cies. It is unlikely that other medium to the bats are more widely distributed and do large mammals await discovery on Pala- not clearly reflect the geological history, as wan, though the ecology of all requires do the non-flying mammal species, which much additional study. in the Philippines are usually restricted to a Biogeography.—As noted above, the Pa- single area that was united by dry land dur- lawan faunal region is part of the Sunda ing the late Pleistocene. From the highly Shelf and may have been connected to diverse fruit bat fauna of Borneo (17 spe- mainland Asia via Borneo (Everett 1889) cies; Payne et al. 1985), only five species during a Pleistocene episode of glacially- extend to the northern continental land- induced sea-level lowering (Heaney 1985, bridge islands of Sabah (Md. Nor 1995). 1986, 1991a), though current data leave this These are the same five non-endemic spe- uncertain. All other islands in the Philip- cies that can be found just to the north on pines are oceanic and have probably never Palawan Island. had a dry-land connection to any mainland We note that the combined totals of na- area (Heaney 1985, 1986, 2001). Of the 58 tive non-volant mammal species on Pala- native species currently known from Pala- wan that either are shared with Borneo and wan Island (tentatively including the small other portions of the Sunda Shelf or that are Crocidura sp.), 13 species arc endemic to endemic to Palawan and have their closest Palawan (and usually to some of the smaller relatives on Borneo or adjacent areas is 22 islands that were included in Pleistocene out of 24 (92%). The apparent exceptions Greater Palawan; Heaney 1986); 12 of are Hylopetes nigripes (related to H. albon- these are non-volant, all of which have their iger of Indochina) and Palawanomys furvus closest relatives on the Sunda Shelf. Eight (an enigmatic genus of unclear phylogenet- of Palawan’s 11 native rodents (73%) are ic position). If this analysis were extended endemic; all three non-endemics are mu- to the entire Palawan faunal region, Axis 298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON calamianensis (related to A. porcinus in In- more diverse on Palawan than in the oce- dochina) would also be included here. Four anic Philippines, and murid rodents notably species are widespread in Southeast Asia less diverse, for an island the size of Pala- (including parts of Wallacea), and no spe- wan. cies is shared with the oceanic Philippines Conservation issues.—The most pressing except for those 4 species (Macaca fasci- issue facing terrestrial wildlife in Palawan cularis, Prionailurus bengalensis, Paradox- is the rapid loss of forest cover, especially urus hermaphroditus, and Viverra tanga- in the primary lowland forests that are tar- lunga). If Sus barbatus, which occurs on geted for logging. Palawan’s forests are less Palawan, the Sunda Shelf, and in the mar- commercially valuable than the diptero- ginal islands of the Sulu Archipelago is carp-dominated forests of the other islands, counted as a widespread Southeast Asian and, consequently, deforestation occurred species, the total rises to five species. These later on Palawan. However, once forests data, in sum, strongly reinforce the conclu- were exhausted on Luzon, Mindanao, Ne- sion of Everett (1889), based on his anal- gros, etc., commercial logging operations ysis of bathymetric features of the ocean began working in lowland forest on Pala- floor and the pattern of relationships of the wan (Environmental Science for Social 18 species then known from Palawan and Change 1999) during the 1970’s and 1980’s associated smaller islands, that the Palawan at unsustainable levels (Quinnell & Balm- faunal region is an extension of the Sunda ford 1988, Kummer 1992). Since a logging Shelf, probably due to a fairly recent dry- ban was imposed in the early 1990s land connection, with only a small portion throughout the Province of Palawan, log- of its fauna shared with the oceanic Phil- ging has declined, but the large commercial ippines. operations appear to have been replaced by Patterns of species richness relative to is- small-scale, illegal commercial logging. We land area are also of interest. Insectivorous have seen that forest continues to disappear bats are so poorly known in the Philippines from the most accessible areas, and forest that it is possible only to say that the 28 edges are gradually creeping higher and species documented here compare favor- higher up the contours, in a manner similar ably with most islands in the Philippines; to that experienced on Leyte (Rickart et al. that is, there is no evidence that Palawan is 1993) and southern Luzon (Heaney et al. species-poor (Heaney et al. 2002). Fruit 1999). Lowland primary forest has been bats, on the other hand, are represented on eliminated from many parts of Palawan and Palawan by only 6 species, and this places the destruction shows few signs of easing. their diversity far below that on much Due to almost complete conversion of the smaller islands in the oceanic Philippines; coastal plain into ricefields, coconut, or oth- Maripipi, for example, has 10 species, but er plantations, distinctive ecosystems such is only 22 km2.Itseems certain that Pala- as freshwater swamp forest and beach forest wan has a depauperate fruit bat fauna, as have virtually disappeared (Widmann well as probably having lower fruit bat den- 1998). Slash and burn agricultural practices sity, as noted above, compared to the oce- have also been very damaging to forests, anic Philippines (Heaney 1991b). Species and Palawan has experienced a population richness of non-flying mammals, on the explosion due to high birth and immigration other hand, at 24 is much above the species/ rates. area curve documented in the oceanic Phil- It should be noted that caves are crucial ippines, though well below that of islands to maintaining the fauna, since approxi- on the primary portion of the Sunda Shelf mately 18 (32%) of Palawan’s mammals are (Heaney 1984, 1986; Heaney et al. 2002). bats that roost in caves. Caves have been It is interesting that carnivores are notably the focus of much destruction in the Phil- VOLUME 117, NUMBER 3 299 ippines; common activities in caves on Pa- gon, Ariel Carino, Leilani Berino, Dr. Ter- lawan include guano mining, general van- esita Salva, Dr. Edgardo Castillo, Dr. Pa- dalism, recreational exploration, and trea- cencia Milan, Dr. Josef Margraf, Indira Lac- sure hunting. Unlike many other parts of erna-Widmann, Siegfred Diaz, Deborah the Philippines (e.g., Heaney et al. 1991, Villafuerte, and the wildlife wardens of 1999; Rickart et al. 1993), we never en- Rasa Island provided valuable logistical and countered any evidence of cave-roosting administrative support. JAE thanks the Ta- bats being hunted on Palawan. We found bugon family for their extraordinary hos- guano mining to be common, usually near pitality during his stay on Palawan. We the mouth of caves. Recreational explora- thank A. C. Alcala, P. C. Gonzales, and P. tion of caves is steadily increasing; many O. Glass for depositing important speci- caves are currently being developed or ad- mens at UMMZ, and P. Myers for access to vertised for this purpose, while many more those specimens. H. Kafka, J. Mead, and R. proposals are in the planning stages. At Thorington provided access to specimens at PPSRNP (Site 15), hundreds of visitors the USNM, and F. Dieterlen kindly loaned may enter a one kilometer stretch of the specimens from SMNH. We are grateful to cave daily. The bats are clearly disturbed P. O. Glass for additional details about spec- by the activity, but the ultimate result of imens he collected. Ding Padilla, Clara such disturbance is unknown. Simpson, and Lisa Kanellos produced the Many medium to large sized mammals map in Fig. 1. Eric Rickart gave sound ad- are under significant hunting pressure on vice during the early stages of the project Palawan for their meat, the live animal and Danilo Balete helped with the transport trade, and medicinal use, as noted above, and identification of voucher specimens; but few data are available on the impact. Eric Rickart and Robert Timm provided The recent and on-going shift from subsis- constructive reviews of the manuscript. The tence to market economies among members Department of Environmental and Natural of the Tagbanua and other ethnic groups Resources, through the Protected Areas and may contribute to the decline of some spe- Wildlife Bureau and the Provincial Envi- cies (Lacerna & Widmann 1999), such as ronment and Natural Resources Office, pro- Sus barbatus, Pteropus vampyrus, and Hys- vided permits and encouragement. LRH trix pumila for meat, Macaca fascicularis thanks the Geological Sciences Department, and Arctictis binturong as pets, and Manis Northwestern University, for use of office culionensis for traditional Chinese medi- space during a sabbatical leave. cine. Literature Cited Acknowledgments Allen, G. M. 1922. Bats from Palawan, Philippine Is- Funding for these studies was provided lands.—Occasional Papers of the Museum of Zoology, University of Michigan 110:1–5. by the Palawan Council for Sustainable De- Allen, J. A. 1910. Mammals from Palawan Island, velopment, United States Peace Corps, Philippine Islands.—Bulletin of the American Philippine Cockatoo Conservation Program Museum of Natural History 28:13–17. through the Loro Parque Fundacion, Ten- Balete, D. S., L. R. Heaney, & R. I. Crombie. 1985. First records of Hipposideros lekaguli Thon- eriffe, Spain, and the Barbara Brown and glongya and Hill 1974 from the Philippines.— Ellen Thorne Smith Funds of the Field Mu- Asia Life Sciences 4:89–94. seum. Jun Saldajeno, Apollo Regalo, Ben- Barbehenn, K., J. P. Sumangil, & J. L. Libay. 1973. igno Maca, Erlito Porka, and Manual Lar- Rodents of the Philippine croplands.—Philip- dizabal made significant contributions to pine Agriculturist 56:217–242. Cabrera, A. 1909. Un nuevo ‘‘Rhinolophus’’ filipi- field work, often under difficult living con- no.—Boletin de la Real Sociedad Espanola de ditions. 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