Herpetologists' League A Checklist of the Herpetofauna of the Palau Islands (Republic of Belau), Oceania Author(s): Ronald I. Crombie and Gregory K. Pregill Source: Herpetological Monographs, Vol. 13 (1999), pp. 29-80 Published by: Herpetologists' League Stable URL: http://www.jstor.org/stable/1467060 Accessed: 05/10/2008 09:49 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=herpetologists. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact support@jstor.org. Herpetologists' League is collaborating with JSTOR to digitize, preserve and extend access to Herpetological Monographs. http://www.jstor.org Herpetological Monographs, 13, 1999, 29-80 ? 1999 by The Herpetologists' League, Inc. A CHECKLIST OF THE HERPETOFAUNA OF THE PALAU ISLANDS (REPUBLIC OF BELAU), OCEANIA RONALD I. CROMBIE1 AND GREGORY K. PREGILL2 'Division of Amphibians and Reptiles, United States National Museum, Smithsonian Institution, Washington D.C. 20560 2Department of Biology, University of San Diego, San Diego, CA 92110-2492 ABSTRACT: The Palau islands of western Micronesia comprise one of the most physiographically varied archipelagos in the Pacific. There are some 350 islands of volcanic and coralline origin that range from Babeldaob at 333 km2 (about 80% of Palau's total land area) to others, unnamed, that are less than one hectare. A barrier reef encircles most of the archipelago creating a lagoon up to 20 km wide. The majority of the islands are uninhabited, accessible with difficulty only by boat at high tide, and are densely vegetated over steep, rocky terrain. Because of their proximity to the Philippines, New Guinea, the Moluccas, and Borneo, the Palau islands host a significantly richer herpetofauna with more unique taxa than is found on other Pacific islands to the east. As with most other Pacific island groups, however, the herpetofauna has been poorly documented. Since the earliest investigations that began in the nineteen century, Palau's herpetofauna has been uncovered piecemeal. No sustained field effort had been conducted until we began a systematic survey of the islands in 1992. Since then we have collected in Palau every month of the year at least once. This checklist of 46 species of reptiles and amphibians includes almost twice as many taxa as were known from Palau (Owen, 1977) at the beginning of our study. Among these are new species or Palauan records of Gehyra, Hemiphyllodactylus, Lepidodactylus, Nactus, Perochirus, Emoia and Sphenomorphus. Because of the difficulties in surveying so many islands we believe that additional taxa and distributional records await discovery; even the smallest island in Palau can turn up sur- prises. Nonetheless, it is appropriate to consolidate what is now known about this intriguing fauna and environment. We provide an historical summary of previous collecting efforts, an overview of the islands' geology and habitats, and for each species account we include distributional information, taxonomic and systematic discussion, natural history observations, and a comprehensive bibliography. Key words: Micronesia, Palau Islands, Amphibians, Reptiles PERHAPS THE MOST zoogeographically significant yet poorly known archipelago in the Oceanic realm is the Palau Island group (now the independent Republic of Belau) in western Micronesia. The earliest herpetological investigations of this area by the Germans in the mid-1800's revealed new taxa in genera (Hypsilurus, Spheno- morphus, and others) otherwise unknown from the oceanic islands of the Pacific Plate, and the few subsequent field studies continued to yield additional novelties. Al- though these early collections suggested a diverse fauna with unexpectedly high lev- els of endemism, they were insufficient for addressing the numerous systematic and zoogeographic problems of this region. Because many other islands in the Pa- cific have relatively depauperate extant vertebrate faunas but much larger lists of recently extinct taxa (e.g., Steadman, 1995), Palau provides an ideal opportunity to see if this pattern also obtains on an island group with a more diverse extant fauna that is closer to potential source ar- eas. Therefore, in 1992 we began extensive surveys of the extant and extinct verte- brates of Palau. Although we have accu- mulated significant collections and data in our field work to date, each additional trip yields important (and often surprising) new specimens and distributional infor- mation. Nevertheless, given the consider- able recent increase of interest in Micro- nesian biology and conservation (e.g., Rodda et al., 1991), we feel it is appropri- ate to present our working list. Modern summaries of the Palauan her- petofauna (e.g., Cassell et al., 1992; Oto- bed and Maiava, 1994) have had to rely on the incomplete and taxonomically outdat- ed list of Owen (1977). Despite the ad- mitted shortcomings of the data, our list reflects current herpetological taxonomy (to August 1998) and it contains nearly twice as many species as listed by Owen 29 30 HERPETOLOGICAL MONOGRAPHS [No. 13 (1977), with many systematic problems duly noted. It is our intention to resolve some of these problems with future field work and collections. The strong zoogeo- graphic affinities between Palau and the Papuan region, and additional influences from the Philippines, Borneo, and Ocean- ia, further complicate systematic studies. This is especially so with respect to the logistics of obtaining pertinent compara- tive material from such a broad area that includes several political entities not known for expansive cooperation with sci- entific endeavours. We hope that our ac- cumulated specimens and data-base for Palau are such that future studies involving this area will be facilitated. The taxonomic aspect of our list is com- prehensive, but we particularly want to stress the incomplete nature of the distri- butional data we present. Although we list all island records currently available for each species, the results of earlier surveys plus our own field work have sampled only a small fraction of the multitude of islands in Palau, with varying degrees of rigor. The logistics of even briefly sampling islands spread across nearly six degrees of latitude are daunting and expensive; the survey work itself is grueling and often danger- ous. The Rock Islands in particular are of- ten very difficult to land on, much less to survey adequately for cryptic and elusive amphibians and reptiles. Even the smallest and seemingly insignificant islands should not be ignored, as demonstrated by the re- cent discoveries of Lepidodactylus pauro- lepis (Ota et al., 1995), Emoia impar, Lep- idodactylus sp. nov., and Nactus sp. (this paper). Our distributional data are intend- ed to convey only the most general pat- terns, many of which will undoubtedly be modified with additional field work. We but hope that the data presented here are not used to generate or justify counterproductive "conservation" efforts that will hinder or prevent the much-need- ed continuing data collection that is nec- essary to make sense of the Palauan her- petofauna. In short, we emphasize that these data are presented for general use, with the major caveat that they are not the "last word" in Palauan herpetology. We think that the following list adequately documents that the discovery phase is far from over. GEOGRAPHY AND ECOLOGY Spelling of Place Names The inconsistent spelling of Palauan place names is attributable to idiosyncratic useage by four separate foreign adminis- trations over the past one hundred years. These administrators made a limited at- tempt at using native names, and often misinterpreted their meaning when they did. Moreover, none of these languages (Spanish, German, Japanese, English) is adequate to fully capture the phonics of the Palauan tongue, such as the clipped nasal-quality ng. Even within Palau it is common to see the same name spelled two or three different ways. There are, for ex- ample, some 15 spellings for Palau itself, with the most common alternate being "Belau". The population center, both the island and the town, called "Koror" by practically everybody who lives there is spelled "Oreor" on many maps. For all place names we follow the spelling found on the USGS Topographic Maps of the Pa- lau Islands (1983-1984), which are the most detailed available and are used by most governmental agencies. We note, however, that some of the spellings on these topographic sheets are at odds with local or casual useage, or are not followed in older (e.g., Anon., 1957; Brigham, 1900; Bryan, 1971) or recent (Karolle, 1993; Motteler, 1986) literature. Finally, most of Palau's smaller islands are unnamed. The following is a list of our island local- ities and their common alternate spellings. Coordinates are for the center of the island/ group (Lat. 0.000 N, Long. 000.000 E) Babeldaob (Babelthuap, Babeldaop) Bablomekang Beliliou (Peleliu) Chemoi Island Group Euidelchol (Ashakasengu) loulomekang Kmekumer Islands (island 46 from Wiles and Conry, 1990) Malakal (Ngemelachel) Mecherchar (Eil Malk) 7.5 134.5 7.14 7.0 7.19 7.19 7.134 7.19 7.33 7.15 134.316 134.25 134.38 134.36 134.304 134.24 134.45 134.36 [ . 1I9 EPTLGCA OORPS3 FIG. 1. The islands of Micronesia and the southwest Pacific. Ngcheangel Atoll (Kayan- gel, Ngerebelas, Nger- iungs, Orak) Ngeanges (Necos) Ngeaur (Angaur) Ngebad (Ngabad) Ngebedangel (Ngoba- sangl) Ngedbus (Ngesebus) Ngelsibel Ngemlis Islands (Baile- chesengel, Iilblau, Ngemlis) Ngercheu (Ngergoi; Carp) Ngerdis (Kobasang) Ngerduais (Nardueis) Ngerechong Ngerechur Ngerekebesang (Araka- besan) Ngermalk Ngeruchubtang (Ngeru- gelbtang) Ngerukeuid Islands (Sev- enty Islands; islands 4, 12, 13, 16, 24, 34 of Wiles and Conry, 1990) Ngeruktabel (Urukthapel) Ngerur Ngesbokel (Gechbokul) Ngetmeduch 8.07 7.206 6.9 7.019 7.27 7.06 7.187 7.12 7.09 7.35 7.355 7.11 7.750 7.35 7.335 7.083 7.17 7.25 7.365 7.508 7.357 134.7 134.370 134.15 134.270 134.33 134.26 134.344 134.25 134.28 134.439 134.582 134.36 134.620 1 34A A.4 Oreor (Koror) Pkulaklim (Orrak) Toiracholodochoel Uchulameradel Uchulangas Ulebsechel (Auluptagel) Ulong (Aulong) Island Group Southwest Islands: Fana Helen Merir Pulo Anna (Palo Anna) Sonsorol Tobi 7.34 7.347 7.339 7.507 7.340 7.31 7.28 5.36 3.0 4.29 4.66 5.31 3.01 Physical Description 134.49 134.565 134.455 134.491 134.460 134.47 134.29 132.22 131.8 132.31 132.95 132.22 131.18 ...t. "" The thousands of islands and atolls in 134.459 Micronesia have a total land area of only 134.327 about 3,227 km2, roughly 0.03% of the sur- face area between 0?-20? N latitude and 135?-170? E longitude. The four major ar- chipelagos of Micronesia, from east to west, are the Gilberts (Kiribati), Marshalls, Carolines, and Marianas (Fig. 1). Palau is 134. 429 the westernmost group of the Caroline Is- 134.488 lands, centered near 7?20' N latitude and 134.496 134? E longitude. The island of Mindanao 19991] H RPETOLOGICAL M NOGRAPHS 31 32 HERPETOLOGICAl, MONOGRAPHS [Vo 12 in the southern Philippines is about 800 km west of Palau, whereas Irian Jaya and the Moluccas are, respectively, to the south and southwest about that same dis- tance; Borneo is an additional 700 km. All of the Gilberts and Marshalls are coral atolls or low islands. The Carolines include both volcanic and coralline islands. Car- tographically, Palau is included in the Car- oline Islands, but the western Carolines (Palau and Yap), as well as the Marianas, are exposed peaks of undersea ridges stretching between Japan and New Guinea and are not located on the Pacific Plate. The Palau islands originated on the now dormant southern section of the Palau- Kyushu Ridge, which formed by about 43 million years ago (mya) in a complex series of rifting and boundary shifts of the Pacific Plate margin during formation of the Phi- lippine Basin (Kroenke, 1984; Seno and Maruyama, 1984). Volcanism along the Pa- lau-Kyushu Ridge ceased in the earliest Miocene about 29 mya. Subsequently, the Palau-Kyushu ridge fractured and the eastern segment ultimately developed into the West Mariana Ridge. Back-arc spread- ing between the the two submerged ridges (Palau-Kyushu and West Mariana) wid- ened the Philippine basin to the east, and for the last 17 mya the basin has been moving northwest with respect to Eurasia. Thus, Palau, Yap and the Mariana Islands rest on the eastern margin of the Philip- pine Plate where it contacts the western edge of the Pacific Plate. The Palau, Yap and Mariana Trenches that mark the sub- duction zone at plate contact are some of the deepest waters on earth. When the is- lands that are now Palau first emerged above sea level is uncertain. The oldest or- ganic limestones are early Miocene (ca. 25 mya), which suggests that the volcanics were emergent sometime before that, per- haps by late Oligocene (ca. 30 mya). The eastern Caroline Islands, by con- trast, are comparatively young and origi- nated on the Pacific Plate. From Kosrae in the east to Woleai (Ulithi Atoll) in the west, the eastern Carolines stretch across the Pacific for 2100 km. The Truk (Chuuk) Islands are estimated to have originated at ca. 12-14 mya, whereas Kosrae is much FIG. 2.-The main islands of the Palau archipelago. younger at ca. 4 mya (Springer, 1982). The Carolines are being carried westward to- ward the Philippine Plate margin. The Republic of Belau includes the Pa- lau Islands proper and a cluster of remote outliers 320 to 480 km to the southwest (Fig. 2). Each of the latter (Tobi, Sonsorol, Fana, Merir, Pulo Anna and Helen) are separated from one another by from 130 to 280 km of open ocean. Few people out- side of Palau are aware of them, and no one in the west has ever bothered to give them a collective name other than the Southwest Islands. The Palau islands themselves are arranged in a 160 km curv- ing chain from Ngeruangel reef in the north to Ngeaur Island in the southwest. The total land area (including mangrove) is approximately 415 km2. Most of Palau's 350 islands are enclosed in a reef: barrier to the west and fringing on the east. Only Ngeaur in the southwest and Ngcheangel Atoll and Ngeruangel reef in the north are [No. 3FERPETOLTOGICAL 199IEPTLGCA OORPS3 Babeldaob 5 km FIG. 3. The island of Babeldaob, Republic of Palau. outside the lagoon. The lagoon is approx- imately 20 km at its widest, covers about 1450 km2, and varies in depth from barely a meter to over 20 m in a few places. Out- side the reef ocean depths drop hundreds of meters almost immediately. Babeldaob (Fig. 3) is the largest island at 333 km2, accounting for about 80% of Palau's total land area. The next largest is- lands are Ngeruktabel (20.2 km2), Beliliou (12.4 km2), Ngeaur (8.3 km2), Mecherchar (8 km2), Oreor (7.5 kin2), Ulebsechel (4.3 km2) and Ngerekebesang (2.3 km2). Most of the other islands are less than 1 km2 and many are no more than a few hectares. About two-thirds of Palau's 15,500 inhab- itants live on Oreor, and practically all of the other residents live in the vicinity, ei- ther on southern Babeldaob, Ngerekebes- ang or Malakal, each of which is connected to Oreor by a causeway. The islands are of four geological types: 1) volcanic, 2) high limestone, 3) low lime- stone, reef or atoll, or 4) a combination of volcanics and limestone. The Military Ge- ology of Palau compiled by the U.S. Army of Corps of Engineers (1956) remains the most thorough description of Palau's geo- logical features. The following summary is based on that compendium. Volcanic is- lands include Babeldaob, western Oreor, Ngerekebesang, Malakal, and numerous tiny islets in the northern end of the la- goon. Contours of the larger volcanic is- lands are generally rolling hills punctuated by steep irregular outcrops or other prom- inent landmarks. Palau's highest point, 242 m Ngerchelchuus in northwest Babeldaob, is one such feature. The largest evident volcanic center is Ngeremeduu Bay on the west coast of Babeldaob. The original cone collapsed leaving an eroded caldera which is now the bay. Poor exposures elsewhere on the volcanic islands obscure most for- mer eruption centers, a possible exception being the volcanic breccias and the incli- nations of pyroclastic rock on Ngerekebes- ang. Coastlines of the volcanic islands are irregular, sometimes steep, and often form small bays. Approximately 125 km of Ba- beldaob's 157 km coastline is bound by mangrove. The extent of calcareous beach sand on the volcanic islands is very limited, and mainly confined to Babeldaob's north- east and east coast. Each of the larger vol- canic islands is dissected by streams, but the only perennial flows are on Babeldaob. On Oreor and Ngerekebesang the radial drainages slow to a trickle when rain ceas- es for more than a week or so. The major streams and rivers of Babeldaob are more or less aligned with the north-south axis of the island, but are linked irregularly with connecting tributaries. The largest river, Ngermeskang, flows for about 16 km into Ngeremeduu Bay from the northeast. The largest body of standing freshwater is Lake Ngerdok (approximately 30 hectares) lo- cated about 2 km from the east-central coast of Babeldaob. The high limestone islands number more then 300, but comprise only about 199 ] HERPETOLOGICAL M NOGRAPHS 33 34 HERPETOLOGICAL MONOGRAPHS [No.~~~~~~~~~~ 13 oNgerur Ngerekebesang O-or J Ngerchaol tu? n Butottoribo Ul. secU Ngebedangel , Nler uce 0pnf Ngeruktabel Euidelchol _, 0 Ngerukeuid lids Mercherchar Bablomekang I % Ngemihs lids 55*. * V * Ngercheu Ngerechong Rock Islands - Palau t-; Ngedbus 5km / l. Ngebad Beliliou FIG. 4.-The Rock Islands of the Palau lagoon. 10% of Palau's land area. They include practically all of the islands in the lagoon. Those in the central and southern part of the lagoon are referred to locally as the Rock Islands (Fig. 4). Ngeruktabel and Mecherchar are the largest. The Rock Is- lands are formed in the Palau Limestone which consists of light-colored, poorly bedded, porous to very dense raised reef and lagoonal coralline deposits laid down in shallow warm water. The age of the Pa- lau Limestone ranges from Miocene in the northern part of the Lagoon and parts of Beliliou (Bloody Nose Ridge) to Pleisto- cene elsewhere. The larger islands are characterized by steep, arcuate ridges flanked by descending skirts of parallel and/or tangential ridges that once marked exposed, successive reef fronts. The small- er rock islands probably derived from scat- tered, irregular deposits in the lagoon. The limestone islands were raised to their pres- ent extent by asymmetrical upwarping fol- lowing the Neogene rifting and faulting that was associated with the tectonic and volcanic activities in the area. Uplift to as much as 200 m elevation occurs on an axis from southwestern Babeldaob to Beliliou. West of this line there is slight westward tilting. Although there are occasional coves and narrow beaches, the shorelines of the limestone islands are often nearly vertical, and almost everywhere are undercut by solution notches forming horizontal over- hangs 1-5 m deep. Undercutting of the smaller islands gives them the appearance of mushrooms. Ashore, solution features of sharp, jagged pinnacles, pits and rubble make for exceedingly treacherous footing. Springs occur on some of the high lime- stone islands but there is no surface drain- age. Caves, sinks, and other subsurface so- lution features are abundant in this highly karstified terrain. On several of the larger Rock Islands numerous interior depres- sions extend below sea level to form saline lakes. The reef islands and atolls of Palau re- sult from minor uplift of reef rocks and wave building accumulations. The largest and best developed of Palau's atolls is Ngcheangel (Kayangel), located about 3 km beyond the north end of the Palau reef. It is more or less oval, about 7 X 3.5 km, and embraces four small islands (beach deposits) on its east side. Several of Palau's islands are a combi- nation of the preceding geological types. High limestone ridges are present on the south end of Babeldaob. In a zone south and southwest of Babeldaob volcanic and high limestone islands occur adjacently or combined. The two volcanic islands, Nger- ekebesang and Malakal, are separated by the high limestone island of Ngerchaol. Western Oreor is underlain by volcanics, whereas the southeast part is a high lime- stone ridge. The two larger islands at the southwest end of the Palau group, Beliliou and Ngeaur, consist of high limestone ridg- es, low platforms, and beach deposits. Today, the Palau islands exist along a continuum of coastal morphologies deter- mined by eustatic changes in sea level dur- ing the Pleistocene. During the last glacial maximum (18,000 years ago), when sea levels may have depressed as much as 120 m, the entire lagoon and most of the ar- chipelago's 350 islands would have formed a single island of approximately 2250 km2, 34 o. 1999] HERPETOLOGICAL MONOGRAPHS 35 more than five times the present area of emergent land. During periods of higher- than-present sea level, for instance in the mid-Holocene (e.g., Allen, 1997), Palau was more fragmented than it is today. The effect of these oscillations on the terres- trial biota was undoubtedly considerable. Indeed, the present distributions of Pacific island amphibians and reptiles in general must be considered in view of these sea level fluctuations (e.g., Gibbons, 1985; Gibbons and Clunie, 1986; see also Alli- son, 1996). Habitat and Land Vertebrates Palau's climate is tropical and rainy. Mean annual temperature is 27.2 C (81?F) and mean daily temperature varies no more than 1 degree throughout the year. On average the islands receive 355 cm (140") annual rainfall with the wetter months (July and August) receiving 25-50 cm, the drier (February to April) 15-20 cm. A survey of vegetation and timber re- sources of Palau was drafted by the U.S. Forest Service (Cole et al., 1987; MacLean et al., 1988), and some of the vascular plants were listed by Fosberg et al. (1980). A soil survey of Palau by Smith (1983) in- cludes 1:10,000 scale aerial photo maps of Babeldaob, Oreor and the causeway is- lands, Beleliou, and Ngeaur. Cole et al. (1987) estimated that three-fourths of Ba- beldaob and the other volcanic islands are covered in native upland forest, in which Calophyllum, Ficus, Elaeocarpus and Campnosperma are common genera. In the past few decades native upland forest has been greatly reduced, and most of the remaining area, especially on Babeldaob, is covered by coarse grasses burned an- nually, along with scattered pandanus and ferns. Mangrove (especially Rhizophora sp.) accounts for about 11% of Palau's veg- etation cover, practically all of which is found around Babeldaob, Oreor, Malakal, Ngerekebesang, and Beleliou. The luxu- rient limestone forests of the Rock Islands form impressive, dense stands with cano- pies reaching 20 m or more on the larger islands. Freshwater swamps and marshes in Palau comprise about 5% of the area. Cultivated lands comprise about 3-4% and yield such crops as bananas, cassava, co- conut, papaya, mango, pineapple, and be- tel nut. Urban areas occupy about 1% (Otobed and Maiava, 1994). Our impres- sions would put the percentage of agricul- tural and urban land considerably higher. Besides amphibians and reptiles, the vertebrates of Palau include a fair diversity of freshwater fishes and birds, but a de- pauperate mammal fauna. The resident freshwater and land birds consist of wide- spread oceanic species, species confined to Micronesia, species found elsewhere only in Indonesia or the Philippines, endemic species, and a few introductions. Palau's modern avifauna has received much more attention than its herpetofauna (Baker, 1951; Engbring, 1983, 1988; Engbring and Pratt, 1985; Pratt et al., 1980), although it would be surprising if these islands have not experienced extinctions of native bird species similar to what has been discov- ered elsewhere in Oceania (Steadman, 1995), including the nearby Marianas (Steadman, 1999). The endemic land mammals known from Palau are the Palau Sheath-tailed Bat, Emballonura semicau- data palauensis, the "Marianas" Fruit Bat, Pteropus marianus pelewensis, and the larger Palau Fruit Bat Pteropus pilosus. The Palau Fruit Bat has not been seen since the 19th century and is presumed to be extinct (Wiles et al., 1997). Introduced taxa include rodents Rattus spp. and Mus musculus (see below), as well as the Asiatic musk shrew Suncus murinus, confined to Ngeaur (see Ruedi et al., 1996 for Suncus zoogeography). The introduced Crab-eat- ing Macaque, Macaca fascicularis, is most common on Ngeaur, and a few individuals also occur on Babeldaob. A review of Palau's freshwater fishes was compiled by Bright and June (1981), but very few of Babeldaob's rivers and streams (particularly in the eastern sectors) have been sampled, much less properly sur- veyed. Undoubtedly endemic, unde- scribed species await discovery (e.g., Par- enti and Maciolek, 1993). Human impact is most obvious in and around Palau's main habitation centers (Oreor, Ngerekebesang, Malakal, Airai) and surrounding settlements and planta- 99] PETOLOGICAL GRAPHS 26 HERPETOLOGICAL MONOGRAPHS [No. 13 tions. Much of Babeldaob has been defor- ested, especially the southern half. Al- though Palau's present human population is relatively small (about 15,500), estimates of around 40,000 native Palauans were be- lieved to be present at the time of first Western contact (Johannes, 1981; Osbor- ne, 1966). There is some dispute of that figure, which is based on accounts of the early traders to the islands (Johannes, 1981). Nonetheless, ancient agricultural terraces and other archaeological indica- tors suggest that the population was sig- nificantly greater than it is today (Craib, 1983; Osborne, 1966). During this century population highs were in the early 1940's when 25,000 Japanese residents and native Palauans lived in the islands (U.S. Army, 1956; see also Bailey, 1991). WWII brought substantial devastation to parts of Palau. Beliliou, the site of one of the bloodiest battles in the Pacific (Hough, 1990; Ross, 1991), was burned over most of its area. The island has gen- erated dense second growth forest over the past 50 years. Ngeaur, Oreor, and southern Babeldaob were also heavily bombed by American forces, but of these only Ngeaur was invaded. Despite the im- pact of human occupation and WWII, Pa- lau still claims some pristine habitat, or that which is but moderately disturbed. The Rock Islands are practically all pri- mary forest due to their steep slopes and brutal topography (Johnson, 1972). Yet even there one of the most odious mea- sures of human disturbance is conspicu- ous-introduced Black Rats (Rattus rat- tus). Our campsite on uninhabited Ulong was overrun by hundreds of rats one night in January 1995. During the same month in 1993 we trapped along a 200 m forest transect in Ngaraard State (northern Ba- beldaob) and had a capture rate of about 73% (19 of 26 traps) in one hour. Rats can also be seen commonly in Koror town by day. The effect of R. rattus as a predator on Palau's herpetofauna and birds has not been documented, but we suspect that it is substantial. Rapidly growing economic interests in Palau probably will adversely affect the na- tive fauna in the next few decades (Anon., 1998; Cassell et al., 1992). The human population is likely to double in 5-10 years (Maiva and BNRD, 1994), and at least one major road is under construction to the north end of Babeldaob where none pres- ently exists. Erosion and siltation of some Babeldaob water courses are already evi- dent. Tourism is burgeoning, and numer- ous hotels are planned or being built in Oreor, southern and eastern Babeldaob, Malakal, and on the Rock Islands. The government of Palau has proudly and jus- tifiably promoted its marine resources as a tourist attraction. We hope that this does not come to the detriment of its terrestrial ecosystems. HISTORICAL SUMMARY The Spaniards were the first Europeans to land on Palau, claiming the islands for the King of Spain in 1710. Their presence was uneventful, particularly biologically except perhaps for the first introduction of rats. Several European ships were wrecked in the treacherous waters around Palau in the 18th and 19th Century and stories of the interactions between the marooned sailors and Palauans have been amply chronicled (Keate, 1802, 1808; Martin, 1980). Not surprising, the early Pacific ex- ploratory expeditions usually avoided Pa- lauan waters in favor of the better harbors on Guam, Yap, and Kosrae, so their natu- ralists and collectors had little or no op- portunity to sample the Palauan fauna. The French ships Astrolabe and Zelee briefly contacted the residents of Beliliou in January 1838 (Rosenman, 1992:162), but we have found no herpetological spec- imens that were collected at that time. The French did, however, make the most sig- nificant early contribution to Micronesian herpetology during the voyage of the Co- quille, 1822-1825. Many of the most com- mon and widespread Pacific lizards were described from the island of Oualan (now Kosrae in the eastern Carolines), four of which also occur in Palau (Gehyra ocean- ica, Emoia atrocostata, Lamprolepis sma- ragdina, Lipinia noctua). Lessons's (1830) zoological report of the Coquil expedition generated confusion on several levels, however. Brygoo (1986) [ o. 3 1999] HERPETOLOGICAL MONOGRAPHS 37 pointed out that the Atlas of plates from the Expedition appeared in 1826 but the text not until 1830, so Lesson's taxa must date from the Atlas. The types of several of the species are no longer extant (Bry- goo, 1986) and Barbour (1911) suggested that Lesson's type locality (Oualan, now Kosrae in the eastern Carolines) was sus- pect. Loveridge (1948) disagreed and crit- icized Barbour's interpretation but we feel that Barbour's suspicion was justified for at least some of the taxa. The situation is complex and a lengthy discussion is not ap- propriate here since it has no direct impact on Palau, although we hope to provide clarification in a future paper. Serious herpetological research in Palau began only after the Germans suceeded Spain in 1899. Unlike the Spanish, Ger- man merchants and diplomats were actu- ally resident in Palau. Particularly notable among them was Carl Semper, who wrote an interesting book on his Palau experi- ences (Semper, 1925, English translation Semper and Craig, 1982). Semper was an experienced field naturalist who had made important collections in the Philippines and elsewhere in Asia. His Palau herpe- tological material was studied by Wilhelm Peters in Berlin, who described many new taxa (Peters, 1867a,b, 1874, 1877, 1879). Most of the Berlin specimens have no lo- cality data other than the island group, var- iously rendered as "Pelew", "Pelew-In- seln" or "Palaos". Additional Palau material (mostly from Angau = Ngeaur) in the Senckenberg Museum, Frankfurt, was reported by Sternfeld (1914, 1920), who (1920:385) mentioned that, "another new lizard from Palau" was present in the Senckenberg collection and that it would "also be described here." However, no new Palauan lizards were described in the 1920 paper or any subsequent publications by Sternfeld before he was executed by the Nazis in 1943, nor are there type spec- imens of any Palauan lizards in the Senck- enberg collection (Mertens, 1967). The material of this new species alluded to by Sternfeld was not located during a visit to Senckenberg by RIC in May of 1986, and we are unsure which of the still unde- scribed Palauan lizards Sternfeld referred to. The Japanese period (1914-1944) pro- duced relatively little herpetological re- search, as summarized in Utinomi's Biblio- graphica Micronesica (Fisher, 1947). No new herpetological taxa were described, and most of the information was published in obscure, generally unavailable journals. Messel and King (1991), in summarizing the crocodilian literature, mentioned two significant papers by Motoda (1937, 1938). However, we have been unable to locate any extant herpetological collections that were made by the Japanese during their 30-year occupation of Palau. U.S. influence in Palauan herpetology began with personnel of the Naval Medi- cal Research Unit (NAMRU II), who were active in Palau soon after the islands were secured from the Japanese. Biological ma- terial was collected from November 1944 until about September 1945 (Baker, 1951; USNM catalogue records). Although col- lecting efforts centered on Beliliou and offshore islands, the team also obtained herpetological material from Ngeaur, Ngercheu, and Ngcheangel Atoll. The NAMRU collections provided the first meaningful distributional data (island and specific localities) for Palau amphibians and reptiles. Although most of the NAM- RU material was deposited in USNM, one member of the team, William Lewellen, contributed most of his Ngeaur specimens (CM 29085-29103, collected between 19 November 1944 and 13 January 1945) to the Carnegie Museum. Until recently, the most extensive and important Palauan herpetological collec- tion was made by H. Adair Fehlmann dur- ing his ichthyological surveys in the mid 1950's (Fehlmann, 1960). Most of the fish data remain unpublished but his discovery of the remarkable endemic skink Lipinia leptosoma appeared early (Brown and Fehlmann, 1958). Fehlmann's herpetolog- ical specimens were deposited in the Stan- ford University collection (now housed at CAS) and they represent important base- line data on distributions before extensive development on Palau. The earliest natural history observations 1999] 38 HERPETOLOGICAL MONOGRAPHS [No. 13 on the endemic Palauan frog (Platymantis pelewensis) was by a Palauan, Kenji Atoda (1950). Palauan citizens and residents have continued to contribute meaningfully to herpetology ever since. Although he was severely criticized by Messel and King (1991) for encouraging exploitation and even extermination of crocodiles in Palau, Robert Owen was the entomologist and Chief Conservation Officer of Palau for many years, and was unfailingly helpful to visiting researchers. He provided data and specimens to many individuals (including us), and meaningfully aided projects in- volving the Palauan flora and fauna during his tenure there. Owen's successors, par- ticularly Demei Otobed and Kammen (Taktai) Chin, have continued this fine tra- dition of cooperation and have made sig- nificant contributions of their own (e.g., Cassell et al., 1992; Fosberg et al., 1980; Otobed and Maiava, 1994). ZOOGEOGRAPHY Although it is premature to discuss Pa- lauan zoogeography in any detail before the relationships of many species are un- derstood, summary papers on SW Pacific biogeography continue to appear (Allison, 1996; Brown, 1997; Holloway, 1997 among others), some of which are hampered by the lack of recent data on Palau. A brief summary is justified. We are confident that analysis of the data in hand will confirm that Palau is an important zoogeographic crossroads, with faunal influences from all four major source areas. Our preliminary data do not confirm any species with un- equivocal Bornean relationships, but the Oceanic (Gehyra oceanica, Lepidodactylus sp. nov., Perochirus sp.) and Philippine (Lepidodactylus moestus, Mabuya sp.) af- finities are more certain. Without ques- tion, the greatest influence on the Palauan herpetofauna is the Greater Papuan region (New Guinea + the Solomons and Moluc- cas, East Wallacea of Brown, 1997 and others). At least 13 Palauan species are de- rivatives of this area, and several of the 13 or 14 others of uncertain origin will likely be added to that total with additional re- search. Critically missing from the puzzle are sufficient data from the Southwest Islands. This chain of islands provides a potential stepping stone pathway between Palau proper and the Greater Papuan region, particularly during periods of lower sea level. The few extant herpetological rec- ords from the islands provide many sur- prises, as much for the species that have not been collected as those that have. Donaldson's (1996) discussion on the strik- ing diversity and complexity of relation- ships in the fish fauna convinces us that the terrestrial fauna will be equally signif- icant. SPECIES ACCOUNTS Forty-six species of amphibians and rep- tiles (39 terrestrial and 7 marine) are treat- ed in the following section (Table 1). Three species are twentieth-century intro- ductions, eight species are certain endem- ics, and ten others are probably endemic. The latter include several species repre- sented by only one or a few specimens and their status cannot yet be determined. Other species are known under existing names but for reasons discussed we be- lieve are distinct species. For each species, the localities listed under Distribution are those islands in Palau where we have col- lected voucher specimens and/or verified existing museum records. Almost all of the island records derive from, or are verified by, our field work through 1998. Appendix I is a checklist of species by island. AMPHIBIA: ANURA Bufonidae Bufo marinus (Linnaeus, 1758) This large toad is very widely distribut- ed, both in its natural range of the Neo- tropics (Zug and Zug, 1979) and in many other parts of the world via introduction (Easteal, 1981; Honegger, 1970). Details of exactly when and whence Bufo marinus was first introduced into Palau are un- known (Easteal, 1981:102) but the first museum specimens were collected in the early 1950's. Although the NAMRU 2 teams did not collect any Bufo during their stay on Beliliou in 1945, it is likely that toads were introduced during U.S. military activity in Palau in the post-war years, pos- RPETOLOGICAL NOGRAPHS [ o. 1-991 R L MONOGRAPHS 39-- - TABLE 1.-Amphibians and reptiles of the Palau Is- lands. Those marked by an asterisk (*) have not been reported previously. FROGS Bufonidae Bufo marinus (introduced) Ranidae Platymantis pelewensis (endemic) LIZARDS Agamidae Hypsilurus godeffroyi (status uncertain) Gekkonidae Gehyra sp.* Gehyra brevipalmata (endemic) Gehyra mutilata Gehyra oceanica Gekko sp. (endemic)* Hemidactylus frenatus Hemiphyllodactylus cf. H. typus* Lepidodactylus sp. nov.* Lepidodactylus lugubris Lepidodactylus moestus Lepidodactylus paurolepis (endemic) Nactus sp. (endemic)* Nactus cf. N. pelagicus* Perochirus sp.* Polychrotidae Anolis carolinensis (introduced) Scincidae Carlia cf. C. fusca (introduced)* Cryptoblepharus sp. (probable endemic)* Emoia atrocostata Emoia caeruleocauda Emoia impar* Emoia jakati Eugongylus sp. (endemic) Lamprolepis smaragdina Lipinia leptosoma (endemic) Lipinia cf. L. noctua* Mabuya sp. (endemic) Sphenomorphus sp. nov. (endemic)* Sphenomorphus scutatus (endemic) Varanidae Varanus cf. V indicus SNAKES Boidae Candoia cf. C. carinata Colubridae Cerberus rynchops Dendrelaphis sp. Hydrophiidae Pelamis platurus Laticaudidae Laticauda colubrina Typhlopidae Ramphotyphlops acuticaudus (endemic) Ramphotyphlops braminus TURTLES Chelidae Gen. & sp. indet.* Bataguridae (?) cf. Chinemys reevesii* TABLE 1.-Continued. Cheloniidae Chelonia mydas Eretmochelys imbricata Dermochelys coriacea Lepidochelys olivacea CROCODILES Crocodylidae Crocodylus porosus sibly from Guam, New Guinea and/or Aus- tralia where Bufo has been well estab- lished since the 1930's. Gressitt (1952) visited Kayangel Atoll in July-November 1951 and the toads were well established by then. The hypothesis of a post-war (i.e., mid-1940's) introduction into Palau is fur- ther supported by USNM 123909, collect- ed in July 1946 on nearby Yap. The field notes of the collector state that, "On Yap (the toad) does not reach the size it does on Guam and other islands ..." but that it was "very common" around Yaptown (now Colonia) at that time. Fisher (1948) did not mention toads from Palau, but he confirmed that they were common on Ponape and all the islands of Yap during his visits there, at some unspecified time before 1948. Although Easteal (1981:117) found mu- seum vouchers for only three islands in Pa- lau-Arakabesan (= Ngerekebesang), Kayangel (= Ngcheangel Atoll), and Ker- rar (= Koror, now Oreor)-Bufo marinus is widely distributed in the archipelago. It is most abundant on the causeway-con- nected islands where agriculture (particu- larly taro patches) and other human activ- ities provide abundant breeding places. The species is not found on most of the Rock Islands where standing water rapidly percolates through the steep, porous lime- stone, thus eliminating potential breeding places. We have sight records from Ulong in January 1995, when the toads were quite common, but several subsequent vis- its to collect vouchers were unsuccessful. Rat density was also startlingly reduced and we initially suspected that a rat poi- soning project might also have extirpated the toads. An El Nino-associated drought in late 1997-early 1998 killed off much of 9 HE PETOLOGICAL 39 40 HERPETOLOGICAL MONOGRAPHS [No. 13 the undergrowth in the disturbed area around the beach that it is unlikely any toads survived this exceedingly dry period on Ulong. Our one vouchered Rock Island record is USNM 531973 from Pkulaklim, a high island just off the Airai Dock area of southeastern Babeldaob. The toad was ac- tive by day in low undergrowth adjacent to a low swampy area just behind a small beach on the west side. Taro had recently been planted in the swamp and the toad(s) may have been intentionally brought in with the agriculture. Distribution.-Babeldaob, Beliliou, Ma- lakal, Ngcheangel Atoll (Ngcheangel), Ngeaur, Ngerekebesang, Oreor, Pkulaklim. Ranidae Platymantis pelewensis Peters, 1867:33 Wilhelm Peters (1867a:33) described Platymantis pliciferus pelewensis from the "Pelewinseln", based on a single specimen collected by Dr. C. Semper. The holotype (ZMB5699) was listed as "not located" in Frost (1985:469) and Peters' extremely brief description provided no morpholog- ical information, merely indicating that the Palau specimen was nearly identical with the Philippine Platymantis plicifera Guin- ther, differing only in the coloration of the side of the head (all black in plicifera, up- per and lower lips flecked with white in pelewensis). The name was overlooked in subsequent reviews of ranids (Boulenger, 1882, 1918), but it reappeared as a valid species in the more recent literature (Gor- ham, 1965; Frost, 1985). Although Ota and Matsui (1995) and Kuramoto (1997) re- cently presented some data on the rela- tionships of the species, no historical re- view has ever been published. We will fill in some of the "lost years". Kampen (1923:190) transferred plicifera to the genus Rana (but maintained Platy- mantis as a subgenus), where the species name became a homonym. He provided the unnecessary replacement name Rana rugata and regarded pelewensis as a strict synonym of it. Loveridge (1948:407) criti- cized Kampen's arrangement, regarded Philippine and New Guinea Platymantis as subspecifically related, and pointed out that the name corrugatus had priority over plicifera. He also commented, ". . . wheth- er Platymantis plicifera pelewensis Peters, 1867, from the Pelew Islands, is a recog- nizable race, scarcely concerns us here." Brown (1952:51) and Inger (1954:351) concluded that the Philippine and New Guinea "corrugatus" were not only specif- ically distinct, but quite unrelated to one another. Inger (1954:355-356) correctly pointed out that both pelewensis and pa- puensis were more similar to the Philip- pine Cornufer meyeri (= Platymantis dor- salis, Brown and Inger, 1964) than to P. corrugatus (see also Tyler, 1979). He re- garded the three taxa as distinct species and provided some data on pelewensis (size, dorsal and ventral skin texture, dig- ital and vocal sac morphology, ova dimen- sions). Atoda (1950) published an excellent em- bryological study of direct development in the Palauan frog but used no scientific name, speculating that it was, "... prob- ably a new species of Rana." Alcala (1961: 721) referred Atoda's material to Cornufer pelewensis (see Zweifel, 1969 for use of the name Platymantis rather than Cornu- fer). Gorham (1965:410-411) confirmed that pelewensis was a valid species and provided the first illustrations of it (sketch- es of the lateral head, ventral view of the hand and a photo of an entire specimen (1965:432, Figs. 3A,B)). Brown (1965:2-3) summarized the known species of "Corn- ufer", also listing pelewensis as a valid spe- cies. The validity of the Palauan Platyman- tis has remained unchallenged since Brown and Tyler (1968). [Tyler's (1979:82, Fig. 6) mention of one species of Rana in Palau is incorrect. This is either a lapsus or perhaps a reference to Atoda's (1950) erroneous remark that P pelewensis as "an undescribed species of Rana", but Atoda's paper is not cited.] Most recently, Ota and Matsui (1995) described the karyotype and Kuramoto (1997) analysed morphological, karyologi- cal, and acoustic information on P pelew- ensis to clarify its relationships. All data unequivocally confirmed that pelewensis is very closely related to P papuensis from New Guinea (e.g., Menzies, 1982a,b) and [ o. 1999] HERPETOLOGICAL MONOGRAPHS 41 quite distant from the Philippine P. dor- salis. We have collected several hundred more P. pelewensis than our Japanese col- leagues, material that includes call record- ings, skeletons, and frozen tissue, but we concur completely with their conclusions. Our larger sample sizes negate the size dif- ferences between pelewensis and papuen- sis noted by Kuramoto (1997) and greatly expand the limited pattern polymorphism he reported. Platymantis pelewensis is ubiquitous in Palau, at least from Kayangel to Ngeaur, but Owen (1977) stated that frogs were not found in the Southwest Islands. This species is abundant in backyards in Koror town and in undisturbed forest on the Rock Islands. Although Kuramoto (1997: 183) characterized the Palau frog as being "found among leaf litter on the forest floor," it is most assuredly not a forest ob- ligate. In fact, based on call records it may be most abundant in patches of thick grass in disturbed areas, where dozens of males can be heard vocalizing among the impen- etrable, moisture-retaining roots. Call rec- ords also confirm the presence of frogs on many of the Rock Islands, even very small pinnacles. We suspect P pelewensis may be salt tolerant as well, since we have heard and collected the species at the edg- es of mangrove swamps and other haline areas. Details on the natural history of the Pa- lau frog have been minimal since Atoda's (1950) description of the development of the clutch he discovered. Unfortunately, he did not provide information on where the eggs were found, if there was an at- tendant frog, or other important details on direct-developing species. No other clutch has ever been found, despite our intense field efforts. We suspect, however, that males call from and defend a calling/nest chamber and attend the clutch. We base this on the fact that calling males are often nearly impossible to find, particularly dur- ing dry periods. On rainy evenings males may call from exposed sites but in general they are well concealed and several have been collected from calling chambers, un- der detritus but not subterranean. One of our more startling discoveries in- volved the ecology of female P. pelewensis. Collections made from the forest floor (e.g., Kuramoto, 1997) are usually strongly skewed towards males and subadults, but we found that females congregate, often in incredible numbers, in the multitude of natural caves and Japanese-made bunkers that are omnipresent in Palau. These bun- kers usually retain moisture and many have an abundance of trash on the floor or crevices in the wall, so we initially sus- pected they may have been an important egg deposition area. However, after thor- oughly searching dozens of bunkers (some repeatedly throughout much of the year) we have yet to find any indication that re- production occurs there. The bunker frogs are 90+% female, many of them larger than maximum previously reported size, and we initially suspected that they might be a second species. However, the few males found in samples from more than 12 "caves" were all immature; calling males have been heard near the mouths of the caves (but never inside) and the calls were all typical pelewensis. Apparently large females use the bunkers as diurnal retreats and venture into the forest at night for reproduction, presumably at the male calling site. The bunkers remain con- sistently moist even during the driest spells, and provide an abundance of food (cave crickets are particularly abundant). On several occasions we have also found juvenile P. pelewensis in the guts of large females from bunkers. Distribution.-ENDEMIC. Babeldaob, Bailechesengel (Ngemlis Islands; call only), Beliliou, Malakal, Ngcheangel Atoll (Ngcheangel, Ngeriungs), Ngeaur, Nger- cheu, Ngerechur, Ngerekebesang, Ngeru- keuid (island 24), Ngeruktabel, Ngetme- duch, Oreor, Ulebsechel, Ulong. REPTILIA: SAURIA Agamidae Hypsilurus godeffroyi Peters, 1867 Originally described from two speci- mens as Lophura (Hypsilurus) Godeffroyi from the "Pelew-Inseln" (Peters 1867b: 707, Fig. 1; lectotype ZMB 5891), this spe- cies has never been collected again any- 99] PETOLOGICAL OGRAPHS 42 HERPETOLOGICAL MONOGRAPHS [No. 13 where in Oceania. Boulenger (1885:295- 296) first associated the species name with populations from the Papuan region. In fact, he did not even include Palau in the distribution of the species, listing only "New Ireland, New Britain, Solomon and Fiji Islands, Northern Queensland". Cog- ger (1975:222) discounted the Australian records (and deleted the species from sub- sequent editions of his book) and listed the distribution as "Widely distributed . . . from the Solomon Islands to the Philip- pines." However, the species (and the ge- nus) has never actually been reported from the Philippines and Cogger (personal com- munication 1997) confirms that "Philip- pines" was probably a lapsus for Palau. Zug (1991:7-8) regarded the Fijian locality as erroneous. The status of this lizard in Palau is prob- lematic. Palauans are utterly unfamiliar with photographs of this large and distinc- tive species. However, McCoy (1980:14) emphasized that, in the Solomons at least, H. godeffroyi can be amazingly cryptic, even where it is common. The only recent evidence of the lizard's presence in Palau comes from a collection of skeletal re- mains found in a cave(s)/gravel pit (now flooded) on Ngeaur by Tom Iliffe in March, 1985, and sent to us for examina- tion. The bones are clearly those of an agamid, and represent a minimum of 11 individuals ranging in approximate snout- vent length of 70-110 mm; McCoy gives an average SVL of 120 mm for Solomon Island godeffroyi. Iliffe recovered the bones along with mollusk shells while sam- pling for archaeological artifacts. The bones show no evidence of deterioration, and appear much like macerated skele- tons, suggesting that they were deposited not too long before they were collected. We think it probable that this species still exists in Palau and it awaits rediscovery in some forested area. If and when additional material from Palau is collected, we be- lieve it equally probable that the lizard will prove not to be conspecific with the pop- ulations in the Solomons and New Guinea. Other possibilities are that the type lo- cality is erroneous. Peters' types were re- ceived from the Godeffroy collection, without a specific locality or collector. The Germans were also active in the New Guinea-Solomons area during this period and the specimens could have been mis- labelled, or Palau may have been where the specimens were shipped to Europe and the shipping port was erroneously in- terpreted as a collecting locality. However, this scenario seems less plausible in view of the bones collected from Ngeaur. Conceivably the species is a recent (i.e., post-European) extinction. It may have been exterminated by rat predation, per- haps coupled with habitat loss in the wake of deforestation, particularly during the Japanese period. Distribution. -unknown (Ngeaur?), probably endemic. Gekkonidae Gehyra sp. On first inspection, the large Palauan Gehyra seem simple taxonomically, i.e., the dark, velvety G. brevipalmata with di- vided terminal lamellae and yellow or or- ange ventral color, versus the pale, ghostly gray G. oceanica with transverse, undivid- ed lamellae and a gray (occasionally yel- low) belly. The reality is considerably more complex. Although the two species are usually easily identifiable in the field, even in areas of absolute syntopy (Ngaraard State, Babeldaob), some small island pop- ulations are more equivocal. One of us (RIC) misidentified the Ngerur population of oceanica as brevipalmata on his first vis- it, despite considerable experience with both species in the field. Judging from the number of misidenti- fied specimens in museum collections, preserved material is more confusing still. The terminal lamellae can appear divided in poorly preserved or dessicated speci- mens of G. oceanica. This apparently caused the misidentification of Saipan oceanica as Peropus papuanus in Dryden and Taylor (1969). Although we are reasonably confident that the Gehyra with transverse lamellae from the core islands of Palau are all oceanica (sensu lato), we are much less sure about the specimens from the South- west Islands. We have no experience in [ . 1999] HERPETOLOGICAL MONOGRAPHS 43 these islands, so no ecological or color data are available. The few specimens known (USNM 348770 from Fana; 348762-63 from Pulo Anna; 348781-82 from Sonso- rol; 348788 from Tobi) have unquestion- ably undivided lamellae, but somewhat more emarginate than is typical for ocean- ica (this may be an artifact of preservation as noted above). The males also have lon- ger preanal-femoral pore series, extending to mid thigh (restricted to the preanal area in Palauan oceanica). King and Horner (1989) described south-coast Papua New Guinea "oceanica" as Gehyra membrana- cruralis but the Southwest Islands speci- mens lack the extensive femoral mem- brane of that species. Perhaps the most pertinent comparisons are with Gehyra from the Moluccas (the most proximate landmass to the Southwest Islands), whence Rooij (1915) records both G. oceanica and marginata. Gehyra margin- ata is a large species with undivided (but slightly emarginate) digital lamellae, well distinguished from oceanica by the exten- sive skin folds along the thighs and the venterolateral body. We have seen no ac- tual Gehyra oceanica from the Moluccas but a series (USNM 237578-83) of ocean- ica-sized Gehyra from Halmahera has strongly divided digital lamellae, unlike any species previously reported from the Moluccas and equally distinct from the Southwest Islands taxon. Clearly the Gehyra "oceanica" from the southwest Pacific are in need of revision. Rather than refer the Southwest Islands material to this catch-all species, we prefer to call it Gehyra sp. to emphasize our im- pressions of its distinctiveness from "oceanica" on the core islands of Palau. Distribution.-Fana, Pulo Anna, Son- sorol, Tobi. Gehyra brevipalmata (Peters, 1874:159) The relationships of this distinctive Pa- lauan endemic [Hemidactylus (Peropus) brevipalmata Peters, 1874; Holotype ZMB7927] are not clear, but we suspect it may be close to some of the poorly known New Guinea species with divided lamellae (e.g., papuana, lampei, interstitialis, bal- iola (Rooij, 1915:38; Bauer, 1994)). Dry- den and Taylor (1969) reported Peropus papuanus (= Gehyra papuana) from Palau based on one specimen from Ngeaur (EHT-HMS = FMNH 6699), citing Bron- gersma's (1934) notes on the holotype of G. papuana as justification for their iden- tification. Brongersma (1934:174) stated that only the basal and terminal lamellae were entire, the rest divided, and that the tail was, "... much depressed ... with a sharp lateral edge." We have reexamined Dryden and Taylor's specimen, which is dessicated and tailless so Brongersma's characters are difficult or impossible to ap- ply (see also comments under Gehyra sp.). However, several of the proximal lamellae (not just the basal ones) appear to be un- divided, similar to the digital structure of G. brevipalmata. King and Horner (1989) characterized G. papuana as a medium- sized species (less than 80 mm SVL) re- stricted to New Guinea. We have collected extensively on Ngeaur, finding only G. brevipalmata and G. mutilata. We are con- fident that Dryden and Taylor's specimen is a misidentified brevipalmata (a species they did not mention), as suggested by Bauer (1994:100). Gehyra brevipalmata is likely one of the "old" Palauan species. It occurs widely throughout the archipelago including for- est on Rock Islands to buildings in dis- turbed areas. Its rich, dark, velvety skin is remarkably fragile and undamaged speci- mens are rare. When grasped, G. brevi- palmata spins and any restraint causes the skin to peel back from the subdermal fas- cia and a completely skinless specimen can result. Gehyra brevipalmata is particularly fond of crevices and the species is found commonly on banyan trees, in Pandanus axils, under bamboo sheaths, and in man- made crevices around buildings (under shutters, behind drain pipes, in corrugated tin roofs). In captivity, the species eats fruit readily and in the wild we have seen it licking at sweets (tree sap, open soft drink cans and bottles). Crombie and Steadman (1987) reported that the stom- achs of several Gehyra oceanica from the Cook Islands were "packed with small seeds" similar to fig, and fruit eating has ] PETOLOGICAL OGRAPHS 44 HERPETOLOGICAL MONOGRAPHS rNo. 13 FIG. 5.-The distribution of Gehyra mutilata in Oceania. (Also occurs in India, Sri Lanka, Indochina, Philippines, Japan, Madacasgar; after Bauer, 1994). been reported for several other species of Gehyra (see Burnett and Nolen, 1996). Distribution.-ENDEMIC. Babeldaob, Bablomekang, Beliliou, Euidelchol, liblau (Ngemlis Islands) Kmekumer Islands, Ma- lakal, Ngeaur, Ngercheu, Ngerduais, Nger- ekebesang, Ngeruktabel, Oreor, Pkulak- lim, Ulebsechel, Ulong. Gehyra mutilata (Wiegmann, 1834) This gecko is very widely distributed in mainland south and southeast Asia, and numerous island groups including Mada- gascar and other Indian Ocean groups, the Indoaustralian Archipelago, the Philippi- nes, and most of Oceania (Fig. 5; Bauer, 1994:92, map 24). Its broad distribution is usually attributed to human transport. Some populations (the Neotropics, New Zealand) are clearly post-European intro- ductions, but the species probably dis- persed easily with indigenous peoples through much of Oceania. Peters (1867a) discussed a "variety" of Gehyra mutilata based on a single 60 mm specimen collected in the "Pelew-Inseln" by C. Semper which he distinguished from typical mutilata by the more extensive dig- ital webbing and the small outer postmen- tal scales. This was a subadult of what he later described as Gehyra brevipalmata (Peters, 1874). This species is ubiquitous in Palau and is found in a wide variety of habitats. In the absence of Hemidactylus frenatus it is often the commonest house lizard, but it is also found in more natural habitats in- cluding undisturbed forest. Distribution.-Babeldaob, Ioulome- kang, Malakal, Ngcheangel Atoll (Ngche- angel), Ngeaur, Ngerechong, Ngereke- besang, Ngetmeduch, Oreor, Tobi (Southwest Islands). Gehyra oceanica (Lesson, 1826) Although King and Horner (1989) found Oceanic populations (Fig. 6) of Ge- hyra oceanica to be homogeneous both morphologically and karyologically, Beck- on (1992) was able to distinguish morpho- logically Micronesian from Polynesian oceanica. Biochemical data confirm the differences between the two populations (R. Fisher, 1997), although no one has yet attempted to sort out the confusing syn- onymy of oceanica to determine which names apply to the two species. Gehyra oceanica has been recorded from Oreor (CAS-SU 23747, 23753, CAS 122393), but we have collected it only in PETOLOGICAL GRAPHS [ o. 19991 HERPETOLOGICAL MONOGRAPHS 45 FIG. 6.-The distribution of Gehyra oceanica (after Bauer, 1994). the extreme north of Babeldaob and on peripheral small islands (e.g., Ngerur). We have not verified its presence on Oreor de- spite intensive effort. In Nagaraard State (northern Babeldaob) it occurs syntopical- ly (often on adjacent trees) with Gehyra brevipalmata in Ulimang Village, but brevipalmata seems to be slightly more abundant. Distribution.-Babeldaob, Chemoi, Ngcheangel Atoll (all islands), Ngeanges, Ngerechur, Ngerur, Oreor. Gekko sp. This large, attenuate gekkonid is an un- described endemic species. It is related to Gekko vittatus of New Guinea and the In- doaustralian Archipelago, and it is fre- quently identified as vittatus in the litera- ture and in museum collections. New Guinea/Indonesian Gekko vittatus are ex- tremely variable and more than one spe- cies is certainly included in vittatus as cur- rently defined. The nomenclatural history of vittatus and its synonyms is plagued by erroneous type localities, missing type specimens, and other problems that pre- clude resolution with currently available data and specimens. A brief summary as it relates to the Palau Gekko is warranted. Basically, Gekko vittatus (in New Guinea and the Solomons at least, data for the In- donesian populations are almost totally lacking) is dimorphic in color pattern, but not sexually correlated (McCoy, 1980: plate 7a,b illustrates both forms). The most common morph has a vivid light ver- tebral stripe outlined in black, and a dis- tinctly banded tail. A more unicolor, ob- scurely striped lizard is often called Gekko vittatus bivittatus (e.g., Boulenger, 1885), but Mertens (1926) pointed out that both morphs occurred at several places in the New Guinea region, and he referred to bivittatus as a "Mutation" rather than a recognizable taxon. Probably because it is obscurely striped, the Palau Gekko has occasionally been re- ferred to G. vittatus bivittatus (Boulenger, 1885; Dryden and Taylor, 1969). However, there are significant meristic and molecu- lar (C. Austin; H. Ota; R. Fisher, personal communication) differences between the unstriped Gekko from Palau and those from the New Guinea region. Although common in Palau, Gekko sp. is secretive and easily overlooked. It does inhabit houses, even in Koror town itself, but it usually haunts the darkest areas, rather than foraging around lights as many 1999] RPETOLOGICAL NOGRAPHS 46 HERPETOLOGICAL MONOGRAPHS [No. 13 FIG. 7.-The distribution of Hemidactylus frenatus in Oceania (after Bauer, 1994). of the smaller species of gekkonids do. More commonly it occurs on tree trunks and leaves, with little preference other than vegetation that provides crevices for diurnal retreats. Palms, Pandanus, ban- yans, and banana (wild or cultivated) are favored. Adults are found after dark on leaves, fronds, trunks or branches, appar- ently awaiting insect or vertebrate prey. Eggs are adherent, laid in crevices, and the clutch size is two. Palm, Pandanus, or ba- nana axils are common deposition sites but cracks in masonry and other man-made sites are also used. Incubation time is about 90 days and hatchlings are as ob- scurely striped as adults. We suspect that Gekko sp. is a major nocturnal predator in Palau, feeding on ar- thropod prey and small vertebrates. Their jaws are quite powerful, capable of inflict- ing a painful bite on human fingers often accompanied by a twisting motion that causes bloody lacerations. Subduing even sizable vertebrate prey would be well with- in their abilities. Captive individuals are highly territorial and aggressive; males will brutalize non-reproductive females and cannibalize juveniles. Fruit is also readily eaten in captivity. These behavioral char- acteristics are strikingly different from our experience with Papuan G. vittatus, which frequently are found in communal groups in the field and live well together in cap- tivity. Distribution.-ENDEMIC. Babeldaob, Bablomekang, Beliliou, Kmekumer Islands (island 46), Malakal, Ngeaur, Ngercheu, Ngerduais, Ngerechur, Ngerekebesang, Ngerukeuid Islands (island 24), Oreor, Ulebsechel, Ulong. Hemidactylus frenatus Dumeril and Bibron, 1836 This widespread Southeast Asian weed species, a quintessential "house lizard" (sensu Mertens 1928), is apparently a re- cent introduction to Palau (Fig. 7). None of the earlier collections included this highly visible human commensal, and the earliest record is a single specimen (CAS- SU 23728) collected near the Conserva- tion/Entomology Lab on Koror by Fehl- mann in July 1955. Dryden and Taylor (1969) next reported it from Ngeaur. When introduced into established com- munities of other gekkonids (native or in- troduced) H. frenatus is often successful in displacing other species from disturbed (lighted) habitats (Case et al., 1994; McCoid, 1996; Petren et al., 1993; Petren [No. 199 EPTLGCA OORPS4 FIG. 8.-The distribution of Hemiphyllodactylus typus in Oceania. (Also occurs in the Mascarenes, through- out southeast Asia including India, Indonesia, and Philippines; after Bauer, 1994.) and Case, 1996). In Palau H. frenatus has not completely displaced Lepidodactylus and Gehyra from buildings. On the Na- tional Museum building in Koror, Gehyra mutilata and two species of Lepidodacty- lus were collected in the mid-1990's, al- though H. frenatus was by far the most common lizard. Perhaps even more in- structive is the situation in Ulimang Vil- lage, Ngaraard State, in a section of north- ern Babeldaob that is not yet connected by road to the southern causeway-con- nected islands. During the time of our field work, Hemidactylus frenatus was al- most completely confined to the Evangel- ical Church, where it was slightly more common than Gehyra mutilata around the light at night. Despite our extensive field work in and around this community, we found only one other individual of H. fre- natus, a single male that vocalized fre- quently in the rafters of a recently con- structed house about one km south of the village. Although we often heard this in- dividual over a span of several years, we never saw it in the characteristic frenatus foraging area around the lights of the house, which were occupied exclusively by Lepidodactylus moestus and occasionally Gehyra brevipalmata. We finally caught several fleeting glimpses of him in shad- owy parts of the house. Although the house was largely constructed from local material, some lumber was brought in from Koror and we suspect this one male came with it. The other population around the church has not dispersed far enough to provide him a mate. Distribution.-Babeldaob, Beliliou, Ma- lakal, Ngeaur, Ngercheu, Ngerekebesang, Ngermalk, Oreor, Uchulangas. Hemiphyllodactylus cf. H. typus Bleeker, 1860 This cryptic, easily-overlooked small species was previously unreported from Palau other than Bauer's (1994:131, map 29) inclusion of Palau in the distribution of H. typus (Fig. 8). The only other Mi- cronesian record (Lamberson, 1987 from Enewetak, Marshall Islands; also mapped in Bauer, 1994) is based on a misidentifi- cation (USNM 205534-35 = Lepidodac- tylus sp.). Although all other Oceanic pop- ulations are all-female, males are present in Palau. This complex of species is under study by G. R. Zug and we defer any tax- onomic conclusions pending completion of his analysis. Hemiphyllodactylus is not a "house liz- 1999] H RPETOLOGICAL M NOGRAPHS 47 48 HERPETOLOGICAL MONOGRAPHS [No. 13 ard" (sensu Mertens, 1928), but it has been found near houses in more sparsely populated areas. When active after dark, the species has been collected on attenu- ate leaves of shrubs (including palm fronds), on small and large trees, and on debris around houses. We also collected it in primary forest (Ngeruktabel). Distribution.-Babeldaob, Ngeanges, Ngeaur, Ngerekebesang, Ngeruktabel, Or- eor, Ulebsechel. Lepidodactylus sp. Bisexual, possibly parental species of the widespread parthenospecies Lepidodacty- lus lugubris are known from Micronesia. One distinctively patterned bisexual spe- cies (Radtkey et al., 1995: plate 1) is known from Takapoto in the Tuamotus and Arno Atoll in the Marshalls, but it has never been reported from Palau. We col- lected one male (USNM 531970, pre- served with everted hemipenes) low on a palm trunk at the high tide line on Ulong that is similar to the illustration in Radtkey et al. (1995). However, two other speci- mens (USNM 531971-72) collected sub- sequently demonstrate to us that it is not conspecific with the Takapoto/Amo popu- lations. Distribution.-Iilblau (Ngemlis Is- lands), Ngeruktabel, Ulong. Lepidodactylus lugubris (Dumeril and Bibron, 1836) This widespread, all-female, partheno- genetic species has been intensively stud- ied (Bolger and Case, 1994; Ineich, 1988; Ineich and Ota, 1992; Radkey et al., 1995, 1996) and several distinct species are cer- tainly included under this name. The spe- cies seems to be a relatively recent arrival in Palau, regardless of the name applied to it. It occupies the most disturbed areas, often syntopic with L. moestus, with which it may hybridize, producing puzzlingly in- termediate specimens (Ota et al., 1995). As noted in the account of Hemidacty- lusfrenatus, this species continues to share the building habitat with frenatus in many areas of Palau. Distribution.-Babeldaob, Beliliou, loulomekang, Kmekumer Islands, Malak- al, Ngeanges, Ngeaur, Ngercheu, Ngerdis, Ngerekebesang, Ngermalk, Ngerur, Nges- bokel, Ngetmeduch, Oreor, Pulo Anna, Uchulangas. Lepidodactylus moestus (Peters, 1867:13) Long mired in the synonymy of the catch-all Lepidodactylus lugubris, this spe- cies (Gecko moestus Peters, 1867; ZMB5610) was recently resurrected by Ota et al. (1995). It is undoubtedly a native species in Palau, but it also occurs in Mi- cronesia as far east as Arno Atoll in the Marshalls (Fig. 9). Lepidodactylus moestus is very widely distributed in Palau includ- ing both disturbed and natural areas. It does quite well even in downtown Koror, surviving on buildings despite the abun- dance of Hemidactylusfrenatus. It also co- exists with L. lugubris and the two species are almost equally common on the picnic pavillion on the Museum grounds. In se- verely disturbed areas (e.g., Klouklubed village on Beliliou), lugubris may be more abundant but we have found only a few instances where this is so. Accumulating evidence (e.g., Radtkey et al., 1995, 1996) strongly suggests that moestus is one of the sexual species that gave rise to the parthe- nogenetic lugubris. Lepidodactylus moes- tus is an adaptable, aggressive, and vocal species that can survive in some surpris- ingly harsh environments. We have col- lected the species on the smallest (un- named) island at the mouth of Ngeremeduu Bay (Ngatpang State, west- ern Babeldaob), a pinnacle barely 4 m2 that supported one Pandanus tree. One adult moestus (USNM531969) and a clutch of viable eggs were found in the Pandanus axil. Radtkey et al. (1995) illustrated the two distinct morphs of moestus: plain and striped. Robert Fisher (personal commu- nication) insists that these morphs are ge- netically identical. Distribution.-Babeldaob, Kmekumer Islands (island 46), Malakal, Ngcheangel Atoll (all islands), Neanges, Ngeaur, Nger- cheu, Ngerduais, Ngerechur, Ngerekebes- ang, Ngermalk, Ngerukeuid Islands (island 4), Ngeruktabel, Ngerur, Ngesbokel, Nget- [ . 1999] HERPETOLOGICAL MONOGRAPHS 49 FIG. 9.-The distribution of Lepidodactyus moestus. meduch, Oreor, Pulo Anna, del, Uchulangas. Uchulamera- Lepidodactylus paurolepis Ota et al., 1995 This recently described species remains known only from the small type series col- lected in the Ngerukeuid Islands. Al- though it is a very distinctive gecko, it would be most surprising if it was indeed restricted to this isolated cluster of small islands. We suspect it will be found on oth- er islands in the Palau lagoon. Distribution.-ENDEMIC. Ngeru- keuid Islands (islands 16, 34) Nactus sp. Prior to our field work the apparent ab- sence of Nactus from Palau was enigmatic because the genus is widespread in the Papuan region and southwest Oceania (see map 40 in Bauer 1994:169). This absence was simply an artifact of inadequate col- lecting. Two species are present in the ar- chipelago (see also Nactus cf. N. pelagi- cus). Nactus sp. is a bisexual gecko that we found only on Ulong (one adult male) and Ulebsechel (two hatchlings), but we sus- pect that its distribution is more extensive. Unlike members of the Nactus pelagicus complex, which are often abundant in beach habitats, this species is very cryptic and secretive. The adult male was found on a tree in scrubby forest just behind the beach, and adjacent to a limestone cliff. No other individuals were seen in 4 hours of collecting that evening, and ca. 30 hours (day and night) subsequently. The two hatchlings from Ulebsechel were found beneath ground litter, one near the beach and the other at the base of a limestone ridge inland. Resolution of the systematics of the Nactus pelagicus complex is still in the preliminary stages (see summary in Bauer, 1994:168; Zug, 1998; Zug and Moon, 1995), and this species appears to be un- described (G. R. Zug, personal communi- cation). More specimens and data are needed to confirm the status and distri- bution of the species in Palau. Distribution.-Ulebsechel, Ulong. Nactus cf. N. pelagicus (Girard, 1858) A single specimen (USNM 348780) from Merir in the Southwest Island Group was referred to N. cf. pelagicus by G. R. Zug (personal communication). The pelag- icus group (Fig. 10) is a complex of bisex- ual and unisexual, parthenogenetic species (see summary in Bauer, 1994:169) but the alpha taxonomy is not yet stabilized. Zug and Moon (1995) recently resurrected N. 1999] 50 HERPETOLOGICAL MONOGRAPHS [No. 13 FIG. 10.-The distribution of the Nactus pelagicus species group (after Bauer, 1994). multicarinatus for the bisexual species of Vanuatu and the southern Solomons, but the situation on New Guinea itself is much more complex. Larger sample sizes from the Southwest Islands are needed to de- termine if they are indeed conspecific with the unisexual pelagicus. The Merir speci- men is a female. Distribution.-Merir. Perochirus sp. The genus Perochirus is a Pacific insular endemic, consisting of three species from scattered island groups in Micronesia and Vanuatu (Brown, 1976), and an extinct form from 'Eua, Tonga (Pregill, 1993). The most widespread species, Perochirus ate- les, occurs close to Palau in the Carolines, Marianas, and Marshalls (Bauer, 1994: 208, map 49). Although Bauer (1994:283) recorded Perochirus ateles from Palau with a question mark, there are no docu- mented records of this species from Palau. A recent collection from the Southwest Islands contains two specimens (USNM 348771-72) of a large Perochirus from Fana. Their proportions and scalation in- vite comparison with Perochirus scutella- tus. Until recently, Perochirus scutellatus was a very poorly known species repre- sented by only a few older specimens from Kapingamarangi Atoll, located more than 2000 km SSE of Palau. In the summer of 1996, Donald W. Buden collected a large series of scutellatus from Kapingamarangi (Buden, 1998a,b). We have not yet had the opportunity to examine these specimens in detail, and the limited material from Palau may be inadequate to demonstrate con- specificity in any case. If the Palau speci- mens are scutellatus the enormous distri- butional hiatus could be explained as the relictual distribution of an old radiation created by local extinctions on the inter- vening islands. Both specimens of Perochirus from Fana were collected on trees. Interesting- ly, no Gekko were collected from any of the Southwest Islands. Gekko is normally a conspicuous species where it occurs, and Perochirus may replace it ecologically in the Southwest Islands. Distribution.-Fana. Polychrotidae Anolis carolinensis Voigt, 1832 This species has been introduced in sev- eral areas of the Pacific, but historical in- formation on the introductions is lacking. Some authors have suggested that the [ . 1999] HERPETOLOGICAL MONOGRAPHS 5' Oceanic Anolis are Cuban in origin (Anolis porcatus or Anolis carolinensis porcatus; Oliver and Shaw, 1953; Vance, 1987) but we agree with McKeown (1996) that Pa- cific populations (including Palau) are typ- ical of A. carolinensis of the southeastern United States. Although Anolis carolinensis has been in Palau for at least 30 years (Owen, 1977), it has not dispersed far from the presumed place of entry, the Port of Palau on Ma- lakal. Robert Owen (personal communi- cation to RIC in 1980) said that the lizards were restricted to the immediate vicinity of the Port and adjacent docks at that time, and their distribution has changed little since then. Our field work 15 years later indicates that the species is now found 50-100 meters east and west of the Port along the road, but it is not abundant. No obvious dispersal barriers confront this species on Malakal, and there is abundant disturbed habitat all over Malakal and ad- jacent Oreor. Our herpetological surveys and transects have covered Malakal from the top of the hill to sea level in several places from 1993-1997, but yielded no specimens or even sight records of Anolis from anywhere other than the narrow coastal area on the west side of the island. Then in July 1996, Chris Austin and RIC collected an adult male in thick, scrubby vegetation in the agricultural area on "Ra- dio Tower Hill", just below the band of forest around the peak, a site we had sam- pled frequently before. The species was also abundant around the newly construct- ed Coral Reef Research Foundation build- ing at the Malakal end of the causeway across to Koror, and in August 1998 RIC found the anole on the edge of Koror town itself. After a long sedentary period, Anolis may finally be dispersing in Palau; it will be interesting to watch its progress. We suspect that Lamprolepis smarag- dina may be somewhat of a controlling agent in the spread of Anolis carolinensis. The two species often occupy the same habitat, and Palauan A. carolinensis is un- characteristically secretive compared to other natural and introduced anole popu- lations we have seen. We have only one direct observation of interactions between Lamprolepis and Anolis, which involved a subadult Anolis we were pursuing in a viney tangle near the Port. The anole at- tempted to escape by leaping to the ground but had barely taken a few steps before an adult male Lamprolepis dashed down the trunk of a nearby sapling and seized it. Distribution.-Malakal, Oreor. Scincidae Carlia cf. C. fusca (Gray, 1845) Despite its abundance in most disturbed areas in Palau, Carlia seems to be a very recent introduction to the islands. The or- igin and taxonomic status of the propagule remains unclear. Carlia has never been re- corded in the literature from Palau and the earliest known specimens were collected by Tom Fritts in 1988, but we are confi- dent that the species was introduced be- fore then. Both the Marianas and Palauan popu- lations of Carlia are clearly members of the widespread and taxonomically unre- solved fusca group, and the logical as- sumption would be that the two areas were synchronously colonized from the same source. However, females and juve- niles in Palau often have strong dorsolat- eral striping, which is rare or nonexistent in the Marianas population. Our impres- sion is that Palau and Marianas Carlia had different origins and may be different spe- cies, neither of which is true 'fusca". Carlia is the dominant terrestrial skink on the causeway-connected islands of Pa- lau, occurring in very high densities in all the disturbed areas of southern Babel- daob, Oreor, and the associated smaller is- lands, but they are not (yet) on any of the adjacent Rock Islands. The more distant islands of Ngeaur and Beliliou have healthy populations of Carlia, but here also the lizards do not penetrate forested areas and are restricted to open grassland or the vicinity of human habitation. The parts of northern Babeldaob not connect- ed by road to the south may be Carlia-free (at least at the moment). Our extensive ex- perience in the Ulimang-Ngesang area yielded a few "possible" Carlia sightings in 1994 and again in 1997, but no vouchers. 9] 1 52 HERPETOLOGICAL MONOGRAPHS [No. 13 Juvenile Mabuya are superficially similar to striped female and juvenile Carlia and difficult to distinguish when they are on the move. Extensive sticky-trapping (40+ trap-hours) in and around Ulimang village in August 1998 yielded many Mabuya but no Carlia. Distribution.-Babeldaob, Beliliou, Ma- lakal, Ngeaur, Ngerekebesang, Ngermalk, Ngetmeduch, Oreor, Uchulangas. Cryptoblepharus sp. Mertens (1931, 1933, 1934b, 1964) re- garded Ablepharus (= Cryptoblepharus) boutonii as a polytypic species with more than 40 subspecies ranging from the In- dian Ocean coast of Africa to the eastern limits of the Pacific. Crombie and Stead- man (1987) cited accumulating evidence that many of these "subspecies" are valid species, and they suggested that the name Cryptoblepharus poecilopleurus was appli- cable to most of the Oceanic populations. Fiji, Vanuatu, and New Caledonia all have distinct species of Cryptoblepharus (Sad- lier, 1986; Zug, 1991 and personal com- munication) but the other Pacific Rim populations have not yet been studied in detail. Peters (1879:37) described the Pa- lauan Cryptoblepharus as Ablepharus ru- tilus (holotype ZMB 7926) and we suspect that it may be a valid species. Both George Zug and Paul Horner are examining vari- ation in Cryptoblepharus and we have made our material available to them to be analysed and compared with broader geo- graphic samples. In many parts of its range Cryptoble- pharus is a beach species, occurring on rocks and wrack in the supratidal zone (e.g., McCoid et al., 1995). Palauan Cryp- toblepharus are much more arboreal, of- ten found well away from the beach on tree trunks, vines and lianas (e.g., USNM 531953-56 from central Oreor). The spe- cies does occur as well in beach situations, frequently on large trees hiding behind ex- foliating bark. On northern Babeldaob more strongly striped (poecilopleurus-like) Cryptoblepharus were seen, but not col- lected, on aerial roots of mangroves. This particular locality has since been destroyed by dredging. More than one species of Cryptoblepharus may be present in Palau but presently available material is inade- quate for a determination. Distribution.-Babeldaob, Beliliou, Iil- blau (Ngemlis Islands), Kmekumer Islands (island 46), Ngeaur, Ngebad, Ngerekebes- ang, Ngetmeduch, Oreor. Emoia atrocostata (Lesson, 1826) This variable "superspecies" (sensu Brown, 1991) occurs from Taiwan and the Ryukyus to New Guinea, the Moluccas, and the southern Malay Peninsula includ- ing Christmas Island in the Indian Ocean, and western Micronesia (Fig. 11). Brown (1991) recognized subspecies from Cape York, Australia (australis) and the Solo- mons/Vanuatu area (freycineti). We agree with Brown that several taxa are probably included in his nominate atrocostata, and that the Micronesian populations are rea- sonably distinctive. Throughout its broad distribution, Emoia atrocostata invariably inhabits rocky beaches and mangrove swamps as report- ed by Alcala and Brown (1967) in the Phil- ippines. Some populations are associated with one habitat over the other, but it is currently unclear if this is merely sampling bias or an ecological preference with sys- tematic implications. In Palau, Emoia atrocostata seems to have a spotty distribution but we are con- fident that this reflects the difficulty of col- lecting individuals amid the multitude of refuges on rocky beaches and in mangrove swamps, rather than actual rarity or a re- stricted distribution. For example, we have seen atrocostata commonly on beach boul- ders near the KB Bridge on Oreor but we have not yet been successful in collecting them there. In other cases something more complex is involved. Near the north- em tip of Babeldaob in Ngaraard State, the species is abundant and easily ob- served in mangrove swamp near the dock west of Ngebuked on the west side of the peninsula. However, in similar swamps south of Ulimang on the east side (less than 3 km airline) not a single atrocostata has been seen in more than 40 person/ hours of searching. The species does occur on the east side of Babeldaob. A single [ . 199 HEPEOLGIA MNORAHS5 FIG. 11.-The distribution of Emoia atrocostata. subadult (USNM 531952) was collected under beach wrack at the high tide line directly in front of our base of operations at Ulimang in February 1996. Six other in- dividuals (USNM 495091-95) also were obtained at Melekeok by Gordon Rodda and Tom Fritts. Although usually charac- terized as a coastal species, Emoia atro- costata can extend well inland along riv- erine mangroves, as documented by USNM 495090 from the Tabecheding Riv- er at Nekkeng, Aimeliik State, Babeldaob. Distribution.-Babeldaob, Beliliou, Ma- lakal, Ngeaur, Ngebad, Ngerdis, Ngerek- ebesang, Ngetmeduch, Oreor, Uchulangas, Ulebsechel. Emoia caeruleocauda (De Vis, 1892) Brown (1991) did not fully analyze var- iation in the "superspecies" Emoia caeru- leocauda. However, many Palau specimens in museum collections are identified as Emoia "pelewensis", apparently an early Brown manuscript name that was never published. Given the broad distribution (Fig. 12) and extensive variation of caeru- leocauda, it is not surprising that Brown (1991) deferred in trying to resolve all the taxonomic problems of this taxon. Palauan caeruleocauda are distinctive in a number of ways. The unicolor ("bronze" morph of Crombie and Steadman, 1987) males in Palau have a profusion of tur- quoise or yellow lateral flecks that are lack- ing in other populations of caeruleocauda that we are familiar with. We feel that Brown's earlier impression was correct, and that additional analysis will demon- strate that Palauan E. caeruleocauda are indeed specifically distinct. Although this species is easily displaced by Carlia in disturbed habitats, it persists in backyard garden plots and other shaded, moist areas even in Koror town itself. The lizard is primarily terrestrial, but juveniles and subadults in particular frequently as- cend low vegetation. Distribution.-Babeldaob, Beliliou, Eu- idelchol, Iilblau (Ngemlis Islands) loulo- mekang, Malakal, Mecherchar, Ngchean- gel Atoll (Ngcheangel, Ngerebelas, Orak), Ngeanges, Ngeaur, Ngebad, Ngebedangel, Ngercheu, Ngerechur, Ngerekebesang, Ngeruchubtang, Ngeruktabel, Ngerur, Ngetmeduch, Oreor, Pkulaklim, Ulebse- chel, Ulong. Emoia impar (Werner, 1898) Originally described from 12 specimens collected on Ralum and Mioko Islands in the Bismarck Archipelago, this species was long considered a synonym of Emoia cy- anura (e.g., Brown, 1991). It is now clear that impar is a distinct species (Ineich and 53 9] RPETOLOGICAL ONOGRAPHS 54 HERPETOLOGICAL MONOGRAPHS [No. 13 FIG. 12.-The distribution of Emoia caeruleocauda "superspecies". Zug, 1991; Guillaume et al., 1994) and some data indicate that even more cryptic sibling species may be recognizable within the broad Pacific distributions of both cy- anura and impar (Bruna et al., 1996). The apparent absence of the ubiquitous cyanura superspecies (sensu Brown, 1991) from the Palau Group was puzzling. How- ever, at least E. impar is present in Palau, albeit with a strange distribution, parallel- ing in some ways that of E. jakati. Why this odd, disjunct distribution appears in two distinct lineages of Emoia is perplex- ing, but we suspect that both impar and jakati are relatively recent adventives to Palau, probably via natural dispersal or ac- cidental transport by Micronesians rather than as post-European introductions. The Palau populations of Emoia impar were apparently restricted to small and pe- ripheral islands at the northern (Ngchean- gel Atoll) and southern (Southwest Is- lands) extremes of the Republic. The species had never been found between those two points (a distance of more than 400 km) until 1996, when Chris Austin and RIC collected a small series on Nges- bokel, the largest of 4-5 tiny islets at the mouth of Ngeremeduu Bay, Ngatpang State, western Babeldaob. Some of these islands were barely emergent at high tide but connected by broad tidal flats at low tide. Ngesbokel has a few hectares of flat plateau 2-3 m above the high tide line. The vegetation is a mixture of thick grass- es, stunted shrubs and scattered Panda- nus. Although we saw a few striped skinks on the plateau, we were unable to verify their identity in the thick vegetation. How- ever, in the wrack above the high tide line in a shaded cove that provides the only access to the plateau, we found Emoia im- par to be abundant at low tide, veritably swarming over driftwood and escaping into the low vegetation on the slope lead- ing up to the plateau. We estimated there were at least a dozen individuals in the small area. On some of the Rock Islands we have found Emoia caeruleocauda in very similar situations. Emoia impar is peripheral on Ngchean- gel Atoll as well, occurring only on the two smallest islands, sympatric with E. caeru- leocauda on Ngerebelas and Orak. The Southwest Islands have been only inciden- tally collected so nothing can be said other than to note that the species occurs on three of the islands. Distribution.-Ngcheangel Atoll (Nger- ebelas, Orak), Ngesbokel, Merir, Pulo Anna, Sonsorol. Emoia jakati (Kopstein, 1926) Although this species is widespread in the Carolines (Fig. 13), often occurring [No. 199IEPTLGCA OORPS5 FIG. 13.-The distribution of Emoiajakati. sympatrically with members of the Emoia caeruleocauda and cyanura "superspecies" (sensu Brown, 1991), its distribution in Pa- lau is decidedly strange. Brown's (1991) treatment of jakati does not lend confi- dence that all populations he referred to that taxon are indeed conspecific. In fact, when Loveridge (1945) examined museum collections for his "Reptiles of the Pacific World" he left many specimens identified with the (still unpublished) manuscript name Emoia mivarti 'fuscolineata," indi- cating that he regarded the Oceanic pop- ulations as distinct from New Guinea "mi- varti," a name misapplied to jakati until Brown's revision. Like Emoia impar, E. jakati was previ- ously known only from Ngcheangel Atoll and the Southwest Islands, but nowhere in between. However, in 1996 Chris Austin and RIC found the species abundant on Ngerechur, the northernmost of two small islands off the northern tip of Babeldaob. At low tide the channel between Ngere- chur and Ngerkeklau is wadable and that between Ngerkeklau and the Babeldaob coast (about 0.5 km) is even shallower. The coastal areas of extreme northern Babel- daob have never been collected herpeto- logically, nor has Ngerkeklau, and E. jakati may indeed occur at either or both, but currently the only record for the species within the fringing reef of Palau is Nger- echur. Based on our experience elsewhere in Micronesia (Pohnpei and surrounding is- lands), Emoiajakati is an inhabitant of the most disturbed and marginal habitats, in- cluding around human habitation. The Ngerechur population confirms this im- pression since the species is extremely abundant around an abandoned house at the south end of the island, an area of mixed strand vegetation and tall Cocos. The northern end of the island is higher, boulder-strewn and well forested, with much evidence of substantial Japanese oc- cupation during WWII. We did not see E. jakati anywhere in the forest. Picknickers and overnight visitors often use the house on the southern end, but there are no per- manent residents at this time. Distribution.-Ncheangel Atoll (Ngche- angel), Helen, Ngerechur, Tobi. Eugongylus sp. Dryden and Taylor (1969) first recorded this large scincid from Palau based on a single specimen from Ngeaur. They re- ferred it to Eugongylus mentovarius, a poorly known species previously reported only from the Moluccas. Based on our pre- liminary comparisons of Palauan Eugon- gylus with one Moluccan E. mentovarius HERPETOLOGICAL M NOGRAPHS 55 199 1 56 HERPETOLOGICAL MONOGRAPHS [No. 13~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ (USNM 237428), we do not consider them conspecific. However, the systematic rela- tionships and content of the genus Eugon- gylus are so chaotic that we defer a deci- sion at this time, but offer the following taxonomic summary of the genus as it cur- rently exists. Despite the contributions of B6hme (1976) and Greer (1974), species limits, relationships, and even the number of spe- cies in Eugongylus (including the "genera" Phoboscincus and Tachygyia) remain largely unresolved. For example, Bohme (1976) did not even mention mentovarius in his list of the recognized species in the genus. He did, however, include sulaensis (Kopstein, 1927) which was distinguished from mentovarius only in lacking the char- acteristic throat markings and in having a shorter tail. Most recently, Sadlier (1986) tacitly redefined Eugongylus by describing a new genus for haraldmeieri (Geoscincus) and recognizing two species of Phoboscin- cus. In so doing, he inferred that Tachy- gyia is also a valid taxon. Eugongylus has been infrequently col- lected in Palau but it seems to be a com- mon, if cryptic, terrestrial skink in a wide variety of habitats. Specimens have been taken in primary forest and mature second growth on rock islands (Wiles and Conry, 1990; our surveys), in backyard gardens in Koror town, and in scrubby forest adjacent to mangrove swamps (our surveys). We found three individuals actively foraging and most of the sticky-trapped individuals (see below) were obviously also active. One specimen (USNM531958) was mov- ing swiftly and silently with an almost ser- pentine motion through thick ground cov- er on the Museum grounds (Koror town) at 0915 h. Two others (USNM 531961-62) from Ulong were collected in the leaf litter in forest on the first terrace at 1415 h, and near the buttress of a large tree near the beach at 0710 h during torrential rain. In- active individuals were found from 1430- 1510 h under logs and in moist piles of coconut husks in beach forest on Babel- daob (Ngesang and Ulimang, Ngaraard State). The secretive habits of Eugongylus give the impression of rarity, but sticky traps are an effective collecting technique that yields specimens even in areas where the species has not been recorded in visual surveys. Wiles and Conry (1990) first not- ed this and our collections amply confirm their observations. Eugongylus are so ef- fective at avoiding detection that most Pa- lauans are incredulous that such a sizable, unfamiliar lizard was collected in their backyard. Sticky traps also yield some in- teresting natural history information on Eugongylus. On at least three occasions we have trapped Eugongylus that clearly had been attracted to smaller skinks (Car- lia and Emoia caeruleocauda) stuck in the trap. Two Emoia and one Carlia stuck near the edge of the trap were neatly sawed off just above the adherent part of their bod- ies, and in another instance a Eugongylus that became stuck trying to reach a Carlia in the center of the trap regurgitated the missing pieces of smaller skinks when he was being removed from the trap. Another Eugongylus was completely stuck along the long axis of the trap with an Emoia at the far end still in his jaws, thoroughly masticated at midbody. When Eugongylus are partially stuck they tend to spin, croc- odile-like, causing the entire trap to wrap around them burrito-fashion. On at least three other occasions we ob- served Eugongylus that did not get stuck trying to pull adherent Emoia or Carlia off the trap, often escaping with a tail or piece of body. We think these data clearly point to Eugongylus being an active, primary predator in the leaf litter community, probably feeding on a wide variety of in- vertebrates and smaller vertebrates. We would not be surprised if Platymantis, Sphenomorphus, typhlopid snakes, and even juvenile Candoia and Dendrelaphis are included in the diet. Distribution. -Babeldaob, Beliliou, Kmekumer Islands (island 46), Ngchean- gel Atoll (Ngcheangel), Ngeaur, Ngercheu, Ngerukeuid Islands (islands 16, 24), Nge- ruktabel, Oreor, Ulebsechel, Ulong. Lamprolepis smaragdina (Lesson, 1826) This widespread western Pacific species was divided into 8 subspecies by Mertens (1929), reflecting the extensive variation 56 [No. 199IEPTLGCA OORPS5 FIG. 14.-The distribution of Lamprolepis smaragdina. throughout its range (Fig. 14). Dryden and Taylor (1969) referred Palauan specimens to the subspecies viridipunctum, but we prefer to retain the binomial pending a more rigorous analysis of variation in the species. Mertens' subspecific arrangement does not accurately reflect the inter- and intra-island variation in Oceania and we suspect that several recognizable species may be subsumed in Mertens' concept of "smaragdina". The large, green tree skink may be the most familiar reptile in Palau, as it often perches conspicuously during the day on large backyard trees, even in Koror town. It is a remarkably adaptable species that also occurs in forest, but we have only a few records from true forest habitat. Per- haps the species is simply more difficult to observe in forest, or it may truly be less abundant there. Our impression is that Lamprolepis is an "edge" species, most suited to the tangle of vegetation at the forest margin. There is considerable vari- ation in habitat preference within the "species", however. Some populations or subspecies (e.g., the Philippine philippin- ica) are more forest-adapted and crypti- cally colored (brown body and dull green head) versus small island Oceanic smarag- dina, which are unmarked, almost electric green, and highly prominent on exposed Cocos trunks. Palauan Lamprolepis are usually bright green with varying amounts of enamel black on the dorsal scale edges. Addition- ally, there is what we have called the "mus- tard morph" which is most abundant in (or possibly restricted to) northern Babeldaob. As the name suggests, "mustard morph" individuals are dull yellow-brown above and lemon yellow below, without a hint of the usual greens. Normal green specimens have been found on trees adjacent to "mustard" perch sites and the morph is not sexually correlated (males, females, and juveniles with this coloration have been collected). We suspect that Lamprolepis is a major predator on small organisms in Palau. It has a broad activity period, forages from the ground to the canopy, and occurs in every habitat that supports trees, even low, scrubby Pandanus on tiny islets. It feeds on a wide variety of invertebrates and on any vertebrate it can subdue, including geckos and skinks (see also Anolis ac- count). Fruit and other sweets are also consumed. We observed Lamprolepis on Ngeaur gorging on ripe papaya. In Uli- mang, northern Babeldaob, which sup- ports a particularly dense population, HERPETOLOGICAL M NOGRAPHS 57 1999] 58 HERPETOLOGICAL MONOGRAPHS [No 13 Lamprolepis regularly came onto the porch of our house to lick residual soft- drink from the lips of cans. Manthey (1985), working mostly with Philippine in- dividuals in captivity, confirmed the spe- cies' fondness for sweets and he even re- ported egg-eating. Reproduction also reflects the broad ecological tolerance of Lamprolepis. Al- though two elongate, leathery eggs are in- variable (at least in Palau), we have found clutches under rocks, inside dead logs, and in the duff of palm axils 3-4 m above the ground. Another aspect of Lamprolepis biology that has received very little attention are the sexually dimorphic heel scales, greatly enlarged in males and often more brightly colored than the plantar surface. Gandolfi (1907) first discussed this structure in Lamprolepis and provided illustrations of sections through the scale, but neither he nor anyone since has suggested a function for them. The field notes of William H. Stickel described MCZ 49312 from Toem, Irian Jaya, New Guinea as, "Heel scale or- ange, which is unusual." (Loveridge 1948: 340-341). Manthey (1985:8) provided a color photo of the plantar surfaces of a live male and female from the Philippines in which the enlarged scale in the male is only slightly more yellowish than the dead- white ventral color. By comparison the scale is usually highly contrasting in the greener populations, described in our notes as "lemon yellow", "yellow-orange" or even bright orange, against the usually yellow-green plantar color. Distribution.-Babeldaob, Bablome- kang, Beliliou, Fana, Kmekumer Islands, Malakal, Merir, Ngcheangel Atoll (all is- lands), Ngeaur, Ngebad, Ngebedangel, Ngercheu, Ngerdis, Ngerechur, Ngereke- besang, Ngermalk, Ngerukeuid Islands (is- lands 12, 16, 24), Ngeruktabel, Ngerur, Ngesbokel, Ngetmeduch, Oreor, Pkulak- lim, Pulo Anna, Sonsorol, Tobi, Ulebse- chel, Ulong. Lipinia leptosoma (Brown and Fehlmann, 1958) Originally described as an endemic ge- nus (Aulacoplax) from Palau, leptosoma was referred to Lipinia by Greer (1974). Austin (1995, 1998) confirmed this generic allocation, but he found the relationships of leptosoma relative to several Papuan species equivocal. Brown and Fehlmann (1958) claimed that this species was an obligate dweller in the leaf axils of a single species of Panda- nus, and they reported specimens from Babeldaob, Oreor, and "Toiracholodoch- oel." The latter is supposedly "a small is- land at the north end of Malakal" (Brown and Fehlmann, 1958:5), but it does not ex- ist on our maps. Gordon Rodda (personal communication) offered that according to Demei Otobed "Toiracholodochoel" is ac- tually the name of a pass between the two islands, the smaller of which (Ngermalk?) is apparently the one to which Brown and Fehlmann (1958) referred. In any case, there is no longer any suitable Pandanus in this area. There are additional museum specimens (CM 29101-03) from Ngeaur collected as long ago as 1944, and we re- cently verified that the species does indeed occur there. Most of the recent Ngeaur specimens were collected from Pandanus near the south end of the island, but sev- eral individuals were seen and collected (e.g., USNM531945, SAM R4800) from palm axils near the northern tip of the is- land. Distribution.-ENDEMIC. Babeldaob, Ngeaur, Oreor, "Toiracholodochoel". Lipinia cf. L. noctua (Lesson, 1826) Zweifel (1979:1) noted "the relative uni- formity of color pattern and scutellation of L. noctua over the Pacific islands, in con- trast to the diversity in the much smaller New Guinea region." He did not list the non-New Guinea material he examined but the AMNH collections have noctua from a broad area of Oceania. Our field experience with noctua in Oceania are at odds with Zweifel's suggestion. Although scutellation is indeed conservative, we have noted significant differences in col- oration, body form, and ecology between Polynesian and Micronesian noctua. Poly- nesian noctua tend to be heavier-bodied, short-legged terrestrial or semi-arboreal skinks with a prominent, light occipital [ o. PETOLOGICAL GRAPHS 1999] HERPETOLOGICAL MONOGRAPHS 59 spot and a distinct broad vertebral stripe. By contrast, Micronesian noctua are more gracile, strongly arboreal skinks (possibly even obligate palm axil dwellers), often with a poorly developed occipital spot and vertebral stripe, but more distinct dorso- lateral striping. This pattern is very similar to Palauan Cryptoblepharus. In Palau, Lipinia noctua is found almost exclusively in or very near the axils of palms, and usually uncovered by peeling dead fronds and fabric from low trees. We have seen a few individuals apparently ac- tive (or at least visible) outside axils but they generally slip back into concealment as soon as they are detected. One excep- tion was a near-neonate that was active on a large dead tree near the beach just south of Ngaramasch, western Ngeaur. The tree had abundant loose bark which provided refuge for numerous (10+) active Cryp- toblepharus, and the Lipinia was mistaken for a juvenile of the former until we ex- amined the specimen more closely when preserving it. Two other adults were col- lected from beneath casuarina bark on northern Ngeaur. We suspect Palauan Lip- inia (and other Micronesian populations) may also occupy the axils of tall palms, similar to the Philippine L. auriculata group (Brown and Alcala, 1980; RIC, per- sonal observation), but we have found no way to sample taller Palauan palms as yet. The preference for palms departs from the preferred habitat of Polynesian L. noc- tua, which are frequently found under rocks and logs (McKeown, 1996; personal observation) or active on a variety of low vegetation, but rarely palms (Crombie and Steadman, 1987). From our limited data on pregnant females, Micronesian noctua usually produce a single offspring, as op- posed to the usual litter of 2 (but with a range of 1-4) in Hawaii (McKeown, 1996). Data from the Cook Islands suggested lit- ter sizes of 1 or 2 (Crombie and Steadman, 1987). Christopher C. Austin is currently re- viewing morphological and molecular var- iation in the genus Lipinia with significant samples from Oceania. His recent (Austin, 1999) mtDNA analysis of the widespread "noctua" suggests a monophyletic eastern and central Polynesian population of hu- man-mediated origin. Micronesian "noc- tua" are genetically diverse and probably represent pre-human colonization of the islands. Although Lesson's type locality (Oualan, now Kosrae in the Carolines) would seem to restrict the name noctua (sensu stricto) to the Micronesian popu- lations, the possible unreliability of the lo- cality and the lack of a type specimen (Brygoo, 1986:75) further complicate no- menclatural resolution of the problem. However, because Lesson's text (1830) gives ecological information (on a building in a sugar cane field) for the type(s) of noc- tua, perhaps the locality is accurate for this species. Distribution.-Babeldaob, Malakal, Ngeaur, Ngercheu, Ngerekebesang, Oreor, Pulo Anna, Ulong. Mabuya sp. Stemfeld (1920:395) first reported Ma- buia multicarinata from Palau. He dis- cussed in detail seven specimens from Feis (= Fais) and one from Mogomay, Ulit (= Ulithi) Atoll, in the western Carolines just north of Palau, and concluded by saying that the Senckenberg Museum also had two specimens from Palau, but no locality or other details were provided. The ma- terial from the Carolines included juve- niles, indicating that the species was estab- lished and reproducing at that time. Sternfeld (1920:395) summarized the distribution of M. multicarinata as Borneo and the Philippines. Brown and Alcala (1980) studied Philippine material of "multicarinata" and described two smaller species, M. cumingi and indeprensa, that had long been confused with it. They also found north-south variation in multicari- nata itself, and named the northern Phil- ippine populations M. multicarinata bo- realis. Ota (1991) extended the northern distributional limits of M. m. borealis to Lanyu Island, off the southeastern coast of Taiwan, noting that it was not found on the main island of Taiwan. However, neither the Bornean nor the Palau/Carolines pop- ulations have been studied in light of Brown and Alcala's (1980) revision of the 1999] PETOLOGICAL OGRAPHS 0 HFRPETOIflGTCAI. MONOGRAPHS [No. 13 group, primarily due to limited samples from these areas. The Palau/Carolines Mabuya represent the eastern distributional limit of the mul- ticarinata group and possibly the eastern limit of the Asian members of the genus. Mabuya multifasciata is found in the Mol- uccas and western New Guinea but its dis- tribution in this area is poorly document- ed; Ingram (1987) concluded that multifasciata was restricted to the western (Irian Jaya) area of New Guinea and that the northern Australian record was based on erroneous locality data or an introduc- tion. Some museum material from Palau and the Carolines has been misidentified as multifasciata. Many of the disjunct populations of Phi- lippine or Papuan vertebrate species in Oceania (e.g., Varanus, Crocodylus, Sun- cus, Macaca) are often attributed to intro- ductions by the Japanese during their pre- World War II occupation of Micronesia and much of the rest of insular East Asia. Mabuya, however, clearly pre-dates the period of Japanese occupation, and the Oceanic populations may represent natu- ral dispersal from the Philippines and/or Borneo. Preliminary data from our mate- rial support this proposal, since Palau/Car- olines Mabuya have the large body size typical of multicarinata but scale counts intermediate between multicarinata and indeprensa. We suspect that these popu- lations represent at least one undescribed species. The problem is currently being studied by Hidetoshi Ota and RIC. Distribution.-Babeldaob, Beliliou, Ngcheangel Atoll (Ngcheangel), Nger- cheu, Oreor, Pulo Anna, Sonsorol. Sphenomorphus sp. nov. This moderate-sized skink has only been found on Koror, Ngermalk, southern Ba- beldaob, and Ngeruktabel, usually in or near moist wooded areas. It is superficially similar to some members of the Spheno- morphus variegatus group, which includes more than 50 named species that do not fit conveniently into other groups. The Pa- lauan species has several distinctive char- acters, including bifurcate hemipenes with long and attenuate terminal elements. We only tentatively refer it to Sphenomorphus pending further study. Interestingly, Tiedemann and Haupl (1980:42) listed 5 "Syntypes" [NMW 14855 (2), and 16638 (3)] of Lygosoma (Hinulia)jagorii Peters from "Palaos", pol- lected by Semper in 1874. Quite aside from the fact that these specimens were supposedly collected 10 years after Peters described the species (1864:54), Peters clearly stated that the types ofjagorii were collected by Jagor at "Borongan, ... Sa- mar" and "Tacloban, ... Leyte". Brown and Alcala (1980:209) list ZMB 4975 and 4610 as syntypes of Sphenomorphus jago- rii. We know of no Philippine locality called "Palaos," but that name is another old permutation for Palau, in which case the above "syntypes" collected by Semper are possibly this species of Sphenomor- phus. The Palauan species vaguely resem- bles some jagorii, which is a poorly re- solved composite of several to many species throughout the Philippine archi- pelago (Brown and Alcala, 1980:211; RIC, unpublished data). However, most of the problematical Philippine populations are in the eastern arc of islands and the far north (greater Luzon landmass and asso- ciated island groups) rather than the area most proximate to Palau (Mindanao), which is occupied by the fairly well de- fined S. jagorii grandis. Thinking that the NMW specimens might actually be types, but of a species other than jagorii (mean- ing that there might be an available name for the Palau species), we carefully exam- ined all of Wilhelm Peters' papers (Bauer et al., 1995) and could find no scincid de- scribed from "Palaos" or any permutation thereof. We characterize this species as a deni- zen of moist woodland, but this requires some comment. The "woodland" need not be primary forest. Most of our specimens have come from the small park adjacent to the National Museum, which barely qual- ifies as woodland. We also collected one specimen (USNM 531965) under a rock in a roadside ditch at the Mechang dock area (Oreor). The ditch was adjacent to the lawn and driveway of a new house and the only "woodland" in the area was a small [ o. 60 E PETOLTOG CAL OGRAPHS 1999] HERPETOLOGICAL MONOGRAPHS 61~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ patch of dry scrub about 10 m away. On the other extreme, the southern Babel- daob specimens (SAM R47736, BPBM 13046) were collected under a sodden ba- nana spathe in a swampy patch of Panda- nus. We also have a single juvenile from Ngermalk (USNM 531966), collected by Kurt Auffenberg in nearly undisturbed vegetation on the top of the small (north of the causeway) part of this rock island that is crossed by the Koror-Malakal cause- way. Although this individual and some of our Babeldaob specimens have a more complex pattern than others, they do not differ in scalation. The pattern differences may be individual or ontogenetic variation. Our only Rock Island record (USNM 531974 from Ngeruktabel) was collected under a log on the trail to German Light- house at the southern end of the island. This trail ascends through lush limestone forest to the lighthouse ruins on the ridge crest. The Japanese had a shore battery and a support base in this area and the ruins are prominent landmarks along the trail. The specimens was collected about halfway between the battery (marked by the enormous gun barrel along the trail) and the ruins of the support base. The Jap- anese surely cleared some of this forest for fields of fire but it is hard to tell now as the trees are huge and the canopy is high, more typical of undisturbed primary for- est. Distribution.-ENDEMIC. Babeldaob, Ngermalk, Ngeruktabel, Oreor. Sphenomorphus scutatus (Peters, 1867) This Palauan endemic [Lygosoma (Hin- ulia) scutatum Peters, 1867; syntypes (2) ZMB5893] is closely related to the small Papuan species of the solomonis subgroup of the S. fasciatus group (Greer and Park- er, 1974). Although S. scutatus and the preceding species are strikingly different in size and scalation, both have the unusu- al, attenuate, bifurcate hemipenes (dis- cussed under Sphenomorphus sp.). Unfor- tunately, hemipenial morphology is unknown for virtually all of the multitude of species currently assigned to Spheno- morphus. The two Palauan species are currently assigned to two different (unre- lated) species groups, but we think it equally plausible that they may represent a Palauan radiation of closely related spe- cies. Although previously rare in museum collections (Greer and Parker, 1974, found only four specimens), S. scutatus is actu- ally a common leaf-litter skink on most is- lands. During the rainy season this skink is often active in the leaf litter by day, but during the dry season it is usually encoun- tered under rocks and logs in shaded ar- eas. It is found both in forest and in dis- turbed areas, including downtown Koror. Distribution.-ENDEMIC. Babeldaob, Malakal, Ngcheangel Atoll (Ngcheangel), Ngeaur, Ngermalk, Ngeruktabel, Ngerur, Oreor, Ulebsechel, Ulong. Varanidae Varanus cf. V indicus (Daudin, 1802) The systematics of the Varanus indicus group (Fig. 15) are still quite chaotic de- spite recent research in the Papuan area that has produced two new species, sub- species elevated to species status, and a drastically revised taxonomy of the group (Bohme et al., 1994; Harvey and Barker, 1998; Sprackland, 1993). None of the re- visions have included Oceanic material, probably because these populations are generally assumed to be introductions by the Japanese before or during WWII (e.g., Owen, 1977). Our experience with Var- anus in Oceania suggests that the situation may not be so simplistic. Uchida (1969) noted that there were museum specimens from Micronesia that long antedated the Japanese occupation, and he concluded that they must have been a German intro- duction. Historical accounts suggest that the Western Caroline (Yap, Ulithi) and Pa- lau populations might be native but that the eastern Caroline (Ifalik, Kosrae, Pohn- pei), Marshall (not represented by any voucher specimens we have located), and Mariana monitors could be recent intro- ductions. One of the earliest zoological re- ports on Micronesia (Chamisso, 1821) mentioned monitors from both Yap and Palau, and commented on their totem sta- tus in local cultures. On the other hand, in 1999] 61 62 HERPETOLOGICAL MONOGRAPHS [No. 12 FIG. 15. The distribution of Varanus indicus. the Northern Marianas Pregill (1998) found Varanus only in recent (post-Euro- pean) strata. We are impressed with the morpholog- ical dissimilarity of Micronesian monitors. The "indicus" on Guam are smaller, paler, and more gracile than Palauan specimens. The common name "mangrove monitor" used for Greater Papuan indicus reflects its arboreality, but the Palauan populations are decidedly terrestrial. They take refuge in trees when pursued but only when ter- restrial retreats are unavailable. In color- ation and pattern, Palauan Varanus are black with prominent yellow dorsal ro- settes and other irregular markings, with a vivid peach-colored throat in both adults and juveniles. They are both longer and bulkier than monitors on Guam. We have seen a reliable photo of an individual col- lected (and eaten) by Filipinos near Uli- mang, northern Babeldaob. It was laid out next to a tape measure that was a bit lon- ger than 6 feet total length. We have seen individuals approaching this length on Ngeaur. Available data on the Marianas/Caro- lines populations suggest that they are usu- ally associated with the most disturbed habitats (Bates and Abbott, 1958; Dryden, 1965). There is no comparable information for Yap and Ulithi, but in Palau Varanus is most definitely not a human commensal. In Palau, we have been unable to verify the earlier Koror records (AMNH 70652, CAS 152219), and have only one sight re- cord from the island, and that from the rugged limestone Ngermeuangel Penin- sula. On Babeldaob we have never seen monitors in the southern part of the island, nor have we received any reports from the area by the Filipinos who work the rural farms. Monitors (bayawok in Tagalog) are a favored food item among Filipinos and if they occurred in southern Babeldaob we are confident we would have confirming information and/or specimens from this valuable source. In sparsely populated northern Babeldaob we have seen but been unable to collect Varanus in the Uli- mang/Ngesang area of Ngaraard State. Gressitt (1952, 1953) made notes on Varanus on Ngcheangel Atoll. Tom Fritts and Gordon Rodda verified that the spe- cies was still present in January 1993, but only on the two larger islands where they were moderately abundant. The only other place Varanus is truly common is Ngeaur, where 20 or more can usually be seen in a few hours walk along the road that cir- cles the island. They do occur in and near Ngaramasch village, but they are most abundant in the rugged limestone of the [No. 3 1999] HERPETOLOGICAL MONOGRAPHS 63 FIG. 16.-The distribution of Candoia carinata. interior where they have multitudes of ref- uges and are extremely difficult to catch. Clearly, the status of Oceanic Varanus needs additional study but collection of pertinent material is hampered by the CITES status of all varanids. Tissue sam- ples from the Oceanic islands would help resolve the status of these important, pri- mary predators, and clarify which popula- tions are introduced versus those that are native (and possibly endemic) species. Distribution. -Babeldaob, Ngcheangel Atoll (Ngcheangel, Ngeriungs), Ngeaur, Oreor. SERPENTES Boidae Candoia carinata (Schneider, 1801) Giinther (1863) described Candoia su- perciliosa from Palau, but the taxon was quickly relegated to the synonymy of the widespread and variable C. carinata (e.g., Boulenger, 1893:107). Our first field im- pressions of Palauan Candoia was how subjectively "different" they were from New Guinea/Solomons carinata. Mc- Dowell (1979) examined a small sample of Palauan Candoia in his survey of New Guinea boids and found them to be an in- teresting mix of his normally allopatric "long tailed" and "short tailed" morphs. Bohme et al., (1998) recently described the rediscovered lectotype in the Zoolo- gisches Forschungsinstitut und Museum Alexander Koenig (ZFMK 35503), and re- ported that the type locality, previously un- known, was Ambon Island ("Amboina" on the specimen label) in the southern Mo- luccas. They examined over 300 specimens from throughout the species' range (Fig. 16), and found morphological uniformity in the populations from the southern Mo- luccas, Irian Jaya, northern Papua New Guinea, New Britain and New Ireland, all of which they regarded as "typical" C. car- inata. Their Palau specimens (n = 29) conformed with these in most respects, but showed several average differences in some scale counts and tail length ratios. Hobart M. Smith has recently taken data on a great many Candoia from throughout the distribution of the genus (including many of our recent Palauan specimens) and has concluded that the Palauan pop- ulations are quite distinct, but most closely related to "true" carinata from the Mol- uccas (personal communication to RIC, June 1998). We expect him to formally recognize Giinther's superciliosa for this species. Wynn and Zug (1985) detected some re- productive differences between Palauan Candoia and their larger sample from New Guinea, which they suggested were 99] 64 HERPETOLOGICAL MONOCRAPHS [No. 13 FIG. 17.-Locality records for Cerberus rynchops (after Gyi, 1970). correlated with latitude. Harlow and Shine (1992) amplified Wynn and Zug's repro- ductive information and added dietary data based on dissection of museum ma- terial and published observations on cap- tive specimens. They further noted that virtually all natural history data on Can- doia were derived from preserved muse- um material, captive specimens, and very few actual field observations, all from the New Guinea region. Candoia is common and widely distrib- uted in Palau and it is found in both dis- turbed habitats (even in Koror town and on Malakal) and in forest. The species is usually reported to be arboreal and noc- turnal, and indeed most of our specimens were so found. Still, several juveniles were found active on the ground in the early morning near the beach in northern Ba- beldaob. An adult also was found sunning on a limestone ledge at 1130 h on Nger- duais (a small island separated from the southeast coast of Babeldaob by mangrove and a stream). Distribution.-Babeldaob, Beliliou, Ma- lakal, Ngeaur, Ngercheu, Ngerduais, Nger- echur, Ngerekebesang, Ngerukeuid Islands (island 13) Ngeruktabel, Ngetmeduch, Or- eor, Pkulaklim. Colubridae Cerberus rynchops (Schneider, 1799) The most recent review of this wide- spread species (Fig. 17) was by Gyi (1970), who characterized it as "usually found in the brackish water of tidal rivers, creeks, and estuaries, but is equally at home in the upper reaches of rivers, and in salt waters along the coasts." Although he apparently did not examine any material from Palau, Gyi included Pa- lau as the eastern distributional limit of the nominate subspecies. He recognized the subspecies novaeguineae based on four specimens from Merauke in the Irian Jaya portion of New Guinea. Cerberus is common in Palau, but often difficult to collect since it is nocturnal and the mangrove swamps it prefers are inhos- pitable after dark. The species is occasion- ally found in the open ocean or along beaches, particularly near river or stream mouths. One specimen (USNM 507563) from northern Babeldaob was found crawling across strand vegetation between the beach and an inland mangrove swamp at 0710 h. Reproduction in Philippine Cerberus rynchlops was described by Gorman et al., (1981). For the Palauan population we have data on a single neonate with an um- bilical scar (USNM 531967, SVL 232 mm, Tail 62 mm, live weight 11.3 gm) collected 29 Jan 1995. This individual emerged from a crevice in the mud bank of a mangrove swamp on Oreor when one of us (RIC) stepped and sank into the ooze. We G [ o. 1999] HERPETOLOGICAL MONOGRAPHS 65 thought that there might be an underwater entrance, but further manipulation of the area produced no further neonates, al- though other juveniles were seen in adja- cent pools of the swamp. Distribution.-Babeldaob, Ngerekebes- ang, Oreor. Dendrelaphis sp. Peters (1867a) described Dendrophis striolatus (ZMB 5450) that was collected from an unspecified locality on Palau; he suggested the new species was most close- ly related to the Australian D. punctulatus. Boulenger (1894:87) synonymized striola- tus with lineolatus and his arrangement was followed by Dryden and Taylor (1969) in their discussion of three specimens (two from Ngeaur, one from Oreor). Meise and Hennig (1932) retained striolatus as a valid subspecies of lineolatus but Mertens (1934a) referred to the Palau species as D. punctulatus striolatus. McDowell (1984:6) also called the Palau snakes Dendrelaphis punctulatus (with lineolatus as a synonym but no mention of striolatus), based on a very small sample size. Despite McDowell's careful analysis of the New Guinea material, we are not con- vinced that his placement of the Palauan populations is correct. Our collections con- tain two distinct forms of Dendrelaphis: the common, greenish, striped variety with a yellow belly occurs in the southern part of the archipelago, but a larger, unmarked blue form with a pale blue venter is found in northern Babeldaob. Our samples do not demonstrate any meristic differences between the two morphs. Occasional "blue" individuals are found among the Philippine Dendrelaphis cau- dolineatus (RIC, personal observation), but there does not seem to be any geo- graphic correlation in the variant. The "melanistic" Dendrelaphis salomonis from Misima Island (McDowell, 1984) may be a similar situation. Palauan Dendrelaphis are very ecologi- cally tolerant, surviving quite well in Koror town as well as in primary forest on the Rock Islands. They are diurnal and pri- marily terrestrial, but usually sleep on fences, low bushes, or (especially in the blue populations) even 3-5 m above ground on palm fronds or in the axils. Distribution.-Babeldaob, Beliliou, Ma- lakal, Ngeanges, Ngeaur, Ngemlis Islands (sight record, Bailechesengel), Ngercheu, Ngerduais, Ngerekebesang, Ngerukeuid Islands (island 24), Ngeruktabel, Oreor, Ulebsechel, Ulong. Hydrophiidae Pelamis platurus (Linnaeus, 1766) This pan-tropical, pelagic species prob- ably rarely if ever enters the waters inside the Palau reef. The only known specimen from the Palau area (CAS 122587) was col- lected "ca. 2 miles off SW side of Anguar (sic) Id.," outside the reef. Pelamis gives birth to live young while swimming at sea so, unlike Laticauda, does not need to make landfall to lay eggs. Owen (1977) re- ported that Pelamis shows up occasionally in Palauan waters after storms. Distribution. -Ngeaur? Laticaudidae Laticauda colubrina (Schneider, 1799) This is the common sea snake in Palau, although some other species could be ex- pected as occasional vagrants. Laticauda colubrina is very common inside the reef and is often seen by divers in a variety of shallow and deep-water situations. Juve- niles are often trapped in tidal pools at low tide, where we have observed reef herons (Egretta sacra) feeding on them. All of our Laticauda vouchers were collected on- shore. As the name implies, sea kraits are not as tied to the ocean as the true sea snakes. Laticauda must come ashore to lay eggs (unlike the live-bearing sea snakes) and they often spend enough time on land to accumulate ticks (Booth et al., 1997; Das, 1992; Pernetta, 1977). Although sea caves and uninhabited, precipitous rocky islands are favoured reproductive areas where large aggregations of snakes may be found (Saint Girons, 1964; Stuebing, 1988; Stuebing et al., 1990), individuals may come ashore singly and climb into beach wrack to shed, rest, and perhaps even bask. Shed skins and inactive individuals have been found in exposed root masses ] 66 HERPETOLOGICAL MONOGRAPHS [No. 13~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ of undercut beach trees in Ngaraard State, northern Babeldaob. In this same area, a large male (USNM 507503) was found ac- tively moving through strand vegetation 10 m from the sea at 2035 h. At this site (just south of Ulimang Village), there is only a narrow strip of "dry land" between the beach and an inland mangrove swamp. La- ticauda has not been found in mangrove swamps in Palau, although, we collected a DOR subadult adjacent to mangrove (USNM 531992; 350 mm SVL) in north Oreor just south of the causeway to Nget- meduch and the KB Bridge. Mangrove is extensive to the east of the road, and the specimen was collected on that side of the road; to the west mangrove is scattered, and opens to the shallows of the lagoon. Distribution.-Babeldaob, Beliliou, Ma- lakal, Ngelsibel, Ngerduais, Ngerechur, Ngerukeuid Islands (islands 13, 24), Oreor, Pkulaklim, Ulong. Typhlopidae Ramphotyphlops acuticaudus (Peters, 1877:416) This species (Typhlops acuticaudus Pe- ters, 1877; ZMB9127) is endemic to Palau but it is very close in coloration and exter- nal morphology to the Papuan R. flaviven- ter (McDowell, 1974). In fact, one muse- um specimen from Ngeaur (CM 29087) was identified as flaviventer by Neil D. Richmond, a specialist in typhlopid snakes, in 1967. Addison H. Wynn has examined most of our recent Palauan typhlopid ma- terial and he confirms the lack of diagnos- tic characters separating acuticaudatus from flaviventer. Van Wallach (personal communication) also dissected and exam- ined the everted hemipenes of one of our acuticaudus specimens (SDSNH 68437) and feels that the species is distinct from flaviventer. However, he (Wallach, 1996) did not discuss acuticaudus in his recent review of the flaviventer group of Ram- photyphlops. We retain acuticaudus as a valid species, related to flaviventer, pend- ing the results of future studies. Although Ramphotyphlops acuticaudus seems to require slightly more mesic con- ditions than R. braminus, R. acuticaudus is certainly not a forest obligate. In fact, we have never collected it in primary for- est, or the more forested parts of the larg- er islands. We have collected it at the for- est edge at Nekkeng, Babeldaob, sympatrically with R. braminus. The two species were collected microsyntopically in sawdust piles on the grounds of the Mu- seum in Koror town (R. Owen, personal communication, based on the following specimens: acuticaudus-USNM 207022, 207024, 207026-28; braminus-USNM 207023, 207025) and we have collected the two species under adjacent rocks in the same area. Unlike Owen's series, bra- minus is now much more abundant every- where than acuticaudus. Although we have never found this spe- cies on any of the many Rock Islands sur- veyed, Ota's record (KUZ 20006-07) for Ngercheu is undeniable. It is possible that this population was accidentally transport- ed there in construction material for the Carp Island Resort. Distribution.-ENDEMIC. Babeldaob, Ngeaur, Ngercheu, Oreor. Ramphotyphlops braminus (Daudin, 1803) This polyploid, unisexual, parthenoge- netic species (Wynn et al., 1987) is the most successful disperser in the snake world, and its broad distribution in the IndoPacific rivals (and parallels in some ways) that of some of the house geckos. Unlike some frogs (e.g., Bufo marinus, Rana catesbeiana) that were intentionally introduced in new places for insect con- trol, Ramphotyphlops braminus is appar- ently a passive disperser, albeit likely with human assistance. The most probable mechanism is in the root balls of orna- mental (more recently) or food (historical- ly) plants transported by humans. Al- though the natural range of braminus is unknown, it occurs from the Indian sub- continent to most of mainland Southeast Asia (Hahn, 1980:40). Presumably the spe- cies originated somewhere in this area. The salinity tolerance of braminus has never been tested, but nonetheless it is a superb island colonizer. It occurs on many Indian Ocean islands and has even reached the coasts of Africa. It is ubiqui- 66 PETOLOGICAL OGRAPHS [ . 3 1999] HERPETOLOGICAL MONOGRAPHS 67 tous on the close-in East Asian archipela- gos (Indonesia, Philippines), and it pre- sumably dispersed from there to the Pacific islands and even to the coast of Mexico. The species' distribution in Oceania is spotty and it did not reach the Neotropics by "island hopping" across the entire Pacific. Until very recently, the Mexican distribution centered around Acapulco, Guerrero (and associated trade routes inland), which was the eastern ter- minus of the "Manila Galleon" route that transported goods from the Philippines to the New World colonies during 18th and 19th Century Spanish occupation (Dixon and Hendricks, 1979). The species contin- ues to disperse in Mexico (Mancilla-Mor- eno and Ramirez-Bautista, 1998). It has even turned up recently in the Caribbean (Censky and Hodge, 1997) and additional range expansion is inevitable. Natural dispersal from the mainland to closer island groups is not out of the ques- tion. Much has been made of "arboreal" behavior in the normally fossorial lepto- typhlopid and typhlopid snakes, but com- mon sense would indicate that these obli- gate ant and termite feeders should be able to utilize above-ground "soil" in the arboreal nests and runways of their prey. Our field observations and anecdotal lit- erature reports provide preliminary data that this "arboreality" may be much more common than previously expected (Das and Wallach, 1998; Gaulke, 1995). Taylor (1922:59-64) suggested that some of the larger endemic Philippine species of Typh- lops may occur exclusively in the extensive root masses of aerial ferns (Asplenium), which can provide substantial "fossorial" habitat in the forest canopy. On several oc- casions we have found Ramphotyphlops braminus climbing a meter or more above the ground on live coconut trees (after dark). We have also found several other species of leptotyphlopid and typhlopid snakes (including R. braminus) 2 m or more above the ground in dead, usually ant- or termite-ridden trees. The punky interior of dead coconut trees knocked into the ocean by storms is an obvious dispersal mechanism. There is insulation from the dehydrating effects of salinity, as well as abundant insect prey for a small snake like R. braminus. Assisted by high-energy storm tracks, a propagule (= a single individual of a parthenogenetic species) could easily cross the 800 km from Mindanao to Palau. In fact, late Ho- locene fossil remains from the Marianas (Pregill, 1988) indicate the likely pre-hu- man (and certainly pre-European) pres- ence of a braminus-like blindsnake. Al- though Dryden and Taylor (1969) described Typhlops pseudosaurus [re- tained as a valid species of Typhlina (= Ramphotyphlops) by Hahn, 1980] from Harmon Village, Guam, RIC examined the holotype of T. pseudosaurus (FMNH 189357) and considers it to be R. brami- nus. The specimen was preserved as it was preparing to shed, and having aberrant head sutures on one side accounted for Dryden and Taylor's (1969) supposed di- agnostic characters. In Palau, the limited historical and eco- logical evidence suggest that Ramphotyph- lops braminus is a recent arrival, and R. acuticaudus is a much older autochthon. The earliest records for Palauan braminus are Fehlmann's (mid-1950's), who found the species commonly on Oreor (25 spec- imens) but nowhere else, despite the fact that he spent far more time on Babeldaob, where he only collected R. acuticaudus. He collected both species on Oreor, in- cluding one locality where both were found syntopically (1 acuticaudus and 11 braminus). Ramphotyphlops braminus is now widespread in Palau and is particular- ly common in disturbed areas on the causeway-connected islands. Still, we have collected it from the northern end of Ba- beldaob (not connected to the south by road in the 1990's), Ngcheangel Atoll, and Ngeaur. There are no records from the Rock Islands or Beliliou, but there is one specimen from the Beliliou satellite of Ngedbus, collected in 1955 by Robert Owen (CAS-SU 19077). Distribution.-Babeldaob, Malakal, Ngcheangel Atoll (Ngcheangel), Ngeaur, Ngedbus, Ngetmeduch, Oreor. REPTILIA: TESTUDINES Chelidae, gen. and sp. indet. Aoki (1977) first reported a freshwater turtle from Palau, but his photograph was 1999] 68 HERPETOLOGICAL MONOGRAPHS rNo. B not adequate to confidently identify it to genus or species and no voucher speci- mens are known to exist. Apparently this turtle is (or was) a re- cent introduction from New Guinea, prob- ably a species of Elseya or Emydura as suggested by Aoki (1977). Nothing is known of the distribution or status of the species on Palau and most Palauans are unaware of its existence. Residents of Ngerekebesang, however, indicated that turtles could be found "commonly" in taro patches on the eastern end of the island. Until a voucher specimen is available to verify the identification, it would be spec- ulative to suggest what impact (if any) this introduction might have if the turtles are indeed established and reproducing. How- ever, since turtles are often top predators in aquatic ecosystems the species could af- fect native aquatic vegetation, inverte- brates, or fishes. Distribution.-unknown. Bataguridae cf. Chinemys reevesii Etpison (1997:211) published photo- graphs of another freshwater turtle from an unspecified locality in Palau. George R. Zug and Carl H. Ernst examined the pho- tograph and they are reasonably sure that it is an Asian batagurid (formerly a sub- family of Emydidae). Hidetoshi Ota (per- sonal communication) suggested that it could be Chinemys reevesii, and we so re- fer it on that basis. Without a voucher specimen no further conclusions can be made. It is unclear if either of the freshwater turtles (see previous account) are estab- lished in Palau, and origin of the introduc- tions is likewise unknown. In Hawaii, sev- eral Asian turtles were imported and released by the Chinese community as a food source, and at least two of them are now established and breeding (McKeown, 1996). Palau has a significant Chinese community and there has been a sizeable recent influx of people and economic sup- port from Taiwan. This seems a plausible possible source of the Etpison (1997) tur- tle. Turtle herpetoculture is not practiced in Palau (but it is a major cause of exotic introductions elsewhere). As noted in the chelid account, exotic turtles are a poten- tially serious threat to the poorly docu- mented endemic freshwater fish and in- vertebrate fauna, and serious effort should be made to document the status of these turtles in Palau. Further import should be prohibited, and extant populations (if any) identified and eradicated while control is still possible. Distribution. -unknown. Cheloniidae Chelonia mydas (Linnaeus, 1758) Eretmochelys imbricata (Linnaeus, 1766) Green sea turtles (Chelonia mydas) and Hawksbills (Eretmochelys imbricata) are common in Palauan waters, and for cen- turies they were exploited by native resi- dents for their meat, eggs, and shells (Jo- hannes, 1981; McCoy, 1981). There are no current censuses of nesting sites within the lagoon. Sandy beaches are limited and nesting sites are restricted accordingly. Hawksbills nest in the Ngerukeuid group (Seventy Islands), and Greens utilize sites on some of the small islands surrounding Babeldaob, on Beliliou, and doubtless oth- er isolated places within the lagoon. The most important nesting site for Greens are on Helen and Merir in the Southwest Is- lands (Pritchard, 1981; Wilson, 1969). Leatherbacks (Dermochelys coriacea) and Olive Ridley turtles (Lepidochelys oliva- cea) also occur in Palau but are uncom- mon compared to Greens and Hawksbills. Distribution in Palau.-Scattered but probably throughout Palau including Southwest Islands. REPTILIA: CROCODYLIA Crocodylus sp. Crocodylus porosus Schneider, 1801 Salt water crocodiles are native to Palau (Motoda, 1937, 1938), and supposedly were much more numerous 30 years ago than they are now (Kimura, 1968a,b; Wil- son, 1964). They occur elsewhere in the Caroline Islands only as rare waifs. A 12- foot individual was caught on Pohnpei in 1971 after three pigs teathered near the beach mysteriously disappeared over a six- [ o. 13 1999] HERPETOLOGICAL MONOGRAPHS 69 month period (Allen, 1974). In Palau, a non-fatal attack in 1964 and the subse- quent capture of a crocodile nearly 15 feet long was followed by the death of a local spear fisherman in December, 1965 (Brown, 1971; Wilson, 1965). Subsequent- ly, a crocodile eradication program was ini- tiated, and during the next 15 years various parties of Japanese, Australians, and Pa- lauans hunted and killed crocodiles. Un- fortunately, records of kills were inade- quately maintained, and there is no accurate figure of the number of croco- diles taken, nor were scientific vouchers saved. Estimates run from a few hundred to several thousand killed during the 1970's according to Messel and King (1991), who reviewed the history of croc- odile populations in Palau and conducted the only systematic biological survey. Mes- sel and King made their survey in June, 1991, and concluded that a generous es- timate of Palau's total number of saltwater crocodiles was less than 150. At that time the majority of their actual sightings were made on Beliliou and various river systems on Babeldoab (tributaries feeding Ngere- meduu Bay, others along the east coast south to Airai) as well as Ngerdok Lake. Crocodiles are also known to occupy the various marine lakes in the Rock Islands, but Messel and King (1991) dismissed the likelihood that they breed there because of insufficient places to lay eggs, and the fact that larger breeding adults would have limited access to these lakes because of the narrow crevices and channels that connect them to the lagoon. Today, a crocodile farm in north Koror harbors about 40 in- dividuals of various size classes of local C. porosus. Supposedly, this enterprise is in place to build a breeding stock for even- tual release of adults into protected areas. Messel and King (1991) also emphati- cally state that C. porosus is the only spe- cies of crocodile in Palau, despite rumors and opinions that have persisted for de- cades that another species, perhaps two, occur (Kimura, 1968a,b; Thyssen, 1988). Crocodylus novaeguineae and the Philip- pine C. mindorensis were imported during the 1930's for a farming venture, but by the end of WWII few if any of these re- mained alive. Escapees apparently are the source of the supposed other species in Palau. We would also like to point out that despite Messel and King's vehement de- nial of a native freshwater croc in Palau, it is hard to imagine that the islands would not have (or had) one on purely zoogeo- graphic grounds. It has long been thought that Crocodylus mindorensis and C. no- vaeguinea are closely related, even con- specific by some authors (see discussion in Hall, 1989). Although palustrine crocodiles can (and do) cross freshwater barriers, most areas (especially islands, even large ones like Borneo) harbour only one fresh- water Crocodylus. If in fact C. mindorensis and C. novaeguinea are sister taxa then only two plausible dispersal routes could explain the distribution: island-hopping through the islands west of New Guinea, or by a direct oversea route between the Philippines and New Guinea. The island- hopping route would have to bypass Bor- neo, which is inhabited by another fresh- water species, C. raninus (Ross, 1992). On the oversea route the only major landfall between New Guinea and the Philippines is Palau. Given the strong Papuan influ- ences (and lesser Philippine ones) in the Palauan herpetofauna, including species that have great difficulty in crossing ma- rine barriers (e.g., frogs), we think that the presence of freshwater crocodiles in Palau is not as unthinkable as Messel and King insist. Note that we do not suggest that the species is there, merely that it is an in- triguing possibility, particularly given the abundance of anecdotal reports and local lore. Unfortunately, it is a moot point since CITES restrictions prevent collection of specimens and data that would resolve the problem. Even more regrettable is that none of the crocodiles killed during the "control" period was saved for science. We made no attempt to search actively for crocodiles, C. porosus or otherwise, but note that ample habitat exists in which crocodiles could breed. We photographed a two-foot juvenile that was brought into the Coral Reef Research Foundation on Malakal in January, 1995 that was taken from nearby Koror. Indeed, our experienc- es with Crocodylus in Palau are somewhat 99] 70 HERPETOLOGICAL MONOGRAPHS [No. 13 at odds with Messel and King's data. With- out even particularly looking for them, we have seen hatchlings, juveniles, and large adults virtually everywhere we have trav- eled in Palau, even quite close to Koror Town itself. Messel and King (1991) re- counted the difficulties in surveying for crocodiles, for example that tidal fluctua- tions limit survey time and that navigating through the slackwater of mangrove is per- ilous at night. We agree, but while we do not diminish the efforts of their survey, it would surprise us if the number of croc- odiles in Palau was not significantly greater than 150. Distribution.-Babeldaob, Beliliou, Or- eor. Acknowledgments.-This project would have been impossible without the cooperation and assistance of numerous Palauan citizens and residents. We have enjoyed more than the usual cordial welcome in Pa- lau, and we owe a huge debt of gratitude to many people. Our governmental contacts in Palau were Robert Owen (early in the planning stages), Demei Otobed, the late David Idib, Noah Idechong, and most recently Kammen (Taktai) Chin, each of whom so greatly facilitated our activities that any success we have had must be credited to their enlightened pol- icies and cooperation. In attempting to locate and verify identifications on all earlier Palau collections, we received cooperation from a great many individuals and institutions. We are unable to acknowledge all of the collections re- sponding to verify that they did not have Palauan ma- terial, but we appreciate the information nonetheless. The following collections and their staffs provided printouts or other lists, loaned specimens and/or made working space available for us (museum acro- nyms follow Leviton et al., 1978): AMNH-R. Zweifel, J. Cole, C. Myers, D. Frost, L. Ford ANSP-E. Malnate BMNH-N. Arnold, C. McCarthy, B. Clambe BPBM-A. Allison, C. Kishinami BYU-W. Tanner CM-J. McCoy, E. Censky FMNH-H. Voris, A. Resetar, C. Redhead KUZ-H. Ota MCZ-E. E. Williams, J. Rosado, J. Cadle MNHN-A. Dubois, R. Bour, E. Brygoo MVZ-H. Greene, D. Wake SAM-M. Hutchinson, C. Austin SMF-K. Klemmer, G. K6hler SMNS-A. Schluter, J. Hallermann UMMZ-A. Kluge, G. Schneider We have not yet had the oportunity to examine the important early Palau collections in Germany and Amsterdam, but Aaron Bauer and Rainer Glinther provided us with a great deal of information on the ZMB collections, and Andreas Schluter provided a list of the Stuttgart holdings. As the repository for the second largest Palauan herpetological collection, The California Academy of Science endured our frequent visits with surprising tolerence. Walter Brown, Bob Drewes, Jacques Gau- thier, Al Leviton, Heidi Robeck and especially Jens Vindum came through with space, assistance and courtesies well above and beyond the call. Jens Vindum and Carol Prince were cordial hosts whose warm hospitality was always appreciated, even if it cast doubt upon their sanity. For basic in-country logistics, Demei Otobed, Tim Taunton, Chris Regel, and the Shiro family of Nga- raard State, Babeldaob deserve special mention. The late Harson Shiro was particularly generous in shar- ing his knowledge of Palau, especially of the Ulimang area. The Coral Reef Research Foundation (funded by the National Cancer Institute) also was a source of immeasurable assistance to us on so many fronts that it is difficult to thank them adequately for it and their companionship. To Pat Colin, Lori Colin, and Larry Sharron, plus their staff Matt, Emilio, Nathan, and Lolita, our appreciation is bottomless. At various times we were assisted in the field or aided in other ways by Tim Asis, Kurt Auffenberg, Chris Austin, Eric Carlson, Ginny Carter, Ken Creighton, Bill Dunson, Mandy Etpison, Robert Fisher, Tom Fritts, Ben Imperial, The New Koror Hotel, George Phocas, Gordon Rodda, Ron Leidich and Sam's Dive Tours, Mary McCutcheon, Susan Schwartz, Dave Steadman, Ken Tighe, and Gary Wiles and the staff of the Guam Division of Aquatic and Wildlife Resources. The Pacific Savings Bank in Koror came to our aid on countless occasions by providing communications, transportation, sanctuary, and numerous other cour- tesies. President Tim Taunton along with Debbie Rengiil and John De Vivo deserve banner acknowl- edgment for their unstinting assistance and good cheer. We apologize for the persistent assumption in Koror that we must be employed by the bank. We are confident, however, that the PSB will survive this slur and continue to prosper. We thank Leslie Overstreet for bibliographic assis- tance, and Fred Kraus and Chris Austin for com- ments on earlier drafts of the manuscript. Angela Kay Kepler provided invaluable information and speci- mens from the Southwest Islands. Hidetoshi Ota shared herpetological records (KUZ) for Ngercheu Island. George Zug and Carl Ernst helped with the photo identification of the batagurid turtle. Gordon Rodda and David Steadman provided expert reviews of the final draft, uncovering several lapses on our part. We are at last grateful to the National Geographic Society for funding this project (NGS 5032-93) and field biology in general. Additional support was gen- erously provided by the USFWS. RIC's museum work and the early permit negotiations in Palau were supported by Smithsonian Research Opportunities funds. . 1999] HERPETOLOGICAL MONOGRAPHS 71 LITERATURE CITED ALCALA, A. C. 1961. Breeding behavior and early de- velopment of frogs of Negros, Philippine Islands. Copeia 1961:679-726. 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Hemidactylus frenatus Hemiphyllodactylus cf. typus Lepidodactylus lugubris Lepidodactylus moestus Carlia cf. C. fusca Cryptoblepharus sp. Emoia atrocostata Emoia caeruleocauda Eugongylus sp. Lamprolepis smaragdina Lipinia leptosoma Lipinia cf. L. noctua Mabuya sp. *Sphenomorphus sp. nov. Sphenomorphus scutatus *Varanus cf. v. indicus (CAS 152219) Candoia carinata Cerberus rynchops Dendrelaphis sp. Laticauda colubrina Ramphotyphlops acuticaudus Ramphotyphlops braminus *Eretmochelys imbricata (CAS 154193) *Crocodylus porosus BABLOMEKANG (ca. 2.5 km W of Mecherchar) Gehyra brevipalmata Gekko sp. Lamprolepis smaragdina BF,.TJTJOU Bufo marinus Platymantis pelewensis *Gehyra brevipalmata Gekko sp. Hemidactylus frenatus Lepidodactylus lugubris Carlia cf. C. fusca *Cryptoblepharus sp. *Emoia atrocostata *Emoia caeruleocauda *Eugongylus sp. (CAS 122407-08) Lamprolepis smaragdina *Mabuya sp. (CAS 122595-97) *Candoia carinata *Dendrelaphis sp. *Laticauda colubrina *Crocodylus porosus CHEMOI GROUP (ca. 0.5 km NW of Mecherchar) Gehyra oceanica EUIDELCHOL ISLAND (ca. 0.5 km N of Mecherchar) Emoia caeruleocauda Gehyra brevipalmata IOULOMEKANG (ca. 0.8 km SSW of Bablomekang) Gehyra mutilata Lepidodactylus lugubris Emoia caeruleocauda KMEKUMER ISLAND GROUP (ca. 2 km W of Ngerukeuid Group) (*specimens from island "46" of Wiles and Conry, 1990) Gehyra brevipalmata *Gekko sp. Lepidodactylus lugubris *Lepidodactylus moestus *Cryptoblepharus sp. *Eugongylus sp. Lamprolepis smaragdina MALAKAL Bufo marinus Platymantis pelewensis Gehyra brevipalmata Gehyra mutilata Gekko sp. Hemidactylus frenatus Lepidodactylus lugubris Lepidodactylus moestus Anolis carolinensis Carlia cf. C. fusca Emoia atrocostata Emoia caeruleocauda Lamprolepis smaragdina Lipinia cf. L. noctua. Sphenomorphus scutatus Candoia carinata Dendrelaphis sp. Laticauda colubrina Ramphotyphlops braminus MECHERCHAR Emoia caeruleocauda NGCHEANGEL ATOLL NGCHEANGEL ISLAND *Bufo marinus *Platymantis pelewensis *Gehyra mutilata *Gehyra oceanica *Lepidodactylus moestus *Emoia caeruleocauda *Emoia jakati 1999] 78 HERPETOLOGICAL MONOGRAPHS No. 13 *Eugongylus sp. (CAS 122409) *Lamprolepis smaragdina *Mabuya sp. *Sphenomorphus scutatus *Varanus indicus *Ramphotyphlops braminus NGEREBELAS ISLAND *Gehyra oceanica *Lepidodactylus moestus *Emoia caeruleocauda *Emoia impar *Lamprolepis smaragdina NGERIUNGS ISLAND *Platymantis pelewensis *Gehyra oceanica *Lepidodactylus moestus *Lamprolepis smaragdina *Varanus cf. v. indicus ORAK ISLAND Gehyra oceanica Lepidodactylus moestus Emoia caeruleocauda Emoia impar Lamprolepis smaragdina NGEANGES Gehyra oceanica Hemiphyllodactylus cf. H. typus Lepidodactylus lugubris Lepidodactylus moestus Emoia caeruleocauda Dendrelaphis sp. NGEAUR Bufo marinus Platymantis pelewensis *Hypsilurus godeffroyi (?) Gehyra brevipalmata Gehyra mutilata Gekko sp. Hemidactylus frenatus Hemiphyllodactylus cf. H. typus Lepidodactylus lugubris Lepidodactylus moestus Carlia cf. C. fusca Cryptoblepharus sp. Emoia atrocostata Emoia caeruleocauda *Eugongylus sp. Lamprolepis smaragdina Lipinia leptosoma Lipinia cf. L. noctua Sphenomorphus scutatus Varanus cf. v. indicus Candoia carinata Dendrelaphis sp. *Pelamis platurus (CAS 122587) *Ramphotyphlops acuticaudus (CM 29087) Ramphotyphlops braminus NGEBAD *Cryptoblepharus sp. *Emoia atrocostata *Emoia caeruleocauda *Lamprolepis smaragdina NGEBEDANGEL (0.1 km NW of Ngeruktabel) Emoia caeruleocauda Lamprolepis smaragdina NGEDBUS (0.6 km N of Beliliou) *Ramphotyphlops braminus CAS 19077 NGELSIBEL (ca. 1 km N of Mecherchar) Laticauda colubrina NGEMLIS ISLANDS BAILECHESENGEL Platymantis pelewensis (calling 1/97) Dendrelaphis sp. (sight record 1/97) IILBLAU Gehyra brevipalmata Lepidodactylus sp. Cryptoblepharus sp. *Emoia caeruleocauda (sight record 1/97) NGERCHEU (4 km N of Beliliou; all specimens KUZ) *Platymantis pelewensis *Gehyra brevipalmata *Gekko sp. *Hemidactylus frenatus *Lepidodactylus lugubris *Lepidodactylus moestus *Emoia caeruleocauda *Eugongylus sp. *Lamprolepis smaragdina *Lipinia of. L. noctua *Mabuya sp. *Candoia carinata *Dendrelaphis sp. *Ramphotyphlops acuticaudus NGERDIS (off Palau Pacific Resort, Ngerekebesang; connected to mainland by jetty) Lepidodactylus lugubris *Emoia atrocostata (CAS 184016) *Lamprolepis smaragdina (CAS 184017) NGERDUAIS (off SE Babeldaob, 100 m SW Garudawisei Point) Gehyra brevipalmata Lepidodactylus moestus Gekko sp. Candoia carinata Dendrelaphis sp. *Laticauda colubrina [No. 1999] HERPETOLOGICAL MONOGRAPHS 79 NGERECHONG (2.3 km S of Mecherchar) Gehyra mutilata NGERECHUR (ca. 2 km off N point Babeldaob) Platymantis pelewensis Gehyra oceanica Gekko sp. Lepidodactylus moestus Emoia caeruleocauda Emoia jakati Lamprolepis smaragdina Candoia carinata Laticauda colubrina NGEREKEBESANG Bufo marinus Platymantis pelewensis Gehyra brevipalmata Gehyra mutilata Gekko sp. Hemidactylus frenatus Hemiphyllodactylus cf. H. typus Lepidodactylus lugubris Lepidodactylus moestus Carlia cf. C. fusca Cryptoblepharus sp. Emoia atrocostata Emoia caeruleocauda Lamprolepis smaragdina Lipinia cf. L. noctua Candoia carinata *Cerberus rynchops Dendrelaphis sp. NGERMALK (between Oreor and Malakal, connected by causeway) Hemidactylus frenatus Lepidodactylus lugubris Lepidodactylus moestus Carlia cf. C. fusca Lamprolepis smaragdina Sphenomorphus sp. nov. Sphenomorphus scutatus NGERUCHUBTANG (4 km SSW of Ngerechong) *Emoia caeruleocauda NGERUKEUID ISLAND GROUP NGERUKEUID ISLAND (Island 24) *Platymantis pelewensis *Gekko sp. *Eugongylus sp. *Lamprolepis smaragdina *Dendrelaphis sp. *Laticauda colubrina ISLAND 4 *Lepidodactylus moestus ISLAND 12 *Lamprolepis smaragdina ISLAND 13 *Candoia carinata *Laticauda colubrina ISLAND 16 *Lepidodactylus paurolepis *Eugongylus sp. *Lamprolepis smaragdina ISLAND 34 *Lepidodactylus paurolepis NGERUKTABEL Platymantis pelewensis Gehyra brevipalmata Hemiphyllodactylus cf. H. typus Lepidodactylus moestus Lepidodactylus sp. Emoia caeruleocauda Eugongylus sp. Lamprolepis smaragdina Sphenomorphus scutatus Sphenomorphus sp. nov. Candoia carinata Dendrelaphis sp. NGERUR (0.8 km W of Ngerekebesang) Gehyra oceanica Lepidodactylus lugubris Lepidodactylus moestus Emoia caeruleocauda Lamprolepis smaragdina Sphenomorphus scutatus NGESBOKEL (Ngeremeduu Bay, west coast of Babeldaob) Lepidodactylus lugubris Lepidodactylus moestus Emoia impar Lamprolepis smaragdina NGETMEDUCH (ca. 0.2 km north of Oreor, connected by causeway) Platymantis pelewensis Gehyra mutilata Lepidodactylus lugubris Lepidodactylus moestus Carlia cf. C. fusca Cryptoblepharus sp. Emoia atrocostata Emoia caeruleocauda Lamprolepis smaragdina Candoia carinata Ramphotyphlops braminus OREOR Bufo marinus Platymantis pelewensis ] 80 HERPETOLOGICAL MONOGRAPHS No. 13 Gehyra brevipalmata Gehyra mutilata *Gehyra oceanica (CAS-SU 23747, 122393) Gekko sp. Hemidactylus frenatus Hemiphyllodactylus cf. H. typus Lepidodactylus lugubris Lepidodactylus moestus Anolis carolinensis Carlia cf. C. fusca Cryptoblepharus sp. Emoia atrocostata Emoia caeruleocauda Eugongylus sp. Lamprolepis smaragdina Lipinia cf. L. noctua *Lipinia leptosoma Mabuya sp. Sphenomorphus sp. nov. Sphenomorphus scutatus *Varanus cf. v. indicus (CAS 154150) Candoia carinata Cerberus rynchops Dendrelaphis sp. Laticauda colubrina Ramphotyphlops acuticaudus Ramphotyphlops braminus Crocodylus porosus *Eretmochelys imbricata (CAS 154194) PKULAKLIM (ORRAK) (ca. 0.6 km SW of Ngerduais) Bufo marinus Gehyra brevipalmata Emoia caeruleocauda Lamprolepis smaragdina Candoia carinata Laticauda colubrina TOIRACHOLODOCHOEL (island name from Brown and Fehlmann, 1958:5 as "a small island at the north end of Malakal") *Lipinia leptosoma UCHULAMERADEL (southernmost of eastern islands, mouth of Ngeremeduu Bay) Lepidodactylus moestus UCHULANGAS (connected to Oreor and Ngermalk by causeway) Hemidactylus frenatus Lepidodactylus lugubris Lepidodactylus moestus Carlia cf. C. fusca Emoia atrocostata ULEBSECHEL Platymantis pelewensis Gehyra brevipalmata Gekko sp. Hemiphyllodactylus cf. H. typus Nactus sp. Emoia atrocostata Emoia caeruleocauda Eugongylus sp. Lamprolepis smaragdina Sphenomorphus scutatus Dendrelaphis sp. ULONG Platymantis pelewensis Gehyra brevipalmata Gekko sp. Lepidodactylus sp. Nactus sp. Emoia caeruleocauda Eugongylus sp. Lamprolepis smaragdina Lipinia cf. L. noctua Sphenomorphus scutatus Dendrelaphis sp. Laticauda colubrina SOUTHWEST ISLANDS FANA ISLAND *Gehyra sp. *Perochirus sp. *Lamprolepis smaragdina HELEN (Helen Reef) *Emoia jakati MERIR ISLAND *Nactus cf. N. pelagicus *Emoia impar *Lamprolepis smaragdina PULO ANNA *Gehyra sp. *Lepidodactylus lugubris *Lepidodactylus moestus *Emoia sp. *Emoia impar *Lamprolepis smaragdina *Lipinia cf. L. noctua *Mabuya sp. SONSOROL ISLAND *Gehyra sp. *Emoia sp. *Emoia impar *Lamprolepis smaragdina *Mabuya sp. TOBI ISLAND *Gehyra sp. *Gehyra mutilata *Emoia jakati *Lamprolepis smaragdina [No.