A Chronology of Middle Missouri Plains Village Sites By Craig M. Johnson with contributions by Stanley A. Ahler, Herbert Haas, and Georges Bonani Smithsonian Institution Scholarly Press Smithsonian Institution Scholarly Press sm i th son i a n con t r i b u t i on s to zoo logy ? n umb e r 6 3 1 Speciation and Dispersal in a Low Diversity Taxon: The Slender Geckos Hemiphyllodactylus (Reptilia, Gekkonidae) George R. Zug SerieS PublicationS of the SmithSonian inStitution Emphasis upon publication as a means of ?diffusing knowledge? was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: ?It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.? 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Manuscripts submitted for series publication are received by the Smithsonian Institution Scholarly Press from authors with direct affilia- tion with the various Smithsonian museums or bureaus and are subject to peer review and review for compliance with manuscript preparation guidelines. General requirements for manuscript preparation are on the inside back cover of printed volumes. For detailed submissions require- ments and to review the ?Manuscript Preparation and Style Guide for Authors,? visit the Submissions page at www.scholarlypress.si.edu. sm i th son i a n con t r i b u t i on s to zoo logy ? n umb e r 6 3 1 Speciation and Dispersal in a Low Diversity Taxon: The Slender Geckos Hemiphyllodactylus (Reptilia, Gekkonidae) George R. Zug washington d.c. 2010 ABSTRACT Zug, George R. Speciation and Dispersal in a Low Diversity Taxon: The Slender Geckos Hemiphyllodactylus (Reptilia, Gekkonidae). Smithsonian Contributions to Zoology, number 631, xi + 70 pages, 25 figures, 7 tables, 2010.?Hemiphyllodactylus is a genus of small geckos occurring widely, although uncommonly seen, throughout the Indo-Pacific islands and South Asia. These geckos consist of both bisexual and unisexual spe- cies. The unisexual species, Hemiphyllodactylus typus, the most widespread of these geckos, apparently at- tained its Polynesian to Mascarene distribution (invasion) through accidental human transport. The bisexual species have much smaller distributions, geographically restricted to island groups or limited continental areas. Until the early 1990s, most bisexual populations were considered subspecies of H. typus. In the last two decades, herpetologists have regularly used species epithets proposed for the region under their inves- tigation. This resurrection of species names has occurred largely without explanation or taxonomic study. This study examines the morphology of Hemiphyllodactylus throughout its known range, using 13 regional samples, first examining the differentiation of unisexual and bisexual populations and individuals, then the possibility of regional differentiation among the different bisexual populations. Variation and consistency in morphology in and among the regional sample identify the existence of a wide-ranging unisexual species, H. typus, and at least eight geographically restricted bisexual species. Available museum specimens for some regions are adequate to characterize eight bisexual species, H. aurantiacus, H. ganoklonis n. sp., H. harterti, H. insularis, H. larutensis, H. margarethae, H. titiwangsaensis n. sp., and H. yunnanensis. Potentially unique bisexual populations occur in Hong Kong, southern Indochina, Borneo, and Sri Lanka, but samples are too small to adequately characterize these populations. The origins and evolution of the species are examined, and the study concludes with a taxonomy for the identified species. Cover image: Palauan slender gecko Hemiphyllodactylus ganoklonis. (Drawing by Molly Dwyer Griffin.) Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS P.O. Box 37012, MRC 957 Washington, D.C. 20013-7012 www.scholarlypress.si.edu Text and images in this publication may be protected by copyright and other restrictions or owned by indi- viduals and entities other than, and in addition to, the Smithsonian Institution. Fair use of copyrighted mate- rial includes the use of protected materials for personal, educational, or noncommercial purposes. Users must cite author and source of content, must not alter or modify content, and must comply with all other terms or restrictions that may be applicable. Users are responsible for securing permission from a rights holder for any other use. Library of Congress Cataloging-in-Publication Data Zug, George R., 1938? Speciation and dispersal in a low diversity taxon : the slender geckos Hemiphyllodactylus (Reptilia:Gekkonidae) / George R. Zug. p. cm. ? (Smithsonian contributions to zoology ; no. 631) Includes bibliographical references and index. 1. Hemiphyllodactylus. I. Title. QL666.L245Z84 2010 597.95'2?dc22 2010042310 ISSN: 0081- 0282 (print); 1943- 6696 (online) The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48?1992. Dedication I dedicate this taxonomic study to Jay M. Savage for the excellence of his half-century of biogeographic and systematic research and in appreciation for his professional support?often ?behind the scenes??and friendship throughout my herpetological career. Contents LIST OF FIGURES  vii LIST OF TABLES  ix PREFACE  xi INTRODUCTION  1 Nomenclatural History  1 MATERIALS AND METHODS 6 CHARACTER ANALYSIS: RESULTS AND DISCUSSION 7 Baseline Estimate of Intra-Observer Variation  7 Recognition of Unisexual and Bisexual Populations  8 Unisexual?Visceral Anatomy  8 Unisexual?Morphometry  9 Unisexual?Scalation  12 Unisexual?Coloration  14 Regional Variation among Bisexual Populations  15 Bisexual?Visceral Anatomy  15 Bisexual?Morphometry  16 Bisexual?Scalation  20 Bisexual?Coloration  25 GEOGRAPHY AND TAXONOMY 29 Regional Patterns of Morphology and Speciation  29 General Observations  29 Morphological Differentiation  29 Taxonomic Decisions and Geography  32 Species Accounts  35 Key to the Species of Hemiphyllodactylus  52 ACKNOWLEDGMENTS 55 v i   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY APPENDIX 1: CHARACTER DEFINITIONS 57 APPENDIX 2: SPECIMENS EXAMINED 59 APPENDIX 3: STATISTICAL ANALYSES 63 REFERENCES 65 INDEX 69 Figures 1. Type localities for the available names of species currently assigned to genus Hemiphyllodactylus  3 2. Visceral pigmentation of Hemiphyllodactylus species  8 3. Contrasting habitus of adult Hemiphyllodactylus  11 4. Principle component graphs of unisexual and bisexual adult females of sunda Hemiphyllodactylus  11 5. Frequency distribution of precloacal?femoral pores of unisexual Hemiphyllodactylus typus  13 6. Dark and light phases of coloration in Hemiphyllodactylus typus  15 7. Morphology of the chin scales in various populations of Hemiphyllodactylus  22 8. Cloacal spur morphology in Hemiphyllodactylus  23 9. Digital lamellae morphology in select species of Hemiphyllodactylus  23 10. Precloacal?femoral pore morphology of Hemiphyllodactylus  24 11. Coloration of select Hemiphyllodactylus taxa  27 12. Types of Bingtang slender gecko  28 13. Holotype of Hemiphyllodactylus typus Bleeker  36 14. Geographic occurrence of Hemiphyllodactylus typus  38 15. Syntypes of Hemiphyllodactylus aurantiacus Beddome  39 16. Geographic occurrence of Hemiphyllodactylus aurantiacus and H. yunnanensis  40 17. Hemiphyllodactylus ganoklonis from Ulebsechel Island, Palau  41 18. Holotype of Hemiphyllodactylus ganoklonis from Ulebsechel Island, Palau  42 19. Geographic occurrence of Hemiphyllodactylus ganoklonis  43 20. Geographic occurrence of Hemiphyllodactylus harterti, H. margarethae, H. titiwangsaensis, and Borneo bisexuals  45 21. Holotype of Hemiphyllodactylus insularis Taylor  46 v i i i   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY 22. Geographic occurrence of Hemiphyllodactylus insularis  47 23. Types of Hemiphyllodactylus margarethae Brongersma  48 24. Types of Hemiphyllodactylus titiwangsaensis  50 25. Lectotype of Gehyra yunnanensis Boulenger  51 Tables 1. Available names for populations and species of Hemiphyllodactylus geckos  6 2. Summary statistics on select characters of unisexual Hemiphyllodactylus samples  10 3. Summary statistics on select mensural characters of adult females of the bisexual Hemiphyllodactylus samples  17 4. Summary statistics on select metric characters of adults of the bisexual Hemiphyllodactylus from southern Asia  19 5. Comparison of some mensural characters of adult females among the Chinese populations of Hemiphyllodactylus yunnanensis  20 6. Summary statistics on select coloration and scalation characters of juveniles and adults of the bisexual Hemiphyllodactylus samples  21 Appendix 1 A1.1. Abbreviations and definitions for characters examined  58 Preface M y fascination with Hemiphyllodactylus began during a study of Fijian lizards. Often, the presence or absence of secreting pre- cloacal and/or femoral pores is used to determine the sex of adult geckos: males possess them, females do not. Although a reliable assumption for some gekkonid and iguanid lizards, a chance observation on an adult Fijian Hemiphyllodactylus typus showed its potential for incorrect sex determination. The Fijian specimen had well-developed pores, yet I remained uncertain of sex even though its gonads appeared to be ovaries. Histology of a gonad revealed developing follicles, thus the specimen was an adult female. Other adult H. typus from Oceania had pores, and examination of their gonads revealed that all were females. This evidence suggested that all Pacific H. typus populations are unisexual (Zug, 1991). Further, this discovery caused me to con- tinue my examination of Hemiphyllodactylus specimens and led to an observa- tion that all individuals from coastal localities from Hawaii and Tahiti westward to New Guinea and those of the Mascarenes share the typus morphotype and are females. Not all Hemiphyllodactylus populations, however, are unisexual. The bi- sexual populations typically occur inland in forested situations from Palau and the Philippines to Sri Lanka and the Eastern Ghats of India. There are a variety of names available for these populations (Kluge, 2001): insularis, harterti, yun- nanensis, aurantiacus (east to west); and other available names not listed by Kluge. My primary goal here is to examine morphological variation among all populations of Hemiphyllodactylus and to address the systematics issues that arise from this study. Speciation and Dispersal in a Low Diversity Taxon: The Slender Geckos Hemiphyllodactylus (Reptilia, Gekkonidae) INTRODUCTION Hemiphyllodactylus are small, inconspicuous geckos but incredibly wide- ranging in the Indo-Pacific realm. The H. typus morphotype occurs from the Mascarenes eastward through southern Asia to eastern Polynesia and Hawaii. Throughout this broad distribution, these geckos are not commonly seen, even by biologists looking for them; thus they have attracted little attention by her- petologists and hobbyists. How does such an inconspicuous gecko attain such a broad occurrence? Human transportation seems the obvious answer, although the subsequent ques- tions of how, why, and when are largely unanswered. The search for a data- supported answer is one of the goals of this study. The major goal is to uncover the diversity of this taxon and to place this diversity in a firm taxonomic setting. NomeNclatural History Bleeker (1860) was the first naturalist to recognize the uniqueness of this small gecko. He described his Sumatran gecko as a new species and genus. His description is adequate, and the survival of the type specimen assures that Hemi- phyllodactylus typus Bleeker is associated correctly with a gecko population today. Although the assignment of the name to a specific taxon is unambigu- ous, confusion exists about the type locality and the nomenclatural status of Ptyodactylus gracilis. These difficulties arise from the last sentence in Bleeker?s description (1860:237): ?Ik bezit eene afbeelding dezer soort; afkomstig van de voormalige Natuurkundige kommissie, voorzien van den naam Ptyodacty- lus gracilis en naast welke aangeteekend is, dat de soort ook op den Goenong Parong (Java) leeft.? My interpretation (based on a translation by T. Ulber, in litt.) is that Bleeker is telling the reader that the yellow underside of the tail George R. Zug, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 162, Washington, D.C. 20013-7012, USA; zugg@si.edu. Manuscript received 31 August 2009; accepted 5 May 2010. 2   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY and other characteristics (from the preceding descriptive sentence) are seen in an illustration of his new species that is labeled P. gracilis; the source of this illustration is presumably an unpublished report of the Natuurkundig Commission. As interpreted by Wermuth (1965) in his gekkonid checklist, Bleeker was not offering a substitute name. Kluge (1968:342) noted this error of interpretation, and he proposed correctly that ?a picture (drawing) of his [Bleeker?s] new species . . . bore the name Ptyodactylus gracilis.? Ulber?s translation shows that Bleeker was not- ing that H. typus also occurred at Goenong Parong (Java) based on an unpublished illustration. Ptyodactylus gracilis is thus a nomen nudum and unavailable. Kluge (1968:342) noted that the association of Goe- nong Parong with P. gracilis ?led Smith (1935) and Wer- muth (1965) to incorrectly consider the type locality of typus as Goenong Parong, Java.? Kluge?s identification of the type locality (Figure 1) with the title of Bleeker?s article is correct, that is, Agam, a locality in central Sumatra (see gazetteer in David and Vogel, 1996). Bleeker was not the only naturalist to recognize the uniqueness of this gecko. Bavay discovered this gecko on buildings during his field work in New Caledonia. He rec- ognized it as a new species, Platydactylus crepuscularis, in his catalog of New Caledonian reptiles (Bavay, 1869), ap- parently unaware of Bleeker?s description. While he noted this gecko?s similarity to Lepidodactylus lugubris, at least to the description of that taxon provided by Dum?ril and Bibron, Bavay?s description explicitly characterized P. cre- puscularis as a Hemiphyllodactylus typus Bleeker; thus P. crepuscularis is a junior subjective synonym. The history of this name and its type specimens is detailed in Bauer?s (1994) comment sections for H. typus and L. lugubris. I wish to examine only one aspect?the holotype or syntypes of P. crepuscularis. Bavay typically gave the dimension of a single specimen in his species accounts, whether describ- ing a new or established taxon. Boulenger (1883:123), however, suggested Bavay had two specimens: ?and two others, male and young, the types of the species, commu- nicated to me by M. Bavay.? Was the ?communicated? a letter with data on the specimens or actual specimens sent to Boulenger? If the latter, they were not cataloged in the British Museum, because Boulenger (1885) listed only the two Benchley specimens that he had mentioned in his 1883 description. There is no evidence that Bavay depos- ited the type(s) in the Paris Museum, because Sauvage?s (1879) subsequent description of a type was based on a specimen of L. lugubris (see Bauer, 1994). Hence the type of P. crepuscularis is lost, but fortunately Bavay?s descrip- tion clearly refers to H. typus Bleeker. Major Beddome, a forestry officer in Madras (pres- ent-day Chennai, India), collected a variety of reptiles and described them in 1870. One of them, Hemidacty- lus aurantiacus Beddome, was a Hemiphyllodactylus spe- cies from mid-elevation in the Shevaroy Hills. Nothing in Beddome?s characterization identifies the new species unequivocally as Hemiphyllodactylus. Boulenger?s (1885) description is more detailed, and his placement with Lepi- dodactylus was a better assessment of aurantiacus? affini- ties. Boulenger also noted that the type series consisted of many adult males, females, and juveniles. G?nther (1872) described a typus gecko from the ?East-Indian archipelago? as Spathodactylus mutilatus. The generic and specific descriptions and the illustration of the fore- and hindfeet readily identify the holotype as Hemiphyllodactylus and likely H. typus. Although G?n- ther did not identify the source of the specimen, Boulenger (1885) did?Dr. Bleeker. This information suggests that the types of H. typus Bleeker and S. mutilatus G?nther are the same specimen; thus the latter name is a junior objective synonym of the former. Because Boulenger in- cluded neither Hemiphyllodactylus nor typus as names in his catalog, it indicates that neither he nor G?nther was aware of Bleeker?s description. Boulenger (1885) did rec- ognize that G?nther?s Spathodactylus was a homonym for a fish and provided a new generic name Spathoscalabotes. Subsequently, Malcolm Smith (1935) listed the type local- ity of S. mutilatus as Agam, Sumatra, in his synonymy of H. typus; this restriction is correct owing to Bleeker?s original source of the specimen, although in the same syn- onymy, Smith incorrectly gave Java as the type locality of H. typus Bleeker. As noted above, Boulenger (1885) was apparently un- aware of Bleeker?s description of Hemiphyllodactylus and H. typus, because these names are absent from his cata- log. He placed crepuscularis, ceylonensis, and aurantiacus in the genus Lepidodactylus and continued to recognize G?nther?s mutilatus as a monotypic taxon although cor- recting the generic homonymy. Boulenger followed the species account of Lepidodac- tylus crepuscularis (=Platydactylus crepuscularis Bavay) with an exceedingly brief description of Lepidodacty- lus ceylonensis. The description identifies the specimen as Hemiphyllodactylus only by Boulenger?s (1885:164) statement: ?This species resembles exactly the preceding [L. crepuscularis] in proportions, pholidosis, and color- ation.? Boulenger?s illustration is suggestive of Hemiphyl- lodactylus, but it would fit other geckos as well. Stejneger (1899) provided the first review of the Ha- waiian terrestrial reptiles. Of the seven lizard species then n um b e r 6 3 1   ?   3 FI G U R E 1 . T yp e lo ca lit ie s fo r th e av ai la bl e na m es (s ee T ab le 1 ) f or th e sp ec ie s cu rr en tly a ss ig ne d to g en us H em ip hy llo da ct yl us . A bb re vi at io ns : d , H . y un na ne ns is d us ha ne ns is ; j , H . y . j in pi ng en si s; l, H . y . l on gl in ge ns is . 4   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY occurring on these islands, only one appeared to represent a new species, Hemiphyllodactylus leucostictus Stejneger. Stej- neger gave a thorough description and included illustrations of the head, pelvic area, and hindfoot of his new species. He stated that the differences of the Hawaiian specimens to H. crepuscularis, H. ceylonensis, and H. typus were slight but real, hence appropriate to recognize a new species. Werner (1900) received a single small gecko from Malakka (now Malaysia) and noted its distinctiveness. He named it Lepidodactylus Harterti after the collector and ornithologist Ernest Johann Otto Hartert. He highlighted the presence of strongly V-shaped lamellae (11 on fourth toe) and proposed that it was closely related to Lepido- dactylus lugubris, although the inner digit was less well developed than in L. lugubris. Boulenger examined a small collection of amphibians and reptiles from the Larut Hills, Perak, Malaysia, many of which were new. His descriptions (Boulenger, 1900) of the new species included a new gecko, Gehyra larutensis. It is unclear why he had shifted his generic placement of typus-like geckos. Using the same generic assignment three years later, Boulenger (1903) described another typus-like gecko from China, Gehyra yunnanensis. In both descrip- tions, he mentioned the low number of chevron-like la- mellae on the digits characteristic of Hemiphyllodactylus geckos. He continued to use Gehyra for these geckos in his book on the Malayan herpetofauna (Boulenger, 1912). More importantly, Boulenger (1912:48) noted that G. larutensis ?may prove to be identical with this species [H. harterti Werner, 1900].? He further noted that Wer- ner?s Malakka type locality should be replaced by Gunong Inas, a site where Hartert collected birds in 1888 (Hartert, 1901, 1902). The nomenclatural significance of this tenta- tive correction [reassignment] is addressed in the Taxo- nomic Decisions and Geography section. In Reptiles of the Indo-Australian Archipelago, de Rooij (1915) recognized Hemiphyllodactylus typus and Lepidodactylus ceylonensis. She used the digital lamellar morphology and rudimentary and clawless first digits as the main diagnostic features for Hemiphyllodactylus. De Rooij was the first researcher to recognize a broad distribution for H. typus and reported its occurrence widely in Sumatra and Java and on Borneo. She used Lepidodactylus ceylonensis as an all-encompassing taxon defined by a rudimentary first finger and with a distribution from Borneo through Java, Sumatra, and Nicobar to Myanmar and Sri Lanka. Taylor (1918) recognized that the Philippine typus gecko as Hemiphyllodactylus insularis. He compared it only to H. leucostictus Stejneger, noting that he was un- certain that the Hawaiian gecko was ?actually distinct.? Presumably, he meant distinct from H. typus; however, he does not mention Bleeker?s taxon or any of the other typus-like taxa. Taylor was the first systematist to discuss variation within a type series (Mindoro) of a typus-like taxon as well as presenting geographic variation of speci- mens from other islands in the Philippines. He was also the first author to provide natural history observations, noting that females lay two adhesive eggs beneath tree bark and that all specimens were beneath bark on seaside trees. In 1924, Barbour received specimens of Hemiphyllo- dactylus yunnanensis from The Reverend John Graham, who had provided the original series of specimens to Boulenger. Barbour was so struck by the morphology of foot lamellae that he proposed a new generic name for this taxon, Cainodactylus. Barbour stated that Dr. Stej- neger agreed with him on the uniqueness of the foot morphology, but Barbour seemingly was so focused on the differences of his specimens to species of Gehyra and Hemidactylus that he overlooked Stejneger?s description and illustrations of H. leucostictus and consequently cre- ated a synonym. Brongersma (1931) described Hemiphyllodactylus margarethae from four Sumatran specimens representing two montane localities. He recognized this species? affin- ity and differentiated it from H. typus and H. ceylonensis. Subsequently, Brongersma (1932) published an evaluation of the nomenclature and characterization of Hemiphyllo- dactylus and its species. He recognized only two species: H. aurantiacus and H. typus. Although he recognized H. aurantiacus, Brongersma examined only two specimens and purposefully kept his remarks brief. Thus his ?Notes? refer mainly to H. typus, in which he synonymizes H. ceylo- nensis, H. crepuscularis, H. leucostictus, H. insularis, and H. margarethae. He noted that M. A. Smith had indepen- dently reached the same conclusions. Brongersma reached his conclusion through the evaluation of three characters regularly used to diagnose Hemiphyllodactylus species: (1) denticulate digits, (2) number of precloacal pores, and (3) if present, number of femoral pores. He concluded that denticulation was slight in most specimens and ?purely in- dividual? (Brongersma, 1932:214). He recognized the dif- ficulty of distinguishing pits and pores, noted the absence of femoral pores in some males, and concluded that the number of pores (precloacal and femoral) ?seems to be of no value in this genus? (Brongersma, 1932:216). M. A. Smith?s conclusions, revealed to Brongersma in a letter, were subsequently promulgated in his work on the lizards of British India (Smith, 1935). Therein, Smith recognized two species of Hemiphyllodactylus: H. typus typus, H. typus aurantiacus; and H. yunnanensis. The n um b e r 6 3 1   ?   5 nominate subspecies included as synonyms all the species mentioned in the preceding paragraphs except H. laruten- sis, thereby giving H. typus a distribution from Sri Lanka eastward into Oceania. Hemiphyllodactylus t. aurantiacus retained a southern India distribution. Hemiphyllodactylus yunnanensis was identified with a Yunnan, northern Laos, and northern Myanmar distribution. In a footnote, Smith (1935:109) proposed that H. yunnanensis was ?perhaps a northern representative of the Malayan Hemiphyllodacty- lus larutensis (Boulenger).? Smith (1933) in an article that likely was preparatory to his 1935 catalog examined the confusion of species assignment to Hemiphyllodactylus. Therein he provided a concise definition of the genus and a list of three included species: typus, yunnanensis, and harterti. He noted that the latter name had appeared three months before Gehyra larutensis Boulenger. Bourret (1937) described Hemiphyllodactylus typus chapaensis from northern Vietnam (Chapa, Tongking). He noted that it resembled H. yunnanensis but that his taxon was not greatly different from H. typus, hence his assignment to subspecific status. His description included five detailed illustrations of the type. After Bourret, Hemiphyllodactylus occurred irregu- larly in the herpetological literature until the 1960s, typi- cally in regional lists, reappearing in Taylor?s lizards of Thailand (Taylor, 1963) with a full description and Wer- muth?s (1965) checklist of all gekkonid lizards. Wermuth recognized three species (larutensis, typus, and yunna- nensis) and three subspecies of H. typus (nominate form, aurantiacus, and chapaensis). Kluge (1968) addressed the relationships of Hemiphyllodactylus as well as com- menting on nomenclatural matters; these matters were discussed above. He considered typus and yunnanensis as full species of Hemiphyllodactylus and left the status of aurantiacus, chapaensis, and harterti for additional inves- tigation. Kluge considered Hemiphyllodactylus as a sister group of Lepidodactylus. This hypothesis returns concep- tually to Boulenger?s catalog treatment, although retaining typus and its congeners as a separate genus (lineage). Wermuth (1966) reexamined a gecko, Platydactylus minutus Giebel 1862, captured in Baltic amber. He pro- posed that the specimen was a Hemiphyllodactylus typus. His Figure 2 of the right forefoot shows subdigital lamel- lae similar to those of H. typus; however, the dorsal view of the entire gecko (Wermuth, 1966: fig. 1) is not typus- like. The head, neck, and body are robust and not elon- gated. The fore- and hindlimbs are large, long, and would overlap one another if laid along the trunk. With this habi- tus, Platydactylus minutus Giebel is not a synonym of H. typus or vice versa. Through the 1970s and 1980s, Hemiphyllodactylus species, mostly H. typus, appeared in assorted publica- tions on regional herpetofaunas. For example, Brown and Alcala (1978) continued the interpretation of H. insularis as a synonym of H. typus in their Philippine gecko catalog. Auffenberg (1980) reviewed the herpetofauna of Komodo and observed that the Komodo H. typus were lightly col- ored and nearly patternless. He described the Komodo population as the subspecies H. t. pallidus. Zhou et al. (1981) examined a large collection of H. yunnanensis from Yunnan, Guizhou, and Guangxi Provinces, China, and observed regional variation in digital lamellae pat- terns. Because the variation was concordant within four regions, they recognized three new subspecies: H. y. du- shanensis, H. y. jinpingensis, and H. y. longlingensis. Lazell (1989) made an unexplained alteration (1989) of leucostictus Stejneger to albostictus in a magazine article. Zug (1991) revealed the unisexual aspect of Fijian and other Oceania populations of H. typus. Bauer?s (1994) checklist of Australian and Oceania gekkonids provided a full syn- onymy of Hemiphyllodactylus typus and an abbreviated review of the various nomenclatural usage and alterations. Manthey and Grossmann (1997) recognized two species (larutensis, typus) of Hemiphyllodactylus in the Sunda area. The two have strikingly different coloration in their illustra- tions and descriptions, confirming the presence of two spe- cies in this area. Their concept of H. typus, however, was as a bisexual species, with males defined by the presence of femoral?precloacal pores. Soon thereafter, Chan-ard et al. (1999) listed four species (harterti, larutensis, typus, and yunnanensis) from Thailand and peninsular Malaysia. Their photographs show variable coloration among the specimens identified as H. harterti and H. larutensis from the Cameron Highlands, Pahang State, Malaysia. Subsequently, Bauer and Das (1999) visited the type locality of Hemidactylus aurantiacus Beddome and cap- tured three adult specimens. Their examination of these specimens and specimens from Malaysia, Philippines, and elsewhere demonstrated that the Shevaroyan geckos had several diagnostic traits that clearly distinguished this midmontane population from other Hemiphyllodactylus typus. On the basis of these consistent differences, they recognized Hemiphyllodactylus aurantiacus as a full spe- cies. Kluge?s most recent gekkonid checklist (Kluge, 2001) similarly returned Hemiphyllodactylus insularis Taylor to specific status but without explanation. Gaulke (2003) briefly examined the nomenclatural history of Philippine Hemiphyllodactylus and, presumably because of the pres- ence of males and females, accepted H. insularis as a dis- tinct taxon from H. typus. The available names for the 6   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY various populations of Hemiphyllodactylus are summa- rized chronologically in Table 1. MATERIALSANDMETHODS Despite the broad distribution of Hemiphyllodacty- lus, the availability of voucher material in museum col- lections is relatively poor. The assembly of an adequate quantity of specimens required access to many collections; the collection names are abbreviated here for subsequent mention in the text. AMNH American Museum of Natural History AMS Australian Museum, Sydney BMNH The Natural History Museum, London BPBM Bernice P. Bishop Museum CAS California Academy of Sciences CM Carnegie Museum of Natural History FMNH Field Museum of Natural History IRSN B Institut royal des Sciences naturelles de Belgique KUZ Kyoto University, Department of Zoology MCZ Museum of Comparative Zoology, Harvard University NMB Naturhistorisches Museum, Basal NMW Naturhistorisches Museum, Wien QM Queensland Museum RMNH Nationaal Natuurhistorisch Museum (formerly Rijkmuseum van Natuurlijke Historie) SAM South Australian Museum SDMNH San Diego Museum of Natural History SMF Natur-Museum u. Forschungs Institut Senckenberg THNHM Thailand Museum of Natural History, National Science Museums UF Florida Museum of Natural History, University of Florida USNM U.S. National Museum (Na- tional Museum of Natural History, Smithsonian Institution) WAM Western Australian Museum WmBeckon William N. Beckon, private collection ZMA Zo?logische Museum, Universiteit van Amsterdam TABLE 1. Available names for populations and species of Hemiphyllodactylus geckos. Type localities presented are from the original descriptions; because many of these localities may not be obvious or known to some readers, the country in which the locality occurs is included in brackets. Date Name Author Type locality 1860 Hemiphyllodactylus typus Bleeker Agam [Sumatra] 1869 Platydactylus crepuscularis Bavay Nouvelle-Cal?donia 1870 Hemidactylus aurantiacus Beddome Shevaroys . . . about Yercaud [India] 1872 Spathodactylus mutilatus G?nther East Indies archipelago 1885 Lepidodactylus ceylonensis Boulenger Ceylon 1899 Hemiphyllodactylus leucostictus Stejneger Kauai, Hawaiian Islands 1900 Lepidodactylus Harterti1 Werner Malakka [Malaysia] 1900 Gehyra larutensis Boulenger Larut Hills [Malaysia] 1903 Gehyra yunnanensis Boulenger Yunnan Fu [China] 1918 Hemiphyllodactylus insularis Taylor Sumagui, Mindoro [Philippine Islands] 1931 Hemiphyllodactylus margarethae Brongserma Fort de Kock, Sumatra 1937 Hemiphyllodactylus typus chapaensis Bourret Chapa [Vietnam] 1980 Hemiphyllodactylus typus pallidus Auffenberg Vai Nggulung, Loho Liang, Komodo 1981 Hemiphyllodactylus yunnanensis dushanensis Zhou et al. Dushan County, Guizhou Province, China2 1981 Hemiphyllodactylus yunnanensis jinpingensis Zhou et al. Jinping County, Yunnan Province, China2 1981 Hemiphyllodactylus yunnanensis longlingensis Zhou et al. Longling County, Yunnan Province, China2 1 Harterti is capitalized as it appears in the original description. 2 Type locality presented in Chinese. n um b e r 6 3 1   ?   7 ZMB Museum f?r Naturkunde, Universit?t zu Humboldt ZMFK Zoologische Forschungsinstitut u. Museum Alexander Koening ZRC Zoological Reference Collection, National University Singapore ZSM Zoologisches Sammlung des Bayerischen Staates I grouped the specimens into 13 regional samples, each representing a putative biogeographic area abbrevi- ated in small capitals and defined as follows: China China and northeastern Myanmar Fiji Fiji and Tonga Hawai Hawaiian Islands India India and Sri Lanka Mascar Mascarenes NCal New Caledonia and Vanuatu NGuin New Guinea and Solomon Islands Palau Republic of Palau Philip Philippine Islands Polyn Polynesia sEasia Thailand (north of Isthmus of Kra) to Vietnam and Hong Kong Sunda Malaysia and Indonesia Taiwan Taiwan and Japan These regional samples vary in size (n = 9?85) and geographic extent. In the latter case, a sample can be ex- amined as two or more subsamples of restricted localities if adequate specimens are available or if intrasample varia- tion indicates a mixed sample. Further, I combined sam- ples and repartitioned specimens when an initial analysis suggested the presence of multiple bisexual taxa in one or more of geographically adjacent samples. Morphological data consist of a combination of mor- phometric and meristic characters. These characters are identified and their abbreviations defined in Appendix 1. Sex and maturity were determined by dissection and examination of the gonads. Maturity criteria were those outlined in Zug et al. (2003). The small size of this taxon seems to have resulted in a high level of inattentiveness to the preservation and positioning of specimens. Contorted specimens, commonly with clenched fore- and hindfeet, made data-gathering challenging and certainly increased the variation in most measurements and scale counts. The data were analyzed by a variety of univariate and multivariate statistics using SYSAT version 11. My goal has been to examine and describe intra- and interpopulational variation as thoroughly as possible considering the variable preservation state of many specimens. The multivariate models were used as exploratory techniques to compare populations and possibly reveal differentiation within and among samples. Explanation of the use of the multivariate analyses and the results are available in Appendix 3. In addition to the abbreviations defined above and in Appendix 1, other symbols and abbreviations used in this publication are defined as follows: alt. altitude CV coefficient of variation DFA discriminant function analysis GPD glycerosphophate dehydrogenase IDH isocitrate dehydrogenase MPI mannose phosphate isomerase PCA principal components analysis SD standard deviation r2 coefficient of determination CHARACTERANALYSIS: RESULTSANDDISCUSSION BaseliNe estimate of iNtra-oBserver variatioN How much of the variation observed in each sample re- sults from the researcher?s data-gathering behavior? Hayek et al. (2001) addressed that question and recommended a repeated measuring protocol to obtain an estimate of this portion of a character?s and a sample?s variation. A single specimen (in this case USNM 563683, female from Palau) was measured and scalation recorded 10 times, each time on a separate day over a period of 6 weeks. Central tendency statistics reveal that for measurements, the coefficients of variation (CVs) range from 0.7% (mean snout?vent length [SVL] = 32.5 mm ? SD 0.24) to 9.3% (mean SnW = 1.1 mm ? SD 0.10). The larger measurements (SVL, TrunkL, HeadL, HeadW?see Appendix 1 for definitions) have the least vari- ation (CV, 0.7?1.9%) relative to the smaller ones (SnEye, NarEye, EyeD, SnW; CV, 3.2?9.3%). The coloration traits were invariant, as were the majority of the scalation char- acters. Ventral (mean = 9.3 mm ? SD 1.16) and 2ToeLm (3.7 mm ? SD 0.48) had the highest CVs (12.5%, 13.1%, respectively); in all other variable traits, CV was <8%. These results provide a baseline for assessing the variation observed in regional samples. Further, these es- timates probably represent the lowest variation for the Hemiphyllodactylus data as they were recorded from a well- preserved and well-positioned specimen and collected under optimal laboratory conditions. While the data 8   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY reported throughout this study were gathered by a single individual (me), they were gathered over two decades (1989?2008) and in a variety of museum situations. As noted above, Hemiphyllodactylus specimens infrequently receive adequate preparation attention when collected. In spite of the inattentive preparations, the subsequent re- sults reveal that intrasample variation is surprisingly low in most characters and samples; nonetheless, the reader is advised to be cautious in over-interpreting reported differ- ences, particularly in small samples and/or where differ- ences are less than 2 times a character?s standard deviation. recogNitioN of uNisexual aNd Bisexual PoPulatioNs Unisexual populations contain only females. Al- though it may be a statement of the obvious, how does one confirm the unisexuality of a population? Because nei- ther data on reproduction of virgin females nor mitochon- drial DNA were available to me, I relied on the absence of males in samples as a hypothesis of unisexuality for popu- lations or sets of populations. On that basis, the Hawaiian (hawai), Polynesian (Polyn), Fijian?Tongan (Fiji), New Caledonian?Vanuatuan (ncal), New Guinean?Solomons (Nguin), Taiwan (taiwan), and Mascarene (Mascar) samples are considered unisexual populations. All other samples contain males, but owing to their manner of as- sembly, some samples, especially the Sundaland (sunda) and Indian?Sri Lankan (india) ones, are likely mixtures of unisexual and bisexual individuals. This situation requires an assumption: Hemiphyllo- dactylus typus Bleeker (BMNH 1946.8.30.83) represents (is) a unisexual species. The holotype is an adult female with precloacal and femoral pore series separated. To test this assumption, the initial character analyses exam- ined variation in and among the Pacific samples (hawai, Polyn, Fiji, ncal, nguin, taiwan) and then when they proved geographically homogeneous compared them to the type specimen. Assuming that this comparison yielded accurate ?diagnostic? data, H. typus specimens were re- moved from the mixed unisexual?bisexual samples prior to examination of bisexual?s within sample and between (interregional) sample variation. Unisexual?Visceral Anatomy Because sex and maturity were determined by dissec- tion, I observed a consistent pattern of differential pigmenta- tion in the viscera of Pacific Hemiphyllodactylus, although I did not report this observation in my study of Fijian lizards (Zug, 1991). All Pacific specimens have the caecum and oviducts heavily pigmented (melanin) (Figure 2A,B). For FIGURE 2. Visceral pigmentation of Hemiphyllodactylus species. All images in ventral view. (A) Pigmented caecum visible externally between pair of oviducal eggs, H. typus, Hawaii (U.S. National Mu- seum [USNM] 570747); (B) pigmented caecum and oviduct, large intestine inflected anteriorly (left side of image), H. typus, Hawaii (USNM 570742); (C) unpigmented caecum and oviduct, H. yun- nanensis, Myanmar (USNM 570734). n um b e r 6 3 1   ?   9 the oviduct the peritoneum sheath likely bears the melanin. The pigmented caecum and oviducts, however, are not con- fined to the unisexual samples. This pigmentation pattern is widespread, but not global, in bisexual samples. The black caecum is often visible through the ventral body wall and skin (Figure 2A). The pigmentation of the oviducts is darkest in virgin females. It is invisible in the greatly stretched oviducts of gravid females. The oviducts become dusky brown once the eggs are expelled. The duct walls retain a flaccid and stretched morphology following the first egg production cycle. The type of Hemiphyllodactylus typus Bleeker was not dissected, so I am unable to confirm the pigmentation of its oviduct or its sex by a direct examination of the reproductive tract. It does not show any thickness at the base of the tail, thereby indicating the absence of hemi- penes. The black caecum is visible through the body wall. Pr?cis. Unisexual Hemiphyllodactylus typus possess darkly pigmented caeca and oviducts. Bisexual Hemiphyllodactylus are variable in this pair of traits. Unisexual?Morphometry hawai is the largest of the Pacific samples and serves as a base to examine levels of variation and differentiation within and among the Pacific samples. Hawai has neither the largest nor smallest adult females. Its mean and me- dian SVL (36.7 and 36.3 mm) match those of Polyn and taiwan and is less than those of the other samples. nguin and Fiji are similar, with mean/median of 40 mm SVL. The smallest adult (29.2 mm SVL) is from Taiwan; the largest (46.1 mm) is from Fiji. The other eight mensural traits show the same pattern of similarity among the samples. Intrasample variance is also similar within and among the Pacific samples, as seen by a comparison of the ranges of the coefficient of variation (CV) with the CV for the hawai datum: SVL 6.5% (hawai datum), 3.0?10.4% (range for the Pacific samples); TrunkL 11.3%, 5.2?11.7%; HeadL 5.0%, 4.6?8.3%; SnEye, 7.0%, 5.0?13.9%; NarEye 7.9%, 6.9?12.8%; EyeD 7.2%, 6.5?11.3%; SnW 10.5%, 5.9?15.5%; HeadW 9.2%, 5.1?12.1%. The higher CVs are associated mainly with taiwan (n = 9), which has the greatest range of adult SVL (29.2?43.6 mm). Mascar is also an all-female sample and presumably represents a unisexual population. Its CVs match those of the Pacific samples (Table 2). Variation and means of the samples, either individually or combined (i.e., total Pacific unisex- ual sample; Pacif), are equivalent. There is no evidence of mensural differentiation among the Pacific insular sam- ples or the distant Mascar sample. Principal components analysis (PCA) and discriminant function analysis (DFA) of the combined Pacific samples, Mascar sample, and the holotype of H. typus similarly reveal a uniform morphol- ogy among these geckos. (See synopsis of PCA and DFA results in Appendix 3.) The Pacific unisexuals and typus holotype are slender, elongate geckos (Figure 3). The proportionately short limbs accentuate the trunk elongation. A proportion of hindlimb length to trunk length would demonstrate this morphol- ogy; however, my preliminary measurements of hindlimb length were extremely variable owing to the difficulty of measuring accurately tiny twisted limbs with fist-like pre- served feet in many specimens. Thus I excluded this trait from subsequent data gathering. The proportion TrunkL/ SVL provides a metric, although a less satisfactory one, for portraying the relative elongation of the trunk. Linear re- gression of the two preceding traits also reveals the degree of elongation through the depression of the slope; how- ever, in contrast to the proportion, variation from linearity (as measured by coefficient of determination, r2) was high, thereby reducing the reliability of the slope as indicator of trunk elongation. The means and standard deviations for TrunkL/SVL (as percentage) among the Pacific typus samples are 52.0% ? 4.0 (hawai), 54.4% ? 3.5 (Polyn), 55.4% ? 4.2 (Fiji), 52.6% ? 1.6 (ncal), 54.0% ? 2.4 (nguin), 52.9% ? 3.9 (Pacif). To place these proportions in the context of other geckos, the mean ? SD TrunkL/SVL for Gehyra oceanica is 42.7% ? 3.3 (n = 113), Hemidac- tylus frenatus is 42.0% ? 2.8 (n = 21), and Lepidodactylus lugubris is 44.1% ? 3.2 (n = 149). Trunk length is over 50% of total SVL in the H. typus samples and distinctly less than half of SVL in other Pacific geckos. Part of this proportional difference derives from a smaller head in H. typus, that is, 21.3% ? 0.9 HeadL/SVL (Pacif), as com- pared to 26.7% ? 1.6 for G. oceanica, 26.4% ? 1.6 for H. frenatus, and 24.2% ? 1.7 for L. lugubris. Combining the head and trunk proportions shows that neck length is pro- portionately shorter in H. typus in contrast to the visual impression of the attenuate habitus of H. typus. Do these proportions permit the segregation of uni- sexual H. typus from bisexual Hemiphyllodactylus? Any of the regional samples might contain both unisexual and bisexual species. Because the Sunda most certainly is mixed, it is the appropriate sample on which to test the morphometrics for differentiation. The sunda sample (n = 83) contained 55 adult females of which 39 were identified as H. typus during data col- lection (based on coloration and pigmentation of caecum and oviducts), 15 males and 7 females as unknowns or uncertains, the holotypes each of H. larutensis (Boulenger) (adult male) and H. harterti (Werner) (adult female), ho- lotype (adult male) and three paratypes (2 adult females, 10   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY TABLE 2. Summary statistics on select characters of unisexual Hemiphyllodactylus samples. Statistical values are mean (mensural traits) or median (scalation) ? standard deviation (SD), range of minimum to maximum, and coefficient of variation (CV); and modes and frequency (%) of occurrence (finger and toe lamellae). Sample sizes (n) are mature females and total sample, respectively; statistics derive solely from adults for mensural traits, and from juveniles and adults for scalation ones. The n value is the total number of specimens examined for each locality sample; the actual statistic may have derived from fewer individuals because not all characters could be mea- sured or counted in all specimens. Character abbreviations are defined in Appendix 1. Sample (n) typus Holotype Hawaiian Islands Total Pacific unisexual sample Mascarene group Character and statistic (1) (37, 42) (99, 118) (8, 11) SVL Mean ? SD 43.3 36.7 ? 2.28 38.0 ? 2.91 38.1 ? 1.00 Range 32.4?42.8 29.2?46.1 38.1?40.9 CV 6.2% 7.6% 2.5% TrunkL Mean ? SD 22.4 19.4 ? 1.94 20.2 ? 2.37 20.7 ? 1.31 Range 15.1?23.9 14.0?28.0 19.2?23.1 CV 10.0% 11.8% 6.3% HeadL Mean ? SD 9.1 7.9 ? 0.38 8.1 ? 0.55 8.3 ? 0.26 Range 7.1?8.8 6.6?9.9 7.8?8.6 CV 4.8% 6.8% 3.2% SnEye Mean ? SD 3.7 3.3 ? 0.22 3.3 ? 0.29 3.5 ? 0.16 Range 2.7?3.7 2.3?4.1 3.3?3.8 CV 6.8% 8.7% 3.5% PostocSpt Median ? SD a 4 ? 0.80 3 ? 0.95 3.2 ? 1.10 Range 1?5 1?5 2?5 CV 22.6% 29.7% 33.7% SnS Median ? SD 3 2 ? 1.01 2 ? 0.81 2 ? 0.50 Range 1?5 1?5 2?3 CV 40.7% 37.7% 22.1% Suplab Median ? SD 11 11 ? 0.85 11 ? 0.94 11 ? 0.82 Range 10?14 9?14 10?13 CV 7.6% 8.3% 7.3% Chin Median ? SD 13 12 ? 0.99 11 ? 1.08 10 ? 0.71 Range 10?14 9?14 10?12 CV 8.4% 9.4% 6.7% Dorsal Median ? SD 13 14 ? 1.53 15 ? 1.64 15.0 ? 1.48 Range 12?18 12?19 13?17 CV 10.6% 11.0% 10.0% CloacS Median ? SD 2 3 ? 0.80 2 ? 0.79 2 ? 1.00 Range 1?5 0?5 1?4 CV 29.6% 34.0% 50.0% 4ToeLm Median ? SD 4 5 ? 0.33 5 ? 0.33 4 ? 0.47 Range 4?5 4?5 4?5 CV 6.8% 6.7% 8.4% FingerLm b Modal values 3-3-4-3 3-4-4-4 3-4-4-4 3-4-4-3 c Frequency 42.5% 47.7% 27.3% ToeLm b Modal values 4-4-4-4 4-4-5-4 4-4-5-4 4-4-5-4 Frequency 51.2% 50% 36.4% a This value/character is unknown in the holotype because of fading. b Lamellae formulae represent the most frequent formula (mode) for each sample and the percent of the sample with this formula (frequency). c Two finger formulae share 27%; the second is 3-4-4-4. n um b e r 6 3 1   ?   1 1 immature male) of H. margarethae Brongersma, and 11 (5 adult females, 6 adult males) other Malaysian ?harterti.? The sunda H. typus included Bleeker?s holotype. A PCA of the body proportions of all sunda females yielded a cluster- ing of H. typus in the bottom quadrant of the PCA scores graph (Figure 4). EyeD/NarEye, NarEye/HeadL, and SnW/ HeadW had the strongest loading (0.63?0.88) on the first component, HeadW/SVL and HeadL/SVL strongest load- ing (0.95, 0.81) on the second component, and SnW/HeadL and SnW/HeadW on third component (0.88, 0.66). The first three components accounted for 68% of the total vari- ance (80%, with the inclusion of the fourth component). These results also place the H. typus holotype (BMNH 1946.8.3) within the H. typus cluster (Figure 4). The bi- sexuals lie principally outside the typus cluster in the upper left quadrant. One of the H. margarethae paratypes (ZMA 11096) is a distant outlier (upper right quadrant) from all other Hemiphyllodactylus females. The other H. marga- rethae paratype (IRNS9338B) is on the outer edge of the typus cluster. Overall, these results indicate a difference in head shape (PCA 1) and relative size (PCA 2) between the unisexual H. typus and the bisexual species. I note that I identified all Bornean females as H. typus and all Bornean males as unknown bisexuals. All Bornean females lie within the typus cluster. The holotype of H. typus (0.288, ?1.362) lies within the lower half of the H. typus cluster (Figure 4B). The assignment of sunda specimens to H. typus was based on the presence of pigmented caeca and oviducts and, to a lesser extent, on dorsal coloration. The unknowns and bisexual type specimens had either unpigmented caeca or both oviducts and caeca unpigmented (Types were not dissected so oviducal pigmentation is usually unknown for FIGURE 4. Distribution of unisexual and bisexual adult females of sunda Hemiphyllodactylus in multivariate (principal components) morphometric space: (A) Females identified by locality (LOC) and (B) females identified by species (ID). Abbreviation by locality: BA, Bali; BO, Borneo; JV, Java; KO, Komodo; MO, Mollucas; MY, Malaysia; SN, Singapore; SU, Sumatra. Abbreviations by species: ha (circle), H. harterti; mg (plus sign), H. margarethae; ty (square), H. typus; un (diamond), unknown/unassigned species. FIGURE 3. Contrasting habitus of adult Hemiphyllodactylus: (A) elongate morphotype, H. typus, Hawaii (USNM 27924); (B) robust morphotype, H. yunnanensis, China (British Museum of Natural History [BMNH] 1904.11.29.10D). (Illustration by J. Kilby.) 12   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY them.). Re-examining the proportions of the outliers and the types versus the typus specimens revealed that most (37 of 39 individuals) sunda H. typus had mean TrunkL/ SVL values ?50% (52.3% ? SD 2.3, range of 46?56%) and a more ambiguous result in the other group. H. mar- garethae paratypes had TrunkL/SVL of 52% and 54%; the outliers were ?50%, although not greatly so except for one Sumatran specimen (RMNH 7371, 40%). sunda H. typus HeadL/SVL matched well the Pacif sample (mean 21.5% ? SD 1.0, range 19?23.8%), showing the small- headed condition; the bisexuals ranged 22.2?26.8%. The three highest loading proportions showed a greater degree of overlap between unisexuals and bisexuals, for example, EyeD/HeadL (highest loading on first component) 75.1% ? 8.6, 61?106% for unisexuals versus 49?81% for bisexu- als; both the 49% and 106% values were extreme outliers of their samples and likely data collection errors. The segregation of the unisexuals (H. typus) and bi- sexuals in the sunda sample allowed a summary of the major statistics for the Sundan H. typus (n = 40, including the holotype): SVL (mean length ? SD, range): 38.7 mm ? 3.19, 32.3?44.1 mm; TrunkL: 20.3 mm ? 1.96, 16.9?24.2 mm; HeadL: 8.3 mm ? 0.62, 7.1?9.4 mm; TrunkL/SVL (mean proportion ? SD, range): 0.52 ? 0.02, 0.46?0.56; HeadL/SVL: 0.21 ? 0.01, 0.20?0.24; HeadW/HeadL: 0.65 ? 0.04, 0.55?0.63. Philip is predominantly a bisexual sample, containing eight unisexuals from Palawan and two from Mindanao. The summary statistics of unisexuals are SVL: 39.0 mm ? SD 2.75 with range of 35.3?41.3 mm; TrunkL: 21.1 mm ? 1.84, 18.4?22.4 mm; HeadL: 8.3 mm ? 0.77, 7.2?9.0 mm; TrunkL/SVL (mean proportion ? SD, range): 0.54 ? 0.02, 0.52?0.57; HeadL/SVL: 0.21 ? 0.01, 0.20?0.22; HeadW/ HeadL 0.70 ? 0.08, 0.66?0.82. The sEasia sample in- cludes only one unisexual (SVL 38.3 mm), confirmed by pigmented oviducts and caecum; this adult female, how- ever, lacks secreting precloacal?femoral pores, thereby questioning its assignment to H. typus. india has four uni- sexual specimens but only in the Sri Lankan component of the sample; two other Sri Lankan specimens are bisexuals. One of the unisexuals is the holotype (BMNH 74.4.1326) of Lepidodactylus ceylonensis Boulenger. Summary statis- tics for the india H. typus are SVL (mean ? SD, range): 34.0 mm ? 10.43, 18.5?40.5 mm; TrunkL: 17.4 mm ? 5.34 9.4?20.5 mm; HeadL: 7.4 mm ? 1.98, 4.5?8.9 mm; TrunkL/SVL (mean proportion ? SD, range): 0.51 ? 0.02, 0.50?0.54; HeadL/SVL: 0.22 ? 0.02, 0.20?0.24; HeadW/ HeadL: 0.66 ? 0.02, 0.63?0.67. Pr?cis. (1) The Oceania H. typus samples are homogeneous within and among samples, representing a single genetic entity and henceforth treated as a single sample (Pacif). (2) Morphometrics weakly differentiate unisexual individuals (Hemiphyllodactylus typus) from bisexual ones. Two proportions, TrunkL/SVL and HeadL/ SVL, appear most useful in this differentiation. Unisexual?Scalation An overview of scalation variation within unambigu- ous unisexual samples is presented in Table 2. Intrasam- ple variation, as estimated by CV, is nearly identical to intersample variation (i.e., CVs of Pacif), resulting from the similarity of means, medians, and ranges of the scala- tion traits; thus the subsequent character survey focuses on the Pacif sample and examines the regional samples only when one or more of the latter samples deviates from Pacif. As noted in the morphometric section, many mu- seum vouchers were poorly positioned at time of preser- vation. Also, Hemiphyllodactylus are small geckos, and many were examined in circumstance of poor lighting and/ or optics. I assumed that such circumstances would cause high variation, but fortunately, ?measurement error? from these data-gathering difficulties is low, and even modest- sized samples (?10 individuals) provide reliable estimates of population parameters. Five traits (CircNa, SnS, CloacS, TotPore, PreclPor) show high variation (CV ? 16%). The high variation for CircNa, SnS, and CloacS results from a high frequency of one character state and a lesser occurrence of the other states. For Pacif, CircNa is three scales for 78% of the sample, and one, two, four, and five scales for the remain- der of the sample. The situation is not as extreme for SnS and CloacS, but with two scales (range 1?5) representing more than 50% of the sample for SnS, and two (~50%) and three (~30%) spurs (range 0?5) for CloacS. TotPore and PreclPor variation results from a broad range (0?26 [median 14], 0?13 [10], respectively) of pore numbers (Figure 5). The majority of adult H. typus have femoral pores, although this trait was not recorded during data collection; it can be calculated by subtracting PreclPor from TotPore. Because the presence of secreting pores in adult fe- males characterizes the unisexual H. typus, PreclPor and TotPore require further examination. Of the 92 Pacif adults examined, 88 individuals possessed PreclPor. The four females without pores ranged in SVL from 35.6 to 42.2 mm, well above the minimum SVL (29.2 mm) at sexual maturity. The condition of the ovaries was not re- corded for the two largest individuals (40, 42 mm SVL). The 39-mm individual had small ovarian follicles, and a n um b e r 6 3 1   ?   1 3 FIGURE 5. Frequency distribution of precloacal?femoral pores in the Pacific sample of unisexual Hemiphyl- lodactylus typus. Size classes are length from snout to vent (SVL) at 2-mm intervals for PreclPor and TotPore with midpoints plotted on x axis: (A) Precloacal pores and (B) total precloacal?femoral pores. 14   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY 35-mm female had mid-vitellogenic follicles. These data indicate that all poreless individuals were mature, al- though the pores may have closed owing to reproductive quiescence or senescence. Both PreclPor and TotPore have broader ranges and greater variation (Figure 5) than for most other gecko species. While gathering data on Hemi- phyllodactylus, I developed the impression that the num- ber of pores associated with body size/maturity. Such an association is not supported by regression or correlation analyses: for example, Pearson R = 0.074, PreclPor to SVL; 0.27, femoral pores to SVL; 0.22, TotPore to SVL. Secreting pores occur only in adult H. typus; perhaps new pores develop with age and/or each successive egg devel- opment event. Precloacal and femoral pores are not con- tinuous in any H. typus. Another pore trait of H. typus is the absence of femoral pores in some sexually mature individuals. There is no obvious pattern to their absence. Femoral pores (?1) occur in the majority of adults, more or less symmetrically on left and right: hawai 37 individu- als with pores, 10 of these lack femoral pores; Polyn 6, 3; Fiji, 13, 3; ncal 9, 1; nguin 15, 5; and taiwan 10, 1. Suplab, Inflab, and Chin display normal levels of vari- ation (i.e., CV < 10%). Suplab has a median of 11 scales (range, 10?12), Inflab 11 (9?11), and Chin 11 (11?12). Dorsal (15, 13?18) and Ventral (11, 10?14) are somewhat more variable (CV = 9.8%, 10.4%, respectively). Subcaud are invariably equal-sized to adjacent caudal scales. The typus digital formulae using the median value of the individual digits are 3-4-4-4 (forefoot) and 4-4-5-4 (hindfoot). Both 4FingLm and 4ToeLm are invariant, 4 and 5, respectively. Digital lamellae vary by only one scale from the median value, either 3 or 4 for 2Fing- to 5FingLm and 2ToeLm, or 4 or 5 for 3Toe- to 5ToeLm. 1FingLm and 1ToeLm have median values of 5 (4?5 1FingLm, 4?6 1ToeLm). The Mascar sample, although considerably smaller than Pacif, shares similar medians and ranges for the scalation traits (Table 2). PreclPor differs slightly (range 4?12) and TotPore has a maximum of 17. These differ- ences likely result from vagaries of sampling. The other difference, probably of similar origin, is the median fore- foot formula of 3-4-4-3. sunda has a much larger H. typus component (n = 40), although as for Mascar, the pattern of variation and ranges match those of Pacif. TotPore and PreclPor share similar ranges, although the TotPore median is less, 6.5 pores. The digital formulae for the fore- and hindfoot are the same. The much smaller H. typus component of Philip (n = 4) and Sri Lanka (n = 4) also largely matches the medians of the Pacif sample. Importantly, pooling adult H. typus from all localities (n = 143) yields coefficients of variation for measurements and scalation (excluding PreclPor and TotPore) that are nearly identical to CVs for the repeated measures sample. This result argues strongly for the genetic homogeneity of H. typus populations across the entire Indo-Pacific distri- bution of this unisexual taxon. Pr?cis. (1) The island and island group sam- ples are homogeneous within and among samples, thereby representing a single genetic entity, Hemiphyllodactylus typus (Pacif). This homogeneity is shared among the uni- sexual components of the other regional samples ? Philip, sunda, india, and Mascar. Unisexual?Coloration Hemiphyllodactylus typus are not brightly colored geckos (Figure 6). Their background color ranges from a dusky tan in the light color phase to a reddish brown in the dark phase. In both phases, a series of narrow, dark brown, transverse bars or blotches lies middorsally from neck to base of tail. These bars are commonly irregularly edged, and they either extend entirely across the back or are bro- ken middorsally. A series of moderately spaced, small light (white to creamy beige) spots lie dorsolaterally on each side of the trunk from the neck to hindlimbs. These spots are a continuation of the pre-orbital light stripe running from the naris to the anterior temporal area, ending in a brighter spot and then continuing as spots on the neck. In most in- stances, a dark brown stripe lies below the light stripe on the lip, fading and disappearing beyond the ear opening. The venter from chin to vent and onto tail is a dusky light tan to yellowish tan; the dusky appearance is created by a multi- tude of tiny dark brown spots, a few in each scale. Dorsally at the base of the tail (postsacral), a dark brown bar and abutting white to beige bar begin the irregular dark?light banding of the tail; this banding quickly becomes progres- sively diffuse and indistinct in most individuals. In life, a few individuals will appear uniformly brown except for faded head markings. This situation is the com- mon condition for preserved specimens and limited the de- velopment of quantitative coloration coding only to two characters (OrbStrp and PostocS; Appendix 1). Neither of these two traits shows any striking varia- tion within or among the Pacific regional samples (Table 2). Similarly there was no variation in coloration among the non-Pacific samples that was not encompassed by the description above. Pr?cis. Hemiphyllodactylus typus are predom- inantly dull-colored geckos. The most striking coloration n um b e r 6 3 1   ?   1 5 features are the lateral head stripe ending in a bright light spot, a series of small light spots dorsolaterally on the trunk, and bright double bar of dark and light at the base of the tail. regioNal variatioN amoNg Bisexual PoPulatioNs The preceding analysis of regional samples identified six samples containing bisexual individuals and popula- tions and revealed the possibility of eight or more different bisexual populations: Palau, Philip, sunda (potentially four bisexuals), sEasia, china, and india (potentially two bisexuals). Bisexual?Visceral Anatomy In the bisexual samples, three (Palau, Philip, india) have females with pigmented caeca and oviducts (see Fig- ure 2B). Females of sEasia and china commonly lack pigmentation on these organs; however, an adult sEasia fe- male (BMNH 1931.11.21.1) from Thailand has pigmented oviducts and caecum. Within the Sunda sample, some fe- males have the pigmented condition and others do not. I observed no differences in extent or intensity of pigmenta- tion among the unisexual and bisexual populations. Oc- casionally, a female of a ?pigmented? population will lack pigmentation on the caecum or oviduct. One each nguin and a sunda H. typus has unpigmented oviducts, and a Philip male and a Sri Lankan female have unpigmented caeca. In pigmented populations, commonly the perito- neum over the testes and epididymides is pigmented, al- though not as densely as oviducal pigmentation in females. Most individuals identified as bisexuals in the sunda sample lack organ pigmentation. All sunda bisexuals have unpigmented caeca except three adult males from Borneo (FMNH 158734, 196268A, 213665), and the last male has pigmented epididymides. Four Sumatran females have pigmented oviducts and two of these are paratypes (IRSN 9338B, ZMA 11096) of Hemiphyllodactylus margarethae Brongersma. Pr?cis. (1) Pigmentation of caecum and oviducts is not confined to unisexual H. typus; this pigmentation FIGURE 6. Dark and light phases of coloration in Hemiphyllodactylus typus from the Pacific population: (A) USNM 310814, Hawaii, Oahu (photograph by G. Zug) and (B) USNM 267979, Fiji, Viti Levu (photograph by J. R. H. Gibbons). 16   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY trait also occurs in females of the bisexual Palau, Philip- pines, and Indian populations, and some Sumatran fe- males. (2) Generally, this pigmentation is absent in bisexual females of mainland Southeast Asia and the Greater Sunda Islands. Bisexual?Morphometry In bisexual populations, sexual dimorphism is the first morphometric issue to examine. A comparison of adult females and males in each regional sample (excluding sunda) reveals size dimorphism in Palau, Philip, SEasia, and china. In all four samples, females average larger than males: 32.8, 35.1, 39.0, 43.3 mm SVL, respectively; 30.6, 31.4, 35.7, 39.9 mm (Student t test, P < 0.025 for significant difference between means of females and males; see Table 3 for female ranges). Disparities in average body size result in significant differences (P < 0.05) for most measurements (not EyeD and SnW) in china and (not SnEye and SnW) in Palau. Philip shows dimorphism in TrunkL, HeadL, and SnEye. sEasia shows dimorphism only for SVL. None of the proportions displays dimor- phism in any sample. Among all samples, average adult male SVL ranges from 30.3 mm (range 28.3?31.6 mm, Palau) to 39.9 mm (35.1?46.4, china). Regional variation in female morphometrics is detailed in Table 3. The two eastern mainland Asian populations (Vietnam and Hong Kong) average strikingly larger than the other mainland bisex- ual populations. The Palauan population has the small- est adults of any bisexual or unisexual populations. It is noteworthy that Palau, Philip, and india adult females average smaller than Hemiphyllodactylus typus (38.1 mm SVL; Table 3). Subsequently, I will provide some evidence for the hybrid origin of H. typus from the Palauan and Philippine bisexual populations. Variation within the bisexual samples is similar to that for the regional H. typus samples. The CVs for mea- surement data (females of Palau, Philip, sEasia, china, india) are: SVL 3.1?11.7%; TrunkL 4.3?15.2%; HeadL 3.6?12.1%; HeadW 4.1?21.4%; SnEye 5.6?13.5%; Nar- Eye 4.3?14.9%; EyeD 5.0?13.6%; SnW 6.2?21.4%. Rela- tive to the repeated-measure CVs, the preceding CVs are greater but only about 3% higher than observed in the repeats. Of these CV ranges, Palau females have the least variation (except SnW), Philip the lowest and SEAsia the greatest. The samples are clustered with one group (Palau, Philip, india) at the low end of the range and SEAsia and china at the high end; this clustering is consistent across all measurements. The high variation of SEAsia and china suggests the possibility of samples containing two or more taxa. Such mixing has a probability for SE- asia owing to its geographic composition extending over 18? of latitude (Hong Kong to Chiang Mai) and 10? of longitude (length of Thailand); mixing will be examined below. india in subsequent mensural discussions consists of only Indian specimens from the vicinity of the type lo- cality. Both Sri Lankan bisexual specimens are adults, a male (26.3 mm SVL) and a female (36.0 mm). This disparity in size is striking. The Sri Lankan fe- male lies within the range of other bisexual female samples (Table 3). In contrast, the male (BMNH 1910.3.16.4; 26.3 mm SVL) is nearly the smallest adult in all bisexual sam- ples. Only one each, Indian and sEasia males (27.2, 25.5 mm SVL, respectively), share small adult size. Even the Palau and Philip males are larger, and these two popula- tions average the smallest in adult size of all Hemiphyl- lodactylus populations. The small size is not a mistake in the recognition of maturity, as this Sri Lankan male has 23 TotPore (16?24, range for Indian males). Although not statistically dimorphic, Indian adult females average slightly larger (35.3 mm SVL, range 33.1?37.9 mm, n = 6) than males (33.5 mm SVL, range 27.2?36.9 mm, n = 8). The proportions are nearly identical in the two sexes, thereby reinforcing the absence of sexual dimorphism within H. aurantiacus. The absence of morphometric differentiation between the Indian and Sri Lankan speci- mens was evident in the high frequency of misclassifica- tion of Indian males and females analyzed (separately) in DF analysis (a synopsis of results is in Appendix 3). The recently collected adult female and two males of H. au- rantiacus (Bauer and Das, 1999) lie within the size ranges noted above. These authors emphasized scalation and col- oration; their findings will be discussed later. sunda has 32 bisexual individuals (29 adults) amidst the total sample (n = 83). This sample visually consists of at least two bisexual taxa, and these bisexuals derive from three geographic areas (adults from peninsular Ma- laysia, n = 17; Sumatra, 8; Borneo, 3; no precise locality, 1). These samples sizes are inadequate to address mensu- ral variation in detail; however, owing to the existence of three names (harterti, larutensis, margarethae), statistical examination of the mensural data is necessary. The Ma- laysian sample consists of nine adult females (mean 48.3, 40.9?62.1 mm SVL) and eight adult males (47.2, 35.3? 56.9 mm); the smallest male (BMNH 1901.3.20.2) is the holotype of Gehyra larutensis Boulenger. This BMNH male is smaller than the other males (36.5?56.9 mm). The smallest female (40.9 mm) is the holotype of Lepidodac- tylus Harterti Werner (ZMB 15360). A series of 17 adults n um b e r 6 3 1   ?   1 7 from the Cameron Highlands and Fraser Hill contains the largest adult Hemiphyllodactylus encountered in my survey of museum collections, females ranging from 42.2 to 62.1 mm (mean ? SD, 50.2 mm ? 6.52; n = 8), males from 36.5 to 56.9 mm (48.1 mm ? 6.44, n = 7). Even though the means of the two sexes differ, there is no sexual dimorphism. This latter sample from the central mountain range (Titiwangsa) attracts attention because the named Malaysian taxa (harterti and larutensis) derive from the western mountain range (Bintang). An exploratory DFA of the two Bintang specimens and the 15 Titiwangsa ones yields 100% overall accuracy in the original classification TABLE 3. Summary statistics on select mensural characters of adult females of the bisexual Hemiphyllodactylus samples. The values are mean ? standard deviation (SD) and range of minimum to maximum. Organization as in Table 2, except sample sizes (n) are adult females, adult males, and total sample, respectively. sunda was excluded because it was a mixed sample of two or more taxa. Character abbreviations are defined in Appendix 1. Sample (n) Character and Palau PhiliP SEaSia a China India b statistic (11, 12, 24) (19, 17, 36) (32, 22, 61) (17, 18, 38) (6, 8, 17) SVL Mean ? SD 32.8 ? 1.01 35.1 ? 2.87 39.0 ? 4.55 43.4 ? 3.89 35.3 ? 2.22 Range 31.1?34.2 29.6?41.3 31.9?50.5 34.9?49.3 33.1?37.9 TrunkL Mean ? SD 17.8 ? 0.76 18.5 ? 1.87 18.5 ? 2.82 20.9 ? 2.89 17.4 ? 1.00 Range 16.8?18.9 15.5?22.4 14.9?25.6 16.1?26.5 16.5?18.8 HeadL Mean ? SD 7.1 ? 0.26 7.7 ? 0.60 9.0 ? 1.09 10.1 ? 1.01 8.1 ? 0.36 Range 6.6?7.5 6.6?9.0 7.4?12.1 7.6?11.5 7.7?8.7 SnEye Mean ? SD 2.8 ? 0.16 3.1 ? 0.34 3.7 ? 0.50 4.3 ? 0.55 3.2 ? 0.37 Range 2.6?3.1 2.6?3.7 3.0?5.0 3.0?5.2 2.7?3.7 SnW Mean ? SD 1.1 ? 0.11 1.3 ? 0.08 1.5 ? 0.33 1.7 ? 0.22 1.4 ? 0.06 Range 1.0?1.3 1.2?1.4 1.0?2.3 1.3?2.2 1.3?1.4 TrunkL/SVL Mean ? SD 0.54 ? 0.023 0.53 ? 0.019 0.47 ? 0.033 0.48 ? 0.030 0.49 ? 0.020 Range 0.50?0.57 0.49?0.57 0.40?0.54 0.43?0.55 0.46?0.50 HeadL/SVL Mean ? SD 0.22 ? 0.006 0.22 ? 0.009 0.23 ? 0.013 0.23 ? 0.009 0.23 ? 0.009 Range 0.20?0.22 0.20?0.24 0.21?0.27 0.21?0.24 0.21?0.24 HeadW/SVL Mean ? SD 0.13 ? 0.006 0.14 ? 0.011 0.16 ? 0.018 0.17 ? 0.017 0.16 ? 0.018 Range 0.12?0.14 0.13?0.17 0.12?0.19 0.14?0.20 0.14?0.18 HeadW/HeadL Mean ? SD 0.59 ? 0.028 0.65 ? 0.053 0.71 ? 0.060 0.72 ? 0.063 0.71 ? 0.078 Range 0.55?0.64 0.58?0.82 0.55?0.80 0.59?0.82 0.58?0.79 SnW/HeadL Mean ? SD 0.15 ? 0.017 0.17 ? 0.011 0.17 ? 0.026 0.17 ? 0.012 0.17 ? 0.009 Range 0.13?0.19 0.15?0.19 0.11?0.22 0.15?0.19 0.16?0.18 OrbD/NarEye Mean ? SD 0.83 ? 0.049 0.83 ? 0.073 0.81 ? 0.075 0.78 ? 0.075 0.81 ? 0.067 Range 0.74?0.90 0.66?0.95 0.70?1.00 0.63?0.90 0.71?0.91 a The sEasia sample includes Hong Kong specimens because of their geographic proximity to the Southeast Asian specimens and their greater geographic separation from the Yunnan specimens largely composing the china sample. b The india sample?s statistics derive exclusively from mainland Indian specimens. 18   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY but only 76% accuracy in the jackknifed one (synopsis of results in Appendix 3). Classification function assignments indicate differences in head shape by identifying HeadL, HeadW, NarEye, Eye D, and SnW as the major variables in the classification function. A DFA of mensural propor- tions yielded lower classification accuracy, indicating no differentiation value. The Sumatran specimens are one adult male (38.8 mm) and seven adult females (41.2, 36.0?46.9 mm). This sample includes the four type specimens of Hemiphyl- lodactylus margarethae Brongersma: two adult females (42.0 mm, ISRN 9338B; 46.9 mm, ZMA 11096), a ju- venile male (38.0 mm, IRSN 9338A), and an adult male (38.8 mm, holotype ZMA 11095). The Bornean sample consists of three adult males (33.2 mm, 27.8?37.3). The final sunda bisexual is an adult male (38.6 mm) of uncer- tain provenance in the East Indies. sunda adults do not display sexual dimorphism in any mensural or proportional trait. The preceding descrip- tion of size differences in the three regional subsamples suggests the possibility of size dimorphism in the Malay- sian sample (i.e., females larger but not statistically sig- nificantly so) and regional differences (smaller adults in Borneo, although the H. larutensis holotype is small also). The sEasia sample shows dimorphism in only SVL of the mensural and proportional traits. In compiling this sample, I assumed that specimens from Thailand eastward through Vietnam and South China represented a single population or taxon, hence potentially recognizable as H. chapaensis (Bourret). The initial analysis of morpho- metrics and the general similarity (means differ but strong overlap of ranges occurs) of the china and sEasia sam- ples (Table 3). indicated that my assumption was incorrect and required testing. Another question on the uniformity of the sEasia sample arises from the broad longitudinal breadth (99??115? E) of the sample, extending from Chi- ang Mai in the west to Hong Kong in the east. The latitu- dinal depth of the sample is much less, approximately 7? (19??12? N). The china sample (assumed to represent H. yunnanensis) is broad but nonetheless not as geographi- cally expansive as sEasia and limited to the ?highlands? (Shan and Yunnan plateaus). To test the preceding assumption of homogene- ity between and within these two samples, I examined them in several ways: (1) variation within a combined china-SEAsia sample and (2) the subdivision of these two samples into various subsamples and comparison of the subsamples. The combined China-sEasia sample contains 48 adult females and 39 adult males. Student t tests of all mensural and proportional traits show that size dimorphism persists in most dimorphic traits (SVL, TrunkL, HeadL, HeadW, SnEye, NarEye) identified in the china sample and additionally the proportions EyeD/ HeadL and EyeD/NarEye. Typically, the p values for the t test are slightly higher in the combined sample (e.g., SVL P = 0.008, 0.019, China versus combined). Even though the SEAsia sample is the larger of the two, its absence of dimorphism did not swamp the average size difference of females larger than males. The combination of these two samples, however, re- sults in a near doubling or more of CV of most mensural traits as compared with the china sample. This increased variation reflects a mixing of two or more phenotypes, presumed here to represent distinct genetic entities. A combination based on topography (china [contains only Yunnan and Shan plateau specimens] and the highland areas of northwestern Thailand [Tak to Mae Hong Song and Chiang Rai]) did not alter combined sample variation (i.e., CVs equivalent to those of china alone). The mean SVL of this china-Thai sample decreased from 43.3 mm (china) to 40.7 mm, but the mean values of all propor- tions of the two samples are less than 1% different, sup- porting the homogeneity of Hemiphyllodactylus from this region. Addition of the northern Vietnam (Chapa area) specimens to the china-Thai sample did not alter means or CVs; similarly, the addition of Hong Kong specimens did not alter the level of variation. These additions of a few individuals to a large (n > 30) sample are, however, unlikely to alter CV unless a striking disparity exists in the added sample. The realignment of china and sEasia specimens suggests the existence of a northern (upland) popula- tion and a southern one (?lowland? Thailand only in the present sample). The latter averages smaller than the former (Table 4). The situation (affinities) of other South- east Asian specimens is unclear owing to small samples (Table 4). The Vietnam chapaensis sample averages larger (SVL) than the four other samples, but its range is within that of the china?NW Thailand sample. Its proportions similarly match the latter sample?s proportions. The Laos sample is the smallest (n = 2) of this comparison, hence difficult to interpret, and a single male from Cambodia is immature. Presently, I note only the low TrunkL/SVL proportion representing a shorter trunk length than in the other Oriental samples. Interestingly, they appear less robust than the Hong Kong Hemiphyllodactylus, whose general appearance matches the stouter habitus (Figure 3) of large adult female H. yunnanensis; yet they have a lon- ger trunk (53% TrunkL/SVL, Table 4) similar to the taxa with the slender habitus. A DFA of Oriental adults using n um b e r 6 3 1   ?   1 9 all 10 proportions yields a poor classification (jackknifed) of these subsamples; all were classified at ?50% (synopsis of results in Appendix 3). Larger samples are essential to address these confusing morphometrics. Zhou et al. (1981) described three subspecies of H. yunnanensis from China without reference to any other species or populations of Hemiphyllodactylus. They had good samples (?20 individuals) for each subspecies, and although they reported some measurements (Table 5), they presented no statistical analysis and no mensural data for the nominate subspecies. The westernmost subspecies (H. y. longlingensis) is the smallest of three forms and shares the range of my Yunnan sample. The two eastern forms (H. y. dushanensis and H. y. jinpingensis) are distinctly larger geckos than the western ones. Their adult SVLs (Table 5) do not overlap with either H. y. longlingensis or the Yunnan sample. TrunkL/SVLs of the east and west forms also do not overlap, but surprisingly HeadL and SnEye do. This overlap suggests that either these forms/ populations have proportionally smaller heads or there was a lack of precision in recording these measurements. All three of these subspecies show strong sexual dimor- phism in SVL with only H. y. longlingensis showing a slight overlap of the largest male and smallest female. The absence of overlap suggests that the specimens examined (and reported in the tables) underwent selection prior to analysis because of the strong overlapping ranges of adult females and males in my china sample, even though the average SVLs are significantly different. As an aside, one Thailand adult or near-adult Hemi- phyllodactylus (BPBM 3502, Sakaerat) is hermaphroditic, with large testes and a pair of vitellogenic follicles (maxi- mum diameter 3.2 mm). All other unisexual and bisexual specimens examined had gonads of only one sex. TABLE 4. Summary statistics on select metric characteristics of adults of the bisexual Hemiphyllodactylus from southern Asia. The val- ues are mean ? standard deviation (SD) and range of minimum to maximum. Realigned and restricted regional samples from the china and sEasia samples: china?northwestern Thailand; Thailand, area exclusive of the northwest; Laos, Phong Saly; Vietnam, chapaensis type locality; and Hong Kong. Sample sizes (n) are total adults, females, and males, respectively. Character abbreviations are defined in Appendix 1. Sample (n) Character and China?nW Thailand Thailand Laos Vietnam Hong Kong statistic (61, 33, 28) (16, 9, 7) (2, 1, 1) (3, 3, 0) (4, 2, 2) SVL Mean ? SD 39.4 ? 4.68 35.1 ? 2.77 37.1 ? 0.78 46.2 ? 3.16 43.0 ? 6.13 Range 25.5?49.3 30.0?39.9 36.5?37.6 42.7?48.8 35.3?50.3 TrunkL/SVL Mean ? SD 0.47 ? 0.031 0.47 ? 0.036 0.42 ? 003 0.49 ? 0.011 0.53 ? 0.032 Range 0.40?0.55 0.42?0.52 0.42?0.43 0.48?0.51 0.51?0.58 HeadL/SVL Mean ? SD 0.23 ? 0.010 0.23 ? 0.014 0.24 ? 0.005 0.23 ? 0.004 0.22 ? 0.007 Range 0.21?0.26 0.21?0.26 0.23?0.24 0.23?0.24 0.21?0.23 HeadW/SVL Mean ? SD 0.17 ? 0.018 0.17 ? 0.017 0.17 ? 0.002 0.17 ? 0.002 0.16 ? 0.010 Range 0.14?0.22 0.14?0.20 0.16?0.17 0.17 0.15?0.17 SnEye/HeadL Mean ? SD 0.41 ? 0.025 0.41 ? 0.026 0.39 ? 0.024 0.42 ? 0.023 0.41 ? 0.0.37 Range 0.34?0.46 0.38?0.46 0.37?0.42 0.40?0.44 0.38?0.46 SnW/HeadL Mean ? SD 0.17 ? 0.020 0.16 ? 0.025 0.21 ? 0.008 0.17 ? 0.002 0.16 ? 0.015 Range 0.11?0.22 0.12?0.20 0.20?0.22 0.16?0.17 0.15?0.18 SnW/HeadW Mean ? SD 0.24 ? 0.036 0.22 ? 0.035 0.30 ? 0.001 0.24 ? 0.007 0.22 ? 0.014 Range 0.15?0.36 0.16?0.29 0.30 0.23?0.24 0.21?0.24 20   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY Pr?cis. (1) Morphometry is only modestly use- ful in the differentiation of intra- and interregional samples of the bisexual populations. (2) Palau, Philip, and china display size dimorphism between adult females and males. Females average significantly larger than males. (3) The Palauan population has the smallest average SVL of all populations. The Philippine and Indian-Sri Lankan popu- lations are also small but average significantly larger than the Palauan one. Average size of the china and sE asia samples is significantly larger than the preceding three samples. (4) The original composition of india, sEasia, and sunda each probably includes representatives of at least two taxa/populations. india contains an Indian com- ponent (H. aurantiacus) and a Sri Lankan one (unnamed). Inclusion of northwestern Thailand specimens in the china population does not increase variation, thus sug- gesting genetic homogeneity among these ?highland? pop- ulations. The remainder of the sEasia sample potentially remains a mixed sample. Presently, there are insufficient museum specimens for a fine-scale geographic analysis. This problem is also shared by sunda, which clearly has multiple taxa therein. Bisexual?Scalation Sexual dimorphism in PreclPor and TotPore is shared by all samples. Adult males have secretory pores; females do not or do so uncommonly. Females with secreting pores occur in all Asian samples. The sample india-in- dia contains a single adult female (BMNH 74.4.1332, a syntype of H. aurantiacus) with three PreclPor, about half the number found in males (Table 6). china has three females with pores, one with only seven precloacal pores, another with nine precloacal pores separated from one femoral pore on each side, and a final individual with a continuous precloacal?femoral series of 19 pores. sEasia also has three pore-bearing females; the holotype of H. t. chapaensis has nine PreclPor, and two Thai females have continuous series of 14 and 15 pores. In sunda, pores occur only in Sumatran females, two of seven individuals (12 PreclPor in one and separate series of 9 and 20 in the other). Generally, but not always, TotPore is distinctly less in females than in males. No other scalation traits show dimorphism in Palau, Philip, sEasia, china, or india. The composite nature of sunda does not permit rigorous testing for dimorphic traits. Most scalation traits are fairly uniform across the six bisexual samples (see Table 6, although Sunda not included). I present medians because the data for scala- tion traits are discontinuous values and integers better portray the actual number of scales than does a decimal value. The uniformity among samples is most evident in the overlapping ranges of minimum and maximum values. Among the six samples, CircNa is identical (three scales) in all samples. SnS ranges strongly overlap amidst the six samples with sEasia individuals typically possessing only two ?internasal? scales, and Palau and india each with four SnS. This latter condition and the three-SnS one rep- resents a large ?supranasal? above the naris on each side separated medially by two or one small scales. Inflab are similar with most samples having ten scales, Palau with nine and india-India with 11. Most samples have 10 Su- plab, with 8 for Palau and 11 for sunda. Although these differences (for Suplab, but other traits as well) are slight TABLE 5. Comparison of some character measurements of adult females among the Chinese populations of Hemiphyllodactylus yun- nanensis. Data for the subspecific samples of H. yunnanensis were derived from Zhou et al. (1981: tbls. 2?4). Character abbreviations are defined in Appendix 1; a dash (-) means no data were available. Length (mm) range for sample or subspecies (n) Yunnan yunnanensis a longlingensis jinpingensis dushanensis Character (12) (0) (19) (10) (10) SVL 37.2?48.8 - 39.0?46.0 49.0?53.5 48.0?51.0 TrunkL 17.4?23.4 - 20.0?23.5 24.0?28.5 25.0?27.0 HeadL 8.8?11.4 - 9.0?10.0 10.5?11.0 10.0?11.0 SnEye 3.7?5.1 - 4.0?5.0 4.5?5.0 5.0?5.5 ForelimbL - - 10.0?12.0 11.0?12.0 11.0?12.0 HindlimbL - - 13.0?16.5 16.0?17.5 16.0?17.5 a Despite a sample of 249 specimens, no measurements were reported by Zhou et al. (1981) in the description of H. y. yunnanensis nor was a table of measurements provided. Measurements also were not included in the descriptions of the new subspecies. n um b e r 6 3 1   ?   2 1 and usually not statistically significant, I suggest the dif- ferences reflect genetic differentiation among the regional populations and are not the result of sampling or measure- ment error. This proposition derives from the relatively low variation (CV) observed for most scalation traits (see the earlier Baseline Estimates section). The re-aligned china?NW Thai sample usually has the same medians as china and similar standard deviations Chin ranges strongly overlap; however, the low medi- ans (8) of sunda, sEasia, and china result from six or seven scales in most individuals and the higher medians of Palau, Philip, and india from most individuals having TABLE 6. Summary statistics on select coloration and scalation characters of juveniles and adults of the bisexual Hemiphyllodactylus samples. Organization as in Tables 2?4. Sample sizes (n) are juveniles, adults, and total sample, respectively. Statistical values are either median ? standard deviation (SD) and range of minimum to maximum (for coloration and scalation characters) or modes and frequency (%) of occurrence (for finger and toe lamellae). Character abbreviations are defined in Appendix 1. Sample (n) Character and Palau PhiliP SEaSia a China India b statistic (11, 12, 24) (15, 19, 35) (32, 22, 61) (17, 18, 38) (6, 8, 17) PostocS Median ? SD 4 ? 0.88 3.5 ? 1.18 1 ? 1.38 2 ? 1.41 0 ? 2.17 Range 2?6 0?6 0?4 0?5 0?7 SnS Median ? SD 4 ? 0.65 3 ? 0.48 2 ? 0.68 3 ? 0.72 4 ? 0.89 Range 3?5 2?4 1?4 2?5 3?6 Suplab Median ? SD 8 ? 0.78 10 ? 1.15 10 ? 0.91 10 ? 0.70 10 ? 0.89 Range 8?11 8?13 8?12 9?12 10?13 Chin Median ? SD 11 ? 0.90 11 ? 1.52 8 ? 1.59 8 ? 1.38 11 ? 1.12 Range 9?12 8?14 6?18 6?11 10?14 Dorsal Median ? SD 15 ? 1.44 15 ? 1.50 14 ? 1.78 13.5 ? 1.80 13 ? 1.87 Range 11?18 12?18 9?18 12?17 11?17 CloacS Median ? SD 2 ? 1.01 2 ? 1.89 1 ? 0.69 1 ? 0.38 2 ? 0.63 Range 1?4 0?3 0?4 1?2 1?3 TotPore c Median ? SD 22.5 ? 8.15 27 ? 5.35 21.5 ? 4.28 19 ? 3.31 21.5 ? 2.91 Range 16?28 17?38 11?26 11?24 16?25 4ToeLm Median ? SD 4 ? 0.38 4 ? 0.41 3 ? 0.80 4 ? 0.42 3 ? 0.46 Range 4?5 4?5 3?5 3?5 2?3 FingerLm d Modal values 3-4-4-3 3-3-3-3 3-3-3-3 3-3-3-3 2-2-2-2 Frequency 55.0% 65.7% 72.2% 36.8% 100% ToeLm d Modal values 3-4-4-4 3-4-4-4 3-3-3-3 3-4-4-4 2-2-3-2 Frequency 45.8% 50.0% 50.1% 57.9% 37.5% a The original composition of china and sEasia samples is retained for consistency of comparison with Table 4 and mensural results described and discussed earlier in the text and tables. b The values are only for the mainland India portion of the India sample. c These values are only for the adult male portion of each sample. d Lamellae formulae represent the most frequent formula (mode) for each sample and the percent of the sample with this formula (frequency). 22   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY 10 or more chin scales. The difference in total number of scales in the chin scale arc results from the enlargement of the two median scales posteriorly bordering the mental (i.e., postmentals). Postmental shape and size are variable within each Asian sample; nonetheless differences in post- mental size serve to delimit southern Asian populations and denote populational differentiation. Beginning in the north with populations identified as H. yunnanensis (china), each postmental is one-third to half the area of the first supralabial (Figure 7D,E); the lat- erally abutting chin scale is commonly one-third to two- thirds the area of the postmental and distinctly larger than chin scales behind and beside it. This anteriormost row, including the postmental, is an arch of larger scales, and posteriorly the chin scales quickly decrease in size to gran- ular scales. In china, the mental is modestly large and roughly an equilateral triangle to a flattened pentagonal shape. In SEAsia, the chin scale condition is as described above, although I have the impression that the next poste- rior scale row to the ?large? scale arc has proportionately larger scales than my china specimens and then a rapid diminution to granular chin scales. I note no substantial difference from the preceding pattern for individuals from Hong Kong to northwestern Thailand. Interestingly, my examination of the holotype of H. chapaensis revealed a low Chin (seven scales), hence a modestly enlarged postmental, but Bourret?s (1937:60) description noted the postmentals as ?une paire de tr?s petites plaques.? His illustration (reproduced in Figure 7F) shows an arc of enlarged chin scales with postmen- tals not much larger than the posterior chin scales. This morphology contrasts to Hong Kong H. chapaensis (so taxonomically labeled by Lazell, 2002), which possess large postmentals followed by a second arc of enlarged (although not as greatly) chin scales. A southern Vietnam- ese specimen (USNM 146161) has very large, rectangular postmentals similar to those depicted for H. margarethae (Figure 7C). The Cambodian male (FMNH 270569) has an arc of nearly equal-sized scales (Chin = 10). All sunda specimens have large, rectangular postmentals. For most and the above-mentioned Vietnamese specimen, the post- mentals broadly contact medially, and posteriorly chin scales rapidly decrease in size (Figure 7). Dorsal and Ventral, each, have overlapping ranges among the six samples. The pattern of variation between these two traits is different. Note that the number of scales for each derives from the EyeD distance; thus the values represent the number of scales within the same area for each specimen, although there may be an increase in vari- ance within a sample owing to the manual measurement of EyeD. The CV is 12?15% and in the same range as Chin and most finger and toe lamellae counts. Dorsal val- ues have identical (or nearly so) medians (14% and 15%) for all samples except india-India, which is distinctly lower (12%; Table 6). For Ventral, the medians are more variable. sunda, sEasia, and china have eight scales, Palau and india 10, and Philip 11. The Sundan to Chi- nese populations have proportionally larger ventral scales compared both to dorsal scales within individuals and to the ventral scales of Pacific and Indian specimens. CloacS ranges from absent to four spurs in the total bisexual sample (Figure 8). Part of this variation is due to sexual dimorphism, with females tending to have fewer or no spurs (statistically significant only in Palau and Philip); nonetheless, there are populational differences, with sEasia and china individuals averaging a single FIGURE 7. Scale morphology of the chin in various populations of Hemiphyllodactylus: (A) H. typus USNM 570742 Hawaii; (B) H. ganoklonis USNM 563682 holotype, Palau; (C) H. margarethae ZMA 11095 holotype, Sumatra (redrawn from Brongersma 1931: fig. 4); (D) H. y. yunnanensis China (from Zhou et al., 1981: fig.4 left); (E) H. y. longlingensis China (from Zhou et al., 1981: fig.4 right); (F) H. chapaensis NMHN 1948.43 holotype, Vietnam (re- drawn from Bourret, 1937: fig. 1b). n um b e r 6 3 1   ?   2 3 spur. This average or median value is not driven by a larger number of females in the samples, as the number of adult females nearly equals the number of males in all samples. In most populations the median row of ventral scales on the tail (Subcaud) equal the size of adjacent scales. Only in a few individuals and only in the sunda, SEAsia, and india samples are Subcaud slightly enlarged over adjacent scales. No Hemiphyllodactylus has enlarged subcaudal scales like those of many Gehyra species. Finger and toe lamellae counts (Table 6; Figure 9) are relatively uniform among all samples except for india. Individuals of the latter population have distinctly fewer enlarged U-shaped subdigital lamellae on the fore- and hindfeet. This difference is strikingly apparent; the fore- and hindfoot lamellae formulae (second through fifth digit of each foot) of india is 2-2-2-2, 2-2-3-2, contrasting to the modal 3-4-4-3, 3-4-4-4 of the other Asian samples. The modality of these formulae, however, is 3-4-4-4 and 4-5-5-4 in the Vietnamese, Hong Kong, and Cambodian specimens and 3-4-4-4 and 4-5-5-5 for the Sunda Titi- wangsa specimens. As with the mensural characters, it is necessary to comment on the Zhou et al. (1981) study of Chinese Hemiphyllodactylus populations. Their scalation data ex- amined seven traits: rostral notched posteromedially, en- larged scale posteriorly bordering ?supranasal,? Suplab, Inflab, chin scale bordering mental posteriorly (postmen- tals)], digital lamellae formulae, and precloacal?femoral pores. The pore condition is examined later. I did not score/ record either of the first two traits. My initial research in unisexual Hemiphyllodactylus populations indicated that a rostral notch was always present although of variable development (i.e., indistinctly notched to cleft one-third length [rarely] of rostral scale); hence this trait was not in- cluded in my character set. The variation in the relative size of the post ?supranasal? scale completely escaped my at- tention. Zhou et al. used the relative size of the postmental scales, in part, to differentiate H. y. yunnanensis and H. y. longlingensis. I have not seen specimens from the range of H. y. longlingensis; however, four Hemiphyllodactylus (USNM 570732?570735) from the western edge of the Shan Plateau (Myanmar) match the enlarged second arc of chin scales shown for H. y. longlingensis (Figure 7) with the exception that in the four Burmese specimens, the chin scales are more uniform in size and the median one is no FIGURE 9. Digital lamellae morphology in select species of Hemiphyllodactylus: (A) Right hindfoot, H. yunnanensis, Thai- land (USNM 310807); (B) right forefoot, H. yunnanensis China (BMNH 1904.11.29.10D); (C) right hindfoot, H. typus Hawaii (USNM 27924); (D) right hindfoot, H. yunnanensis China (BMNH 1904.11.29.10D). Illustrations by J. Kilby. FIGURE 8. Cloacal spur morphology in Hemiphyllodactylus: (A) Pinted state, H. yunnanensis, Thailand (USNM 310807); (B) rounded stated, H. typus, Hawaii (USNM 279240). Illustration by J. Kilby. 24   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY larger than its laterally abutting scales in the arc. Because my china sample derives largely from Kunming, my re- sults match the Zhou et al. concept of H. y .yunnanensis. My results for Suplab and Inflab similarly match their data. Their data for digital lamellae, however, contrast sharply to mine. Foremost is low variation in their counts compared with mine. Deciding which lamellae is the prox- imalmost one on each pad can be challenging owing to a gradual reduction in size proximally. My protocol to in- crease the consistency (precision) of my counts is/was to count only the distinctly U-shaped lamellae. Comparison of my China formulae to their H. y. yunnanensis formulae yields five forefoot formulae (2-3-3-3, 3-3-3-3, 3-4-4-3, 3-4-3-3, and 3-3-4-3; the second and third formulae rep- resent 37 and 21%, respectively, of the sample) versus two formulae in Zhou et al. (3-4-4-3, 3-4-4-4; they do not present the frequency of occurrence of each), I recorded nine different hindfoot formulae (ranging from 3-3-3-3 to 4-5-5-5) with 3-4-4-4 in 60% of the sample, occurrence of others is ?10%, usually much less, versus an invari- ant 3-4-4-4 reported by Zhou et al. I, obviously, believe that my data more accurately reflect the natural variation of the Yunnan population. My data cause me to question the discriminatory potential that Zhou et al. attributed to these formulae for the differentiation of populations within their broader China sample. Digital formulae do discriminate populations (e.g., compare india to the other five bisexual samples [Table 6]), but the validity of the Zhou et al. data requires an independent study. Precloacal?femoral pore morphology is similar in me- dians (19?22.5) and range of (11?28) of TotPore (Table 6) for Palau, sEasia, china, and india. Philip has a dis- tinctly higher median TotPore and a partially overlapping range, although extending well beyond the upper limits of the four former samples. The medians and ranges of PreclPor for the preceding five samples are more similar to one another (median range 7.5?11), presumably owing to all geckos having proportionally the same pubic or es- cutcheon area for pore development. Higher or lower Tot- Pore counts derive mainly from an increase or decrease in the length of the femoral pore series. Number of pores, however, disguises two pore mor- phologies (Figure 10): (1) continuous series of precloacal and femoral pores and (2) separate longitudinal patch of precloacal and femoral pores. The continuous pore mor- phology occurs in all males of SEAsia and China samples and the separate-patch morphology in all males of Palau, Philip, and india (including a single Sri Lankan male) samples. The relative size of the pore scales appears identi- cal in the two morphologies. In some sEasia males (e.g., USNM 146161, southern Vietnam), the pore scales are separated by one (usually) or two smaller (one-third or less pore scale area) scales. From the low frequency of this condition and the very few specimens from southern Viet- nam through southern Thailand, it is uncertain whether this condition is a variant or populational state. Pore morphology in sunda shows regional differen- tiation with the occurrence of both continuous-patch and FIGURE 10. Schemata of precloacal?femoral pore morphology of Hemiphyllodactylus: (A) Separate pore series and (B) continuous pore series. (A modified from and B reproduced from Taylor, 1918: fig. 4; H. insularis.) n um b e r 6 3 1   ?   2 5 separate-patch states, each state apparently occurring at a specific location. Even though sample sizes of the various islands or sites are small, the differences among localities are suggestive of genetic differentiation of populations. Sumatran and Bornean adult males (n = 1, 3, respectively; median TotPore 26, 29) have the separate-patch state; fur- ther as noted in discussion of sexual dimorphism, some adult females from these two islands possess precloacal pores and occasionally femoral pores. Malaysian males have continuous precloacal?femoral series with a lower median TotPore (21, 17?39; Titiwangsa specimens only, n = 7). The presence of these two states in SEAsia is no- menclaturally important because of three available names. The H. margarethae holotype has the separate-patch state, the male G. larutensis holotype a continuous state, and the female L. Harterti holotype lacks pores. Pr?cis. (1) Sexual dimorphism of scalation occurs only in the presences (males) and absences (fe- males) of precloacal and femoral pores. (2) Most scala- tion traits are uniform throughout the Asian and Pacific Rim populations. Difference among samples is evident in the characters of Chin, Dorsal, Ventral, digital lamellae, and precloacal?femoral pore. (3) Chin shows two states: enlarged chin scales (postmentals) in sunda, sEasia, and china and no postmentals in Palau, Philip, and india. (4) india has large dorsal trunk scales, only slightly smaller than the ventral ones; all other samples display a greater size difference between dorsal and ventral scales and typically more scales per unit area. (5) Digital lamellae formula is strikingly lower in india than in the other five regional samples, and the formulae in the latter samples are the same. (6) Pore morphology differs with china, sEasia, and some sunda populations with continuous se- ries of precloacal?femoral pores and Palau, Philip, some sunda, and india with separate patches of pores. Bisexual?Coloration The drab, presumably cryptic, coloration of Hemi- phyllodactylus makes characterization of coloration dif- ficult. I recorded two external coloration characters: PostocS and OrbStrp. The results show regional differ- ences. For example, Palau and Philip have higher median PostocS (Table 6) in contrast to the median of one or two postocular spots for SEAsia, china, and india; however, the ranges for these five localities are the same. The ?iden- tical? ranges reflect the difficulties of seeing these spots in older or poorly preserved specimens. If an actual dif- ference exists, it will require documentation with living specimens or recently preserved ones. OrbStrp is less likely to disappear with the age of a specimen owing to its dark pigmentation; nonetheless, old specimens fade to unicolor and this trait can be lost also. My data (Table 5) suggest the near universal presence of OrbStrp in bisexual speci- mens, at least to the level of the ear. With my attempt at quantification of coloration ill- founded for museum specimens, I offer a comparative descriptive approach using the major pattern features identified in the H. typus coloration section. These de- scriptive data derive from my notes on museum speci- mens, descriptions and illustrations from herpetological literature, and field notes of mine and others sources are noted in brackets. Coloration is variable locally within populations and broadly throughout the distribution of bisexual popula- tions. Dorsal ground color for all populations is variously described as tan to grayish brown. The intensity of the ?brown? background depends on the color phase, light or dark, assumed at death and preservation or when pho- tographed. The phase also affects the conspicuousness of markings, whether they are light (usually off-shade whites to tans) or dark (brown to nearly black-brown). The venter also lightens and darkens, hence ranges from near-white to dusky; because each scale normally shows tiny spots (melanophores), the venter is never white. The following descriptions will emphasize five areas: (1) loreal or snout area, (2) side of head and neck (ignoring the light- ening resulting from accumulation of calcium carbonate in the endolymphatic system), (3) dorsal and lateral trunk, (4) sacrum and base of tail, and (5) sides and top of tail. Several traits are shared by most individuals of the bisexual populations. Intensity and color vary from popu- lation to population. A dark brown lateral stripe extend- ing from the loreal area to at least the anterior quarter of the neck occurs widely, although its thickness and lo- real development are variable. For example, in a series of northwestern Thai specimens, the loreal stripe ranges from a dark spot immediately in front of the eye through a well-defined stripe from naris to eye to a fully dark brown loreal area. Similarly behind the eye, the lateral stripe (OrbStrp) ranges from narrow to broad (always above the ear opening), terminating at head?neck juncture or any- where from this juncture to the shoulder. A dark lateral stripe is variously evident on the trunk, and if present, it is typically moderately broad but irregularly edged above and below, occasionally bordered above by a broad lighter area. Often this stripe breaks into a series of dark blotches. Dorsally, the head is unicolor or nearly so with a faded and diffuse mottling. The back is similar, ranging from unicolor through small, paired parasagittal dark spots to 26   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY broad, irregular dark brown bars (continuous across the midline or interrupted). Most individuals display a signa- ture mark posteriorly from the sacrum onto base of the tail. This mark contains a (supracloacal) middorsal dark blotch bordered posteriorly by a (supravent) light mark that can have anterior extension on each side of the mid- dorsal dark blotch with the anterior arm usually ending at the top of the ilium, hence a U-shaped light mark. This contrasting dark and light postsacral mark is often the brightest area of coloring of a Hemiphyllodactylus. Poste- riorly the top and sides of the tail range from light to me- dium brown, nearly unicolor to distinctly banded. In life, the underside of the tail is variously reported as shades of red or orange. Palau Hemiphyllodactylus are generally drab lizards with the exception of the bright postsacral signature mark. Ground color is light gray to tan. The dorsal pattern ranges from a sparse speckling of dark brown spots to small, ir- regular V- or trident-shaped bars. A dorsolateral row of widely spaced pinkish yellow spots extends from the rear of the head to the postsacral mark; the spots on the pos- terior third of the trunk are the most prominent ones. The postsacral mark has a small dark middorsal spot (roughly triangular) encompassed by a broad-based U-shaped light pink mark, which is bordered laterally on each side by a diffuse dark dorsolateral stripe (Figure 11). The tail ranges from yellow to tan with diffuse midline dark marks and narrow light pink transverse bands. In the loreal region, a preorbital stripe is always present but generally only mod- erately developed, ranging from a dark preorbital spot to a narrow stripe extending midway to the naris. The post- orbital stripe is narrow, always fragmented, and often only with a few fragments on the side of head, somewhat more extensive on the neck to the shoulder. (Coloration is reported from Ronald I. Crombie?s [RIC] unpublished 1993?2002 field data notes and my personal observation of color in RIC?s images and preserved specimens.) Philip Hemiphyllodactylus are similar to Palau ones with light to medium brown backgrounds. The dorsal pattern ranges from a faint dark flecking or reticulation through paired, parasagittal dark longitudinal dashes to small dark blotches. Dorsolaterally, a row of brick red, dark-edged spots occurs on each side from behind eye to and onto tail. The postsacral mark has a brown V-shaped anterior border to the broader red U-shaped portion; the latter is usually strongly edged in brown laterally and pos- teriorly. Tail is lightly colored, presumably tannish and commonly with small dark paired spots to tip. A lateral stripe, variously fragmented, extends from the loreal area to anterior neck; the preorbital portion is usually sharply defined (Taylor, 1918; Brown and Alcala, 1978; G. R. Zug [GRZ], personal observation of museum specimens). china and sEasia geckos tend to have darker back- grounds than individuals from the Pacific Rim samples. Although Asian geckos have a light phase, the background in this phase is gray to brownish gray. The overall im- pression of a darker background is heightened by more extensive dorsal and lateral markings in many individuals (Figure 11). Dorsally, the neck and trunk vary from faint reticulations of medium brown through diffuse wavy and brown fragmented crossbars or chevrons to broad, dark brown transverse blotches (usually paired); a moderate pattern seems to be the average condition. Dorsolaterally, a row of light tan to nearly white spots occurs on each side from behind the head to and seemingly forming the anterolateral arms of the postsacral mark. This mark is variable in contrast intensity with the anteromedial dark brown portion, ranging from a large triangular blotch to a moderate transverse bar. The lighter portion of the postsacral mark is typically shades of cream to light yel- low. The postsacral mark is broad and deep and extends well on to the tail base; the anterolateral arms are usu- ally weakly developed and short (inconspicuous). The tail is strikingly lighter than trunk with some dark transverse spots or bars (Boulenger, 1903; Bourret, 1937; GRZ, pers. obs. of color images and specimens). Hong Kong is a geographic outlier from other speci- mens comprising the china and SEAsia samples. Their coloration differs from the preceding in two major ways. First, the dorsolateral spotting is commonly so faded that it is nearly invisible. The immediate postsacral area is light brown and a median dark edge or bar lies above the vent, and the lighter area is the first band of the diffuse light and dark tail banding (Chan et al., 2006; GRZ, pers. obs. of specimens). india H. aurantiacus is a boldly colored gecko of dark brown bars and blotches on a light to medium brown background (Figure 11A). A dorsal pattern of dark, nar- row, wavy crossbars begins on the shoulders and contin- ues to the sacral area; these crossbars can be variously fragmented and/or narrowly connected to anterior or pos- terior crossbars. A dorsolateral series of light (presumably white to light tan) spots extend from the neck to the in- guen; these spots are not evident in all individuals. The tail base pattern is similar to one described above for Hong Kong Hemiphyllodactylus. The remainder of the tail is distinctly banded in brown and dark brown; the lighter bands are about twice the width of the dark ones. The loreal area usually has a dark stripe from naris to eye. A dark supraorbitial stripe extends from the rear edge of the n um b e r 6 3 1   ?   2 7 eye to the anterior trunk. A dark postorbital stripe extends from the eye to the axilla. Both of these stripes are usu- ally unfragmented (Beddome, 1870; Bauer and Das, 1999; GRZ, pers. obs of specimens). Coloration for the sunda gecko ?species? cannot be unequivocally resolved with the data at hand. My notes and Boulenger?s (1900) description of the H. larutensis type are minimal. Its ground color was originally gray- ish brown, although now (in preserved state) it is faded to light tan. A paired series of small dark spots extend from the neck onto the base of the tail. A few dark spots occur on the sides of the trunk. A dark loreal and a post- orbital stripe are present. My notes do not indicate the presence or absence of a dorsolateral series of light spots FIGURE 11. Coloration of select Hemiphyllodactylus taxa: (A) H. aurantiacus (India, Yercaud; AMBauer 5749) (photograph by I. Das); (B), H. ganoklonis (Palau; no field/museum number) (photograph by R. I. Crombie); (C) H. harterti (Malaysia, Bukit Larut; KUZ 21264) (photograph by H. Ota); (D) H. typus (Tonga, ?Eua; USNM 322119) (photograph by G. Zug); (E) H. typus (Fiji, Viti Levu; USNM 267978) (photograph by G. Zug); (F) H. titiwangsaensis (Malaysia, Cameron Highlands, Tanah Rata; KUZ, no field/museum number) (photograph by H. Ota); (G) H. yunnanensis (Thailand, Loei Province; no field/museum number) (photograph by P.-P. van Dijk); (H) H. yunnanensis (Myan- mar, Pyin Oo Lin; USNM 570734) (photograph by G. Zug). 28   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY or a postsacral mark. Werner?s (1900:196) description of H. harterti reports ?gleicht in F?rbung and Zeichnung ganz den L. lugubris.? I interpret this coloration as a medium brown background with dark wavy crossbars. Photographs (Figure 12) of the type show it faded to near unicolor tan. Brongersma (1931) did not include a de- scription of coloration for H. margarethae. Although I examined his type series, I have no notes on their col- oration. Presumably, all four type specimens are faded. That is certainly the condition of the two ZMA syntypes photographed (Figure 12). Excellent color illustrations of Sundan Hemiphyllo- dactylus are available in some recent field guides. Addi- tionally, color photographs were provided by colleagues and an examination of recently preserved specimens from the Malaysian Cameron Highlands. These images and observations allow me to conclude that the three named populations have different colorations. Manthey and Grossmann (1997, hereinafter M&G) depicted a H. larutensis (fig. 172; adult male, Cameron Highlands, Palang, western Malaysia) and a H. typus (fig. 173; adult male, Berastagi, northern Sumatra) (In this para- graph, I am using the species identifications provided by the authors for their illustrated specimens.) The same indi- vidual (my assumption) of H. typus is portrayed in the Cox et al. (1998:85) field guide and in Malkmus et al. (2002: fig. 275). A different individual of M&G?s H. typus is shown in Chan-ard?s et al. (1999:130) (photographic checklist; adult female, Bukit Larut, Perak, Malaysia); another H. typus (Chan-ard et al., 1999:130) (adult male, KhaoYai Natl. Park, Nakhon Rachasima, Thailand) is darker. Chan-ard et al. (1999:128?130) also included six images (Cameron Highlands, Malaysia) of H. larutensis (Chan-ard?s species identification); the male (Chan-ard et al., 1999:130) is the same individual of H. larutensis as presented in M&G?s field guide. Chan-ard et al. (1999:128) also provided an image of H. harterti (adult male, Cameron Highlands). Without specimens available for examination, I cannot confirm the identification of the depicted specimens; nonetheless, a few comments on their specific identification and coloration seem useful although speculative. First, the Berastagi ?H. typus? does not have a typi- cal H. typus coloration, and, indeed, it is quite striking from other populations described above. Because it lacks the dark dorsal crossbars and the dorsolateral series of light spots, I do not believe the specimen was a H. typus. Further, if it was really an adult male, it could not be a H. typus, because H. typus is a unisexual species. Could it be a H. margarethae? Possibly, because it derived from the same mountain range as the latter, although about 3? latitude northward of the Fort de Koch (now Bukittinggi) type locality. The two Chan-ard figured H. typus possess the near uniform mid-dorsa of the Berastagi one, and the Bukit Larut one has the same overall coloration, although darker, including the continuous dark dorsolateral lateral FIGURE 12. Types of Bingtang slender gecko. (A?C) Lepidodacty- lus Harterti Werner 1900 (ZMB 15360): (A) dorsal view of whole body, (B) ventral view of throat and chin, and (C) ventral view of pelvic area (photographs by M.-O. R?del); (D?F) Gehyra laruten- sis Boulenger 1900 (BMNH 1901.3.20.2): (D) dorsal view of whole body, (E) ventral view of throat and chin, and (F) ventral view of pelvic area (photographs by G. Zug). n um b e r 6 3 1   ?   2 9 stripes and bright anterolateral arms of the postsacral mark of the Berastagi gecko. Chan-ard?s Thai H. typus has a dorsolateral series of small light spots and well-de- veloped postorbital stripe; nevertheless, I do not believe it is a H. typus, owing to light anterolateral arms of a post- sacral mark. It is labeled as an adult male and appears to have a hemipenial bulge on the base of the tail. The images labeled H. harterti and H. larutensis represent individu- als from the Cameron Highlands, Malaysia. They do not have the typus-style coloration and might represent one or two bisexual species. Certainly, the pattern of several of the specimens is like Lepidodactylus lugubris, as noted in Werner?s Lepidodactylus Harterti description. Using these images, recent specimens, and colleagues? images, I propose that the coloration of three Sundan populations is distinct for each and offer the following hypotheses. H. margarethae is a lightly marked gecko with a striking contrast between body and tail color. The break between the darker trunk and head coloration is at the postsacral ?mark.? Dorsally, the head and body are a light brown with faded darker brown dorsolateral and lateral stripes. The nape and neck can have a brown mid- dorsal stripe. A light canthal stripe extends from the naris to eye and across the temporal area. The dark postsacral mark is small to absent, but the light arms extend on the posterior trunk. These beige colored arms are continu- ous with a nearly unicolor tail. (The preceding description was based on Manthey and Grossman, 1997: fig. 173.) Because Larut Hills (Bukit Bintang range) and Cameron Highlands (including Fraser?s Hill; both of Banjaran Titi- wangsa range) specimens do not share a common color- ation in any images available to me, I propose them as separate genetic entities; the former equals H. harterti and H. larutensis, and the latter represents an unnamed species. Hemiphyllodactylus harterti has contrasting body?tail coloration as in H. margarethae but is over- all darker. The center of the postsacral mark is a narrow dark brown V, which is continuous with a dark brown dorsolateral stripe on each side of the trunk, extending anteriorly to mid neck. The light colored arms (border) extend to above the hips and posteriorly rapidly fade into the unicolor olive of the tail. Light canthal stripes are not evident. (The preceding coloration was based on Chan- ard et al. (1999:130) Bukit Larut ?typus.?) The Titiwan- gsa gecko is strongly marked. Its background ranges from light tan to olive brown, overlain by a dense dark brown reticulation. This reticulation lacks the regularity of the dark ladder pattern of Lepidodactylus lugubris, although it is reminiscent of that pattern. A narrow dark postsacral V with light-colored (to over hips) is usually present. Tail background color generally matches the light color of the postsacral V; additionally, the tail bears light and dark transverse bars, commonly equal sized. A dorsolateral dark stripe extends from mid-loreal area to axilla. One coloration variant is scattered dark blotches dorsally and laterally on neck and trunk. Pr?cis. (1) Although generally drab, bisexual Hemiphyllodactylus display some bright markings that seemingly are unique for different populations. (2) Color- ful postsacral marks occur in Palau and Philip specimens. The postsacral marks are distinctly U- or V-shaped in the two preceding samples, whereas the dorsolateral arms are weakly developed in most individuals of sE asia and china. The postsacral mark is the beginning of a lighter (than trunk) tail in Sumatran and Larut Hills individuals and less distinct in Titiwangsa specimens. The postsacral mark for india and Hong Kong geckos is the first con- trasting band of the tail. GEOGRAPHYANDTAXONOMY regioNal PatterNs of morPHology aNd sPeciatioN General Observations The present study does not address the generic sta- tus of the various species currently assigned to the genus Hemiphyllodactylus. I recognize the presence of two adult body types among the species currently assigned to this genus (e.g., attenuate body in Hemiphyllodactylus typus and robust body in H. yunnanensis). I, however, weigh the lamellar morphology of the fore- and hindfeet much more strongly and am comfortable with Hemiphyllodactylus as a small monophyletic clade, at least until a robust molecu- lar data set demonstrates otherwise. Morphological Differentiation Several morphological features allow the recognition of unique populations (samples) and sets of populations. Foremost among these features is the absence of males in the Hemiphyllodactylus populations that are the most geographically widespread. Another striking trait is the presence or absence of gonadal, peritoneal, and caecal pigmentation among both unisexual and bisexual popula- tions. Two body forms are displayed among populations, at least in the older and/or mature individuals. Further, av- erage adult SVL differs among populations, even though size at maturation appears similar among all populations. Three features of scalation show populational dif- ferences. Chin scales in contact with mental and anterior 30   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY infralabial scales show two patterns: small (subequal) or slightly larger than the posteriorly adjacent chin scales; or distinctly enlarged with median pair largest (= postmen- tal scales). Secreting precloacal and femoral pores occur consistently in adult females of some Hemiphyllodactylus populations and rarely or not at all in other populations. Also, two precloacal?femoral pore arrangements exist (each state uniform within a population): continuous pore series or discontinuous with precloacal pores separated from left and right femoral pores series. In all popula- tions the shape of the digital lamellae is distinctly U- or lyre-shaped. These U-shaped lamellae match Russell?s morphological definition (Bergmann and Russell, 2003) of scansors, and they distinctly show a dense microvillous surface as the pad surface dries. The number of lamellae on the digits and hence the digital lamellae formulae varies among populations. Although these geckos are mostly dull in coloration and this somberness is accentuated by their small size, color and pattern also offer evidence of populational dif- ferentiation. The overall dorsal pattern of dark blotches and stripes ranges from nearly nonexistent or diffuse to bold stripes and bars. Intensity and color of the dorsolat- eral spots varies among populations, as does the presence and coloration of the postsacral mark. Beginning with reproductive biology, unisexuality is universal among most Pacific-area populations and coastal ones from southern Melanesia through Sunda and Southeast Asia to Sri Lanka. These populations and the Mascarene ones represent the nominate H. typus. All these populations share a densely pigmented intestinal caecum, which is fre- quently visible through the body wall and the pigmentation of the peritoneum covering the gonads and gonadal ducts (specifically the oviducts and epididymides). This pigmen- tation is not unique to the unisexual populations but also occurs in bisexual ones: Palau, Philip, and india. I propose that this unique visceral pigmentation (Fig- ure 2) reflects close phylogenetic relatedness among the unisexual H. typus and the three bisexual species (justi- fication of the specific status for each Palau, Philip, and india below). I further propose that H. typus arose from the hybridization of individuals from the Philippines and Palau. An electrophoretic study (T. D. Schwaner, unpub- lished data, 1990) examined individuals from northern Thailand (n = 13), Philippines (3), Fiji (5), and Hawaii (9) using nine isozymic loci. The Philippine and two Pa- cific samples share allozymes for all nine loci; although for GPD and IDH, the Philippine sample has two alleles for each of these two isozymes and MPI of the Pacific samples has two alleles. The Thai sample shares alleles for only three isozyme loci for the Philippine and Pacific samples. The similarity of shared alleles supports the Phil- ippine species as one of the parental species of H. typus. Similarities in morphology of Palau and Philippine indi- viduals suggest the former as the other parental species. When and where the hybridization occurred is far more speculative, although a few reasonable conjectures are possible based on the present observations. First, the hy- bridization event(s) occurred somewhere other than on the parental island groups. Hemiphyllodactylus typus occurs infrequently in the Philippine sample and not at all in the Palauan one. Additionally, survival of hybrids without a competitive (numerical) swamping by bisexuals and the opportunity to develop sufficient population density for subsequent dispersal seem more likely to occur off the pa- rental islands than on them. This supposition derives from my observation on rarity of H. typus on the islands of the south central Pacific and their rarity on human-made structures in the presence of other gecko species. When other geckos occur with H. typus, it occurs in the dark- est area, well removed from Gehyra and Lepidodactylus lugubris, and this segregation occurred before the Hemi- dactylus frenatus invasion of the late twentieth century. Unlike L. lugubris in which the unisexuals have displaced the bisexuals to marginal habitats (Ineich, 1999), H. typus has not displaced bisexuals or has displaced them only in human habitats. I further speculate that the hybridization event occurred in a location outside the distribution of a bisexual Hemiphyllodactylus and likely subsequent to the arrival of humans to the Pacific islands. The preceding ori- gin hypothesis derives from my assumption that the entire distribution of H. typus derives from human transport? shipboard, possibly during the eighteenth and early nine- teenth centuries. Phenetic similarities (i.e., primitive advance state po- larities not determined) suggest the relatedness of the Pa- lauan and Philippine populations. They share the elongate habitus, small body size (medians, 33 and 35 mm SVL, respectively; Table 3), separated precloacal and femoral pore series, no enlarged chin scales (Table 6), usually two cloacal spurs on each side, and similar coloration traits of a U-shaped postsacral mark and series of small, but bright, spots dorsolaterally on the trunk. Of the preceding traits, the two Pacific bisexual populations share habitus, body size, pore morphology, and chin scale and cloacal spur morphologies with the Indian sample. The presumed hybridization of the Pacific bisexuals yielded a unisexual hybrid (H. typus Bleeker) with a larger average body size (>36 mm SVL) but otherwise identical or very simi- lar to the bisexual parental species. H. typus is the only n um b e r 6 3 1   ?   3 1 Hemiphyllodactylus taxon in which most (>90%) adult females have secreting precloacal pores. The pores of the unisexual adults appear generally smaller than those of the males of the bisexual taxa. Dorsolateral spots occur in unisexual individuals, although the spots are white to light tan, and the postsacral mark has lost the anterolat- eral arms. Even though similar, Palauan and Philippine popula- tions have morphological differences that I interpret to demonstrate speciation. The dorsolateral spots and post- sacral mark are red in the Philippine population (H. insu- laris Taylor) in contrast to pink spots and yellow suffused with pink postsacral mark of the Palauan population (un- named). These two populations also differ in average size, with H. insularis slightly larger; more than half the Philip sample exceeds the maximum SVL of Palau specimens. The size distribution of Palau seems unlikely to result from sampling owing to the number of individuals in the sample and the thoroughness, recency, and duration of the Palauan inventory effort (Crombie and Pregill, 1999). Pa- lauan geckos have a proportionately broader head than H. insularis ones (65 versus 59% HeadW/HeadL; Table 3, and see also SnW/HeadL therein). In scalation, Palauan geckos average more TotPore than did the Philippine pop- ulation (27 versus 22.5; Table 6); as noted in the Results section, an increase in TotPore reflects the addition of more femoral pores to the left and right series. The Indian sample (H. aurantiacus Beddome) is a dis- tant geographic outlier to the Pacific Hemiphyllodactylus taxa. The Indian geckos also differ strikingly from them by a bolder coloration. The dark transverse dorsal markings are broader and more continuous across the dorsum, and the pre-postorbital stripe is also broader and continuous from naris to shoulder. The postsacral mark is dark yel- low to gold and the anterolateral arms are variously devel- oped. The average size of Indian geckos (Table 3) matches that of H. insularis, but the relative trunk length (TrunkL/ SVL) is shorter (49%) in H. aurantiacus than the Pacific taxa (53% Philip, 54% Palau). H. aurantiacus has a sig- nificantly broader head (76 versus 59 and 65% HeadW/ HeadL) than the Pacific taxa, although relative snout width (SnW/HeadL) is the same for all three. The most striking difference of H. aurantiacus is its digital lamellae formulae of 2-2-2-2 forefoot and 2-2-3-2 hindfoot. No other popu- lation of Hemiphyllodactylus has such a low number of lamellae on its digits. The two Sri Lankan non-typus speci- mens examined have very different formulae: 2-2-2-2 and ?-3-3-2 adult male (BMNH 1910.3.16.4); 3-4-4-4 and 5-5- 6-5 adult female ([NMB 8552). The male is potentially a H. aurantiacus. Its locality data are Hambonota, Ceylon; however, given that other early BMNH specimens with Ceylon locality data are not members of the Sri Lankan herpetofauna (R. Somaweera, personal communication, March 2007), its status is questionable until a broader sampling of Sri Lankan Hemiphyllodactylus is available. The NMB female?s locality is only Ceylon; it is possibly a H. typus with an unusual hindfoot lamellae formula. The broader geographic ranges of the eastern Asian samples and prior recognition of multiple taxa makes the resolution of populational and taxonomic boundaries dif- ficult. Bisexual Hemiphyllodactylus populations occur from west-central Myanmar (CAS 231030, Chin State) to eastern Guangxi (Zhou et al., 1981: fig. 7). Eastward, a gap of ~1000 km exists from the Guangxi occurrence to the population in the Hong Kong area (Lazell, 2002; Chan et al., 2006). Bisexual populations occur southward from Yunnan-Guangxi through Malayan Peninsula and the Greater Sunda Islands. The occurrence of populations in Indo-China/SE Asia appears very spotty. It is uncertain whether this spottiness is actual occurrence or simply a result of few inventory surveys and/or inadequate vouch- ering of specimens. I must note that finding Hemiphyl- lodactylus even with intense field surveys is uncommon. For example, in the Pyin-Oo-Lwin area (Myanmar), six researchers inventoried a site multiple times over 2 weeks (mid-monsoon, August 2003) and found only two indi- viduals, one each on two consecutive evenings. The Chinese populations of H. yunnanensis seemingly contrast sharply with the preceding statement of rarity. Late nineteenth century sampling in the Kunming (=Yun- nan-fu) area resulted in series of H. yunnanensis in many European and North American museums. More recently, Zhou et al. (1981) amassed a collection of 640 specimens of this taxon from nine localities within Yunnan and Gui- zhou. This number does not indicate rarity within this area and possibly reflects the absences of microsympatry with other geckos, particularly Hemidactylus (see Zhao and Adler (1993:304) for list of Yunnan and Guizhou geckos). This abundance also supports the low competitiveness hy- pothesized for H. typus earlier in this section. Hemiphyllodactylus yunnanensis and the other sam- ples and populations of mainland eastern Asia share the adult robust body form, absence of caecal and gonadal pig- mentation, a pair of enlarged postmental chin scales, and a continuous series of precloacal?femoral pores in adult males. The robust morphology (Figure 3) is most apparent in larger individuals and perhaps in adult females, although my measurements and proportions are inadequate to test this proposition. Juveniles and subadults retain a slender trunk, hence a more elongate appearance than adults. The 32   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY enlargement of chin scales is more variable within and among populations. None of the eastern Asian populations lack the enlargement of the pair of chin scales abutting the mental scale; however, the degree of enlargement ranges from barely to strongly among populations. The relative enlargement is consistent within each population. Zhou et al. (1981) showed two patterns of postmental enlarge- ment (Figure 7D,E) in their H. yunnanensis sample. In their H. y. yunnanensis, the postmentals are large with a rapid decrease in size of the arc of chin scales bordering the in- fralabials; the next arc of chin scales are distinctly smaller and nearly subequal to the remainder of chin scales. The postmentals of H. y. longlingensis are moderately large, the infralabial abutting arc of chin scales slightly enlarged as also are the scales in the second arc, inwardly the chin scales quickly become smaller and ?typical.? Are these two patterns discrete or a continuum? I favor the latter inter- pretation, although individuals from single-site subsamples that I have examined can usually be assigned to a single state, a few individuals grade into the opposite state. As noted in the Results section, my chin character does not discriminate between the two postmental states of Zhou et al., but all eastern Asian samples share the low count state, including the holotype of Hemiphyllodactylus typus chapaensis Bourret. I single out this latter specimen for two reasons: First, Bourret noted small chin scales abutting the mental and infralabials, and his illustration (reproduced in Figure 7F) is ambiguous on this condition; second, if the chapaensis population has no enlarged postmental, it differs from the other eastern Asian populations. In this respect, Lazell (2002; also later publications) has identified the Hong Kong population as H. chapaensis. Hong Kong geckos match the general eastern Asian morphology, per- haps with a proportionately longer trunk (53 versus <49% for all other Asian subsamples). They have enlarged, al- though not proportionately larger, postmentals than other Asian mainland population; however, the second arc and some of third arc of chin scales seem proportionately larger than in the other populations. In a contrasting situation, a southern Vietnamese specimen (USNM 146161) has strik- ingly large postmentals, similar to those depicted for H. margarethae in Figure 7C. In contrast, the single southern Cambodian specimen has an arc of small scales. There is no pattern/differentiation in precloacal? femoral pores visible within the populations and samples from mainland southeastern Asia. These populations share the continuous pattern of precloacal and femoral pore series in contrast to discontinuous pattern of uni- sexual H. typus and the bisexual populations of India, Philippines, and Palau. The continuous pattern contin- ues through the peninsular populations of Thailand and Malaysia. In contrast, the bisexual males of Sumatra and Borneo have discontinuous (separated) pore series. Taxonomic Decisions and Geography Which came first, the gecko or the egg? This twisted banality highlights the necessity of addressing the geogra- phy of unisexual Hemiphyllodactylus separately from that of the bisexual species. Because squamate unisexuality ap- pears to be universally derived from hybridization (Zug et al., 2001), I examine the geographic patterns of occur- rence and taxonomy of the bisexual species first. Among the bisexual species, H. aurantiacus (Bed- dome) was the first to be described and is the easiest of the Hemiphyllodactylus populations to be addressed taxo- nomically. Small adult body size and uniquely low fore- and hindfoot lamellar formulae readily identify it as a distinct genetic entity and phylogenetic lineage. Assuming habitus, visceral pigmentation, and precloacal?femoral pore morphology reflect phylogenetic affinity, H. auran- tiacus is a member of the typus clade. The two other bisexual members of the typus clade are geographically distant?Palau and Philippine islands ?from H. aurantiacus. Taylor (1918) recognized the distinctiveness of the Philippine populations by several minor differences in scalation from Stejneger?s description of Hemiphyllodactylus leucostictus. He even noted that it might not be distinct from the latter species, although he highlighted some differences in coloration. Crombie and Pregill (1999) were the first biologists to document the occurrence of Hemiphyllodactylus in Palau. Early in their herpetofaunal survey of this island group, they rec- ognized the differentiation of the Palauan population from H. typus, first because of the presence of males and second because of its distinct coloration. The latter trait and a few others demonstrate its uniqueness to me, and I describe it as a new species in the following Species Account section. The bisexual populations of mainland and islands Asia are less readily delimited. This difficulty results largely from inadequate sampling in number of individu- als from most localities, too few localities, and localities geographically distant. The uniqueness of H. yunnanensis (Boulenger) is unquestionable. Its traits (robust body, un- pigmented caeca and gonads, and continuous precloacal? femoral pores) serve to identify a China?Indochina?Sunda clade. I propose that the populations from north central Burma through south central China and the adjacent northern half of Thailand, Laos, and northern Vietnam represent a single taxon H. yunnanensis. Some regional populational differentiation occurs in this widely distrib- uted species, although morphological data suggest minimal n um b e r 6 3 1   ?   3 3 genetic differentiation. This hypothesis thus relegates the Zhou et al. subspecies to the synonymy of a monotypic H. yunnanensis. Hemiphyllodactylus typus chapaensis Bour- ret similarly becomes a synonym of H. yunnanensis (Bou- lenger). Geographically, the H. t. chapaensis type locality is only 300 km south of Kunming (type locality of H. yun- nanensis) at the southern terminus of a continuous range of mountains. This mountain range also contains the type locality of H. y. jinpingensis Zhou et al., and this locality is less than 100 km north of Chapa. Zhou et al. (1981) did not compare the morphology of their samples to H. t. chapaensis Bourret. I noted earlier Lazell?s use of H. chapaensis for Hong Kong Hemiphyllodactylus. This usage must be discontin- ued because the morphology of the Hong Kong slender geckos is distinct from that of H. typus chapaensis Bourret. Does the Hong Kong population represent an outlier of H. yunnanensis or is it a unique Hemiphyllodactylus popula- tion? My present data suggest the latter interpretation, but they are insufficient to address this hypothesis rigorously. Presently, I am unaware of vouchered records of Hemi- phyllodactylus in Guangxi and Guangdong provinces of southern China. A broad distributional gap also exists for geckos of the Hemidactylus bowringii group. Initially, we (Zug et al., 2007) proposed that Hong Kong ?bowringii? was an exotic species, accidentally introduced from Burma or India. A more detailed study (McMahan and Zug, 2007) subsequently demonstrated the uniqueness of the Hong Kong population. This latter study urges caution, so I am hesitant to postulate the specific status of the Hong Kong Hemiphyllodactylus without enlarging my sample and lo- cating vouchers from the Guangxi?Guangdong corridor. The preceding restriction (in Morphological Differ- entiation subsection) of H. yunnanensis to slender gecko populations from southern China and adjacent northern Indochina leaves the taxonomy of southern Indochina populations unresolved. Again, the inadequacy of mu- seum vouchers does not permit a satisfactory resolution. Presently, the data are adequate to declare that Malay peninsular populations are distinct; however, it is uncer- tain whether they are confined to the peninsula or extend into southern Indochina, likely the former owing to their restricted occurrence to montane rainforests. The single specimen (USNM 146161) from southern Vietnam sug- gests this possibility, although morphologically, it appears more similar to the Sumatran population. Also, a sample (THNHM 4910-4917) from Kaeng Krachan National Park, Thailand, just north of the Isthmus of Kra is more similar to other southern Thailand geckos than to Malayan ones. For the present, I recommend labeling southern In- dochina and Hong Kong specimens as ?H. yunnanensis.? Two names are available for Malayan Hemiphyllodac- tylus populations. H. larutensis derives from Bukit Larut (=Maxwell?s Hill or Larut Hills), a mountain at the south- ern end of the Bintang range. Although the type locality of H. harterti was given as ?Malakka? by Werner (1900), Boulenger (1912) noted that this locality name referred to a general locality and not a specific site (i.e., Malakkahal- binsel [German] equals Malaya [English]). Furthermore, because Ernest Hartert (the donor of the specimen to the Berlin museum) collected birds at Gunong Inas in 1888 (Hartert, 1901?1902; AMNH Department of Ornithol- ogy records), Boulenger (1912) tentatively recommended changing the type locality to Gunong Inas. Gunong Inas is ~40 km north of Bukit Larut, and both are mountains within the western Malayan mountain range (Bukit Bin- tang). I have examined personally only one specimen (G. larutensis type) and have data and images from a sec- ond specimen (L. Harterti type), and there are no traits suggesting that these two represent different taxa. Thus I accept Boulenger?s restriction of the type locality to Gu- nong Inas. I note only that Werner?s (1900:196) ? gleicht in F?rbung und Zeichnung ganz dem L. lugubris? bet- ter matches the pattern of geckos of the central Malayan mountain range (Bunjaran Titiwangsa); nevertheless, I can find no evidence that Hartert collected in the latter moun- tain range or received specimens from there. The similarity of morphology of the two type specimens and the type re- striction of H. harterti to the Bintang range makes H. har- terti the senior synonym and valid name for the Bintang taxon. The preceding nomenclatural decision results in the populations of Bunjaran Titiwangsa, at least those from the Cameron Highlands southward to the Fraser Hill area, being nameless. These populations have coloration and minor morphological differences from H. harterti, their occurrence in a mountain range with a different geological history and long separation by a lowland valley leads me to propose them as a separate phylogenetic lineage. The sample of Sumatran bisexual geckos is modest but adequate to demonstrate the morphological distinc- tiveness of these geckos. Thus, in spite of Brongersma?s belated change of mind, H. margarethae is a unique lin- eage and presumably Sumatran endemic species. Like H. harterti and the central mountain taxon, it appears to be a montane species, confined to cooler and moister habitats than the invasive H. typus, whose type locality occurs in Sumatra as well. I identified most Bornean Hemiphyllodactylus speci- mens as unisexual typus. As I analyzed and reexamined my data, I discovered that my identifications resulted in all female specimens as H. typus and the three males as unknowns. All Bornean slender geckos have pigmented 34   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY caeca; all males have pigmented testes-epididymides peri- toneum, all females except one with pigmented oviducal peritoneum. Were my identifications driven by knowledge of sex? The evidence definitely suggests a bias. A reexami- nation of the specimens is not possible at this stage of the analysis; however, a reexamination of the data suggests that most females were correctly identified as H. typus. Tentatively, bisexual individuals can be differentiated from unisexual ones by a higher lamellar formula. The presence of males with normal appearing gonads indicates the presence of a bisexual species as well as the unisexual H. typus on Borneo (entire island, not just Indonesian Bor- neo). Formal recognition of this bisexual species requires a reexamination of all Bornean specimens and comparison with the Philippine H. insularis. Elsewhere in the Sundas, all Hemiphyllodactylus are unisexual typus. A peculiar situation occurs in Komodo where all individuals are pale, hence Auffenberg?s (1980) decision to describe them as a subspecies (pallidus) of H. typus. Pale H. typus are observed elsewhere as one end of a color-phase shift. Because Auffenberg had only two specimens and presumably observed no additional ones, I suggest that by chance he collected and preserved both in their light phase. Presumably, selection for lighter indi- viduals can occur in clonal vertebrates, but the Komodo environment does not seem greatly different from other island habitats with resident populations of H. typus. Phylogeny and biogeography: specula- tions. This study was not designed to address phy- logenetic relationships within Hemiphyllodactylus or its relationships with other gekkonid genera. Morphological similarities and differences among the recognized species taxa led to my hypothesis of two subclades?the typus and the yunnanensis species groups?and the monophyly of Hemiphyllodactylus. Accepting these two items suggests an ancient origin (?Miocene or earlier) of Hemiphyllodactylus and an early divergence of the elongate and robust-bodied species groups. Kluge (1968) implicitly proposed Hemi- phyllodactylus as a member of the Lepidodactylus clade, although in his study, he examined nomenclatural issues, not phylogenetic ones. His subsequent studies of gekkonid relationships did not address directly either the interspecific or intergeneric relationships of Hemiphyllodactylus. Simi- larly, this genus has been absent from most other gekkonid phylogenetic studies, although the broad taxon-based stud- ies of A. Bauer and colleagues likely will provide informa- tion on the relationships of Hemiphyllodactylus and some of the species in this genus. Their initial evaluation (Han et al., 2004) leaves its relationships unresolved. Having proposed the existence of two sister groups (subclades) of Hemiphyllodactylus, I wish to delineate and define these species groups. The typus species group con- tains four, and possibly five, species: H. typus Bleeker, H. aurantiacus (Beddome), H. insularis Taylor, the Palauan population described in the following species accounts, and possibly a fifth, the populations in Borneo. The yun- nanensis species group contains four, and likely more, spe- cies: H. yunnanensis (Boulenger), H. harterti (Werner), a central Malaysian taxon, H. margarethae Brongersma, and possibly a separate species in South China?Hong Kong and southern Indochina. The typus species group members have an elongated habitus, accentuated by a long trunk (TrunkL/SVL >50%) with short limbs and proportionately small head, caecum and gonadal ducts darkly pigmented, and precloacal and femoral pore series separated. The yunnanensis species group members have a stouter adult habitus although still elongated (TrunkL/ SVL ~50%) and proportionately larger head, caecum not pigmented and gonadal ducts rarely so, and precloacal and femoral pore series separated or continuous. Continuing my speculations on origins and biogeog- raphy of the bisexual populations, I suggest that the elon- gate clade derived from the more robust clade and that its origin occurred in northern precursor-Sunda with disper- sal eastward into the Philippines and hence into the west- ern Pacific, and westward to peninsular India, now with only relict populations remaining. Hemiphyllodactylus aurantiacus represents a surviving member of this early ?dispersal? to India with a long period of isolation. Pres- ently, the situation for the Sri Lankan bisexual population is undecipherable owing to paucity of data. The robust- bodied Hemiphyllodactylus has remained and differenti- ated within the area of the group?s origin and northward in southwestern China. I suggested previously (Morphological Differentia- tion section) that the origin of the hybrid H. typus was an extralimital hybridization of H. insularis and the Palauan species. When the hybridization occurred is unknown. A human-induced event seems most likely, as also does an ?off the native island? event. The event could have occurred during the early human migrations into and through the Pacific islands or, my preference, with the Euro-American exploration and commercialization of the Pacific. ?Off-island? hybridization seems likely owing to presumed low interspecific competitiveness of the unisex- ual H. typus in contrast to the displacement of parental bisexuals by clones of the unisexual Lepidodactylus lugu- bris (Ineich, 1999). I postulate the possibility of onboard ship hybridization then colonization and population ex- pansion in central Pacific. While such an origin could have occurred during the initial human migration into the Pa- cific, onboard hybridization seems more likely aboard the n um b e r 6 3 1   ?   3 5 larger European sailing ships. Whose ships? The broad distribution and the early occurrence in the Mascarenes hint at an association with French exploration; however, the impact of whaling vessels for the dispersal of Pacific lizards has been overlooked. Whaling vessels were notori- ous for their catholic (trashy) cargo and for their regular and widespread shore leave. Thus whaling ships provide a ?safe-haven? habitat for hybridization and the initial survival of the hybrids; then they could serve for the trans- fer of hybrids to diverse islands and to other whalers to broaden the dispersal of the hybrids. Other phylogenetic and biogeographic questions re- main for this low diversity taxon. First and foremost is why are there so few species in this gekkonid clade? Its putative sister clade, Lepidodactylus, has at least 4 times as many species, and its unisexual species L. lugubris (multiple clones, multiple origins) has ?outcompeted? its bisexual parent in many Pacific island ecosystems. The Hemiphyl- lodactylus clade displays several distributional anomalies. The widespread occurrence of unisexual H. typus con- trasts sharply with the bisexual species. In one aspect, this feature is shared with L. lugubris, although its low density at most invasive sites differs greatly from mourning gecko occurrences. Also, in contrast to L. lugubris, it dispersed more widely. Another peculiarity is the dominance of the stout-bodied Hemiphyllodactylus in southeastern Asia and the division of the slender body clade into extreme east- ern and western outposts. The stout-bodied geckos show a southern China distributional hiatus, absent from the China?Indochina border then occurring in the Hong Kong area. I earlier noted a similar pattern for the Hemidactylus bowringii species group, hence my reluctance to attribute the Hong Kong occurrence to human introduction. The documentation of the distributional patterns of tropical Asian amphibians and reptiles is rudimentary. This situ- ation derives from our poor knowledge of tropical Asian diversity, particularly among mainland taxa, and this poor knowledge results from the continuing recognition of pan- Asian species, when few such species exist (Stuart et al., 2006; Zug, In press). sPecies accouNts Hemiphyllodactylus typusBleeker Indo-Pacificslendergecko Hemiphyllodactylus typus Bleeker, 1860:327 [type locality: ?Agam? (Suma- tra); holotype, BMNH 1946.8.30.83]. Platydactylus crepuscularis Bavay, 1869:8 [type locality: ?Nouvelle-Cal?do- nie? (locality implied from title of publication); holotype lost (Brygoo, 1990:49)]. Spathodactylus mutilatus G?nther, 1872:594 [type locality: ?East-Indies ar- chipelago?; holotype, BMNH 1946.8.30.83]. Lepidodactylus ceylonensis Boulenger, 1885:164, Pl. XIII, fig. 3 [type local- ity: ?Gampola? (Ceylon); holotype, BMNH 74.4.29.1326]. Hemiphyllodactylus leucostictus Stejneger, 1899:800 [type locality: ?Kauai, Hawaiian Islands?; holotype, USNM 23500]. Hemiphyllodactylus typus pallidus Auffenberg, 1980:72 [type locality: ?along Vai Nggulung, Loho Liang, Komodo, 30 mm?; holotype, UF 28985]. Hemiphyllodactylus albostictus Lazell, 1989:126 [spelling error]. comments. ?Bleeker?s types were sold to the British Museum in 1863 (or at least years before 1879, . . .). The type of H. typus is the same as that of Spathodactylus mutilatus Gthr. (Boulenger l.c).? The preceding statement is from Brongersma?s review (1932:212[footnote 2]) of Hemiphyllodactylus. Taylor?s (1963) description of H. typus was based on a male from Fraser?s Hill, Malaysia. In addition to the speci- men being a male, it had a continuous precloacal?femoral pore series; hence the description is not representative of H. typus. description. An all-female taxon of geckos (Gekkoninae) with elongate, slender habitus, slightly compressed, elongated appearance accentuated by short limbs and small head (see Figures 3, 6, 13), tail round in cross section and commonly shorter than SVL. Adults 29.4?46.1 mm SVL (mean ? SD, 38.4 mm ? 2.91; n = 143), 14?36 mm TailL (28.5 mm ? 5.03), 15.0?28.0 mm TrunkL (20.3 mm ? 2.11), 6.6?9.9 mm HeadL (8.2 mm ? 0.56), 3.7?6.6 mm HeadW (5.2 mm ? 0.52), 2.3?4.1 mm SnEye (3.4 mm ? 0.30), 1.8?3.4 mm NarEye (2.6 mm ? 0.28), 1.5?2.4 mm EyeD (2.1 mm ? 0.17), 0.9?1.7 mm SnW (1.3 mm ? 0.17). Adult proportions 40?65% TrunkL/SVL (mean ? SD, 52.9% ? 3.2), 18?24% HeadL/ SVL (21.3% ? 0.9), 10?16% HeadW/SVL (13.7% ? 1.1), 51?77% HeadW/HeadL (64.1% ? 4.8), 34?48% SnEye/ HeadL (41.1% ? 2.4), 24?40% NarEye/HeadL (32.2% ? 2.8), 20?32% EyeD/HeadL (25.3% ? 1.9), 11?21% SnW/ HeadL (16.2% ? 1.9), 61?106% EyeD/NarEye (79.0% ? 7.9), 16?34% SnW/HeadW (25.3% ? 3.3). Scalation predominantly granular from head onto tail, both dorsally and ventrally; ventral trunk scales slightly larger than dorsal ones, 12?19 Dorsal (median ? SD, 15 ? 1.6) and 8?14 Ventral (10 ? 1.3); similarly, subcaudal scales slightly larger than dorsal caudal scales but not plate- like. Cloacal spurs present, modest sized, 1?5 CloacS (2 ? 0.8). Larger scales on lips and snout, rostral largest, rect- angular to pentagonal, often slightly concave on dorsome- dial edge with slight cleft; 1?5 CircNa (3 ? 0.6), 1?5 SnS (2 ? 0.7); labial scales enlarged from rostral to below eye, 36   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY becoming progressively smaller in subocular rictus, 9?14 Suplab (11 ? 1.0), 7?13 Inflab (10 ? 0.9); 9?14 Chin (11 ? 1.1), those behind mental slightly or not enlarged; ear opening distinct with no bordering enlarged scales. Each digit with expanded pad, terminal two phalanges free, aris- ing from within pad on second to fifth digits of fore- and hindfoot and each clawed; pads of these digits each with large triangular apical lamella bordered proximally by lyre- shaped lamellae (scansors); modal digital formulae 3-4-4-4 (forefoot) and 4-4-5-4 (hindfoot) for scansors; first digit of fore- and hindfeet compressed, usually 5 rectangular lamel- lae (4-5 fore, 5-6 hind) ventrally, terminal phalanx not free with or without minute claw. Adults usually with precloa- cal pore series (0?14 PreclPor, median ? SD = 10 ? 3.9) always separated from femoral pore series (0?12 combined left and right femoral series), 0?26 TotPore (12 ? 7.2). Dusky tan to reddish brown ground color dorsally and laterally from head to tail, usually with dark ocular stripe from loreal area to anterior trunk; series of widely spaced small white spots, often darkly edged, dorsolater- ally from temporal area to inguina; dorsal postsacral dark brown blotch bordered posteriorly by transverse bar of white or beige. Underside dusky from chin to vent, pale yellowish orange on tail. Major diagnostic features are as follows: all-female taxon; pigmented caecum and gonadal ducts; if present (uncommonly), femoral pore series separate from pre- cloacal pore series; chin scales bordering mental and first infralabial not greatly enlarged; digital lamellae formulae 3-4-4-4 (forefoot) and 4-4-5-4 (hindfoot); average adult SVL ~38 mm; series of white spots dorsolaterally on trunk and bright postsacral bar of white and dark brown. Description of holotype: An adult female (Figure 13), 43.3 mm SVL, 40.0 mm TailL, 22.4 mm TrunkL, 9.1 mm HeadL, 5.8 mm HeadW, 3.7 mm SnEye, 2.9 mm NarEye, 2.1 mm EyeD, and 1.1 mm SnW. Proportions: 52% TrunkL/SVL, 21% HeadL/SVL, 13% HeadW/SVL, 64% HeadW/HeadL, 41% SnEye/HeadL, 32% NarEye/ HeadL, 23% EyeD/HeadL, 12% SnW/HeadL, 72% EyeD/ NarEye, 19% SnW/HeadW. Scalation: 3 CircNa, 3 SnS, 11 Suplab, 11 Inflab, 13 Chin (anteromedial ones only slightly larger than adjacent ones), 13 Dorsal, 8 Ventral, 2 CloacS, Subcaud not enlarged, 15 PreclPor, 23 TotPore with no contact between precloacal and femoral, digital formulae 3-3-4-3 fore and 3-4-4-4 hind. Pigmented cae- cum, pigmentation unknown for oviducts. Body ground color faded to a uniform orangish tan, no lateral spotting evident, dark lateral stripe from in front of eye to axilla, broken dark reticulations on rear of head and nape, dark chevron at tail base. Etymology. Bleeker offered no explanation of his selection of the name typus. The name is a Latin noun for impression, shape, Figure, or example. I assume that he chose typus because this species represented the type species for his new genus Hemiphyllodactylus. I propose a standard English name at variance to the commonly used Indo-Pacific tree gecko. Whereas this gecko is occasionally found on trees, it more commonly occurs in the leaf whorls of Pandanus and on human- made structures. Slender refers to the elongate and attenu- ate appearance of H. typus and most of its congeners. Variation. The means or medians and ranges are detailed in the preceding Description section. No sig- nificant or striking variation was seen in any of the char- acters among the samples from throughout the range of this taxon. The variation observed within each character is equivalent to that observed in the repeated measures data of a Palauan bisexual female. Where character variation is greater, the variation is likely attributed to data-gath- ering variance arising from the poor preservation qual- ity of many specimens. I suggest that this low variation among the widely separated population is evidence that all populations of H. typus derive from a single hybridiza- tion event and subsequent dispersion of this single clonal FIGURE 13. Holotype of Hemiphyllodactylus typus Bleeker, 1860 (BMNH 1946.8.30.83): (A) dorsal view of whole body, (B) ventral view of throat and chin, and (C) ventral view of pelvic area. n um b e r 6 3 1   ?   3 7 population. The lightness of Auffenberg?s Komodo speci- mens is attributed to chance. distribution. Broadly, if somewhat spottily, distributed from Hawaii and French Polynesia in the cen- tral Pacific westward to Pacific Rim islands and coastally through New Guinea, Sunda, and Indochina to Sri Lanka; also occurs in Mascarene Islands (Figure 14). Hemiphyllodactylus aurantiacusBeddome SouthernGhatsslendergecko Hemidactylus aurantiacus Beddome, 1870:33 [Type locality: ?Shevaroys, un- der stones about Yercaud and elsewhere at an elevation of 4000 feet? (Tamil Nuda, India); syntypes: BMNH 74.4.29.1332?1337, ZMB 10233; lectotype, BMNH 74.4.29.1333]. comments. Beddome?s description is am- biguous on the number of specimens available to him as he composed the description. His description is based on a single specimen (~30?31 mm SVL), presumably imma- ture, sex uncertain (no precloacal pores). The syntypic series (BMNH 74.429.1332?1337) consists of nine in- dividuals, two adult males, three females (all with adult SVLs but two with immature ovaries), and four juveniles. I designate the male BMNH 74.429.1333 as the lectotype of Hemidactylus aurantiacus Beddome, owing to my re- luctance to use an immature specimen as a type and the absence of an immature specimen matching Beddome?s dimensions. Additionally, I accept Bauer and G?nther?s (1991) assessment that ZMB 10233 is a syntype; it is a mature male (32-mm SVL) with precloacal and femoral pores, hence also not the specimen on which Beddome based his description. description. A bisexual taxon of geckos (Gekkoninae) with elongate, slender habitus, slightly com- pressed, elongated appearance accentuated by short limbs and modest head (see Figures 3, 11, 15), tail somewhat elliptical in cross section and regularly shorter than SVL. Adults 27.2?37.9 mm SVL (mean ? SD, 34.3 mm ? 2.80; n = 14), 26?33 mm TailL (29.2 mm ? 2.87), 13.8?18.8 mm TrunkL (16.7 mm ? 1.27), 6.0?8.7 mm HeadL (7.9 mm ? 0.64), 4.3?6.5 mm HeadW (5.6 mm ? 0.72), 2.3? 3.7 mm SnEye (3.1 mm ? 0.36), 1.9?2.8 mm NarEye (2.4 mm ? 0.25), 1.7?2.1 mm EyeD (2.0 mm ? 0.13), 1.2?1.5 mm SnW (1.3 mm ? 0.10). Adult proportions 44?51% TrunkL/SVL (mean ? SD, 48.7% ? 2.0), 21?26% HeadL/ SVL (23.0% ? 1.0), 14?19% HeadW/SVL (16.5% ? 2.9), 57?79% HeadW/HeadL (71.3% ? 7.1), 34?42% SnEye/ HeadL (39.2% ? 2.6), 27?33% NarEye/HeadL (30.6% ? 1.8), 22?28% EyeD/HeadL (25.0% ? 1.7), 14?20% SnW/ HeadL (16.7% ? 1.4), 69?96% EyeD/NarEye (82.0% ? 7.7), 21?30% SnW/HeadW (23.5% ? 2.7%). Scalation is predominantly granular from head onto tail, both dorsally and ventrally; ventral trunk scales slightly larger than dorsal ones, 11?17 Dorsal (median ? SD, 13.0 ? 1.87) and 8?12 Ventral (10.0 ? 1.51); similarly, subcaudal scales slightly larger than dorsal caudal scales but not plate-like. Cloacal spurs present, modest sized, 1?3 CloacS (2 ? 0.6). Larger scales on lips and snout, ros- tral largest, rectangular to pentagonal, often slightly con- cave on dorsomedial edge with slight cleft; 2?4 CircNa (3 ? 0.5), 3?6 SnS (4 ? 0.9); labial scales enlarged from rostral to below eye, becoming progressively smaller in subocular rictus, 10?13 Suplab (10 ? 1.0), 8?12 Inflab (11 ? 1.0); 10?14 Chin (11 ? 0.8), those behind mental slightly or not enlarged; ear opening distinct with no bordering en- larged scales. Each digit with expanded pad, terminal two phalanges free, arising from within pad on second to fifth digits of fore- and hindfoot and each clawed; pads of these digits each with large triangular apical lamella bordered proximally by lyre-shaped lamellae (scansors); modal digi- tal formulae 2-2-2-2 (forefoot) and 2-2-3-2 or 3 (hindfoot) for scansors; first digit of fore- and hindfeet compressed, usually 4 rectangular lamellae (3-4 fore, 4-5 hind) ven- trally, terminal phalanx not free with or without minute claw. Adult females rarely with precloacal pore series (0?3 PreclPor), males always with precloacal pores (median ? SD, 7 ? 1.6; range, 6?11) always separated from femoral pore series, 16?25 TotPore (21.5 ? 2.92). Dusky brown ground color dorsally and laterally from head to tail, dark ocular stripe from loreal area to axilla thereafter interrupted and part of zigzag dorsal markings; also narrow dorsolateral dark stripe from rear of eye to axilla, where it also breaks into pieces of the dorsal zigzag marks; small white spots dorsolaterally on trunk but overwhelmed by dark trunk markings; dorsal postsacral mark, anteriormost broad dark brown traverse bar bordered behind by light golden bar then tan and sub- sequently by irregular edged dark brown bars separated by tan interspaces. Major diagnostic features are as follows: bisexual taxon; pigmented caecum and gonadal ducts; in adult males femoral pore series separated from precloacal pore series (TotPore typically ?20-?25), always absent in fe- males; chin scales bordering mental and first infralabial not greatly enlarged; digital lamellae formulae 2-2-2-2 (fore- foot) and 2-2-3-2 or 2-2-3-3 (hindfoot); average adult SVL ~33?35 mm; bold body pattern of contrasting dark brown and dusky brown background and dorsal postsacral dou- ble bar of dark brown and light orange (Figure 15). 38   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY FIGURE 14. Geographic occurrence of Hemiphyllodactylus typus. Not all localities in the same area are plotted. (A) Asia records [Asia and Islands] and (B) Oceania records. Symbols: solid circle = specimen(s) represented by museum vouchers and specific identity con- firmed; open square = locality from published records or museum records but specimen not examined. n um b e r 6 3 1   ?   3 9 Description of lectotype: An adult male: 34.5 mm SVL, broken TailL, 17.5 mm TrunkL, 7.8 mm HeadL, 5.6 mm HeadW, 3.7 mm SnEye, 2.8 mm NarEye, 2.0 mm EyeD, and 1.2 mm SnW. Proportions: 51% TrunkL/SVL, 23% HeadL/SVL, 16% HeadW/SVL, 72% HeadW/HeadL, 40% SnEye/HeadL, 30% NarEye/HeadL, 26% EyeD/ HeadL, 15% SnW/HeadL, 87% EyeD/NarEye, 21% SnW/ HeadW. Scalation: 3 CircNa, 5 SnS, 13 Suplab, 11 Inflab, 12 Chin (anteromedial ones only slightly larger than adja- cent ones), 16 Dorsal, 12 Ventral, 1 CloacS, Subcaud not enlarged, 7 PreclPor, 22 TotPore with no contact between precloacal and femoral, digital formulae 2-2-2-2 (forefoot) and 2-3-3-3 (hindfoot). Pigmented caecum, pigmentation unknown for testis epididymis. Body ground color brown, no lateral light spotting ev- ident, dark dorsolateral stripe from eye to shoulder, lateral stripe from in front of eye to axilla, these stripes broken on trunk and form lateral parts of dorsal chevron or zigzag markings of trunk, dark chevron at tail base. Etymology. Beddome offered no explanation for his choice of aurantiacus as the epithet for his new species. He did mention the orange color of the tail base, and because aurantium is a new Latin noun for the orange (fruit), I suggest his choice derived from the association between the color of the orange and the gecko?s tail. Variation. The means or medians and ranges are detailed in the preceding Description subsection. Males are somewhat smaller than females, but the differ- ence in average size is slight and not statistically signifi- cant. None of the mensural traits shows significant sexual dimorphism and neither do any meristic traits other than precloacal and femoral pores. One large female possesses three secreting precloacal pores; no other females have se- creting pores. All adult males possess both precloacal and femoral pore series. distribution. This gecko is an endemic of the southern tip of India (Tamil Nadu) (Figure 16). It occurs mainly in association with evergreen forest at FIGURE 15. Syntypes of Hemiphyllodactylus aurantiacus Beddome, 1870: (A) lectotype (BMNH 74.4.29.1333) and (B) syntypic series (from top row, left to right, BMNH 74.4.29.1332?1337 and three unnumbered juveniles). (Photographs by G. Zug.) 40   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY mid-elevations in the Nilgiri and Anaimalai Hills (Western Ghats) and Shevaroy and Kolli Hills (Eastern Ghats). Its occurrence in Banglore is edificarian and appears to rep- resent a recent accidental transport. (Distributional sum- mary largely from Bauer and Das, 1999.) Hemiphyllodactylus ganoklonisZug, newspecies Palauanslendergecko holotype. USNM 563682, adult male from Palau Islands, extreme northern tip of Ulebsechel Island, 50 m south of channel between Ngermalk and Ulebsechel, collected by Ronald I. Crombie, 2 August 1998. Paratypes. All subsequent specimens are from Republic of Palau (= Palau Islands); that datum is re- moved from each subsequent locality for brevity. SAM R47715, Babeldaob Island, 4 mi [6.4 km] north on west coast road to Aimeliik, 7?24?N 134?30?E, collected by Christopher C. Austin, 2 June 1996. USNM 495065, Babel daob Island, east of Nekkeng, 0.5 km west of Ta- becheding River (Aimeliik State) 7?27?25?N 134?30?40?E, collected by Gordon H. Rodda, 17 January 1993; USNM 495066, Babeldaob Island, 1 km east of Ngerchaech Mountain, Koksai Radio Station (Ngetpang State) 25 m 7?26?42?N 134?31?48?E, collected by Gordon H. Rodda and Renee J. Rondeau, 18 January 1993; USNM 563663, Babeldaob Island, access road to Palau airport, near turn- off into airport itself (Airai State), collected by Ronald I. Crombie and Christopher C. Austin, 7 July 1996; USNM 563664?665, Babeldaob Island, just north of Ulimang vil- lage on road to Galap village (Ngaraard State), collected by Ronald I. Crombie and Gregory K. Pregill, 14 Janu- ary 1995; USNM 563666, Babeldaob Island, south of Ulimang village (Ngaraard State), collected by Ronald I. Crombie, 6 April 1995; USNM 563667, Ngeanges Is- land, 7?12?26?N 134?22?21?E, collected by Ronald I. Crombie, 9 January 1998; USNM 563668, Ngeaur Is- land, Ngaramasch village, collected by Ronald I. Crom- bie, Gregory K. Pregill, and G. Ken Creighton, 30 January FIGURE 16. Geographic occurrence of Hemiphyllodactylus aurantiacus and H. yunnanensis. Not all localities in the same area are plotted. Symbols: circle, specimens represented by museum vouch- ers and specific identity confirmed; square, localities from published records or museum records but specimen not examined; diamond or square below mid-Indochina boundary (dotted line), speci- mens and records tentatively assigned to H. yunnanensis. n um b e r 6 3 1   ?   4 1 1993; USNM 563669?71, Ngercheu Island group, Carp Island, 7?05?36?N 134?16?44?E, collected by Ronald I. Crombie, 18 July 2001; USNM 563672?673 same as pre- ceding, except 13 August 2001; USNM 563674 same as preceding, except 15 August 2001; USNM 563675 Nger- ekebesang Island, southwest (by road) of Meyungs village, just northeast of turnoff to Echang village, collected by Ronald I. Crombie and Gregory K. Pregill, 9 June 1994; USNM 563676 Ngerektabel Island, approx. 0.5 km (air) northwest of Rael Di, at a sandy beach labelled Oimade- ruul on topographic map, collected by Ronald I. Crombie, 4 August 1998; USNM 563677, Oreor Island, southwest of Ngermid village at Ngerunguikl, Hotel Nikko Palau, collected by Ronald I. Crombie, 22 April 1992; USNM 563678, Ulebsechel Island, Snake Dick Point, midpoint of east coast, 7?19?19?N 134?29?15?E, collected by Ronald I. Crombie and Artemio B. Asis, 28 December 1997; USNM 563679?681, Ulebsechel Island, same data as holotype; USNM 563683, Ulebsechel Island, same data as holotype, except 12 February 2002. Adult females: USNM 495066, 563663?664, -666? 667, -672?673, -675, -679, -681, -684; adult males: SAM R47713, USNM 495065, 563665, -668?671, -674, -676, -678, -682; juveniles: USNM 563677. description. A bisexual taxon of geckos (Gekkoninae) with elongate, slender habitus, slightly com- pressed, elongated appearance accentuated by short limbs and small head (see Figures 3, 11, 17, 18), tail round in cross section and usually shorter than SVL. Adults dimor- phic, females larger than males: 31.1?34.2 mm (mean ? SD, 32.8 ? 1.01; n = 11 females), 28.3?31.6 mm (30.3 mm ? 1.11, n = 12 males) SVL; TailL ~2/3?3/4 of SVL; 16.8? 18.9 mm (17.8 mm ? 0.76), 14.4?17.4 mm (16.0 mm ? 0.95) TrunkL; 6.6?7.5 mm (7.1 mm ? 0.26), 6.3?7.0 mm (6.7 mm ? 0.24) HeadL; 3.8?4.4 mm (4.2 mm ? 0.17), 3.5?4.1 mm (3.8 mm ? 0.20) HeadW; 2.6?3.1 mm (2.8 mm ? 0.16), 2.4?3.0 mm (2.6 mm ? 0.18) SnEye; 2.1?2.3 mm (2.2 mm ? 0.09), 1.8?2.3 mm (2.1 mm ? 0.17) NarEye; 1.7?1.9 mm (1.8 mm ? 0.09), 1.5?1.9 mm (1.7 mm ? 0.15) EyeD; 1.0?1.3 mm (1.1 mm ? 0.11), 1.0?1.1 mm (1.0 mm ? 0.05) SnW. Adult proportions not dimorphic, 49?57% TrunkL/SVL (mean % ? SD, 53.5% ? 2.3), 20?23% HeadL/ SVL (21.8% ? 0.6), 12?15% HeadW/SVL (12.6% ? 0.7), 53?65% HeadW/HeadL (58.0% ? 3.2), 36?45% SnEye/ HeadL (39.2% ? 2.0), 28?34% NarEye/HeadL (31.0% ? 1.4), 23?28% EyeD/HeadL (25.5% ? 1.5), 13?19% SnW/ HeadL (15.6% ? 1.2), 73?95% EyeD/NarEye (82.4% ? 5.6), 23?32% SnW/HeadW (27.0% ? 2.0%). Scalation is predominantly granular from head onto tail, both dorsally and ventrally; ventral trunk scales slightly larger than dorsal ones, 11?18 Dorsal (median ? SD, 15 ? 1.4) and 9?12 Ventral (10 ? 1.0); similarly, subcaudal scales slightly larger than dorsal caudal scales but not plate-like. Cloacal spurs present, modest sized, 1?4 CloacS (2 ? 1.0). Larger scales on lips and snout, rostral largest, rectangular to pentagonal, often slightly concave on dorsomedial edge with slight cleft; 2?4 CircNa (3 ? 0.6), 3?5 SnS (4 ? 0.7); labial scales enlarged from rostral to below eye, becom- ing progressively smaller in subocular rictus, 8?11 Suplab FIGURE 17. Hemiphyllodactylus ganoklonis from Ulebsechel Is- land, Palau (USNM 563680). (Illustration by M. D. Griffin.) 42   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY (8 ? 0.8), 8?10 Inflab (9 ? 0.6); 9?12 Chin (11 ? 0.9), those behind mental slightly or not enlarged; ear opening distinct with no bordering enlarged scales. Each digit with expanded pad, terminal two phalanges free, arising from within pad on second to fifth digits of fore- and hindfoot and each clawed; pads of these digits each with large tri- angular apical lamella bordered proximally by lyre-shaped lamellae (scansors); modal digital formulae 3-4-4-3 (fore- foot) and 3-4-4-4 (hindfoot) for scansors; first digit of fore- and hindfeet compressed, usually 4 rectangular lamellae (3-5 fore, 3-5 hind) ventrally, terminal phalanx not free with or without minute claw. Adult females lack precloacal pore series, males always with precloacal pores (median ? SD, 8 ? 0.9; range, 6?9) always separated from femoral pore series, 16?28 TotPore (22.5 ? 4.01). In life, light dusky tan to reddish beige ground color dorsally and laterally from head to hips, narrow dark brown ocular stripe on posterior edge of loreal area and continuing behind eye either slightly or to edge of temporal region (jowl), a narrow dark lateral stripe on neck broken and reduced to complete stripe, no dark lateral stripe but in some individuals widely spaced series of dark spots or lines; dorsally on head some dark brown bilateral spotting, continuing onto trunk as parasagittal row of small dark marks; dorsolateral series of yellow spots on each side from jowl to anterior arm of postsacral mark; bright postsacral mark with middorsal dark brown spot bordered behind and laterally by light orange to pinkish yellow V, arms of which extending to posterior edge of abdomen. Tail dis- tinctly lighter (yellowish) than trunk, widely spaced narrow orangish rings and middorsal diffuse dark spots in brown interspaces. Venter similar to dorsal ground color, a shade lighter; underside of tail light to bold yellow. In alcohol, pattern same as above, generally ground color darker from light to medium brown; dark stripes and other markings remain distinct, lighter marks lose color becoming white to light tan, including the orange border of postsacral mark; venter strikingly lighter than dorsum, retaining a dusky appearance owing to one or two small brown spots in most ventral scales. Major diagnostic features are as follows: bisexual taxon; pigmented caecum and gonadal ducts; in adult males femoral pore series separate from precloacal pore series, absent in females; chin scales bordering mental and first infralabial not greatly enlarged; digital lamellae for- mulae usually 3-4-4-3 (forefoot) and 3-4-4-4 (hindfoot); average adult SVL ~31?32 mm; mute coloration of light brown background with small scattering of dark brown spots, contrasting with a bright postsacral marking with small dark brown center with light orange V-shaped bor- der and series of widely spaced dorsolateral orange spots. Description of holotype: An adult male (Figure 18), 31.2 mm SVL, 23 mm TailL (regenerated), 16.9 mm TrunkL, 7.0 mm HeadL, 4.0 mm HeadW, 2.8 mm SnEye, 1.9 mm NarEye, 1.7 mm EyeD, and 1.2 mm SnW. Propor- tions: 53% TrunkL/SVL, 22% HeadL/SVL, 12% HeadW/ SVL, 57% HeadW/HeadL, 38% SnEye/HeadL, 30% Nar- Eye/HeadL, 25% EyeD/HeadL, 16% SnW/HeadL, 84% EyeD/NarEye, 28% SnW/HeadW. Scalation: 3 CircNa, 5 SnS, 8 Suplab, 9 Inflab, 12 Chin (anteromedial ones only slightly larger than adjacent ones), 16 Dorsal, 11 Ventral, 4 CloacS, Subcaud not enlarged, 9 PreclPor, 25 TotPore with FIGURE 18. Holotype of Hemiphyllodactylus ganoklonis (USNM 563682) from Ulebsechel Is- land, Palau. (Photograph by G. Zug.) n um b e r 6 3 1   ?   4 3 no contact between precloacal and femoral, digital formulae 3-4-4-4 forefoot and 4-4-5-4 hindfoot. Pigmented caecum, testes lightly pigmented, no pigment on anterior portion of epididymis, heavily pigmented posterior two-thirds. In alcohol, body ground color light brown with me- dium to dark brown markings, dorsolateral light spotting from eye to inguina (4 between eye and axilla, 9 between axilla and inguina (right side), partial dark dorsolateral stripe from eye to neck, no lateral stripes elsewhere, series of widely spaced small dark dashes and spots parasagittally and fewer laterally, postsacral dark brown chevron mid- dorsally at tail base edged laterally by broad white border. Tail background slightly darker than trunk with widely scattered dark flecks. Ventrally dusky cream from chin to vent because many ventral scales with central dark spot. Etymology. The name ganoklonis derives from the Greek adjective and noun, ganos for bright or brightness and klonis for buttock or rump. The bright rump refers to the bright yellow chevron (postsacral) mark at the base of the tail. The name is proposed as an adjective. Variation. The means or medians and ranges are detailed in the preceding Description section. Hemi- phyllodactylus ganoklonis is the smallest Hemiphyllo- dactylus taxon, yet it possesses a slight, but significant, size dimorphism with females averaging 2 mm larger than males. This larger size in females also is reflected in other mensural traits: TrunkL, HeadL, HeadW, NarEye, and EyeD. Body proportions and scalation are not dimorphic. Overall, variation within each trait is low and often the lowest of all regional samples. This low variation re- flects the high quality of preservation of the Palauan sam- ple and is not due to a limited geographic sampling. The Palauan sample derives from seven different islands and multiple sites on a few islands. distribution. Hemiphyllodactylus ganoklonis occurs throughout the major islands of Palau (Figure 19). Hemiphyllodactylus hartertiWerner Bintangslendergecko Lepidodactylus Harterti Werner, 1900:196 [type locality: ?Malakka? (Ma- laysia), restricted to ?Gunong Inas? (Perak, Malaysia); holotype, ZMB 15360]. Gehyra larutensis Boulenger, 1900:188 [type locality: ?Larut Hills, . . . , at 3500 feet altitude? (Malaysia); holotype, BMNH 1901.3.20.2]. comment. The type locality of H. harterti was tentatively restricted by Boulenger in a footnote (1912: 48): ?Dr. Hartert collected on Gunong Inas, the type locality of G. larutensis, and this should, perhaps be substituted for ?Malacca,? over 200 miles distant.? Following my review of E. Hartert?s publications and the locality data from his bird collection, it is evident that Boulenger?s assessment is correct and his restriction should be followed. The type of Gehyra larutensis Boulenger and other specimens from Boulenger?s descriptions of new frogs and reptiles from the Larut Hills, Perak, are listed as in/ from the Selangor Museum. Presumably, that is their ori- gin, but it appears that Boulenger retained them because the catalog number (1901.3.20.2) of G. larutensis indi- cates that it became part of the British Museum collection in 1901. My identification of the holotype relies on the FIGURE 19. Geographic occurrence of Hemiphyllodactylus ganok- lonis in Palau Islands. Circles indicate specimens represented by mu- seum vouchers. (Image modified from map by D. Dalet.) 44   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY BMNH?s labeling of the specimen as type, and in most features my measurements and counts match those of Boulenger. My count of the continuous precloacal?femo- ral pore series, however, is six pores less than Boulenger?s count of 42. I have seen only a single specimen of H. harterti, that is, the holotype of H. larutensis. In spite of the insuffi- ciency of my observations, I recognize this taxon owing to a unique coloration (from published images) and the values of several mensural and meristic traits lying on the edges of the ranges of those traits for geckos of the Titi- wangsa mountain range. description. A bisexual taxon of geckos (Gekkoninae) with robust habitus, slightly compressed trunk and moderately large head (Figures 3, 11, 12), tail round to elliptical in cross section and somewhat shorter than SVL. An adult male (holotype of H. larutensis; Figure 12), 35.3 mm SVL, 31 mm TailL, 17.8 mm TrunkL, 8.3 mm HeadL, 6.3 mm HeadW, 3.5 mm SnEye, 2.6 mm Na- rEye, 2.1 mm EyeD, and 1.8 mm SnW. Proportions: 50% TrunkL/SVL, 24 % HeadL/SVL, 18% HeadW/SVL, 75% HeadW/HeadL, 42% SnEye/HeadL, 31% NarEye/HeadL, 25% EyeD/HeadL, 22% SnW/HeadL, 81% EyeD/NarEye, 29% SnW/HeadW. Scalation: 2 CircNa, 3 SnS, 10 Suplab, 10 Inflab, 9 Chin (anteromedial ones strongly enlarged, ?2? larger than adjacent ones that are also enlarged), 15 Dorsal, 6 Ventral, 2 CloacS, Subcaud not enlarged, 36 TotPore precloacal and femoral pores continuous with demarcation between two series, digital formulae 3-3-3-3 (forefoot) and 3-3-4-3 (hindfoot). Caecum not pigmented, pigmentation unknown for testis epididymis. Aside from coloration, female morphological traits are detailed in the holotype description below. Head to trunk dorsally and laterally a yellowish to dusky tan ground color; tail distinctly brighter (lighter) than body and limbs. Light spots in a dorsolateral series from neck to postsacral mark or dark brown dorsolateral stripe from neck merging into dark median border of post- sacral mark. Tail uniformly colored. Major diagnostic features are as follows: bisexual taxon; caecum and (likely) gonadal ducts not pigmented; precloacal?femoral pore series continuous; chin scales bordering mental and first infralabial distinctly enlarged; digital lamellae formulae 3-3-3-3 (forefoot) and 3-3-4-3 (hindfoot); adult SVL < 40 mm; dorsum of head and trunk either nearly uniform tan or with narrow dark dorsolat- eral stripes and contrasting with lighter tail, outer edge of postsacral mark continuous with caudal color. Description of holotype: An adult female (Figure 12) , 40.9 mm SVL, 39 mm TailL, 21.5 mm TrunkL, 9.7 mm HeadL, 6.3 mm HeadW, 4.0 mm SnEye, 3.2 mm Na- rEye, 2.9 mm EyeD, and 2.0 mm SnW. Proportions: 53% TrunkL/SVL, 24% HeadL/SVL, 15% HeadW/SVL, 65% HeadW/HeadL, 41% SnEye/HeadL, 33% NarEye/HeadL, 30% EyeD/HeadL, 21% SnW/HeadL, 91% EyeD/Nar- Eye, 32% SnW/HeadW. Scalation: 3 CircNa, 2 SnS, 10 Su- plab, 11 Inflab, 6 Chin (anteromedial ones enlarged, twice as large as adjacent ones that are also enlarged), ? Dorsal, ? Ventral, 1 CloacS, Subcaud not enlarged, 0 PreclPor, 0 TotPore, digital formulae not known. No pigmentation on caecum or oviducts. Specimen faded to uniform beige dorsally and later- ally, somewhat lighter ventrally. Evidence of a dark post- sacral mark. Etymology. Werner (1900) noted that a single specimen (holotype) of this gecko was collected in Malakka by Hartert and deposited in the Berlin collec- tion. Presumably, the Hartert referred to by Werner is Er- nest Johann Otto Hartert, an ornithologist who served as the bird curator in L. W. Rothschild?s private museum at Tring, UK, between 1892 and 1929. Prior to his employ- ment at Tring, Hartert visited Asia and elsewhere and col- lected birds, insects, and other animals. He reported his research travels in a popular book, Aus den Wanderjahren eines Naturforschers (Hartert, 1901?1902). distribution. Presently known from Larut Hills and Gunong Inas, Perak (Figure 20); presumably, it occurs throughout the forest of the Bukit Bintang mountains. Hemiphyllodactylus insularisTaylor Philippineslendergecko Hemiphyllodactylus insularis Taylor, 1918:237 [type locality: ?Sumagui, Mindoro? (Philippines); holotype, CM 2052]. description. Adults dimorphic, females larger than males: 29.6?37.3 mm (mean ? SD, 33.9 ? 1.82; n = 15 females), 28.8?34.4 mm (31.3 mm ? 1.86, n = 19 males) SVL; 15.5?19.3 mm (17.8 mm ? 1.08), 14.3?17.7 mm (15.8 mm ? 1.05) TrunkL; 6.6?8.6 mm (7.6 mm ? 0.51), 6.2?8.2 mm (7.2 mm ? 0.51) HeadL; 3.8?5.7 mm (4.8 mm ? 0.48), 4.0?5.8 mm (4.8 mm ? 0.46) HeadW; 2.6?3.7 mm (3.0 mm ? 0.28), 2.1?3.4 mm (2.8 mm ? 0.28) SnEye; 2.0?2.6 mm (2.3 mm ? 0.20), 1.9?2.6 mm (2.2 mm ? 0.17) NarEye; 1.8?2.1 mm (2.0 mm ? 0.11), 1.6?2.1 mm (1.9 mm ? 0.14) EyeD; 1.2?1.4 mm (1.3 mm ? 0.08), 1.0?1.5 mm (1.3 mm ? 0.16) SnW. Adult proportions not dimorphic: 45?58% TrunkL/SVL (mean ? SD, 50.1% ? 2.9), 21?24% HeadL/SVL (22.7% n um b e r 6 3 1   ?   4 5 ? 0.1), 12?18% HeadW/SVL (14.9% ? 0.1), 54?82% HeadW/HeadL (65.7% ? 5.9), 26?44% SnEye/HeadL (39.4% ? 3.1), 27?42% NarEye/HeadL (31.2% ? 2.5), 23?32% EyeD/HeadL (26.1% ? 1.6), 14?21% SnW/ HeadL (17.2% ? 1.8), 74?95% EyeD/NarEye (84.0% ? 5.2), 20?37% SnW/HeadW (26.3% ? 3.2%). Scalation is predominantly granular from head onto tail, both dorsally and ventrally; ventral trunk scales slightly larger than dorsal ones, 13?18 Dorsal (median ? SD, 16 ? 1.4) and 8?14 Ventral (11 ? 1.6); similarly, sub- caudal scales slightly larger than dorsal caudal scales but not plate-like. Cloacal spurs present, modest sized, 0?3 CloacS (1 ? 0.9). Larger scales on lips and snout, rostral largest, rectangular to pentagonal, often slightly concave on dorsomedial edge with slight cleft; 1?4 CircNa (3 ? 0.8), 2?4 SnS (3 ? 0.6); labial scales enlarged from rostral to below eye, becoming progressively smaller in subocular rictus, 9?13 Suplab (10 ? 1.2), 9?11 Inflab (10 ? 0.6); 8?14 Chin (11 ? 1.5), those behind mental slightly or not enlarged; ear opening distinct with no bordering enlarged scales. Each digit with expanded pad, terminal two pha- langes free, arising from within pad on second to fifth dig- its of fore- and hindfoot and each clawed; pads of these digits each with large triangular apical lamella bordered proximally by lyre-shaped lamellae (scansors); modal digital formulae 3-3-3-3 (forefoot) and 3-4-4-4 (hindfoot) for scansors; first digit of fore- and hindfeet compressed, usually 4 or 5 rectangular lamellae (2?5 fore, 3?6 hind) ventrally, terminal phalanx not free with or without min- ute claw. Adult females never with precloacal pores, males always with precloacal pores (median ? SD, 9 ? 1.5; range, 6?13) always separated from femoral pore series, 17?38 TotPore (27 ? 5.35). In alcohol, light to medium brown ground color dor- sally and laterally from head to tail; top of head with scat- tering of small dark brown blotches, lateral dark brown stripe from loreal to shoulder variously developed (barely visible to sharply defined); dorsally on trunk, dark blotches FIGURE 20. Geographic occurrence of Hemiphyllodactylus harterti, H. margarethae, H. titiwan- gsaensis, and bisexual specimens from Borneo. Symbols: circle, H. harterti; diamond, H. titiwang- saensis; solid square, H. margarethae; open square, bisexuals. 46   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY variously arranged from series of parasagittal elongate spots through randomly arranged spots creating irregular narrow transverse bars to nearly absent; dark lateral stripe on trunk typically series of lateral dark spots or blotches; dorsolateral series of light spots from temporal area to in- guina, posteriormost one enlarged and forming anterior end of lateral light arm of postsacral mark; well-defined postsacral mark in all individuals with large pentagonal middorsal dark brown mark bordered behind and on sides by light (whitish) base and arms, which are edged laterally and caudally in dark brown. Tail usually lighter brown than trunk with amorphous dark smudges middorsally. Venter, chin to tail, dusky cream owing to dark spot in most ventral scales. In life, light dusky tan to reddish brown ground color dorsally and laterally from head to hips; pattern of mark- ings as described for preserved individuals. Light dorso- lateral spots and light area of postsacral mark brick red. Major diagnostic features are as follows: bisexual taxon; pigmented caecum and oviducts; no precloacal? femoral pores in females, present in males, precloacal and femoral pore series separated; chin scales bordering mental and first infralabial not greatly enlarged; digital lamellae formulae 3-3-3-3 (forefoot) and 3-4-4-4 (hindfoot); aver- age adult SVL ~34, 31 mm (females, males); series of red spots dorsolaterally on trunk and bright postsacral bar of red and dark brown. Description of holotype: An adult male (Figure 21), 30.2 mm SVL, 14.7 mm TrunkL, 6.9 mm HeadL, 2.6 mm HeadW, 4.0 mm SnEye, 1.9 mm NarEye, 1.8 mm EyeD, and 1.2 mm SnW. Proportions: 49% TrunkL/ SVL, 23% HeadL/SVL, 13% HeadW/SVL, 58% HeadW/ HeadL, 38% SnEye/HeadL, 28% NarEye/HeadL, 26% EyeD/HeadL, 17% SnW/HeadL, 95% EyeD/NarEye, 30% SnW/HeadW. Scalation: 3 CircNa, 3 SnS, 10 Suplab, 11 In- flab, 11 Chin (anteromedial ones only slightly larger than adjacent ones), 15 Dorsal, 11 Ventral, 3 CloacS, Subcaud not enlarged, 9 PreclPor, 27 TotPore precloacal and femo- ral series separated, digital formulae 3-3-3-3 (forefoot) and 3-4-4-4 (hindfoot). Pigmented caecum, pigmentation un- known for testis epididymis. Faded, body ground color brown. Etymology. Taylor (1918) did not explain his choice of the epithet insularis, presumably because he assumed the name was self-explanatory; insularis is a Latin adjective for of islands. FIGURE 21. Holotype of Hemiphyllodactylus insularis Taylor, 1918 (CM 2052): (A) dorsal view of whole body, (B) ventral view of head, and (C) ventral view of posterior half of trunk. (Photo- graphs by M. McNaugher.) n um b e r 6 3 1   ?   4 7 Variation. The means or medians and ranges are detailed in the preceding Description section. Males are smaller, statistically significantly so, than females, but the difference in average size is slight (~2.5 mm). None of the meristic traits shows significant dimorphism among adults, other than presence of precloacal and femoral pores in males and their absence in females. Variation of most scalation traits is modest to low with the values of most traits equaling the median. Chin scales are usually small, although a few individuals have a modest enlarge- ment of those touching the mental and first supralabial. There are six digital formulae each for forefoot and hind- foot. Forefoot formulae range from 3-3-3-3 (66%) to 3-4- 4-4 with only 3-3-4-3 (17%) also occurring in more than two individuals. Hindfoot formulae range from 3-3-4-3 to 4-5-5-4; 3-4-4-4 is the most frequent (49%), followed by 3-4-4-3 (17%) and 4-4-5-4 (14%). distribution. Hemiphyllodactylus insularis occurs throughout the Philippine Islands (Figure 22) from Mindoro to Mindanao and westward on both the Palawan and Sulu Archipelago arcs. The presence on both these lat- ter island groups recommends a reexamination of the bi- sexual Bornean Hemiphyllodactylus. Hemiphyllodactylus margarethaeBrongersma Sumatranslendergecko Hemiphyllodactylus margarethae Brongersma, 1931:11 [type locality: ?Fort de Kock, Sumatra? (Bukittinggi, Sumatera Barat); holotype, ZMA 11095]. comment. Brongersma (1932:218 [footnote]) noted that while the H. margarethae description was in press and after he had examined additional Hemiphyl- lodactylus specimens, he attempted to suppress the new name in page proofs, but his recommended changes were not made. description. A bisexual taxon of geckos (Gekkoninae) with robust habitus, slightly compressed trunk and moderately large head (see Figures 3, 23), tail round in cross section and subequal to SVL. Adults not di- morphic: 36.0?46.9 mm SVL (mean ? SD, 40.8 mm ? 3.6], n = 8), 14.5?25.4 mm TrunkL (20.6 mm ? 3.3), 8.2?10.4 mm HeadL (9.6 mm ? 0.87), 5.8?8.2 mm HeadW (6.8 mm ? 0.86), 3.3?4.7 mm SnEye (4.1 mm ? 0.56), 2.7?3.6 mm NarEye (3.2 mm ? 0.40), 1.6?2.5 mm EyeD (2.1 mm ? 0.35), 1.4?1.9 mm SnW (1.7 mm ? 0.23). Adult propor- tions: 40?54% TrunkL/SVL (mean ? SD, 50.3% ? 4.8), 21?26% HeadL/SVL (23.5% ? 1.6), 15?19% HeadW/ SVL (16.7% ? 1.3), 66?79% HeadW/HeadL (71.2% ? 4.4), 39?46% SnEye/HeadL (42.6% ? 2.4), 29?35% Nar- Eye/HeadL (33.0% ? 2.0), 16?25% EyeD/HeadL (22.0% ? 3.6), 15?19% SnW/HeadL (17.3% ? 1.4), 47?81% EyeD/NarEye (67.0% ? 12.6), 22?28% SnW/HeadW (24.3% ? 1.9%). Scalation is predominantly granular from head onto tail, both dorsally and ventrally; ventral trunk scales slightly larger than dorsal ones, 11?17 Dorsal (median ? SD, 12.5 ? 2.3) and 6?12 Ventral (7.5 ? 2.3); similarly, subcaudal scales slightly larger than dorsal caudal scales but not plate- like. Cloacal spurs present, modest sized, 1?2 CloacS (2 ? 0.5). Larger scales on lips and snout, rostral largest, rectan- gular to pentagonal, often slightly concave on dorsomedial FIGURE 22. Geographic occurrence of Hemiphyllodactylus insu- laris in the Philippine Islands. Not all localities in the same area are plotted. Circles indicate specimens represented by museum vouchers and whose specific identity is confirmed. 48   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY edge with slight cleft; 2?3 CircNa (3 ? 0.4), 2?4 SnS (3 ? 0.6); labial scales enlarged from rostral to below eye, be- coming progressively smaller in subocular rictus, 10?13 Suplab (11 ? 1.2), 9?12 Inflab (10 ? 0.9); 6?11 Chin (7.5 ? 2.0), those behind mental distinctly enlarged; ear opening distinct with no bordering enlarged scales. Each digit with expanded pad, terminal two phalanges free, arising from within pad on second to fifth digits of fore- and hindfoot and each clawed; pads of these digits each with large tri- angular apical lamella bordered proximally by lyre-shaped lamellae (scansors); modal digital formulae likely 4-4-4-4 (forefoot) and 4-5-5-5 (hindfoot) for scansors; first digit of fore- and hindfeet compressed, usually 5 rectangular la- mellae (4?8 fore, 4?7 hind) ventrally, terminal phalanx not free with or without minute claw. Adult females often with precloacal pores series (0?12 PreclPor), males (only one adult in sample) always with precloacal pores (median 11) always separated from femoral pore series, 0?29 (female) 26 (male) TotPore. Coloration, no images of living specimens and pre- served specimens pattern indistinct owing to fading. Major diagnostic features are as follows: bisexual taxon; unpigmented caecum, oviducts pigmented or not; if present, femoral pore series separate from precloacal pore series; chin scales bordering mental and first infralabial distinctly enlarged; digital lamellae formulae 4-4-4-4 (fore- foot) and 4-5-5-5 (hindfoot); average adult SVL ~41 mm. Description of holotype: An adult male (Figure 23), 38.8 mm SVL, 40 TailL, 18.7 mm TrunkL, 9.9 mm HeadL, 7.5 mm HeadW, 4.3 mm SnEye, 3.4 mm Nar- Eye, 1.6 mm EyeD, and 1.9 mm SnW. Proportions: 48% TrunkL/SVL, 25% HeadL/SVL, 19% HeadW/SVL, 76% HeadW/HeadL, 43% SnEye/HeadL, 34% NarEye/HeadL, 16% EyeD/HeadL, 19% SnW/HeadL, 47% EyeD/NarEye, 25% SnW/HeadW. Scalation: 3 CircNa, 3 SnS, 10 Suplab, 11 Inflab, 6 Chin (anteromedial ones distinctly enlarged, see Figure 7C), 13 Dorsal, 7 Ventral, 2 CloacS, Subcaud not enlarged, 11 PreclPor, 26 TotPore, separate precloacal and femoral series, digital formulae 4-5-5-5 (forefoot) and 5-5-6-5 (hindfoot). Caecum and testis epididymis presum- ably unpigmented. Body ground color faded to pinkish beige, small paired dark blotches dorsally on trunk; paratype (ZMA 11096) has a hint of a postsacral mark. Etymology. The origin of the epithet marga- rethae is unknown. Variation. The means or medians and ranges are detailed in the preceding Description section. Sample is small and specimens old and generally poorly preserved; hence comments on variation are inappropriate. distribution. With a single exception, the H. margarethae specimens derive from the central moun- tain ranges of central and northern Sumatra (Figure 20), extending from Bukittinggi in the south to Takengon in the north. The Nias island locality is the exception and requires confirmation. Hemiphyllodactylus titiwangsaensis Zug,newspecies Titiwangsanslendergecko holotype. ZRC 2.4782, adult male from Malaysia, Pahang Province, Cameron Highlands, Gunong Brinchang (= Berincang) summit area, collected by H. H. Tan and others, 25 June 2000. Paratypes. All subsequent specimens are from Malaysia, Pahang Province; that datum is removed from each subsequent locality for brevity. ZFMK 32284?286, Cameron Highlands, Tana Ratah (=Tanah Rata), col- lected by Dietmar Kiehlmann, July 1980; ZRC 2.4780? 781, 2.4783?785, collecting data same as holotype; ZRC 2.4832, Cameron Highlands, Tanah Rata, Bala?s Lodge, collected by H. H. Tan, 4 May 2000; ZRC 2.5165, Cam- eron Highlands, Parit Falls, T. M. Leong, and L. J. Lim, 30 July 2001; ZRC 2.5419, Cameron Highlands, Parit Falls, collected by B. L. Lim and K. K. P. Lim, 10 May 2002; FIGURE 23. Types of Hemiphyllodactylus margarethae Brong- ersma, 1931: left, ZMA 11095 holotype, male; right, 11096 para- type, female. (Photograph by G. Zug.) n um b e r 6 3 1   ?   4 9 ZRC 2.5942, Cameron Highlands, Telom Valley, Kuala Terla 4000?4500?, collector unknown, March 1935; ZRC 2.5943, Cameron Highlands, Telom Valley, Gunong Siku at ~4500?, collector unknown, March 1935. Adult fe- males: ZFMK 32284, 32286; ZRC 2.4780?4781, 2.4785, 2.4832, 2.5165, 2.5943; adult males: ZFMK 32385; ZRC 2.4783?4784, 2.5419, 2.5942. description. Bisexual taxon of geckos (Gek- koninae) with robust habitus, slightly compressed trunk and moderately large head (see Figures 3, 11, 24), tail round to elliptical in cross section and somewhat shorter than SVL. Adults not dimorphic: 36.5?62.1 mm (mean ? SD, 49.2 mm ? 6.34; n = 15) SVL; 18.1?32.5 mm (23.3 mm ? 3.69) TrunkL; 8.9?13.8 mm (12.0 mm ? 1.18) HeadL; 6.1?10.2 mm (8.2 mm ? 0.98) HeadW; 3.4?5.8 mm (4.9 mm ? 0.60) SnEye; 2.6?4.4 mm (3.6 mm ? 0.42) NarEye; 2.4?3.6 mm (3.0 mm ? 0.31) EyeD; 1.4?2.6 mm (2.0 mm ? 0.32) SnW. Adult proportions: 42?50% TrunkL/SVL (mean ? SD, 46.4% ? 3.4), 23?27% HeadL/ SVL (24.6% ? 1.4), 16?19% HeadW/SVL (16.9% ? 1.0), 64?73% HeadW/HeadL (68.5% ? 3.5), 37?43% SnEye/ HeadL (40.7% ? 2.1), 26?32% NarEye/HeadL (29.4% ? 2.1), 21?30% EyeD/HeadL (25.6% ? 2.5), 14?21% SnW/ HeadL (16.5% ? 2.5), 68?94% EyeD/NarEye (85.4% ? 9.6), 21?31% SnW/HeadW (23.8% ? 3.2%). Scalation is predominantly granular from head onto tail, both dorsally and ventrally; ventral trunk scales slightly larger than dorsal caudal scales, 14?19 Dorsal (median ? SD, 16 ? 1.9) and 7?9 Ventral (7 ? 1.0); similarly, subcau- dal scales slightly larger than dorsal ones but not plate-like. Cloacal spurs present, modest sized, 1?4 CloacS (3 ? 1.0). Larger scales on lips and snout, rostral largest, rectangular to pentagonal, often slightly concave on dorsomedial edge with slight cleft; 3 CircNa (3 ? 0.0), 1?3 SnS (3 ? 0.8); la- bial scales enlarged from rostral to below eye, becoming progressively smaller in subocular rictus, 9?11 Suplab (10 ? 0.7), 8?10 Inflab (9 ? 0.7); 8?9 Chin (9 ? 0.5), those be- hind mental distinctly enlarged; ear opening distinct with no bordering enlarged scales. Each digit with expanded pad, terminal two phalanges free, arising from within pad on second to fifth digits of fore- and hindfoot and each clawed; pads of these digits each with large triangular api- cal lamella bordered proximally by lyre-shaped lamellae (scansors); modal digital formulae 3-4-4-4 (forefoot) and 4-5-5-5 (hindfoot) for scansors; first digit of fore- and hind- feet compressed, usually 5 or 7 rectangular lamellae (4?6 fore, 5?8 hind) ventrally, terminal phalanx not free with or without minute claw. Adult females never with precloacal pores; males always with continuous precloacal?femoral pore series 17?39 TotPore (median ? SD, 21 ? 7.95). In life, dorsal and lateral ground color ranges from light grayish tan to medium brown, head to tail occasion- ally distinctly lighter than neck and trunk (Figure 11). This lightness is emphasized by absence or diffuseness of dark markings on head. Neck and trunk bear numerous transverse dark brown irregularly shaped bars, lighter in- terspaces typically narrower than dark bars. Bars extend onto sides; dorsolaterally in shoulder area bars are darker, creating an impression of dark dorsolateral stripe. Dark lateral stripe from loreal to neck, occasionally to midneck. Dark stripe bordered above by cream to beige stripe from canthus rostralis to shoulder, often continuing as series of spots or dashes on trunk and at inguina becoming narrow arm of postsacral mark; center dark spot of mark absent to small. Tail usually lighter than trunk and distinctly banded in light and dark, relative size of which very variable. (Col- oration from images by H. Ota and Chan-ard et al., 1999.) Coloration in alcohol is muted, although dark and light pattern usually persists. Ventrally from chin onto tail, uniform light cream in most individuals, brown in a few. Females seem to be more boldly patterned than males. Major diagnostic features are as follows: bisexual taxon; caecum and gonadal ducts not pigmented; pre- cloacal?femoral pore series continuous in males (TotPore 17?39), absent in females; chin scales bordering mental and first infralabial distinctly enlarged; digital lamellae formulae usually 3-4-4-4 (forefoot) and 4-4-5-5 or 4-5-5-5 (hindfoot); average adult SVL ~49 mm; dorsal and lateral trunk pattern of dark brown irregular transverse bands, muted postsacral bar of narrow white arms onto hips. Description of holotype: An adult male (Figure 24), 56.9 mm SVL, 48 mm TailL (regenerated), 24.1 mm TrunkL, 13.2 mm HeadL, 9.6 mm HeadW, 5.4 mm SnEye, 3.9 mm NarEye, 3.0 mm EyeD, and 2.4 mm SnW. Propor- tions: 42% TrunkL/SVL, 23% HeadL/SVL, 17% HeadW/ SVL, 73% HeadW/HeadL, 41% SnEye/HeadL, 28% Na- rEye/HeadL, 23% EyeD/HeadL, 18% SnW/HeadL, 81% EyeD/NarEye, 25% SnW/HeadW. Scalation: 3 CircNa, 3 SnS, 10 Suplab, 9 Inflab, 8 Chin (anteromedial ones en- larged), 16 Dorsal, 7 Ventral, 3 CloacS, Subcaud not en- larged, precloacal and pore series continuous, 30 TotPore, digital formulae 4-4-5-4 (forefoot) and 4-5-5-5 (hindfoot). No pigmentation on caecum or oviducts. Specimen brown dorsally and laterally with scattered indistinct dark brown markings, somewhat lighter ven- trally. Postsacral mark indistinct, small median dark spot on first tail segment, anterior arms muted. In life, the ven- tral surface of the tails (type series) were orangish pink. Etymology. These geckos occur in the south central region of the Banjaran Titiwangsa; hence the 50   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY taxon is a resident (likely endemic) of Titiwangsa and so named. Variation. The means or medians and ranges are detailed in the preceding Description section. Adults are not dimorphic, and both sexes have broad ranges of adult sizes, females 42.2?62.1 mm SVL and males 36.5?56.9 mm. This broad range yields a modest increase in variation among the mensural traits (CV = 8?17%). Larger (?46 mm SVL) adults are distinctly robust-bodied geckos. Scalation has low variation with the exception of pha- langeal formulae. Seven forefoot formulae range from 2-3- 3-3 to 4-5-5-5; however, 3-4-4-4 is the mode and median with a uniform and sharp drop in number of individu- als on each side of the mode. Hindfoot formulae (8) are dominated by 4-4-5-5 (n = 3) and 4-5-5-5 (7); the other six formulae observed have low (n = 1 each) representation; 3-3-3-3 is the lowest. Coloration has two patterns. The transverse bar pat- tern (somewhat Lepidodactylus lugubris-like) described above occurs in the majority of individuals, and a few individuals show gradation from this pattern into a ragged spotted one. distribution. All specimens derive from lo- calities within Banjaran Titiwangsa (Figure 20) and par- ticularly from Fraser Hill and Cameron Highland areas. At both sites, H. titiwangsaensis occurs in forest and on and around buildings. I assume that these geckos occur more broadly in moist evergreen forest of the Titiwangsa mountain range and that the present vouchers represent the most accessible areas. Hemiphyllodactylus yunnanensis(Boulenger) Asianslendergecko Gehyra yunnanensis Boulenger, 1903:429 [type locality: ?Yunnan Fu? (= Kumning, Yunnan Province, China); lectotype, BMNH 1904.1.26.1]. Hemiphyllodactylus typus chapaensis Bourret, 1937:60 [type locality: ?Chapa? (Sa P? [also Lao Cai], Vietnam); holotype, MNHN 1948.43]. Hemiphyllodactylus yunnanensis longlingensis Zhou and Yang in Zhou et al., 1981:203 [type locality: Longling Junior High School in Longling County, Yunnan Province (alt. 1530 m) (China) (original in Chinese)]. FIGURE 24. Types of Hemiphyllodactylus titiwangsaensis: left to right, ZRC 2.4780?2.4785; 2.4782 is the holotype. (Photograph by G. Zug.) n um b e r 6 3 1   ?   5 1 Hemiphyllodactylus yunnanensis jinpingensis Zhou and Yang in Zhou et al., 1981:204 [type locality: Jinpling Junior High School in Jinping County, Yunnan Province (alt. 1260 m) (China) (original in Chinese)]. Hemiphyllodactylus yunnanensis dushanensis Zhou and Yang in Zhou et al., 1981:206 [type locality: Dushan Junior High School in Dushan County, Guizhou Province (alt. 970 m) (China) (original in Chinese)]. comments. Readers are reminded that H. yunnanensis is considered here to include the highland populations of Hemiphyllodactylus across southern China and adjacent northern Southeast Asia from Myanmar to Vietnam. The taxonomic status of the ?lowland? popu- lations of Southeast Asia and Hong Kong remains unre- solved owing to the sparsity of vouchers in numbers and geography. Boulenger (1903) identified two syntypes ?male and young.? Both specimens (BMNH 1904.1.26.1?2; Figure 25) are extant. I designate the adult male (BMNH 1904.1.26.1) as the lectotype of Gehyra yunnanensis Boulenger. Bourret said in the type description: ?J?ai pris ? Chapa une femelle ? queue reconstitu?e (S130) . . .? Brygoo (1990:44) and I interpret this statement as a description based on a single female specimen. This interpretation con- flicts with Guibe?s (1954) type catalog listing of two speci- mens; the sex of neither is identified by Guibe, although one is noted to be damaged and 56 mm long. Brygoo noted that of the two specimens labeled syntypes, both possess Bourret?s registration numbers and one (S130) is unambiguously the holotype of H. typus chapaensis Bour- ret. Also, the holotype is 43 mm SVL (Brygoo 1990:44) and 42.7 mm (my measurement), and 33 mm (Bourret, 1937:60). This disparity, yet similarity, suggests that Bour- ret accidentally entered 33 instead of 43. description. A bisexual taxon of geckos (Gekkoninae) with robust habitus, slightly compressed trunk and moderately large head (Figures 3, 11, 25), tail round in cross section and typically shorter than SVL. Adults dimorphic, females larger than males: 31.9? 49.3 mm (mean ? SD, 40.7 mm ? 4.44; n = 33), 25.5? 46.4 mm (37.9 mm ? 4.58, n = 28) SVL; 15.4?26.5 mm (19.5 mm ? 2.93), 12.8?22.5 mm (17.9 mm ? 2.30) TrunkL; 7.6?11.5 mm (9.5 mm ? 0.98), 6.7?10.3 mm (8.8 mm ? 0.89) HeadL; 5.4?8.4 mm, (6.8 mm ? 0.95), 4.8? 7.4 mm (6.3 mm ? 0.76) HeadW; 3.0?5.2 mm (3.9 mm ? 0.55), 2.4?4.7 mm (3.7 mm ? 0.50) SnEye; 2.1?4.0 mm (3.0 mm ? 0.44), 1.9?3.4 mm (2.8 mm ? 0.37) NarEye; 1.9?3.2 mm (2.4 mm ? 0.31), 1.6?3.0 mm (2.2 mm ? 0.27) EyeD; 1.0?2.2 mm (1.7 mm ? 0.25), 0.9?2.0 mm (1.5 mm ? 0.25) SnW. Adult proportions not dimorphic: 40?55% TrunkL/SVL (mean ? SD, 47.4% ? 3.1), 21?26% HeadL/ SVL (23.3% ? 1.0), 14?22% HeadW/SVL (16.7% ? 1.8), 59?83% HeadW/HeadL (71.9% ? 6.5), 34?46% SnEye/ HeadL (41.4% ? 2.5), 26?35% NarEye/HeadL (31.4% ? 2.1), 22?29% EyeD/HeadL (25.1% ? 1.7), 11?22% SnW/ HeadL (17.3% ? 2.0), 63?100% EyeD/NarEye (80.3% ? 6.8), 15?36% SnW/HeadW (24.3 ? 3.6%). Scalation is predominantly granular from head onto tail, both dorsally and ventrally; ventral trunk scales slightly larger than dorsal ones, 9?18 Dorsal (median ? SD, 13 ? 1.8) and 6?12 Ventral (8 ? 1.1); similarly, sub- caudal scales slightly larger than dorsal caudal scales but not plate-like. Cloacal spurs usually present, mod- est sized, 0?2 CloacS (1 ? 0.3). Larger scales on lips and snout, rostral largest, rectangular to pentagonal, often slightly concave on dorsomedial edge with slight cleft; 2?4 CircNa (3 ? 0.2), 2?5 SnS (3 ? 0.7); labial scales enlarged from rostral to below eye, becoming progres- sively smaller in subocular rictus, 8?13 Suplab (10 ? 1.0), 8?12 Inflab (10 ? 1.1); 6?11 Chin (8 ? 1.1), those be- hind mental moderately to distinctly enlarged; ear open- ing distinct with no bordering enlarged scales. Each digit with expanded pad, terminal two phalanges free, arising from within pad on second to fifth digits of fore- and hindfoot and each clawed; pads of these digits each with large triangular apical lamella bordered proximally by FIGURE 25. Lectotype of Gehyra yunnanensis Boulenger, 1903 (BMNH 1904.1.26.1): (A) dorsal view of whole body, (B) ventral view of throat and chin, and (C) ventral view of pelvic area. (Photo- graphs by G. Zug.) 52   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY lyre-shaped lamellae (scansors); modal digital formulae 3-3-3-3 (forefoot) and 3-4-4-4 (hindfoot) for scansors; first digit of fore- and hindfeet compressed, usually 4 and 5 rectangular lamellae (4?6 fore, 4?7 hind) ventrally, terminal phalanx not free with or without minute claw. Adult females rarely with precloacal?femoral pore series (7?9 PreclPor, n = 2; 0?19 TotPore), males always with continuous precloacal?femoral pore series 11?25 TotPore (median ? SD, 20.0 ? 3.26). In life, light grayish brown to medium reddish brown ground color dorsally and laterally from head to tail base; top of head with scattering of small dark brown marks, lateral dark brown stripe from loreal to shoulder bordered above by white to tan stripe to end of head; dorsally on trunk, narrow dark brown transverse lines to squiggles, dorsolateral medium-sized light spots from neck to above hindlimb and laterally dark lateral stripe occasionally across neck to anterior trunk and thereaf- ter fragmented to dark dashes or diffuse brown marks; typically, area between dorsolateral light spots and dark lateral stripe lighter than dorsal ground color; postsacral mark small to large blotch bordered behind by rectangu- lar white spot, no anterior extensions; tail ground color lighter than trunk with series of transverse blotches of narrow dark brown bordered behind by broader area of light tan. Venter dusky, tail base pinkish to light orange blush. In alcohol, pattern as above and fading toward a uni- form light to medium brown with brown marks. Major diagnostic features are as follows: bisexual taxon; caecum and gonadal ducts not pigmented; pre- cloacal?femoral pore series continuous in males (TotPore 11?25), usually absent in females; chin scales bordering mental and first infralabial distinctly enlarged; digital la- mellae formulae usually 3-3-3-3 (forefoot) and 3-4-4-4 (hindfoot); average adult female SVL ~41 mm, males ~39 mm; dorsal trunk pattern of narrow dark brown irregular transverse bands bordered dorsolaterally by longitudinal series of light spots, postsacral bar of dark and light with no anterior extensions dorsolaterally. Description of lectotype: An adult male, 40.3 mm SVL, ~41 regenerated TailL, 20.1 mm TrunkL, 9.9 mm HeadL, 7.8 mm HeadW, ~4.3 mm SnEye, ~3.3 mm Nar- Eye, ~2.6 mm EyeD, and ~1.8 mm SnW. Proportions: 50% TrunkL/SVL, 25% HeadL/SVL, 19% HeadW/SVL, 79% HeadW/HeadL, ~44% SnEye/HeadL, ~34% NarEye/ HeadL, ~26% EyeD/HeadL, ~18% SnW/HeadL, ~79% EyeD/NarEye, ~26% SnW/HeadW. Scalation: 3 CircNa, 3+ SnS, 9 Suplab, 9 Inflab, ?10 Chin (anteromedial ones distinctly larger than adjacent ones), ~16 Dorsal, ~8 Ventral, 2 CloacS, Subcaud not enlarged, precloacal and femoral pore series continuous 36 TotPore, digital formu- lae (estimate) 3-3-3-3 (forefoot) and 3-4-4-4 (hindfoot). Pigmentation of caecum and testis epididymis unknown, likely no pigmentation. Body ground color grayish brown above and below, scattered dark spots and small dark spots middorsally, no lateral spotting on trunk; dark dorsolateral stripe from eye to shoulder, lateral stripe from in front of eye to end of head. Caecum not visible through body wall; not dissected so unable to confirm gonadal pigmentation. Etymology. The name yunnanensis identifies this species as the gecko from Yunnan, the type locality of Boulenger?s new species. Variation. The means or medians and ranges are detailed in the preceding Description section. Adults are dimorphic in size, and both sexes have broad ranges of adult sizes. The variation in mensural and meristic traits is examined in the bisexual portion of the Character Analy- sis section. distribution. Highlands of southwestern China and adjacent uplands from the western edge of the Shan Plateau in Myanmar, across northern Thailand, Laos, and Vietnam (Figure 16). The southern limits of the distribution are ill-defined owing to limited sampling of these geckos through much of Asia. KEY TO THE SPECIES OF HEMIPHYLLODACTYLUS 1. Chin scales bordering mental scale posteriorly distinctly enlarged [Figure 7C,D] appear as a pair of scales labeled post- mentals in other geckos; caecum and gonadal peritoneum white [Figure 2C] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 1?. Chin scales bordering mental scale posteriorly slightly or not enlarged [Figure 7A,B], their size nearly same as more medial chin scales; caecum and gonadal-duct peritoneum pig mentation usually black [Figure 2B] . . . . . . . . . . . . . . . . . . . . . . 2 2. Adult females with actively secreting precloacal and femoral pores; unisexual species, all individuals are females; adult size often >36 mm SVL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. typus 2?. Adult females with no or fewer than five secreting precloacal pores; populations of fe males and males; adult size seldom >38 mm SVL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 n um b e r 6 3 1   ?   5 3 3. Usually two U-shaped digital lamellae under fourth digit of forefoot; dorsal trunk pattern bold, transverse dark blotches, longitudinal series of white dorsolateral spots and post sacral mark of dark brown and orange [Figure 11A] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. aurantiacus 3?. Three or four U-shaped digital lamellae under fourth digit of forefoot; dorsal trunk pattern muted, faded and small dark blotches or widely separated dark spots [Figure 11F]; post sacral mark with U- or V-shaped outer edge of yellow or red; dorsolateral spots yellow or red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Total number precloacal and femoral pores (TotPore) usually <24 (16?28) in males; fore foot digital lamellar formula usu- ally 3-4-4-3; postsacral mark outer edge yellow to pinkish yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. ganoklonis 4?. Total number precloacal and femoral pores (TotPore) usually >24 (17?38) in males; fore foot digital lamellar formula usu- ally 3-3-3-3; postsacral mark outer edge red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. insularis 5. Precloacal and femoral pore series separate; females commonly with precloacal pores; forefoot digital lamellar formula usually 4-4-4-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. margarethae 5?. Precloacal and femoral pore series continuous; females usually lack precloacal pores.: fore foot digital lamellar formula 3-3-3-3 or 3-4-4-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Adults large, usually >45 mm SVL; precloacal?femoral pore series usually >22 (17?39) pores; hindfoot digital lamellar formula usually 4-4-5-5 or 4-5-5-5; postsacral mark with anterior arms . . . . . . . . . . . . . . . . . . . . . . H. titiwangsaensis 6?. Adults moderate size, usually <42 mm SVL; hindfoot digital lamellar formula usually 3-3-3-3 or 3-4-4-4, occasionally higher; postsacral mark without anterior arms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Trunk usually with distinct dark dorsolateral stripe; precloacal and femoral pore series continuous with >30 pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. harterti 7?. Trunk usually without dark dorsolateral stripe; precloacal and femoral pore series continu ous with <26 pores . . . . . . 8 8. Precloacal and femoral pore series usually >18 pores; hindfoot digital lamellar formula usually 3-4-4-4; postsacral mark of anterior dark blotch and posterior larger light bar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. yunnanensis 8?. Precloacal and femoral pore series usually <18 pores; hindfoot digital lamellar formula usually 4-5-5-4; postsacral mark absent or muted dark transverse bar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hong Kong population Acknowledgments I offer my appreciation to numerous colleagues who have ?kept the faith? in my dogged effort to study the relationships of Oceania lizards. Thank you all. There are more taxonomic studies in progress. I express my highest appreciation to another group of colleagues and collections management staffs worldwide. They have been generous in their assistance and courtesies during my visits to their collections and have permitted me to borrow specimens and retain some for an inordinately long time. I thank D. Kizirian, R. Pasco- cello, R. Zweifel (AMNH); R. Sadlier, A. Greer (AMS); C. Mc Carthy (BNMH); C. Kishinami, K. Imada (BPBM); J. Vindum, R. Lucas, H. Brignall (CAS); M. McNaugher and S. Rogers (CM); A. Resetar, C. Redhed, H. Voris (FNMH); G. Leng let, G. Coulon, M. Lang (IRSN); T. Hikida, M. Matsui, H. Ota (KUZ); J. Rosado (MCZ); I. Ineich (MNHN); E. Kramer (NMB); F. Tiede mann, R. Gemel (NMW); J. Covacevich, P. Couper (QM); M. Hoogmoed, J. Arntzen (RMNH); M. Hutchinson (SAM); G. Pregill (SDMNH); A. Schulter, K. Kramer (SMF); T. Chan-ard, S. Mekchai (THNMH); D. Auth, K. Krysko, W. King (UF); R. How, L. Smith (WAM); private collection of William Beckon (WmBeckon); L. van Tuijl (ZMA); R. G?nther, M.-O. R?del (ZMB); W. B?hme (ZMFK); K. Lim (ZRC); U. Gruber (ZSM). Additionally, I thank the collection staff of the National Museum of Natural History, Smithsonian Institution, who regu- larly support my specimen-based research: F. Blasdell, R. Crombie, S. Gotte, T. Hartsell, K. Tighe, and R. Wilson. Numerous individuals have aided my Hemiphyllodactylus research. R. Wil- son did the initial georeferencing of Hemiphyllodactylus specimens and type lo- calities. B. Trimmer assisted with an early phase of data verification. T. Schwaner, performed an electrophoretic analysis of Thai, Philippine, and Hawaiian Hemi- phyllodactylus. A. Bauer, H. Ota, and I. Das have kept my interest in mind and regularly apprised me of newly collected specimens and observations. R. Crom- bie made a special effort to obtain specimens from Palau and Hawai?i and pro- vided field notes and photographs of Palauan specimens. P.-P. van Dijk provided notes and images of northern Thailand H. yunnanensis. H. Ota gave sketches, notes, and images of Asian Hemiphyllodactylus. I. Das provided Hemiphyllo- dactylus images from India and Borneo, and J. R. H. Gibbons gave me images 56   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY of Fijian H. typus. M.-O. R?del generously took data and photographs of the type of Lepidodactylus Harterti Wer- ner. L. Grismer challenged me to examine more closely the nomenclatural status of harterti and larutensis. S. Rodgers arranged for the photography of the type of H. insularis. Jennifer Kilby [n? Westhoff] of Luray Zoo and Molly Grif- fin of Redlands lent their artistic skills. T. Ulber translated Bleeker?s type description of H. typus, and a volunteer from Smithsonian Volunteer Services translated the Hemi- phyllodactylus section from V. V. Bobrov and D. V. Se- menov?s (2008) Lizard Fauna of Vietnam. My wife, Pat, contributed to this research by her regular assistance in data gathering, entry, and organization and by her review of an early draft of the manuscript. Other colleagues? A. Bauer, M. Cota, R. Crombie, R. Fisher, F. Kraus, H. Ota, J. Vindum?read and commented on drafts of the manuscript; their comments significantly improved the quality of the current monograph. I am responsible for remaining errors and misinterpretations. I thank all of the above for their time and effort on my behalf. Appendix 1: Character Definitions M any of the characters examined and recorded in this study are used broadly in other systematics studies of geckos. I use the abbreviations proposed by me previously (Zug, 1998) for con- ciseness and for permitting quick identification of the charac- ters. Most characters are defined in Zug et al. (2003). Any not defined there or that are defined differently for Hemiphyllodactylus are presented in Table A1.1 below. All measurements were recorded in millimeters to the nearest 0.1 mm and from the right side; bilateral meristic characters were also recorded from right side. 58   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY TABLE A1.1. Abbreviations and definitions for characters examined. Character class and abbreviation Character name Definition Mensural characters EyeD Orbit diameter Maximum horizontal diameter of exposed eyeball HeadL Head length HeadW Head width NarEye Nares?eye length SnEye Snout?eye length SnW Snout (internarial) width SVL Snout?vent length TailL Tail length TrunkL Trunk length Meristic characters of scalation Chin Chin scales Number of scales touching internal edge of infralabials and mental from juncture of 2nd and 3rd infralabials on left and right CircNa Circumnasal scales CloacS Cloacal spurs Dorsal Dorsal scales Number of scales longitudinally at midbody on dorsum contained within one EyeD Inflab As for Suplab PoreC Femoral and precloacal pores series FemPor and PreclPor series continuous or separated PreclPor SnS Scales between supranasals Number of scales touching rostral scale between left and right supranasals Subcaud Scales subequal to dorsal scales or enlarged into plates Suplab Supralabial scales Number of enlarged scales from rostral to top of mouth curve, usually equivalent to end of orbit TotPore Total number of secreting pores Total number of left and right femoral pores and PreclPor Ventral Ventral scales Number of scales longitudinally at midbody on venter contained within one EyeD 1FingLm First digit lamellae Number of lamellae (wider than long) on 1st digit of forefoot 1ToeLm First digit lamellae Number of lamellae (wider than long) on 1st digit of hindfoot 2-5FingLm, 2-5ToeLm Second to fifth digit lamellae Number of entire, U-shaped subdigital lamellae (=scansors) on enlarged pad of 2nd to 5th digit, single apical lamella not counted, only large U-shaped lamellae touching edge of pad Meristic characters of coloration CaecMel Pigmentation of caecum Caecum pigmented or not OrbStrp Postorbital stripe Dark lateral stripe from eye to mid-neck or beyond, absent or present OvidMel Pigmentation of oviduct Oviduct pigmented or not PostocS Postocular spots Number of light spots above OrbStrp from behind eye to front of shoulder Appendix 2: Specimens Examined LOCALITYSAMPLES The specimens are segregated by the geographic-specific samples. These samples are arranged from east to west and, for those within similar longitu- dinal bands, from north to south. The number of specimens listed below for a locality can exceed the number of individuals in a locality sample [small cap name in brackets] because data were not collected on all specimens due to size or state or preservation, although the specimen?s specific identity was confirmed. Catalog numbers for primary type specimens are in bold. Hemiphyllodactylus typus Hawaiian islands [Hawai]. No island given: AMNH 22340, MCZ R20268, R154043, USNM 21220. Hawaii: USNM 23459?460, 310815?816, 518722, 570745?748. Kauai: USNM 163573, 23485, 23499, 23500, 279241. Lanai: USNM 570736?744. Molokai: BPBM 1576, 6595, 6715?17. Maui: BPBM 11557?560, MCZ R1093, R174988. Oahu: BMNH 1903.2.21.5?7, BPBM 0863?64, 6158, 6567, FMNH 42251, 212245, USNM 23509, 58969, 59482, 59493?496, 59722?723, 279238?240. Polynesia [Polyn]. Cook Islands, Mangaia: SDNHM 67822?824. French Polynesia, Marquesas: BMNH 1926.1.20.38, 1926.1.20.50, FMNH 17914, MNHN 1988.3034; Society Islands: MNHN without number, USNM 68047. Henderson Island: BMNH 1913.1.17.1?17.3. Fiji and Tonga [Fiji]. Vanua Levu: AMS R107894, USNM 322442; Viti Levu: AMNH 41689, BMNH 1938.8.2.7, QM J048853, J048898, USNM 230185, 267928, 267978?979, 310810?814, 345104, WmBeckon 80?82, 148, 169, 173, 175. Samoa: USNM 345102. ?Eua: USNM 268045?046, 322119; Ton- gatapu: CAS 49971, USNM 268044, 322120; Vava?u: USNM 333617; Tonga- Ata: BMNH 91.11.13.1. new Caledonia and VanuaTu [nCal]. New Caledonia: AMS R125697, R125699, R125787?788, BMNH 71.4.16.30A?B, 85.11.16.8, CAS 172739, MNHN 1887.270, NMB 6978. Vanuatu: FMNH 69613, ZSM R110. 60   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY taiwan and Japan [taiwan]. Ryukyu Is- lands: KUZ 018095?096. Taiwan: CM 118859, KUZ 009612, USNM/field 123689, USNM 291710?712, 291807?810. Philippines [Philip]. Palawan: CAS-SUR 28692?696, FMNH 52003, MCZ R150339. Saub: MCZ R26082. new guinea and solomon islands [nguin]. New Guinea: AMNH 59045, 95887, 100200?206, CAS 192984, 192986?987, MCZ R49273, R140954?140955, R145981, USNM 119246, 203865. Solomon Islands: MCZ R79198, USNM 287441. islands of indonesia and Malaysia [sunda]. Malaysia: KUZ 18094, MCZ 43480, 140968. Singapore: BMNH 96.6.25.11, ZRC 2.2291, 2.3282, 2.3378, 2.3469, 2.5361, 2.5385?86, 2.5415?16, 2.5622, 2.6021?22, 2.6596. Borneo: BMNH 912287, 1959.115A?B, FMNH 63661?662, 138545, 158734, 196268A, 213665, 239661, 243789, KUZ 8732, 8746, MCZ R43478, USNM 313965, ZMB 11355, ZRC 2.5671?72, 2.5675?78, 2.5955. Sumatra: BMNH 91.10.27.2, 1931.5.5.2, 1946.8.30.83, MCZ R38971, NMW 179171, RMNH 4172 (3), 7161, SMF 23125, 30326, ZMA no number (3). Java: BMNH 85.12.3.17, NMW 179172, RMNH 28008?014, SMF 8930, 22611, USNM 44202, ZMB 31280, ZRC 2.1392. Bali: SMF 23126, WAM 109012. Komodo: UF 28878, 28985. East Indies: RMNH 3991. southeast asia [sEasia]. Thailand: ZRC 2.5367. india and sri Lanka [india]. Sri Lanka: BMNH 74.4.29.1326, 90.11.8.5, 1908.7.2.1, 1910.3.16.4, 1972.2108, NMB 8552. Mascarenes [Mascar]. Mauritius: BMNH 1926.1.20.38, 1926.1.20.50, IRSN 24309, MCZ R51642? 643, USNM 149760, 565090?093, ZFMK 25350, ZMA 14717, 14766. Rodriquez: BMNH 1975.416. Hemiphyllodactylus aurantiacus india and sri Lanka [india]. India: BMNH 74.4.29.1332, 74.4.29.1333, 74.4.29.1334? 1337, 91.11.27.1?3, 94.8.30.2, NMB 2900, 9682, NMW 14753, ZMB 10233, ZRC 2.4601, 2.4678?680. Hemiphyllodactylus ganoklonis Republic of Palau [Palau]. Babel- daob: SAM R47715, USNM 495065?066, 563663?666. Ngeanges: USNM 563667. Ngeaur: USNM 563668. Ngercheu: USNM 563669?674. Ngerekebesang: USNM 563675. Ngeruktabel: USNM 563676. Oreor: USNM 563677. Ulebsechel: USNM 563678?681, 563682, 563683. Hemiphyllodactylus harterti islands of indonesia and Malaysia [sunda]. Malaysia: BMNH 1901.3.20.2, ZMB 15360. Hemiphyllodactylus insularis Philippines [Philip]. Bohol: CAS-SU 25107. Borocay: CAS 127889, 127965?971. Cancuman: MCZ R26600. Cebu: CAS-SU 27310, 125228, 132632, 136844, 138320, 145922?929, CAS-SU 28451, 28602. Great Govenen: CAS 60605. Mantique: CAS-SU 28987. Mindoro: BMNH 26085, CAS 62065, CM 2052, 2053, MCZ R26084, R26601, ZMA no number. Negros: AMNH 86598?599, 115512, BMNH 1976.1681, CAS 131795, 137652, 137654, 137659, 156017, 156019, 185989, CAS-SU 19373?374, 24832, MCZ 37700, RMNH 18009, USNM 310791?793. Palawan: CAS 139142, MNHN A951. Panay: CAS 137581?583. Poro: CAS 124517?518. Semirara: CAS 127855?857. Siquijor: CAS-SU 26450, 26597?607. Tabalas: CAS 137203?206, MCZ 26083. Hemiphyllodactylus margarethae islands of indonesia and Malaysia [sunda]. Sumatra: AMS R129492, BMNH 91.9.24.9, IRSN 2375A?B, RMNH 7341 ZMA no num- ber (1), 11095, 11096. Hemiphyllodactylus titiwangsaensis islands of indonesia and Malay- sia [sunda]: Malaysia: AMS R135270, MCZ R166921, ZFMK 32284?286, ZRC 2.4780?81, 2.4782, 2.4783?85, 2.4832, 2.1565, 2.5419, 2.5942?5943. Hemiphyllodactylus yunnanensis china [china]. Myanmar: BMNH 1933.7.8.11, USNM 310819, 570732?735. Yunnan: BMNH 1904.1.26.1, 1904.11.29.1?9, 1904.11.29.10A? N, CMS 8153, FMNH 7716?17, MCZ R18967, MNHN 1912.293, 1912.295A?B, 1912.296, NMB 9541. Laos: n um b e r 6 3 1   ?   6 1 FMNH 14451?452. Thailand: BMNH 1931.11.21.1, BPBM 3502, FMNH 178328, 180867, 215988?994, QM 4820, THNHM 0153?54, 5943?949, USNM10621?622, 310798?808. Vietnam: MNHN 1948.43?44, RMNH 28007, USNM 310797. Hemiphyllodactylus ?yunnanensis? southeast asia [sEasia]. Cambodia: FMNH 270569. Thailand: THNHM 075, 4714?715, 4910?17, 8620, ZRC 2.3567. Thailand, country only: BPBM 3502 hermaphrodite with large testes and pair of vitellogenic follicles (diameter 3.2 mm). Vietnam: USNM 146161. Hemiphyllodactylus [speciesindeterminate] islands of indonesia and Malaysia [sunda]. Borneo: Brunei: ZRC 2.5672, 2.5675?78; Kalimatan: KUZ R8723, R8746, USNM 313965; Sabah: BMNH 95.9.11.5A&B, 1929.12.22.87, FMNH 63661? 662, 239661, 243789, MCZ R43478; Sarawak: FMNH 138545, 158734, 196268A, 213665, ZRC 2.5671, 2.5955. china [china]. Hong Kong: MCZ R182874? 876, MNHN 1912.293. india and sri Lanka [india]. Sri Lanka: BMNH 91.03.16.4, NMB 8552. TYPESPECIMENS Catalog numbers for the primary type specimens here are identified in bold in the preceding locality samples. Hemiphyllodactylus typus Bleeker: BMNH 1946.8.30.83. Hemidactylus aurantiacus Beddome: BMNH 74.4.29.1332?1337, ZMB 10233. Spathodactylus mutilatus G?nther: BMNH 1946.8.30.83. Lepidodactylus ceylonensis Boulenger: BMNH 74.4.29.1326. Hemiphyllodactylus leucostictus Stejneger: USNM 21220, 23459?460, 23485, 23499?500, 23509. Lepidodactylus Harterti Werner: ZMB 15360. Gehyra larutensis Boulenger: BMNH 1901.3.20.2. Gehyra yunnanensis Boulenger: BMNH 1904.1.26.1?26.2. Hemiphyllodactylus insularis Taylor: CAS 62065, CM 2052?53. Hemiphyllodactylus margarethae Brongersma: ZMA 11095?096, IRSN 2375A?B. Hemiphyllodactylus typus chapaensis Bourret: MNHN 1948.43?44. Hemiphyllodactylus typus pallidus Auffenberg: UF 28878, 28985. Hemiphyllodactylus ganoklonis Zug: SAM R47715, USNM 495065?066, 563663?683. Hemiphyllodactylus titiwangsaensis Zug: ZFMK 32284?286, ZRC 2.4780? 4785, 2.4832, 2.1565, 2.5419, 2.5942?5943. ADDITIONALLOCALITYRECORDS The following localities derive from distributional re- cords appearing in publications and from museum speci- men records for specimens that I did not examine directly. The specific identity provided by the museum or in the publication is the one usually followed; however, where information was adequate and contrary to author?s species determination, I have re-identified the specimen. Hemiphyllodactylus typus Mascarene islands. La R?union: D?so et al. (2007); Rodriques: Schr?der and R?ll (2004). nicobar islands. Great Nicobar Island: Biswas and Sanyal (1980). sumatra. Pulau Enggano: MVZ 39345? 39346, 239586. thailand. Kanchanaburi, Nakhon Ratcha- sima, Narathiwat, Phang-Nga, Phuket, Ranong, Trang, and Trat provinces: Pauwels and Sumontha (2007). Vietnam. Southernmost mapped locality: Bo- brov and Semenov (2008). china. Hainan Island: MVZ 42817?42818. Ryukyu. Iriomotejima: Ota (1990). taiwan. Main island: Ota (1989). Papua new guinea. Milne Bay Province, Pini Range: Kraus and Allison (2004). solomon islands. Guadalcanal: McCoy (2006). hawaiian islands. All major islands: Mc- Keown (1996). Marshall islands. Enewetak: R. I. Crom- bie (unpublished manuscript, ?Pacific amphibian and rep- tile distributions,? 1994). cook islands. Rarotonga: Gill (1998). French Polynesia. Marquesas: Elao, Hivo Oa, Mohotani: Ineich and Blanc (1989); Pitcairn: Ineich (1992); R. I. Crombie (unpublished manuscript, 1994). Hemiphyllodactylus aurantiacus india. Anaimalai Hills, Bangalore, Kolli Hills, Nilgiri Hills, Shevaroy Hills: Bauer and Das (1999); 62   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY Bangalore: Daniels (1994); Andhra Pradesh, Visakhapat- nam District, Araku Valley and Madhygulu: Sanyal et al. (1993). Hemiphyllodactylus yunnanensis china. Yunnan?Changyuan, Chengjiang, Chuxiong, Gejiu, Jinping, Lijang, Longling,Yao?an; Gui- zhou?Anlong, Dushan, Huishui, Xingyi; Guangxi?Day- aoshan: Zhou et al. (1981). Vietnam. All mapped localities N of 15?N: Bobrov and Semenov (2008, op. cit.); Tam Dao: MVZ 226500. Hemiphyllodactylus ?yunnanensis? cambodia. Phnom Tumpor: Grismer et al., (2008); Koh Rongnieur and Koh Khlee-Ay islands, Me- gong R.: Bezuijen et al. (2009). thailand. Khao Yai National Park: Chan-ard et al. (1999). Hemiphyllodactylus [speciesindeterminate] (THESE LOCALITIES NOT MAPPED) sri Lanka. Southwestern Sri Lanka: So- maweera and Somaweera (2009). Appendix 3: Statistical Analyses I used standard univariate statistics to summarize variation of the charac- ters within each sample. These data are the main ones presented for the comparison and description of samples, the means for mensural data, and the medians for meristic data. I tested all samples with adequate numbers of adult females and males for sexual dimorphism with the Student t test, sig- nificance at P ? 0.05. A repeats protocol provided an estimate of the variation derived from my data gathering (see detailed explanation in Baseline Estimate of Intra-Observer Variation subsection). Multivariate statistics (discriminant function analysis [DFA] and principal components analysis [PCA]) were used mainly to explore the homogeneity of samples. My goal was to discover which mensural traits best differentiated be- tween unisexual and bisexual individuals within large regional samples. I did not use scalation traits in these analyses, although I did use proportions in some analyses but did not mix measurements. Some authors (e.g., Atchley et al., 1976) have argued against the use of proportions in multivariate tests and dem- onstrated problems with proportional data through simulation studies. Other authors have shown that proportions and nontransformed measurements do not yield significantly different results in data sets from museum specimens such as frogs (Heyer, 1978) and turtles (Iverson, 1981). I justify my use of proportions herein because I was neither testing differences between or among groups or relying on statistical significance in the assignment of specimens to taxonomic group. The following paragraphs present a synopsis of the statistical results from the comparison using PCA and DFA. They are arranged in the same sequence as in the text. Uniformity among the unisexual samples: A DFA was employed to test uniformity (homogeneity) of the adults (n = 119) of the Pacific samples including the holotype of H. typus using the eight mensural traits (EyeD, HeadL, HeadW, NarEye, SnEye, SnW, SVL, TrunkL; TailL was not examined in this test nor any of the subsequent ones). The adjusted classification matrix (jackknifed) yielded an average accuracy of 23% assignment. The first three predictor variables (SVL, TrunkL, HeadL) had eigenvalues of 0.373, 0.317, and 0.180 and accounted for 64   ?   SM I TH SON I AN CONTR I B U T I ON S TO ZOOLOGY 36.5%, 31.0%, and 17.6% of the total variance, respec- tively. A PCA (covariance matrix) of the same combined samples and traits produced a compact clustering on the first two components. The first three components had ei- genvalues of 24.72, 1.064, and 0.156, respectively, with the first component accounting for 94.6% and the second com- ponent 4.1% of total variance. SVL, TrunkL, and HeadL had the strongest loading (eigenvalues of 0.826, 0.534, and 0.131). Principal components analysis of body proportions in sunda sample: Principal components analysis (corre- lation matrix) results for adult females (n = 72) are sum- marized in the text with the exception of eigenvalues for the first four components: 2.712, 2.400, 1.639, 1.206; these components accounted for 79.7% of total variance. A PCA of adult males (n = 9) identified HeadL/SVL and SnW/HeadW as the highest loading variables on the first component, NarEye/HeadL on the second, TrunkL/SVL on the third, and SnW/HeadL on the fourth. In total, these four components accounted for 91% of total variance, with 48.1% for the first component. Eigenvalues for the first four components were 4.826, 2.567, 0.948, and 0.826. Dispersion on a plot of first and second components was broad, although a regional clustering occurred with Suma- tran males in the upper left quadrant, Bornean males in the lower third of the upper right quadrant, and Malaysian males in the lower left quadrant. Discriminant function analysis examination of india sample: A DFA of india males (n = 6 India, 1 Sri Lanka) yielded 71% accuracy (jackknifed classification) using eight mensural traits and a considerably lower accuracy (14%) with the 10 proportional traits. The accuracy for the unad- justed classification was 100% for both character sets. The eigenvalues for the two analyses were 13.073 and 7.029, respectively. Using the 10 proportional traits, classification accuracy was better (80% jackknifed) in the adult females (n = 4 India, 1 Sri Lanka); mensural traits were not exam- ined in females. Discriminant function analysis examination of Malay- sian Hemiphyllodactylus samples: The distinctiveness of the H. harterti sample (n = 2 adults) and a central moun- tain range sample (15 adults) was explored with two data sets, the eight mensural traits, and a subset of eight meristic traits (CircNa, SnS, Suplab, Inflab, Chin, CloacS, Subcaud, TotPore). The latter subset had been identified as the best set of discriminators by an earlier DFA of all 22 scalation characters. Of the mensural set, HeadL, SnEye, and OrbD were assigned the largest classification functions. The unad- justed classification matrix yielded 100% accuracy of group assignment and the jackknifed matrix only 76% total accu- racy, with a single eigenvalue (3.045) reported. The meristic set yielded 100% classification accuracy in the unadjusted matrix and 94% in the jackknifed matrix, with a single ei- genvalue (13.241) reported. References Atchley, W. R., C. T. Gaskins, and D. Anderson. 1976. 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R., J. V. Vindum, and M. S. Koo. 2007. Burmese Hemi- dactylus (Reptilia, Squamata, Gekkonidae): Taxonomic Notes on Tropical Asian Hemidactylus. Proceedings of the California Academy of Sciences, 58(19):387?405. Zug, G. R., L. J. Vitt, and J. P. Caldwell. 2001. Herpetology. Introductory Biology of Amphibians and Reptiles. 2nd ed. San Diego, Calif.: Academic Press. Zug, G. R., D. Watling, T. Alefaio, S. Alefaio, and C. Ludescher. 2003. A New Gecko (Reptilia: Squamata: Genus Lepido- dactylus) from Tuvalu, South-Central Pacific. Proceedings of the Biological Society of Washington, 116:38?46. Index of Taxa Names Page numbers in bold italic font indicate the start of the taxonomic account. Cainodactylus, 4 Gehyra, 4, 23, 30 larutensis, 4?6, 16, 25, 28, 33, 43, 50, 61 oceanica, 9 yunnanensi, 4, 6, 51, 61 Hemidactylus, aurantiacus, 6 bowringii, 33, 35 frenatus, 9, 30 Hemiphyllodactylus, ii, v, vii, xi, 1?6, 8?12, 14, 16?19, 21, 23?32, 34?35, 47 albostictus, 5, 35 aurantiacus, ii, vii, xi, 2?5, 16, 20, 26?27, 31?32, 34, 37, 39?40, 53, 60?61 ceylonensis, 4 chapaensis, 18?20, 22, 32?33 crepuscularis, 4 ganoklonis, ii, vii, 22, 27, 40, 41?43, 53, 60?61 harterti, ii, vii, xi, 3?5, 9, 11, 16?17, 27?29, 33?34, 43, 44?45, 53, 60, 64 insularis, ii, vii, viii, xi, 3?6, 24, 31, 34, 44, 46?47, 53 60?61 larutensis, ii, 3, 5, 9, 16?18, 28?29, 33, 44 leucostictus, 4, 6, 32, 35, 61 margarethae, ii, vii, viii, 3?4, 6, 11?12, 15?16, 18, 22, 25, 28?29, 32?34, 45, 47, 48, 53, 60?61 titiwangsaensis, ii, vii, viii, 27, 45, 48, 50, 53, 60?61 typus, ii, vii, xi, 1?6, 8?16, 22, 23, 25, 27?34, 35, 36?37, 52, 59, 61, 63 aurantiacus, 4?5 chapaensis, 5?6, 32?33, 50?51, 61 pallidus, 3, 5?6, 34?35, 61 typus, 4?5 yunnanensis, ii, vii, viii, xi, 3?5, 8, 11, 18?20, 22?23, 27, 29, 31?34, 40, 50, 51, 53, 60, 62 dushanensis, 3, 5?6, 19?20, 51 jinpingensis, 3, 5?6, 19?20, 33, 51 longlingensis, 3, 5?6, 19?20, 22?23, 32, 51 yunnanensis, 20, 22?24, 50 70   ?   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