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s m i t h s o n i a n c o n t r i b u t i o n s t o z o o l o g y • n u m b e r 6 3 1

Speciation and Dispersal in 

a Low Diversity Taxon: 

The Slender Geckos 

Hemiphyllodactylus

(Reptilia, Gekkonidae)

George R. Zug
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s m i t h s o n i a n c o n t r i b u t i o n s t o z o o l o g y • n u m b 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 species. The unisexual species, Hemiphyllodactylus typus, the most widespread of these geckos, apparently attained 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 investigation. 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

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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 • smithsonian contributions 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 • smithsonian contributions 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

My fascination with Hemiphyllodactylus began during a study of 

Fijian lizards. Often, the presence or absence of secreting precloacal 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 continue my examination of Hemiphyllodactylus specimens and led to an observation 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 bisexual 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, yunnanensis, 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 wideranging 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 herpetologists and hobbyists.

How does such an inconspicuous gecko attain such a broad occurrence? 

Human transportation seems the obvious answer, although the subsequent questions of how, why, and when are largely unanswered. The search for a datasupported 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 Hemiphyllodactylus typus Bleeker is associated correctly with a gecko population 

today. Although the assignment of the name to a specific taxon is unambiguous, 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 Ptyodactylus 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 • smithsonian contributions 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 noting 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 Goenong Parong with P. gracilis “led Smith (1935) and Wermuth (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 recognized it as a new species, Platydactylus crepuscularis, in 

his catalog of New Caledonian reptiles (Bavay, 1869), apparently 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 Duméril and 

Bibron, Bavay’s description explicitly characterized P. crepuscularis 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 describing 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, communicated 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 deposited 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 description clearly refers to H. typus Bleeker.

Major Beddome, a forestry officer in Madras (present-day Chennai, India), collected a variety of reptiles 

and described them in 1870. One of them, Hemidactylus aurantiacus Beddome, was a Hemiphyllodactylus species 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 Lepidodactylus was a better assessment of aurantiacus’ affinities. Boulenger also noted that the type series consisted of 

many adult males, females, and juveniles.

Günther (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 Günther 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 Günther 

are the same specimen; thus the latter name is a junior 

objective synonym of the former. Because Boulenger included neither Hemiphyllodactylus nor typus as names in 

his catalog, it indicates that neither he nor Günther was 

aware of Bleeker’s description. Boulenger (1885) did recognize that Günther’s Spathodactylus was a homonym for 

a fish and provided a new generic name Spathoscalabotes.

Subsequently, Malcolm Smith (1935) listed the type locality 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 synonymy, Smith incorrectly gave Java as the type locality of 

H. typus Bleeker. 

As noted above, Boulenger (1885) was apparently unaware of Bleeker’s description of Hemiphyllodactylus and 

H. typus, because these names are absent from his catalog. He placed crepuscularis, ceylonensis, and aurantiacus

in the genus Lepidodactylus and continued to recognize 

Günther’s mutilatus as a monotypic taxon although correcting the generic homonymy.

Boulenger followed the species account of Lepidodactylus crepuscularis (=Platydactylus crepuscularis Bavay) 

with an exceedingly brief description of Lepidodactylus 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 coloration.” Boulenger’s illustration is suggestive of Hemiphyllodactylus, but it would fit other geckos as well.

Stejneger (1899) provided the first review of the Hawaiian terrestrial reptiles. Of the seven lizard species then 
n u m b e r 6 3 1

• 

3

FIGURE 1. Type localities for the available names (see Table 1) for the species currently assigned to genus Hemiphyllodactylus. Abbreviations: d, H. yunnanensis dushanensis; j, 

H. y. jinpingensis; l, H. y. longlingensis.
4 • smithsonian contributions to zoology

occurring on these islands, only one appeared to represent a 

new species, Hemiphyllodactylus leucostictus Stejneger. Stejneger 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 Lepidodactylus 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 descriptions, he mentioned the low number of chevron-like lamellae 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 Werner’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 tentative correction [reassignment] is addressed in the Taxonomic 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 uncertain 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 specimens 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 Hemiphyllodactylus 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. Stejneger 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 created a synonym.

Brongersma (1931) described Hemiphyllodactylus 

margarethae from four Sumatran specimens representing 

two montane localities. He recognized this species’ affinity and differentiated it from H. typus and H. ceylonensis. 

Subsequently, Brongersma (1932) published an evaluation 

of the nomenclature and characterization of Hemiphyllodactylus 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. ceylonensis, H. crepuscularis, H. leucostictus, H. insularis, and 

H. margarethae. He noted that M. A. Smith had independently 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 individual” (Brongersma, 1932:214). He recognized the difficulty 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 u m b e r 6 3 1 • 5

nominate subspecies included as synonyms all the species 

mentioned in the preceding paragraphs except H. larutensis, 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 Hemiphyllodactylus 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 irregularly in the herpetological literature until the 1960s, typically in regional lists, reappearing in Taylor’s lizards of 

Thailand (Taylor, 1963) with a full description and Wermuth’s (1965) checklist of all gekkonid lizards. Wermuth 

recognized three species (larutensis, typus, and yunnanensis) and three subspecies of H. typus (nominate form, 

aurantiacus, and chapaensis). Kluge (1968) addressed 

the relationships of Hemiphyllodactylus as well as commenting 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 investigation. Kluge considered Hemiphyllodactylus as a sister 

group of Lepidodactylus. This hypothesis returns conceptually 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 proposed that the specimen was a Hemiphyllodactylus typus. 

His Figure 2 of the right forefoot shows subdigital lamellae similar to those of H. typus; however, the dorsal view 

of the entire gecko (Wermuth, 1966: fig. 1) is not typuslike. The head, neck, and body are robust and not elongated. The fore- and hindlimbs are large, long, and would 

overlap one another if laid along the trunk. With this habitus, 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 publications 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 colored 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 patterns. Because the variation was concordant within four 

regions, they recognized three new subspecies: H. y. dushanensis, 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 synonymy 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 illustrations and descriptions, confirming the presence of two species 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 captured 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 species. 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 presence of males and females, accepted H. insularis as a distinct taxon from H. typus. The available names for the 
6 • smithsonian contributions to zoology

various populations of Hemiphyllodactylus are summarized chronologically in Table 1.

MATERIALS AND METHODS

Despite the broad distribution of Hemiphyllodactylus, the availability of voucher material in museum collections 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

IRSNB 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 (National Museum of Natural History, 

Smithsonian Institution)

WAM Western Australian Museum

WmBeckon William N. Beckon, private collection

ZMA Zoölogische 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-Calédonia

1870 Hemidactylus aurantiacus Beddome Shevaroys . . . about Yercaud [India]

1872 Spathodactylus mutilatus Günther 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 u m b e r 6 3 1 • 7

ZMB Museum für Naturkunde, Universität 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 abbreviated 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 examined as two or more subsamples of restricted localities 

if adequate specimens are available or if intrasample variation indicates a mixed sample. Further, I combined samples 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 morphometric 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

CHARACTER ANALYSIS:

RESULTS AND DISCUSSION

Baseline Estimate of Intra-Observer Variation

How much of the variation observed in each sample results 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 variation (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 characters. 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 estimates 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 • smithsonian contributions 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 results reveal that intrasample variation is surprisingly low 

in most characters and samples; nonetheless, the reader is 

advised to be cautious in over-interpreting reported differences, particularly in small samples and/or where differences are less than 2 times a character’s standard deviation.

Recognition of Unisexual and Bisexual Populations

Unisexual populations contain only females. Although it may be a statement of the obvious, how does 

one confirm the unisexuality of a population? Because neither data on reproduction of virgin females nor mitochondrial DNA were available to me, I relied on the absence of 

males in samples as a hypothesis of unisexuality for populations 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 assembly, 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: Hemiphyllodactylus 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 examined 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 removed 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 dissection, I observed a consistent pattern of differential pigmentation 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 Museum [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. yunnanensis, Myanmar (USNM 570734).
n u m b e r 6 3 1 • 9

the oviduct the peritoneum sheath likely bears the melanin. 

The pigmented caecum and oviducts, however, are not confined 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 hemipenes. The black caecum is visible through the body wall.

Précis. 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 median 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 unisexual sample; Pacif), are equivalent. There is no evidence 

of mensural differentiation among the Pacific insular samples 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 morphology 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 morphology; however, my preliminary measurements of hindlimb 

length were extremely variable owing to the difficulty of 

measuring accurately tiny twisted limbs with fist-like preserved 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 regression of the two preceding traits also reveals the degree 

of elongation through the depression of the slope; however, 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), Hemidactylus 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 compared 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 proportionately shorter in H. typus in contrast to the visual 

impression of the attenuate habitus of H. typus.

Do these proportions permit the segregation of unisexual 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 collection (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), holotype (adult male) and three paratypes (2 adult females, 
1 0 • smithsonian contributions 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 measured 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%

aThis 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).

cTwo finger formulae share 27%; the second is 3-4-4-4.
n u m 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 clustering 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 loading (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 variance (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 bisexuals 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. margarethae 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.)
1 2 • smithsonian contributions 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. margarethae 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 smallheaded 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 bisexuals; 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 bisexuals 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 includes only one unisexual (SVL 38.3 mm), confirmed by 

pigmented oviducts and caecum; this adult female, however, lacks secreting precloacal–femoral pores, thereby 

questioning its assignment to H. typus. India has four unisexual 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 statistics 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.

Précis. (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 unambiguous unisexual samples is presented in Table 2. Intrasample 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 scalation 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 museum vouchers were poorly positioned at time of preservation. 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 modestsized 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 remainder 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 females 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 recorded for the two largest individuals (40, 42 mm SVL). 

The 39-mm individual had small ovarian follicles, and a 
n u m b e r 6 3 1 • 1 3

FIGURE 5. Frequency distribution of precloacal–femoral pores in the Pacific sample of unisexual Hemiphyllodactylus 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.
1 4 • smithsonian contributions to zoology

35-mm female had mid-vitellogenic follicles. These data 

indicate that all poreless individuals were mature, although 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 Hemiphyllodactylus, I developed the impression that the number 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 development event. Precloacal and femoral pores are not continuous 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 individuals 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 variation (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 differences likely result from vagaries of sampling. The other 

difference, probably of similar origin, is the median forefoot 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 distribution of this unisexual taxon.

Précis. (1) The island and island group samples are homogeneous within and among samples, thereby 

representing a single genetic entity, Hemiphyllodactylus 

typus (Pacif). This homogeneity is shared among the unisexual 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 broken 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 instances, 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 multitude 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 progressively diffuse and indistinct in most individuals.

In life, a few individuals will appear uniformly brown 

except for faded head markings. This situation is the common condition for preserved specimens and limited the development of quantitative coloration coding only to two 

characters (OrbStrp and PostocS; Appendix 1).

Neither of these two traits shows any striking variation 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.

Précis. Hemiphyllodactylus typus are predominantly dull-colored geckos. The most striking coloration 
n u m 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 populations 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 Figure 2B). Females of SEAsia and China commonly lack 

pigmentation on these organs; however, an adult SEAsia female (BMNH 1931.11.21.1) from Thailand has pigmented 

oviducts and caecum. Within the Sunda sample, some females have the pigmented condition and others do not. I 

observed no differences in extent or intensity of pigmentation among the unisexual and bisexual populations. Occasionally, 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 peritoneum over the testes and epididymides is pigmented, although 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.

Précis. (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).
1 6 • smithsonian contributions to zoology

trait also occurs in females of the bisexual Palau, Philippines, and Indian populations, and some Sumatran females. (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 dimorphism 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 bisexual populations. The Palauan population has the smallest 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 measurement 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%; NarEye 4.3–14.9%; EyeD 5.0–13.6%; SnW 6.2–21.4%. Relative 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 SEAsia 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 locality. 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 female 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 samples. 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 populations average the smallest in adult size of all Hemiphyllodactylus 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 specimens was evident in the high frequency of misclassification 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. aurantiacus (Bauer and Das, 1999) lie within the size ranges 

noted above. These authors emphasized scalation and coloration; 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 Malaysia, n = 17; Sumatra, 8; Borneo, 3; no precise locality, 

1). These samples sizes are inadequate to address mensural variation in detail; however, owing to the existence of 

three names (harterti, larutensis, margarethae), statistical 

examination of the mensural data is necessary. The Malaysian 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 Lepidodactylus Harterti Werner (ZMB 15360). A series of 17 adults 
n u m 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.
1 8 • smithsonian contributions 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 proportions 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 Hemiphyllodactylus margarethae Brongersma: two adult females 

(42.0 mm, ISRN 9338B; 46.9 mm, ZMA 11096), a juvenile 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 uncertain provenance in the East Indies.

Sunda adults do not display sexual dimorphism in 

any mensural or proportional trait. The preceding description of size differences in the three regional subsamples 

suggests the possibility of size dimorphism in the Malaysian sample (i.e., females larger but not statistically significantly 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 morphometrics and the general similarity (means differ but strong 

overlap of ranges occurs) of the China and SEAsia samples (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 Chiang Mai in the west to Hong Kong in the east. The latitudinal 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 geographically expansive as SEAsia and limited to the “highlands” 

(Shan and Yunnan plateaus). 

To test the preceding assumption of homogeneity 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, results 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 proportions of the two samples are less than 1% different, supporting 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) population 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 Southeast 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 longer trunk (53% TrunkL/SVL, Table 4) similar to the taxa 

with the slender habitus. A DFA of Oriental adults using 
n u m 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 dimorphism 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 Hemiphyllodactylus (BPBM 3502, Sakaerat) is hermaphroditic, 

with large testes and a pair of vitellogenic follicles (maximum 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 values 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
2 0 • smithsonian contributions to zoology

Précis. (1) Morphometry is only modestly useful 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 populations are also small but average significantly larger than 

the Palauan one. Average size of the China and SE­Asia

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 component (H. aurantiacus) and a Sri Lankan one (unnamed). 

Inclusion of northwestern Thailand specimens in the 

China population does not increase variation, thus suggesting genetic homogeneity among these “highland” populations. 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-India 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 scalation 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 represents 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 Suplab, 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 yunnanensis. 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 u m b e r 6 3 1 • 2 1

and usually not statistically significant, I suggest the differences reflect genetic differentiation among the regional 

populations and are not the result of sampling or measurement 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 medians (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).
2 2 • smithsonian contributions 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 postmental 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 laterally abutting chin scale is commonly one-third to twothirds 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 granular 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 posterior 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 très 

petites plaques.” His illustration (reproduced in Figure 

7F) shows an arc of enlarged chin scales with postmentals 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 Vietnamese 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 postmentals 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 variance 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 values 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 Chinese 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 (redrawn from Bourret, 1937: fig. 1b).
n u m 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 foreand 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 Titiwangsa 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 examined seven traits: rostral notched posteromedially, enlarged scale posteriorly bordering “supranasal,” Suplab, 

Inflab, chin scale bordering mental posteriorly (postmentals)], 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 included in my character set. The variation in the relative size 

of the post “supranasal” scale completely escaped my attention. 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, Thailand (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.
2 4 • smithsonian contributions to zoology

larger than its laterally abutting scales in the arc. Because 

my China sample derives largely from Kunming, my results 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 proximalmost one on each pad can be challenging owing to a 

gradual reduction in size proximally. My protocol to increase 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 represent 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 invariant 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 medians (19–22.5) and range of (11–28) of TotPore (Table 6) 

for Palau, SEAsia, China, and India. Philip has a distinctly 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 escutcheon area for pore development. Higher or lower TotPore counts derive mainly from an increase or decrease in 

the length of the femoral pore series. 

Number of pores, however, disguises two pore morphologies (Figure 10): (1) continuous series of precloacal 

and femoral pores and (2) separate longitudinal patch of 

precloacal and femoral pores. The continuous pore morphology 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 identical 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 Vietnam through southern Thailand, it is uncertain whether 

this condition is a variant or populational state.

Pore morphology in Sunda shows regional differentiation 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 u m 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; further 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 nomenclaturally 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.

Précis. (1) Sexual dimorphism of scalation 

occurs only in the presences (males) and absences (females) of precloacal and femoral pores. (2) Most scalation 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 series of precloacal–femoral pores and Palau, Philip, some 

Sunda, and India with separate patches of pores.

Bisexual—Coloration

The drab, presumably cryptic, coloration of Hemiphyllodactylus makes characterization of coloration difficult. I recorded two external coloration characters: 

PostocS and OrbStrp. The results show regional differences. 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 “identical” ranges reflect the difficulties of seeing these spots 

in older or poorly preserved specimens. If an actual difference 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 specimens, at least to the level of the ear.

With my attempt at quantification of coloration illfounded for museum specimens, I offer a comparative 

descriptive approach using the major pattern features 

identified in the H. typus coloration section. These descriptive data derive from my notes on museum specimens, 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 populations. 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 photographed. 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 lightening 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 population to population. A dark brown lateral stripe extending from the loreal area to at least the anterior quarter 

of the neck occurs widely, although its thickness and loreal 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 anywhere 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 
2 6 • smithsonian contributions to zoology

broad, irregular dark brown bars (continuous across the 

midline or interrupted). Most individuals display a signature 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 middorsal 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. Posteriorly the top and sides of the tail range from light to medium 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, irregular 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 posterior 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 moderately developed, ranging from a dark preorbital spot to 

a narrow stripe extending midway to the naris. The postorbital 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 posteriorly. 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 backgrounds 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 impression 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 yellow. The postsacral mark is broad and deep and extends 

well on to the tail base; the anterolateral arms are usually 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 specimens 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, narrow, wavy crossbars begins on the shoulders and continues to the sacral area; these crossbars can be variously 

fragmented and/or narrowly connected to anterior or posterior crossbars. A dorsolateral series of light (presumably 

white to light tan) spots extend from the neck to the inguen; 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 u m 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 usually 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 grayish 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 postorbital 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 (Myanmar, Pyin Oo Lin; USNM 570734) (photograph by G. Zug).
2 8 • smithsonian contributions to zoology

or a postsacral mark. Werner’s (1900:196) description of

H. harterti reports “gleicht in Färbung 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 description of coloration for H. margarethae. Although I 

examined his type series, I have no notes on their coloration. 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 Hemiphyllodactylus are available in some recent field guides. Additionally, 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 paragraph, I am using the species identifications provided by 

the authors for their illustrated specimens.) The same individual (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 typical 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) Lepidodactylus 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. Rödel); (D–F) Gehyra larutensis 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 u m 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-developed postorbital stripe; nevertheless, I do not believe it 

is a H. typus, owing to light anterolateral arms of a postsacral 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 individuals 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 middorsal 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 continuous 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 Titiwangsa range) specimens do not share a common coloration 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 overall 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 Chanard et al. (1999:130) Bukit Larut “typus.”) The Titiwangsa 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.

Précis. (1) Although generally drab, bisexual 

Hemiphyllodactylus display some bright markings that 

seemingly are unique for different populations. (2) Colorful 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 SE­Asia 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 contrasting band of the tail. 

GEOGRAPHY AND TAXONOMY

Regional Patterns of Morphology and Speciation

General Observations

The present study does not address the generic status 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 molecular 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 populations. Two body forms are displayed among populations, 

at least in the older and/or mature individuals. Further, average adult SVL differs among populations, even though 

size at maturation appears similar among all populations.

Three features of scalation show populational differences. Chin scales in contact with mental and anterior 
3 0 • smithsonian contributions 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 (= postmental 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 populations 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 differentiation. 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 dorsolateral 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 frequently visible through the body wall and the pigmentation 

of the peritoneum covering the gonads and gonadal ducts 

(specifically the oviducts and epididymides). This pigmentation is not unique to the unisexual populations but also 

occurs in bisexual ones: Palau, Philip, and India.

I propose that this unique visceral pigmentation (Figure 2) reflects close phylogenetic relatedness among the 

unisexual H. typus and the three bisexual species (justification 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, unpublished data, 1990) examined individuals from northern 

Thailand (n = 13), Philippines (3), Fiji (5), and Hawaii 

(9) using nine isozymic loci. The Philippine and two Pacific 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 Philippine species as one of the parental species of H. typus. 

Similarities in morphology of Palau and Philippine individuals 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 hybridization 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 parental 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 darkest area, well removed from Gehyra and Lepidodactylus 

lugubris, and this segregation occurred before the Hemidactylus 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 origin hypothesis derives from my assumption that the entire 

distribution of H. typus derives from human transport—

shipboard, possibly during the eighteenth and early nineteenth centuries. 

Phenetic similarities (i.e., primitive advance state polarities not determined) suggest the relatedness of the Palauan 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 similar to the bisexual parental species. H. typus is the only 
n u m 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 anterolateral arms.

Even though similar, Palauan and Philippine populations have morphological differences that I interpret to 

demonstrate speciation. The dorsolateral spots and postsacral mark are red in the Philippine population (H. insularis Taylor) in contrast to pink spots and yellow suffused 

with pink postsacral mark of the Palauan population (unnamed). 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). Palauan 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 population (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 distant 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 yellow to gold and the anterolateral arms are variously developed. 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 significantly 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 population of Hemiphyllodactylus has such a low number of 

lamellae on its digits. The two Sri Lankan non-typus specimens 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 difficult. 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 vouchering of specimens. I must note that finding Hemiphyllodactylus 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 individuals, 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 (=Yunnan-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 Guizhou. 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 hypothesized for H. typus earlier in this section.

Hemiphyllodactylus yunnanensis and the other samples and populations of mainland eastern Asia share the 

adult robust body form, absence of caecal and gonadal pigmentation, 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 
3 2 • smithsonian contributions 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 enlargement (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 infralabials; 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 interpretation, 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, perhaps with a proportionately longer trunk (53 versus <49% 

for all other Asian subsamples). They have enlarged, although 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 strikingly 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 unisexual H. typus and the bisexual populations of India, 

Philippines, and Palau. The continuous pattern continues 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 geography of unisexual Hemiphyllodactylus separately from that 

of the bisexual species. Because squamate unisexuality appears to be universally derived from hybridization (Zug 

et al., 2001), I examine the geographic patterns of occurrence and taxonomy of the bisexual species first. 

Among the bisexual species, H. aurantiacus (Beddome) was the first to be described and is the easiest of 

the Hemiphyllodactylus populations to be addressed taxonomically. Small adult body size and uniquely low foreand 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. aurantiacus 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 recognized 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 individuals from most localities, too few localities, and localities 

geographically distant. The uniqueness of H. yunnanensis

(Boulenger) is unquestionable. Its traits (robust body, unpigmented 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 distributed species, although morphological data suggest minimal 
n u m 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 Bourret similarly becomes a synonym of H. yunnanensis (Boulenger). Geographically, the H. t. chapaensis type locality 

is only 300 km south of Kunming (type locality of H. yunnanensis) 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 discontinued 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 population? My present data suggest the latter interpretation, but 

they are insufficient to address this hypothesis rigorously. 

Presently, I am unaware of vouchered records of Hemiphyllodactylus 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 locating vouchers from the Guangxi–Guangdong corridor.

The preceding restriction (in Morphological Differentiation 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 museum vouchers does not permit a satisfactory resolution. 

Presently, the data are adequate to declare that Malay 

peninsular populations are distinct; however, it is uncertain 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 suggests 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 Indochina and Hong Kong specimens as “H. yunnanensis.”

Two names are available for Malayan Hemiphyllodactylus populations. H. larutensis derives from Bukit Larut 

(=Maxwell’s Hill or Larut Hills), a mountain at the southern 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., Malakkahalbinsel [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 Ornithology 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 Bintang). I have examined personally only one specimen 

(G. larutensis type) and have data and images from a second 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 Gunong Inas. I note only that Werner’s (1900:196) “gleicht 

in Färbung und Zeichnung ganz dem L. lugubris” better 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 mountain range or received specimens from there. The similarity 

of morphology of the two type specimens and the type restriction of H. harterti to the Bintang range makes H. harterti 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 distinctiveness of these geckos. Thus, in spite of Brongersma’s 

belated change of mind, H. margarethae is a unique lineage 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 specimens 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 
3 4 • smithsonian contributions to zoology

caeca; all males have pigmented testes-epididymides peritoneum, all females except one with pigmented oviducal 

peritoneum. Were my identifications driven by knowledge 

of sex? The evidence definitely suggests a bias. A reexamination 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 Borneo). 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 individuals 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: speculations. This study was not designed to address phylogenetic 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 Hemiphyllodactylus 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. Similarly, this genus has been absent from most other gekkonid 

phylogenetic studies, although the broad taxon-based studies of A. Bauer and colleagues likely will provide information 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 contains 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 yunnanensis species group contains four, and likely more, species: 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 biogeography of the bisexual populations, I suggest that the elongate clade derived from the more robust clade and that its 

origin occurred in northern precursor-Sunda with dispersal eastward into the Philippines and hence into the western 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. Presently, the situation for the Sri Lankan bisexual population 

is undecipherable owing to paucity of data. The robustbodied Hemiphyllodactylus has remained and differentiated within the area of the group’s origin and northward 

in southwestern China.

I suggested previously (Morphological Differentiation 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 unisexual H. typus in contrast to the displacement of parental 

bisexuals by clones of the unisexual Lepidodactylus lugubris (Ineich, 1999). I postulate the possibility of onboard 

ship hybridization then colonization and population expansion in central Pacific. While such an origin could have 

occurred during the initial human migration into the Pacific, onboard hybridization seems more likely aboard the 
n u m 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 notorious 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 transfer of hybrids to diverse islands and to other whalers to 

broaden the dispersal of the hybrids.

Other phylogenetic and biogeographic questions remain 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 Hemiphyllodactylus clade displays several distributional anomalies. 

The widespread occurrence of unisexual H. typus contrasts 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 eastern 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 situation derives from our poor knowledge of tropical Asian 

diversity, particularly among mainland taxa, and this poor 

knowledge results from the continuing recognition of panAsian species, when few such species exist (Stuart et al., 

2006; Zug, In press).

Species Accounts

Hemiphyllodactylus typus Bleeker

Indo-Pacific slender gecko

Hemiphyllodactylus typus Bleeker, 1860:327 [type locality: “Agam” (Sumatra); holotype, BMNH 1946.8.30.83].

Platydactylus crepuscularis Bavay, 1869:8 [type locality: “Nouvelle-Calédonie” (locality implied from title of publication); holotype lost (Brygoo, 

1990:49)].

Spathodactylus mutilatus Günther, 1872:594 [type locality: “East-Indies archipelago”; holotype, BMNH 1946.8.30.83].

Lepidodactylus ceylonensis Boulenger, 1885:164, Pl. XIII, fig. 3 [type locality: “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 specimen 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 platelike. Cloacal spurs present, modest sized, 1–5 CloacS (2 ± 

0.8). Larger scales on lips and snout, rostral largest, rectangular to pentagonal, often slightly concave on dorsomedial edge with slight cleft; 1–5 CircNa (3 ± 0.6), 1–5 SnS 

(2 ± 0.7); labial scales enlarged from rostral to below eye, 
3 6 • smithsonian contributions 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, 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 lyreshaped 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 lamellae (4-5 fore, 5-6 hind) ventrally, terminal phalanx not free 

with or without minute claw. Adults usually with precloacal 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, dorsolaterally 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 precloacal 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 caecum, 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 humanmade structures. Slender refers to the elongate and attenuate appearance of H. typus and most of its congeners.

Variation. The means or medians and ranges 

are detailed in the preceding Description section. No significant or striking variation was seen in any of the characters 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-gathering variance arising from the poor preservation quality 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 hybridization 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 u m b e r 6 3 1 • 3 7

population. The lightness of Auffenberg’s Komodo specimens is attributed to chance.

Distribution. Broadly, if somewhat spottily, 

distributed from Hawaii and French Polynesia in the central 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 aurantiacus Beddome

Southern Ghats slender gecko

Hemidactylus aurantiacus Beddome, 1870:33 [Type locality: “Shevaroys, under 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 ambiguous 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 immature, sex uncertain (no precloacal pores). The syntypic 

series (BMNH 74.429.1332–1337) consists of nine individuals, 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 reluctance to use an immature specimen as a type and the 

absence of an immature specimen matching Beddome’s 

dimensions. Additionally, I accept Bauer and Günther’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 compressed, 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, rostral largest, rectangular to pentagonal, often slightly concave 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 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 lyre-shaped lamellae (scansors); modal digital 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) ventrally, 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 subsequently 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 females; chin scales bordering mental and first infralabial not 

greatly enlarged; digital lamellae formulae 2-2-2-2 (forefoot) 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 double bar of dark brown and light orange (Figure 15).
3 8 • smithsonian contributions 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 confirmed; open square = locality from published records or museum records but specimen not examined.
n u m 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 adjacent 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 evident, 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 difference in average size is slight and not statistically significant. 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 secreting 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.)
4 0 • smithsonian contributions 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 represent a recent accidental transport. (Distributional summary largely from Bauer and Das, 1999.)

Hemiphyllodactylus ganoklonis Zug,

new species

Palauan slender gecko

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 removed 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, 

Babeldaob Island, east of Nekkeng, 0.5 km west of Tabecheding 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 turnoff 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 village on road to Galap village (Ngaraard State), collected 

by Ronald I. Crombie and Gregory K. Pregill, 14 January 1995; USNM 563666, Babeldaob Island, south of 

Ulimang village (Ngaraard State), collected by Ronald I. 

Crombie, 6 April 1995; USNM 563667, Ngeanges Island, 7°12′26′′N 134°22′21′′E, collected by Ronald I. 

Crombie, 9 January 1998; USNM 563668, Ngeaur Island, Ngaramasch village, collected by Ronald I. Crombie, 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 vouchers 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), specimens and records tentatively assigned to H. yunnanensis.
n u m 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 preceding, except 13 August 2001; USNM 563674 same as 

preceding, except 15 August 2001; USNM 563675 Ngerekebesang 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 Oimaderuul 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 compressed, 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 dimorphic, 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, becoming progressively smaller in subocular rictus, 8–11 Suplab 

FIGURE 17. Hemiphyllodactylus ganoklonis from Ulebsechel Island, Palau (USNM 563680). (Illustration by M. D. Griffin.)
4 2 • smithsonian contributions 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 triangular apical lamella bordered proximally by lyre-shaped 

lamellae (scansors); modal digital formulae 3-4-4-3 (forefoot) and 3-4-4-4 (hindfoot) for scansors; first digit of foreand 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 distinctly 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 formulae 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 border 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. Proportions: 53% TrunkL/SVL, 22% HeadL/SVL, 12% HeadW/

SVL, 57% HeadW/HeadL, 38% SnEye/HeadL, 30% NarEye/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 Island, Palau. (Photograph by G. Zug.)
n u m 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 medium 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 middorsally 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. Hemiphyllodactylus ganoklonis is the smallest Hemiphyllodactylus 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 reflects the high quality of preservation of the Palauan sample 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 harterti Werner

Bintang slender gecko

Lepidodactylus Harterti Werner, 1900:196 [type locality: “Malakka” (Malaysia), 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 origin, but it appears that Boulenger retained them because 

the catalog number (1901.3.20.2) of G. larutensis indicates 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 ganoklonis in Palau Islands. Circles indicate specimens represented by museum vouchers. (Image modified from map by D. Dalet.)
4 4 • smithsonian contributions 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–femoral 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 insufficiency 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 Titiwangsa 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 NarEye, 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 postsacral 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 dorsolateral 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 NarEye, 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/NarEye, 32% SnW/HeadW. Scalation: 3 CircNa, 2 SnS, 10 Suplab, 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 laterally, somewhat lighter ventrally. Evidence of a dark postsacral mark.

Etymology. Werner (1900) noted that a 

single specimen (holotype) of this gecko was collected in 

Malakka by Hartert and deposited in the Berlin collection. Presumably, the Hartert referred to by Werner is Ernest 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 employment at Tring, Hartert visited Asia and elsewhere and collected 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 insularis Taylor

Philippine slender gecko

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 u m 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, subcaudal 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 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 

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 minute 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 dorsally and laterally from head to tail; top of head with scattering 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. titiwangsaensis, and bisexual specimens from Borneo. Symbols: circle, H. harterti; diamond, H. titiwangsaensis; solid square, H. margarethae; open square, bisexuals.
4 6 • smithsonian contributions 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 inguina, 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 markings as described for preserved individuals. Light dorsolateral 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); average 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 Inflab, 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 femoral series separated, digital formulae 3-3-3-3 (forefoot) and 

3-4-4-4 (hindfoot). Pigmented caecum, pigmentation unknown 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. (Photographs by M. McNaugher.)
n u m 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 enlargement of those touching the mental and first supralabial. 

There are six digital formulae each for forefoot and hindfoot. 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 latter island groups recommends a reexamination of the bisexual Bornean Hemiphyllodactylus.

Hemiphyllodactylus margarethae Brongersma

Sumatran slender gecko

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 Hemiphyllodactylus 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 dimorphic: 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 proportions: 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% NarEye/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 platelike. Cloacal spurs present, modest sized, 1–2 CloacS (2 ± 

0.5). Larger scales on lips and snout, rostral largest, rectangular to pentagonal, often slightly concave on dorsomedial 

FIGURE 22. Geographic occurrence of Hemiphyllodactylus insularis 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.
4 8 • smithsonian contributions 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, becoming 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 triangular 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 lamellae (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 preserved 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 (forefoot) 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 NarEye, 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 presumably 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 margarethae 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 mountain 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, new species

Titiwangsan slender gecko

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), collected 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, Cameron 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 Brongersma, 1931: left, ZMA 11095 holotype, male; right, 11096 paratype, female. (Photograph by G. Zug.)
n u m 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 females: 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 (Gekkoninae) 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, subcaudal 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); labial 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 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 triangular apical 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 hindfeet 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 occasionally 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 interspaces 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. (Coloration 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; precloacal–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. Proportions: 42% TrunkL/SVL, 23% HeadL/SVL, 17% HeadW/

SVL, 73% HeadW/HeadL, 41% SnEye/HeadL, 28% NarEye/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 enlarged), 16 Dorsal, 7 Ventral, 3 CloacS, Subcaud not enlarged, 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 ventrally. Postsacral mark indistinct, small median dark spot 

on first tail segment, anterior arms muted. In life, the ventral surface of the tails (type series) were orangish pink.

Etymology. These geckos occur in the south 

central region of the Banjaran Titiwangsa; hence the 
5 0 • smithsonian contributions 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 phalangeal 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 individuals 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 pattern (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 localities within Banjaran Titiwangsa (Figure 20) and particularly 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)

Asian slender gecko

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 Pá [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 u m 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” populations of Southeast Asia and Hong Kong remains unresolved 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 à Chapa 

une femelle à queue reconstituée (S130) . . .” Brygoo 

(1990:44) and I interpret this statement as a description 

based on a single female specimen. This interpretation conflicts with Guibe’s (1954) type catalog listing of two specimens; 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 Bourret. 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 Bourret 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, subcaudal scales slightly larger than dorsal caudal scales 

but not plate-like. Cloacal spurs usually present, modest 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 progressively smaller in subocular rictus, 8–13 Suplab (10 ± 1.0), 

8–12 Inflab (10 ± 1.1); 6–11 Chin (8 ± 1.1), those behind mental moderately to 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 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. (Photographs by G. Zug.)
5 2 • smithsonian contributions 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 thereafter 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 rectangular 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 uniform light to medium brown with brown marks.

Major diagnostic features are as follows: bisexual 

taxon; caecum and gonadal ducts not pigmented; precloacal–femoral pore series continuous in males (TotPore 

11–25), usually absent in females; chin scales bordering 

mental and first infralabial distinctly enlarged; digital lamellae 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 NarEye, ~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 formulae (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 Analysis 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 postmentals 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 pigmentation 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 females and males; adult size seldom 

>38 mm SVL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
n u m 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 postsacral 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]; postsacral 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; forefoot digital lamellar formula usually 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; forefoot digital lamellar formula usually 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.: forefoot 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 continuous 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. Pascocello, R. Zweifel (AMNH); R. Sadlier, A. Greer (AMS); C. McCarthy (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. Lenglet, G. Coulon, M. Lang (IRSN); T. Hikida, M. Matsui, H. Ota (KUZ); 

J. Rosado (MCZ); I. Ineich (MNHN); E. Kramer (NMB); F. Tiedemann, 

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. Günther, M.-O. Rödel (ZMB); W. Böhme (ZMFK); 

K. Lim (ZRC); U. Gruber (ZSM). Additionally, I thank the collection staff of 

the National Museum of Natural History, Smithsonian Institution, who regularly 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. Wilson did the initial georeferencing of Hemiphyllodactylus specimens and type localities. B. Trimmer assisted with an early phase of data verification. T. Schwaner, 

performed an electrophoretic analysis of Thai, Philippine, and Hawaiian Hemiphyllodactylus. A. Bauer, H. Ota, and I. Das have kept my interest in mind and 

regularly apprised me of newly collected specimens and observations. R. Crombie made a special effort to obtain specimens from Palau and Hawai’i and provided 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 Hemiphyllodactylus images from India and Borneo, and J. R. H. Gibbons gave me images 
5 6 • smithsonian contributions to zoology

of Fijian H. typus. M.-O. Rödel generously took data and 

photographs of the type of Lepidodactylus Harterti Werner. 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 [né Westhoff] of Luray Zoo and Molly Griffin 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 Hemiphyllodactylus section from V. V. Bobrov and D. V. Semenov’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

Many 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 conciseness and for permitting quick identification of the characters. 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.
5 8 • smithsonian contributions 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

LOCALITY SAMPLES

The specimens are segregated by the geographic-specific samples. These 

samples are arranged from east to west and, for those within similar longitudinal 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; Tongatapu: CAS 49971, USNM 268044, 322120; Vava’u: USNM 333617; TongaAta: 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.
6 0 • smithsonian contributions to zoology

Taiwan and Japan [Taiwan]. Ryukyu Islands: 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]. Babeldaob: 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 number (1), 11095, 11096.

Hemiphyllodactylus titiwangsaensis

Islands of Indonesia and Malaysia [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 u m 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 [species indeterminate]

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.

TYPE SPECIMENS

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 Günther: 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.

ADDITIONAL LOCALITY RECORDS

The following localities derive from distributional records appearing in publications and from museum specimen 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 Réunion: Déso et 

al. (2007); Rodriques: Schröder and Röll (2004).

Nicobar Islands. Great Nicobar Island: 

Biswas and Sanyal (1980).

Sumatra. Pulau Enggano: MVZ 39345–

39346, 239586.

Thailand. Kanchanaburi, Nakhon Ratchasima, Narathiwat, Phang-Nga, Phuket, Ranong, Trang, 

and Trat provinces: Pauwels and Sumontha (2007). 

Vietnam. Southernmost mapped locality: Bobrov 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: McKeown (1996).

Marshall Islands. Enewetak: R. I. Crombie (unpublished manuscript, “Pacific amphibian and reptile 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); 
6 2 • smithsonian contributions to zoology

Bangalore: Daniels (1994); Andhra Pradesh, Visakhapatnam District, Araku Valley and Madhygulu: Sanyal et al. 

(1993).

Hemiphyllodactylus yunnanensis

China. Yunnan–Changyuan, Chengjiang, 

Chuxiong, Gejiu, Jinping, Lijang, Longling,Yao’an; Guizhou–Anlong, Dushan, Huishui, Xingyi; Guangxi–Dayaoshan: 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, Megong R.: Bezuijen et al. (2009).

Thailand. Khao Yai National Park: Chan-ard 

et al. (1999).

Hemiphyllodactylus [species indeterminate]

(These localities not mapped)

Sri Lanka. Southwestern Sri Lanka: Somaweera and Somaweera (2009).
Appendix 3: Statistical Analyses

I used standard univariate statistics to summarize variation of the characters 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, significance 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 between 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 demonstrated 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 
6 4 • smithsonian contributions to zoology

36.5%, 31.0%, and 17.6% of the total variance, respectively. 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 eigenvalues of 24.72, 1.064, and 0.156, respectively, with the 

first component accounting for 94.6% and the second component 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 (correlation matrix) results for adult females (n = 72) are summarized 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 Sumatran 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 unadjusted 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 examined in females.

Discriminant function analysis examination of Malaysian Hemiphyllodactylus samples: The distinctiveness 

of the H. harterti sample (n = 2 adults) and a central mountain 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 unadjusted classification matrix yielded 100% accuracy of group 

assignment and the jackknifed matrix only 76% total accuracy, 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 eigenvalue (13.241) reported.
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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
7 0 • smithsonian contributions to zoology

Lepidodactylus, 5, 34–35

aurantiacus, 2

ceylonensis, 2, 4, 6, 12, 35, 61

crepuscularis, 2

Harterti, 4, 6, 16, 25, 28–29, 33, 43, 

61

lugubris, 2, 4, 9, 28–30, 33–35, 50

Platydactylus,

crepuscularis, 2, 6, 35

minutus, 5

Pytodactylus, 

gracilis, 1, 2

Spathodactylus, 2

mutilatus, 2, 6, 35, 61

Spathoscalabotes, 2
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