A NEW SPECIES OF BENT-TOE GECKO (GEKKONIDAE: CYRTODACTYLUS

A NEW SPECIES OF BENT-TOE GECKO (GEKKONIDAE:CYRTODACTYLUS) FROM SULAWESI ISLAND, EASTERN INDONESIA

CHARLES W. LINKEM1,6, JIMMY A. MCGUIRE

2, CHRISTOPHER J. HAYDEN3, MOHAMMED IQBAL

SETIADI4, DAVID P. BICKFORD

5, AND RAFE M. BROWN1

1Department of Ecology and Evolutionary Biology and Natural History Museum and Biodiversity Institute, University ofKansas, Dyche Hall, 1345 Jayhawk Blvd, Lawrence, KS 66045, USA

2Museum of Vertebrate Zoology, University of California Berkeley, Berkeley, CA 94720, USA3Museum of Natural Science, 119 Foster Hall, Louisiana State University, Baton Rouge, LA, 70803, USA

4McMaster University, 1280 Main St., West Hamilton, Ontario, L8S 4K1, Canada5Conservation Ecology Laboratory, National University of Singapore, Block S2 14 Science Drive 4, Singapore 117543

ABSTRACT: A new species of Cyrtodactylus is described from Lore-Lindu National Park, Sulawesi Island,Indonesia. It is distinguished from all other Cyrtodactylus by a unique suite of scalation characters and adistinctive color pattern. The new species is the fourth Cyrtodactylus known from the island of Sulawesi andone of two new species found in 2004. These recent discoveries suggest that the diversity of the herpetofaunain Wallacea, a poorly studied biological ‘‘hotspot,’’ may be far richer than previously thought.

Key words: Cyrtodactylus; Gekkonidae; Indonesia; Lore-Lindu National Park; New species; SoutheastAsia; Squamata; Sulawesi

THE GENUS Cyrtodactylus contains 95 de-scribed species distributed throughout theIndo-Australian Archipelago westward to In-dia (Bauer and Henle, 1994). Although manyspecies recently have been reassigned to othergenera such as Tenuidactylus, Cyrtopodion,Nactus, and Geckoella (Golubev and Szczer-bak, 1985; Kluge, 1983, 1991, 1993, 2001;Macey et al., 2000; Szczerbak and Golubev,1984, 1986), the number of species currentlyor formerly in this genus continues to grow.New species have been recently describedfrom Myanmar (Bauer, 2002, 2003), Sri Lanka(Batuwita and Bahir, 2005), Malaysia (Gris-mer, 2005; Grismer and Leong, 2005; You-mans and Grismer, 2006), Thailand (Bauer etal., 2002, 2003; Pauwels et al., 2004), Vietnam(Heidrich et al., 2007; Orlov et al., 2007;Quang et al., 2007; Ziegler et al., 2002), andsouthern Laos (David et al., 2004).

For the island of Sulawesi, Boulenger(1897) and de Rooij (1915) listed three speciesof Cyrtodactylus: C. fumosus, C. jellesmae,and C. marmoratus. Based on overlap in porecharacters, Brongersma (1934) synonomizedC. fumosus with C. marmoratus therebyreducing the number of species on the islandto two. Hayden et al. (2008) recently de-scribed a third species of Cyrtodactylus from

the southwestern peninsula of Sulawesi.Herein we describe a fourth species ofCyrtodactylus from Sulawesi that differsdramatically from all known congeners.

MATERIALS AND METHODS

A herpetological biotic survey was conduct-ed on Sulawesi between September andDecember 2004. Specimens were tissued,preserved in 10% buffered formalin andtransferred to 70% ethanol approximatelytwo months later. The following measure-ments (after Bauer, 2002) were made onpreserved specimens with dial calipers to thenearest 0.1 mm: snout–vent length (SVL);trunk length (TrunkL); crus length (CrusL);tail length (TailL); tail width (TailW); headlength (HL); head width (HW); head height(HH); ear length (EarL); forearm length(ForeaL); orbit diameter (OrbD); nares toeye distance (NarEye); eye to ear distance(EyeEar); internarial distance (Internar); in-terorbital distance (InterOrb). Bauer’s (2002)snout to eye distance (SnEye) is referred to asrostrum length (RostL) to reflect the pre-ferred definition of rostrum as the portion ofthe head anterior to the orbit. In contrast,snout length (SnL) is defined as the portion ofthe head anterior to the nares.

Sex was determined by gonadal inspectionand scoring of prominent secondary sexual6 CORRESPONDENCE: e-mail, [email protected]

Herpetologica, 64(2), 2008, 224–234

E 2008 by The Herpetologists’ League, Inc.

224

characteristics. We scored the following scalecounts following Grismer (2005): postmentals(and their degree of medial contact); suprala-bials; infralabials; number of longitudinaltubercle rows; number of paravertebral tu-bercles; number of ventral scales; number andtype of subdigital lamellae on fourth toe.Additional scale counts in this manuscript are:number of spines on ventrolateral fold, countof spines between fore- and hind limb alongthe ventrolateral fold; number of caudalannuli, count of annuli down length of tail.

For the recognition of the new species, weadopted the General Lineage Species Con-cept of de Queiroz (1998, 1999) as the naturalextension of the Evolutionary Species Con-cept (Wiley, 1978). Application of lineage-based species concepts to island endemics isstraightforward because of the known historyof isolation of island populations (Brown andGuttman, 2002; Brown and Diesmos, 2002).We consider as new species morphologicallydiagnosable forms for which the hypothesis ofconspecificity can be rejected.

Images of specimens were taken using acopy stand and a manual-focus digital camera.Four images were taken at different focallengths and then combined using the programCombineZE to create one image with fulldepth of field.

RESULTS

Cyrtodactylus spinosus sp. nov.

Holotype.—MZB 7024 (BSI-FS 1694)(Fig. 1, 2, 3), adult male, collected 8 Novem-ber 2004 at 19:35 h, from Indonesia: SulawesiIsland: Sulawesi Tenggah Province: Kabupa-ten Donggala: Kecematan Kulawi: Desa Ma-taue: Lore-Lindu National Park (01.44883 S,119.99483 E) at 696 m. Collected by CJH,RMB, JAM and CWL.

Paratopotypes.—MZB 7025–9 (BSI-FS1408–11) collected 3 November 2004: 7025adult male, 7026–8 adult females, 7028 withtwo eggs. Specimen MZB 7029 collected 6November 2004: adult male. All other data arethe same as for the holotype.

Diagnosis.—Cyrtodactylus spinosus is dis-tinguished from all other Cyrtodactylus spe-cies by the following characters: a row ofspines along ventrolateral body fold; six lateralrows of small, unkeeled body tubercles, withmost ventral row intermixed with spines; twospines on temporal region of head; 31 spine-adorned annuli encircling original tail; tuber-cles on fore- and hind limbs; spines onpostantefemoral portion of hind limb. Addi-tional characters distinguishing this speciesinclude: proximal subdigital lamellae trans-versely expanded; 19–21 subdigital lamellae

FIG. 1.—Photo of holotype, MZB 7024, showing pattern and tail curling behavior.

June 2008] HERPETOLOGICA 225

on toe IV; 38–44 mid body ventral scales; mostscales in femoral region small, granular; 7–12enlarged femoral series scales lacking pores;presence of pre-cloacal groove in males(absent in females); presence of pre-cloacalpores (12–13) in a chevron-shaped groove;subcaudals not transversely expanded; threechevron-shaped dark bands on a grayish-brown background.

Description of the holotype.—Adult maleSVL 70.9 mm. Head moderately long (HL/SVL 0.3), wide (HW/HL 0.6), somewhatdepressed (HH/HL 0.3), distinct from neck,and spade-shaped in dorsal profile; loresweakly raised, prefrontal region concave,canthus rostralis rounded and granular; ros-trum short (RostL/HL 0.4) and narrow indorsal profile. Eye large (OrbD/HL 0.3); pupilvertical with crenulated margin. Ear opening

tear-shaped, small (EarL/HL 0.05); eye-to-eardistance slightly greater than diameter of eye.Rostral scale rectangular, width twice height,partially divided dorsally, bordered posteriorlyby large left and right supranasals and twosmall internasals; external nares borderedanteriorly by rostral, dorsally by large super-nasal, posteriorly by three small postnasals,and ventrally by the first supralabial. Supra-labials square, 11/12 extending to center ofeye, first supralabial larger than remainder.Infralabials 11/11 extending to posterior oforbit; first five scales of series largest. Scales ofrostrum, lores, top of head, and occiput smalland granular; scales of occiput and top of headwith infrequent, large tubercles; spines pre-sent on temporals and gular regions. Dorsaland ventral supraciliaries circular; mentaltriangular, bordered laterally by infralabials

FIG. 2.—Illustration of holotype features. (A) Dorsal view of head showing temporal spines, (B) lateral view of head,(C) ventral view of head showing complete separation of post-mentals, variable within the species. (D) Palmar surface ofright pes showing differential size between proximal and distal subdigital lamellae.

226 HERPETOLOGICA [Vol. 64, No. 2

and posteriorly by left and right squarepostmentals; postmentals not in contact,separated by large gular scale. One slightlyenlarged and elongate row of sublabialsextending posteriorly to the third infralabial;gular scales small and granular, gradingposteriorly into slightly larger, flatter, imbri-cate pectoral and ventral scales (Fig. 2).

Body relatively elongate (TL/SVL 0.4) withmoderate ventrolateral folds that containlarge, semiregular, spines; dorsal scales smalland granular interspersed with moderate-sized, triangular, semiregularly arrangedkeeled tubercles; tubercles extending fromocciput to anterior portion of tail; tubercles onocciput irregularly spaced, those on nape andanterior of body largest; approximately sixlongitudinal rows of tubercles; 40 flat, imbri-cate ventral scales between ventrolateral bodyfolds, ventral scales larger than dorsals; tworows of enlarged pre-cloacal scales borderingpre-cloacal groove; 12 pre-cloacal pores oc-curring within groove.

Forelimbs slender, relatively short (ForeL/SVL 0.2); granular scales of forearm slightlylarger than those of body; tubercles present;scales on palmar surface slightly elevated;digits well developed, inflected at basalinterphalangeal joints; subdigital lamellaetransversely expanded proximal to joint inflec-tions, digits narrow distal to joints; claws welldeveloped, sheathed by dorsal and ventralscales; relative lengths of fingers: IV . III .II . V . I.

Hind limbs more robust than forelimbs,moderately long (CrusL/SVL 0.2), covereddorsally with flat, granular scales interspersedwith larger, raised tubercles; ventral scales offemora oval and larger than dorsals; ventraltibial scales granular, imbricate; nine enlargedscales in pore-bearing femoral series, lackingpits; scales on plantar surface oval, imbricate,elevated; toes well developed, inflected atbasal interphalangeal joints; subdigital lamel-lae transversely expanded proximal to inflect-ed joints, digits narrow distal to joints; nineexpanded subdigital lamellae, 12 unexpandedsubdigital lamellae on right toe IV; claws welldeveloped, sheathed by dorsal and ventralscales. Relative lengths of toes: IV . V 5 III. II . I (Fig. 2D).

Tail 90 mm long, complete, 4.3 mm inwidth at base, tapering to a point; dorsalsand caudals granular; spines oriented withrecurved points facing posteriorly, numberingfour per caudal annulus, situated at theposterior edge of each annulus; two small,oval, smooth, postanal scales on either side oftail base.

Coloration in life.—Dorsal ground color ofhead, neck, trunk, limbs and tail brown withdark mottling. Three chevron-shaped inter-leaved black and tan bands on the dorsalsurface between limbs (Fig. 1); pattern con-tinues down tail, transitioning to stripes; trunklacking a pattern, but having numerous whitespines along ventrolateral body fold; nuchalregion covered with a black triangle, bordereddistally by a tan nuchal stripe extending fromeye to first tan chevron on nape; two tan-whitespines along tan nuchal stripe at apex ofpostocular region; a postocular black patchextends to insertion of forelimb ventral tonuchal stripe; labial region mottled anteriorly,transitioning posteriorly to two white subocu-

FIG. 3.—Pre-cloacal region of the holotype andparatopotype (MZB 7026) showing the sexual dimorphismin pre-cloacal pores. The male holotype (A) has a pre-cloacal groove and pore-bearing scales. The female (B)lacks a pre-cloacal groove and pores.


lar bands separated by a black band; canthusrostralis dominated by black scales; withmottled brown/yellow scales on both sides ofcanthal ridge; postnasal region yellow; supra-cilliary scales banded yellow and dark brown;eye golden-brown with dark brown venation.Limbs uniform brown with darker mottling;digits banded with dark and light; most bodyspines tan-to-white, becoming lightest at theirapices. Caudal spines match overlying band-ing pattern coloration (i.e., dark bands withblack spines); ventral ground color gray withdark mottling; scales flecked with black spotsthroughout ventral surface.

Variation.—The paratopotypes closely re-semble the holotype. All specimens exceptMZB 7029 are adults. There is sexualdimorphism in pre-cloacal pore structure.Males posses large, pore-bearing scales in a

chevron shape with a large groove; femalesposses differentiated scales in the pore-bearing series (arranged in a chevron shape)but without a groove and lacking pores(Fig. 3). The numbers of differentiated scalesin the pore-bearing series do not differbetween the sexes. Specimen MZB 7028 hastwo large eggs. Coloration is similar among allspecimens, with each possessing three chev-rons across the back. Chevrons on MZB 7029are less complete than others. Meristicvariation in the type series is presented inTable 1.

Natural History.—Our specimens werefound in secondary-growth forest (neighbor-ing selectively logged first growth forest) ofLore Lindu National Park. The first speci-mens were found at night on shrubs (#1.5 mabove the ground), although we searched in

TABLE 1.—Meristic characteristics of the holotype and five paratopotypes. Abbreviations of measurements are the sameas in methods. Indications of character states in the table are presence of a character state (1), and absence of a character

state (0).

MZB

MeanStandarddeviation7024 7025 7026 7027 7028 7029

Sex Male Male Female Female Female Male — —SVL 70.9 70.0 79.2 78.3 83.2 58.4 73.3 8.9Postmentals 2 2 2 2 2 2 — —Degree of postmental contact 0 25% 25% 0 25% 25% — —Supralabials 11 9 10 8 9 9 — —Infralabials 11 8 9 8 8 7 — —Longitudinal rows of tubercles 8 8 8 8 8 8 — —Ventrolateral spines 9 12 12 11 11 11 — —Paravertebral tubercles 26 25 30 25 25 26 — —Ventral scales 40 44 40 38 41 42 — —Expanded lamellae on 4th toe 9 7 8 8 8 7 — —Narrow lamellae on 4th toe 12 12 12 13 13 12 — —Enlarged pre-cloacal scale patch 1 1 1 1 1 1 — —Pre-cloacal groove 1 1 0 0 0 1 — —Pre-cloacal pores 1 1 0 0 0 1 — —Number of pre-cloacal pores 12 13 13 12 12 13 — —Femoral pore-like scales 9 12 12 7 9 9 — —TrunkL 30.7 31.1 34.9 34.4 40.9 27.5 33.3 5.0CrusL 14.0 11.2 13.2 13.9 14.6 10.2 12.9 1.7TailL 90.1 58.7 95.8 79.6 NA NA 54.0 8.2TailW 4.4 5.1 5.3 4.7 4.3 NA 4.0 0.5HL 22.1 20.8 23.2 23.1 23.6 18.2 21.8 2.2HW 14.3 13.4 15.4 16.4 16.1 11.5 14.5 2.0HH 8.6 8.2 9.1 9.8 9.6 6.8 8.7 1.2EarL 1.1 0.9 1.0 1.4 1.5 1.4 1.2 0.2ForeaL 10.4 9.5 11.5 11.4 11.7 8.6 10.5 1.3OrbD 6.0 4.8 6.4 5.9 5.9 4.3 5.6 0.8NarEye 7.0 6.7 7.5 8.2 7.7 5.8 7.2 0.9RostL 8.5 9.0 9.5 9.7 10.1 7.5 9.1 1.1EyeEar 5.7 5.5 7.2 7.1 7.3 4.8 6.3 1.1Internar 2.4 2.5 2.9 2.7 2.9 2.2 2.6 0.3Interorb 7.7 7.6 9.1 8.9 9.2 6.2 8.1 1.3


the same habitat for several more nights anddid not encounter any other lizards. Fournights later, one individual was found on thetrunk of a tree at a height of 4–5 m. Althoughthe new species appears to be primarilyarboreal (possibly a canopy specialist thatwas driven to lower forest strata by heavyrains prior to collection), limited samplingprecludes confident inference of the species’preferred microhabitat.

Distribution.—This species is only knownto occur at the type locality of Lore LinduNational Park, Kecematan Kulawi, KabupatenDonggala, Sulawesi Tenggah Province, Sula-wesi Island, Indonesia (Fig. 4). It is possiblethat this species occurs outside of this area,but not likely given the low abundance ofquality forest in Sulawesi. This species is likelyan island endemic and should be consideredthreatened due to its limited range.

Etymology.—The name is derived from theLatin word spina which means ‘‘spiny’’ or‘‘thorny’’ in reference to the unique spinespossessed by this species.

Comparison to coastal Sunda Shelf, Philip-pine, Indonesian and Papuan species.—Cy-rtodactylus spinosus can be distinguishedfrom all other congeners by the followingsuite of characteristics: the presence of spinesalong the ventrolateral fold, temporal region,tail and postantefemoral portion of hind limbs

distinguish it from all other species; amaximum SVL of 83 mm distinguishes it fromthe larger species: C. aurensis, C. consobri-nus, C. darmandvillei, C. derongo, C. irian-jayensis, C. lateralis, C. louisiadensis, C.mimikianus, C. novaeguineae, C. pequensis,C. pulchellus, and C. tiomanensis; a minimumSVL of 70 mm distinguishes C. spinosus fromthe smaller species: C. annulatus, C. elok, C.laevigatus, C. papuensis and C. semenanjun-gensis; the presence of pre-cloacal poresdistinguishes C. spinosus from C. darmand-villei, C. jellesmae, C. laevigatus, C. semenan-jungensis, C. sermowaiensis and C. thira-khupti; the presence of a pre-cloacal groovedistinguishes it from all species except C.annulatus, C. aurensis, C. cavernicolous, C.marmoratus, C. papuensis, C. philippinicus,C. pubisulcus, C. pulchellus, C. redimiculus,C. semenanjungensis and C. tiomanensis;absence of a continuous series of pre-cloacaland femoral pores distinguishes it from: C.angularis, C. chanhomeae, C. jarunjini, C.loriae, C. louisiadensis, C. novaeguinea, C.pulchellus, C. seribuatensis, C. thiakhupti;absence of enlarged subcaudal scales distin-guishes it from: C. angularis, C. aurensis, C.baluensis, C. chanhomeae, C. consobrinus, C.d’armandvillei, C. derongo, C. ingeri, C.intermedius, C. irianjayensis, C. jarunjini, C.louisiadensis, C. malayanus, C. mimikianus,C. peguensis, C. pulchellus, C. redimiculus, C.sumonthai and C. thirakhupti; the presence ofbroad subdigital lamellae distinguish it from:C. tigroides and C. yoshii; the presence ofmore than 17 subdigital lamellae distinguishesit from: C. angularis; the presence of fewerthan 22 subdigital lamellae distinguishes itfrom: C. cavernicolous, C. consobrinus, C.derongo, C. ingeri, C. irianjayensis, C. loui-siadensis, C. marmoratus, C. matsuii, C.mimikianus, C. noveaguinea, C. philippinicus,C. sermowaiensis, and C. yoshii; and thepresence of enlarged scales in the pore-bearing femoral series distinguishes it from:C. angularis, C. annulatus, C. aurensis, C.cavernicolous, C. elok, C. ingeri, C. interme-dius, C. irianjayensis, C. jellesmae, C. laevi-gatus, C. lateralis, C. malayanus, C. matsuii,C. papuensis, C. peguensis, C. philippinicus,C. pubisulcus, C. quadrivirgatus, C. semenan-jungensis, C. sermowaiensis, C. sumonthai, C.

FIG. 4.—Map of Sulawesi Island showing the typelocality of Cyrtodactylus spinosus.


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00

00

00

00

01

0la

evig

atus

K43

0?

?0

030

01

140

00

00

00

00

10

late

ralis

L85

11

11

60–6

40

121

,22

??

00

113

00

01

0lo

riae

8H?

11

11

144

–49

01

20–2

41

153

–70

01

53–7

01

00

10

loui

siad

ensi

s6

110–

122

11

11

130

–38

11

23–3

11

136

–64

01

36–6

41

01

00

mal

ayan

usC

70–7

31

11

11

58–6

21

121

–23

01

00

18–

100

11

00

mar

mor

atus

E76

11

11

040

–50

01

23,

241

13–

101

112

–16

00

01

0m

atsu

iiC

105

11

11

151

01

221

00

00

70

10

10

mim

ikia

nus

H95

11

11

034

–48

11

22–2

8?

110

–12

01

7–14

00

10

0no

vaeg

uine

aeJ

115–

129

11

11

135

–46

01

28–3

31

138

–42

01

38–4

21

01

00

papu

ensi

sJ

61–6

51

11

11

??

1?

11

01

18,

90

01

10

pequ

ensi

sC

851

11

11

29–3

81

116

–18

?0

00

17–

90

11

00

phili

ppin

icus

74–9

21

11

11

36–4

20

122

–24

00

01

18–

100

01

00

pubi

sulc

usC

59–7

41

01

11

43–5

50

117

–22

00

01

17–

90

00

10


sworderi, C. thirakhupti, C. tiomanensis andC. yoshii (Table 2).

DISCUSSION

We consider herpetological diversity on theisland of Sulawesi to be underestimated owingto a lack of attention by taxonomists andcomprehensive survey work (Brown et al.,2000; Evans et al., 2003a,b; Iskandar and Tjan,1996; McGuire et al., 2007). Recent bioticsurvey and inventor efforts have found newspecies on Sulawesi and adjacent islandsincluding two new species of flying lizard,genus Draco (McGuire et al., 2007) andadditional species that are in need of descrip-tion. Genetic analyses of some groups: Lim-nonectes (Evans et al., 2003a), Bufo celebensis(Evans et al., 2003b) and Lamprolepis (un-published data) have revealed cryptic lineag-es. Exploration of the northern, central andsouthwestern regions of the island haveresulted in the discovery of two new speciesof Cyrtodactylus—a new species described byHayden et al. (2008) and C. spinosus. Wesuspect that additional species of Cyrtodacty-lus await discovery and several taxa currentlymasquerade under the widespread species C.jellesmae.

Among SE Asian members of the genus,Cyrtodactylus spinosus is morphologicallydistinct, owing to the presence of spines,unique scalation, and distinct color pattern; itis consequently unlikely to be confused withany other species. Indeed, this species is sounique we cannot speculate on its closestrelatives within the genus.

Discovery of this new species within Lore-Lindu National Park further emphasizes theneed for conservation efforts targeting thismajor center of SE Asian biodiversity. Despitethe park’s having been designated as aprotected area, farmers in the area continueto encroach the park’s boundary, using slash-and-burn methods to clear land for agricul-ture. Both small-scale timber poaching andlarge-scale logging operations continue todegrade the park, especially along its un-guarded southern boundary (Bickford et al.,2007). Improved security of the park bound-ary is needed to protect this stronghold ofbiodiversity that likely houses a large diversityof endemic species yet to be discovered.

Tax

onn

SV

L1

23

45

67

89

1011

1213

1415

1617

1819

20

pulc

hellu

sC

115

11

11

133

–35

11

19,

201

114

–18

11

6–8

10

10

0qu

adri

virg

atus

C51

–71

11

11

134

–42

01

19,

200

10

01

0–4

00

00

1re

dim

icul

us78

??

??

??

1?

20–2

4?

?8–

91

15–

8?

??

??1

sem

enan

jung

ensi

sC

59–6

91

11

1?

48–5

30

117

–21

10

01

10

00

11

0se

ribu

aten

sis

C75

11

11

128

–39

01

19–2

21

142

–45

01

42–4

51

00

10

serm

owai

ensi

s68

–93.

51

11

10

30–3

80

122

00

00

00

00

01

0su

mon

thai

M71

11

11

033

–36

11

18?

00

01

20

01

00

swor

deri

N63

–77

11

11

42–4

80

120

11

00

15,

60

00

10

thir

akhu

pti

72–8

01

11

10

37–4

01

120

,21

11

00

10

11

10

0ti

groi

des

F83

11

1?

134

00

19–2

3?

05–

70

18–

90

01

00

tiom

anen

sis

O84

11

11

036

–40

01

20–2

21

10

11

3–5

00

10

0yo

shii

C75

–96

11

11

150

–58

00

25–3

00

00

00

8–12

00

01

0

TA

BL

E2.

—C

onti

nu

ed.


Acknowledgments.—We thank N. Andayani (Univ. ofIndonesia; Wildlife Conservation Society), J. Supriatna(Univ. of Indonesia; Conservation International), Mum-puni (Museum Zoologicum Bogoriense, LIPI), and A.Riyanto (Museum Zoologicum Bogoriense, LIPI) for theirassistance in obtaining permits to work in Indonesia, aswell as with other logistical issues. We thank the followinggovernment officials who provided research and exportpermits, as well as specimen loans: A. Budiman (formerlyof LIPI), S. Prijono (former Director of MuseumZoologicum Bogoriense). For assistance and companion-ship in the field, we are grateful to F. Chain, D. Halliwel,A. Rosyid, Shobi, and Ted Townsend (plus others). Fordiscussion and/or comments on the manuscript, we thankL. Trueb. This work was supported by the NationalScience Foundation (DEB-BSI 0328700).

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APPENDIX

Specimens Examined

Codes for institutions are CAS, California Academy ofSciences; KU, University of Kansas Natural HistoryMuseum; LSUMZ, Louisiana State University Museumof Zoology; MVZ, Museum of Vertebrate Zoology,University of California, Berkeley; MZB, Museum Zool-ogicum Bogoriense, Bogor Indonesia; NMPNG, NationalMuseum of Papua New Guinea; UPNG, University ofPapua New Guinea; CCA, Christopher C. Austin; RMB,Rafe M. Brown; JAM, Jimmy A. McGuire. Field collectionnumbers are in parentheses.

Cyrtodactylus agusanensis: PHILIPPINES: MINDANAO:Province of Davao del Norte: 70 km S Bislig: LSUMZ41601, 41602 (males), LSUMZ 41603 (female), LSUMZ41604, 41605 (males); DINAGAT: Province of Surigao delNorte: Municipality of Loreto: KU 305564–5; SAMAR:Province of Eastern Samar: Municipality of Taft: BarangaySan Rafael: KU 305569. Cyrtodactylus annulatus: PHI-LIPPINES: MINDANAO: Province of Davao del Norte:70 km S Bislig: LSUMZ 41606 (female), 41608, 41609(males). Cyrtodactylus baluensis: EAST MALAYSIA:SABAH: Kina Balu Peak: CAS 25626 (male). Cyrtodactyluscavernicolus: EAST MALAYSIA: SARAWAK: Niah Cave:CAS 23726 (male). Cyrtodactylus consobrinus: EASTMALAYSIA: SARAWAK: Bintulu District: Sungei Seran:CAS 105993 (male); Kapit District: Sungai Mengiong:MVZ 11782 (male); BRUNEI: Temburang District: SungaiBelalang: LSUMZ 55833 (male); INDONESIA: PROPINSI

SUMATERA UTARA: Bukit Lawang: Bahorak: MZB 4355(male), 4356 (juvenile). Cyrtodactylus darmandvillei:INDONESIA: PROPINSI NUSA TENGGARA BARAT: PulauLombok.LSUMZ 81732 (JAM 3176) (male). Cyrtodacty-lus derongo: PAPUA NEW GUINEA: WESTERN PROVINCE:Derongo: NMPNG R23452 (female). Cyrtodactylus


jellesmae: INDONESIA: PROPINSI SULAWESI SELATAN:Anabanua: JAM 5628, 5631 (male); Harapan: JAM 5643(female); Takalasi: JAM 5670, 5671, 5678 (male), 5677(female); Maroangin: JAM 5680, 5683, 5684, 5686, 5688,5704 (males), 5681, 5682, 5685, 5687 (females); Enrekang:JAM 5705, 5747, 5749, 5768, 5769, 5771 (males), 5770,5772, 5773 (females); Tapung: JAM 5783, 5784 (females);Pecinong: JAM 5850, 5851 (females), 5852 (juvenile);Mariorilau: JAM 5892, 5895, 5897, 5899 (males), 5893,5896, 5898 (females), 5900 (juvenile); PROPINSI SULAWESI

BARAT: Kabiraan: JAM 6341–6343, 6346 (males), 6339,6340, 6344, 6345, 6347 (females), 6338 (juvenile);PROPINSI SULAWESI TENGGAH: Donggala Kabupaten:LSUMZ 8403 (male), 8400 (female), 8401 (juvenile); PosoKabupaten: LSUMZ 8402 (male); Luwuk Kabupaten:LSUMZ 8405 (male), 8404, 8406 (females). Cyrtodactylusloriae: PAPUA NEW GUINEA: CHIMBU PROVINCE:Karimui: CAS 117964 (male), CAS 118018 (female),NMPNG R23625, R8347, R8348; PAPUA NEW GUIN-EA: UPNG 5687 (male); WESTERN HIGHLANDS PROVINCE:Kaironk: UPNG 0976 (female); MOROBE PROVINCE: Wau.:NMPNG 24730. Cyrtodactylus louisiadensis: PAPUANEW GUINEA: BOUGAINVILLE: Kunua: KU 98481; MILNE

BAY PROVINCE: Halowina: CCA 4096 (female), 4426, 4427,4582, (males), 4428 (juvenile). Cyrtodactylus malayanus:INDONESIA: PROPINSI KALIMANTAN TIMUR: Maruwai:MZB 2927–2929 (males); PROPINSI KALIMANTAN BARAT,

Taman Nasional Bentuang Karimun: MZB 3920 (female).Cyrtodactylus marmoratus: INDONESIA: JAVA: DjurungSriti: Mount Merapi: KU 153796; PROPINSI JAVA BARAT:Sukabumi: LSUMZ 81868 (male), LSUMZ 81869–81870(females), LSUMZ 81871–81872 (males), LSUMZ 81873(female), LSUMZ 81874 (subadult male), LSUMZ 81875(female), LSUMZ 81876 (male). Cyrtodactylus novaegui-neae: PAPUA NEW GUINEA: SANDAUN PROVINCE: Utai:CCA 3142 (female), 3415 (juvenile); Bewani Station: CCA3674 (male). Cyrtodactylus quadrivirgatus cf.: INDO-NESIA: SUMATRA: Propinsi Bengkulu: 46 km East ofBengkulu:.MVZ 239338 (male), 239578 (female). Cyrto-dactylus philippinicus: PHILIPPINES: LUZON: Camar-ines del Sur Province: Municipality of Naga: KU 305571;NEGROS: Negros Oriental Province: Municipality ofDumagete, Baragay Valencia: KU 305572; LUBANG:Occidental Mindoro Province: Municipality of Lubang,Barangay Vigo: KU 303846, 303844. Cyrtodactylussermowaiensis: PAPUA NEW GUINEA: SANDAUN PROV-

INCE: Utai: CCA 3108, 3143, 3407, 3411, 3448–3451, 3506,3507, 3517 (males), 2895, 2911, 3034, 3144, 3188, 3224,3233, 3358, 3359, 3490, 3530 (females); Bewani Station:CCA 3603 (male), 3604 (juvenile); MADANG PROVINCE:Sapi Creek: NMPNG R22712; South Naru: NMPNGR24803–24806; EAST SEPIK PROVINCE: Maprik: NMPNGR22796; SANDAUN PROVINCE: Idam River: UPNG 3914–3916.


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A NEW SPECIES OF BENT-TOE GECKO (GEKKONIDAE: CYRTODACTYLUS