Herpetologica, 64(2), 2008, 224–234 E 2008 by The Herpetologists’ League, Inc.

A NEW SPECIES OF BENT-TOE GECKO (GEKKONIDAE: CYRTODACTYLUS) FROM SULAWESI ISLAND, EASTERN INDONESIA CHARLES W. LINKEM1,6, JIMMY A. MCGUIRE2, CHRISTOPHER J. HAYDEN3, MOHAMMED IQBAL SETIADI4, DAVID P. BICKFORD5, AND RAFE M. BROWN1 1

Department of Ecology and Evolutionary Biology and Natural History Museum and Biodiversity Institute, University of Kansas, Dyche Hall, 1345 Jayhawk Blvd, Lawrence, KS 66045, USA 2 Museum of Vertebrate Zoology, University of California Berkeley, Berkeley, CA 94720, USA 3 Museum of Natural Science, 119 Foster Hall, Louisiana State University, Baton Rouge, LA, 70803, USA 4 McMaster University, 1280 Main St., West Hamilton, Ontario, L8S 4K1, Canada 5 Conservation 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 a distinctive color pattern. The new species is the fourth Cyrtodactylus known from the island of Sulawesi and one of two new species found in 2004. These recent discoveries suggest that the diversity of the herpetofauna in Wallacea, a poorly studied biological ‘‘hotspot,’’ may be far richer than previously thought. Key words: Cyrtodactylus; Gekkonidae; Indonesia; Lore-Lindu National Park; New species; Southeast Asia; Squamata; Sulawesi

THE GENUS Cyrtodactylus contains 95 described species distributed throughout the Indo-Australian Archipelago westward to India (Bauer and Henle, 1994). Although many species recently have been reassigned to other genera such as Tenuidactylus, Cyrtopodion, Nactus, and Geckoella (Golubev and Szczerbak, 1985; Kluge, 1983, 1991, 1993, 2001; Macey et al., 2000; Szczerbak and Golubev, 1984, 1986), the number of species currently or formerly in this genus continues to grow. New species have been recently described from Myanmar (Bauer, 2002, 2003), Sri Lanka (Batuwita and Bahir, 2005), Malaysia (Grismer, 2005; Grismer and Leong, 2005; Youmans and Grismer, 2006), Thailand (Bauer et al., 2002, 2003; Pauwels et al., 2004), Vietnam (Heidrich et al., 2007; Orlov et al., 2007; Quang et al., 2007; Ziegler et al., 2002), and southern Laos (David et al., 2004). For the island of Sulawesi, Boulenger (1897) and de Rooij (1915) listed three species of Cyrtodactylus: C. fumosus, C. jellesmae, and C. marmoratus. Based on overlap in pore characters, Brongersma (1934) synonomized C. fumosus with C. marmoratus thereby reducing the number of species on the island to two. Hayden et al. (2008) recently described a third species of Cyrtodactylus from 6

CORRESPONDENCE: e-mail, [email protected]

the southwestern peninsula of Sulawesi. Herein we describe a fourth species of Cyrtodactylus from Sulawesi that differs dramatically from all known congeners. MATERIALS AND METHODS A herpetological biotic survey was conducted on Sulawesi between September and December 2004. Specimens were tissued, preserved in 10% buffered formalin and transferred to 70% ethanol approximately two months later. The following measurements (after Bauer, 2002) were made on preserved specimens with dial calipers to the nearest 0.1 mm: snout–vent length (SVL); trunk length (TrunkL); crus length (CrusL); tail length (TailL); tail width (TailW); head length (HL); head width (HW); head height (HH); ear length (EarL); forearm length (ForeaL); orbit diameter (OrbD); nares to eye distance (NarEye); eye to ear distance (EyeEar); internarial distance (Internar); interorbital distance (InterOrb). Bauer’s (2002) snout to eye distance (SnEye) is referred to as rostrum length (RostL) to reflect the preferred definition of rostrum as the portion of the head anterior to the orbit. In contrast, snout length (SnL) is defined as the portion of the head anterior to the nares. Sex was determined by gonadal inspection and scoring of prominent secondary sexual 224

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

characteristics. We scored the following scale counts following Grismer (2005): postmentals (and their degree of medial contact); supralabials; infralabials; number of longitudinal tubercle rows; number of paravertebral tubercles; number of ventral scales; number and type of subdigital lamellae on fourth toe. Additional scale counts in this manuscript are: number of spines on ventrolateral fold, count of spines between fore- and hind limb along the ventrolateral fold; number of caudal annuli, count of annuli down length of tail. For the recognition of the new species, we adopted the General Lineage Species Concept of de Queiroz (1998, 1999) as the natural extension of the Evolutionary Species Concept (Wiley, 1978). Application of lineagebased species concepts to island endemics is straightforward because of the known history of isolation of island populations (Brown and Guttman, 2002; Brown and Diesmos, 2002). We consider as new species morphologically diagnosable forms for which the hypothesis of conspecificity can be rejected. Images of specimens were taken using a copy stand and a manual-focus digital camera. Four images were taken at different focal lengths and then combined using the program CombineZE to create one image with full depth of field.

RESULTS Cyrtodactylus spinosus sp. nov. Holotype.—MZB 7024 (BSI-FS 1694) (Fig. 1, 2, 3), adult male, collected 8 November 2004 at 19:35 h, from Indonesia: Sulawesi Island: Sulawesi Tenggah Province: Kabupaten Donggala: Kecematan Kulawi: Desa Mataue: 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-FS 1408–11) collected 3 November 2004: 7025 adult male, 7026–8 adult females, 7028 with two eggs. Specimen MZB 7029 collected 6 November 2004: adult male. All other data are the same as for the holotype. Diagnosis.—Cyrtodactylus spinosus is distinguished from all other Cyrtodactylus species by the following characters: a row of spines along ventrolateral body fold; six lateral rows of small, unkeeled body tubercles, with most ventral row intermixed with spines; two spines on temporal region of head; 31 spineadorned annuli encircling original tail; tubercles on fore- and hind limbs; spines on postantefemoral portion of hind limb. Additional characters distinguishing this species include: proximal subdigital lamellae transversely expanded; 19–21 subdigital lamellae

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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 of right pes showing differential size between proximal and distal subdigital lamellae.

on toe IV; 38–44 mid body ventral scales; most scales in femoral region small, granular; 7–12 enlarged femoral series scales lacking pores; presence of pre-cloacal groove in males (absent in females); presence of pre-cloacal pores (12–13) in a chevron-shaped groove; subcaudals not transversely expanded; three chevron-shaped dark bands on a grayishbrown background. Description of the holotype.—Adult male SVL 70.9 mm. Head moderately long (HL/ SVL 0.3), wide (HW/HL 0.6), somewhat depressed (HH/HL 0.3), distinct from neck, and spade-shaped in dorsal profile; lores weakly raised, prefrontal region concave, canthus rostralis rounded and granular; rostrum short (RostL/HL 0.4) and narrow in dorsal profile. Eye large (OrbD/HL 0.3); pupil vertical with crenulated margin. Ear opening

tear-shaped, small (EarL/HL 0.05); eye-to-ear distance slightly greater than diameter of eye. Rostral scale rectangular, width twice height, partially divided dorsally, bordered posteriorly by large left and right supranasals and two small internasals; external nares bordered anteriorly by rostral, dorsally by large supernasal, posteriorly by three small postnasals, and ventrally by the first supralabial. Supralabials square, 11/12 extending to center of eye, first supralabial larger than remainder. Infralabials 11/11 extending to posterior of orbit; first five scales of series largest. Scales of rostrum, lores, top of head, and occiput small and granular; scales of occiput and top of head with infrequent, large tubercles; spines present on temporals and gular regions. Dorsal and ventral supraciliaries circular; mental triangular, bordered laterally by infralabials

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

and posteriorly by left and right square postmentals; postmentals not in contact, separated by large gular scale. One slightly enlarged and elongate row of sublabials extending posteriorly to the third infralabial; gular scales small and granular, grading posteriorly into slightly larger, flatter, imbricate pectoral and ventral scales (Fig. 2). Body relatively elongate (TL/SVL 0.4) with moderate ventrolateral folds that contain large, semiregular, spines; dorsal scales small and granular interspersed with moderatesized, triangular, semiregularly arranged keeled tubercles; tubercles extending from occiput to anterior portion of tail; tubercles on occiput irregularly spaced, those on nape and anterior of body largest; approximately six longitudinal rows of tubercles; 40 flat, imbricate ventral scales between ventrolateral body folds, ventral scales larger than dorsals; two rows of enlarged pre-cloacal scales bordering pre-cloacal groove; 12 pre-cloacal pores occurring within groove.

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Forelimbs slender, relatively short (ForeL/ SVL 0.2); granular scales of forearm slightly larger than those of body; tubercles present; scales on palmar surface slightly elevated; digits well developed, inflected at basal interphalangeal joints; subdigital lamellae transversely expanded proximal to joint inflections, digits narrow distal to joints; claws well developed, sheathed by dorsal and ventral scales; relative lengths of fingers: IV . III . II . V . I. Hind limbs more robust than forelimbs, moderately long (CrusL/SVL 0.2), covered dorsally with flat, granular scales interspersed with larger, raised tubercles; ventral scales of femora oval and larger than dorsals; ventral tibial scales granular, imbricate; nine enlarged scales in pore-bearing femoral series, lacking pits; scales on plantar surface oval, imbricate, elevated; toes well developed, inflected at basal interphalangeal joints; subdigital lamellae transversely expanded proximal to inflected joints, digits narrow distal to joints; nine expanded subdigital lamellae, 12 unexpanded subdigital lamellae on right toe IV; claws well developed, sheathed by dorsal and ventral scales. Relative lengths of toes: IV . V 5 III . II . I (Fig. 2D). Tail 90 mm long, complete, 4.3 mm in width at base, tapering to a point; dorsals and caudals granular; spines oriented with recurved points facing posteriorly, numbering four per caudal annulus, situated at the posterior edge of each annulus; two small, oval, smooth, postanal scales on either side of tail base. Coloration in life.—Dorsal ground color of head, neck, trunk, limbs and tail brown with dark mottling. Three chevron-shaped interleaved black and tan bands on the dorsal surface between limbs (Fig. 1); pattern continues down tail, transitioning to stripes; trunk lacking a pattern, but having numerous white spines along ventrolateral body fold; nuchal region covered with a black triangle, bordered distally by a tan nuchal stripe extending from eye to first tan chevron on nape; two tan-white spines along tan nuchal stripe at apex of postocular region; a postocular black patch extends to insertion of forelimb ventral to nuchal stripe; labial region mottled anteriorly, transitioning posteriorly to two white subocu-

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TABLE 1.—Meristic characteristics of the holotype and five paratopotypes. Abbreviations of measurements are the same as in methods. Indications of character states in the table are presence of a character state (1), and absence of a character state (0). MZB

Sex SVL Postmentals Degree of postmental contact Supralabials Infralabials Longitudinal rows of tubercles Ventrolateral spines Paravertebral tubercles Ventral scales Expanded lamellae on 4th toe Narrow lamellae on 4th toe Enlarged pre-cloacal scale patch Pre-cloacal groove Pre-cloacal pores Number of pre-cloacal pores Femoral pore-like scales TrunkL CrusL TailL TailW HL HW HH EarL ForeaL OrbD NarEye RostL EyeEar Internar Interorb

7024

7025

7026

7027

7028

7029

Mean

Standard deviation

Male 70.9 2 0 11 11 8 9 26 40 9 12 1 1 1 12 9 30.7 14.0 90.1 4.4 22.1 14.3 8.6 1.1 10.4 6.0 7.0 8.5 5.7 2.4 7.7

Male 70.0 2 25% 9 8 8 12 25 44 7 12 1 1 1 13 12 31.1 11.2 58.7 5.1 20.8 13.4 8.2 0.9 9.5 4.8 6.7 9.0 5.5 2.5 7.6

Female 79.2 2 25% 10 9 8 12 30 40 8 12 1 0 0 13 12 34.9 13.2 95.8 5.3 23.2 15.4 9.1 1.0 11.5 6.4 7.5 9.5 7.2 2.9 9.1

Female 78.3 2 0 8 8 8 11 25 38 8 13 1 0 0 12 7 34.4 13.9 79.6 4.7 23.1 16.4 9.8 1.4 11.4 5.9 8.2 9.7 7.1 2.7 8.9

Female 83.2 2 25% 9 8 8 11 25 41 8 13 1 0 0 12 9 40.9 14.6 NA 4.3 23.6 16.1 9.6 1.5 11.7 5.9 7.7 10.1 7.3 2.9 9.2

Male 58.4 2 25% 9 7 8 11 26 42 7 12 1 1 1 13 9 27.5 10.2 NA NA 18.2 11.5 6.8 1.4 8.6 4.3 5.8 7.5 4.8 2.2 6.2

— 73.3 — — — — — — — — — — — — — — — 33.3 12.9 54.0 4.0 21.8 14.5 8.7 1.2 10.5 5.6 7.2 9.1 6.3 2.6 8.1

— 8.9 — — — — — — — — — — — — — — — 5.0 1.7 8.2 0.5 2.2 2.0 1.2 0.2 1.3 0.8 0.9 1.1 1.1 0.3 1.3

lar bands separated by a black band; canthus rostralis dominated by black scales; with mottled brown/yellow scales on both sides of canthal ridge; postnasal region yellow; supracilliary 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 body spines tan-to-white, becoming lightest at their apices. Caudal spines match overlying banding pattern coloration (i.e., dark bands with black spines); ventral ground color gray with dark mottling; scales flecked with black spots throughout ventral surface. Variation.—The paratopotypes closely resemble the holotype. All specimens except MZB 7029 are adults. There is sexual dimorphism in pre-cloacal pore structure. Males posses large, pore-bearing scales in a

chevron shape with a large groove; females posses differentiated scales in the porebearing series (arranged in a chevron shape) but without a groove and lacking pores (Fig. 3). The numbers of differentiated scales in the pore-bearing series do not differ between the sexes. Specimen MZB 7028 has two large eggs. Coloration is similar among all specimens, with each possessing three chevrons across the back. Chevrons on MZB 7029 are less complete than others. Meristic variation in the type series is presented in Table 1. Natural History.—Our specimens were found in secondary-growth forest (neighboring selectively logged first growth forest) of Lore Lindu National Park. The first specimens were found at night on shrubs (#1.5 m above the ground), although we searched in

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

the same habitat for several more nights and did not encounter any other lizards. Four nights later, one individual was found on the trunk of a tree at a height of 4–5 m. Although the new species appears to be primarily arboreal (possibly a canopy specialist that was driven to lower forest strata by heavy rains prior to collection), limited sampling precludes confident inference of the species’ preferred microhabitat. Distribution.—This species is only known to occur at the type locality of Lore Lindu National Park, Kecematan Kulawi, Kabupaten Donggala, Sulawesi Tenggah Province, Sulawesi Island, Indonesia (Fig. 4). It is possible that this species occurs outside of this area, but not likely given the low abundance of quality forest in Sulawesi. This species is likely an island endemic and should be considered threatened due to its limited range. Etymology.—The name is derived from the Latin word spina which means ‘‘spiny’’ or ‘‘thorny’’ in reference to the unique spines possessed by this species. Comparison to coastal Sunda Shelf, Philippine, Indonesian and Papuan species.—Cyrtodactylus spinosus can be distinguished from all other congeners by the following suite of characteristics: the presence of spines along the ventrolateral fold, temporal region, tail and postantefemoral portion of hind limbs

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distinguish it from all other species; a maximum SVL of 83 mm distinguishes it from the larger species: C. aurensis, C. consobrinus, C. darmandvillei, C. derongo, C. irianjayensis, C. lateralis, C. louisiadensis, C. mimikianus, C. novaeguineae, C. pequensis, C. pulchellus, and C. tiomanensis; a minimum SVL of 70 mm distinguishes C. spinosus from the smaller species: C. annulatus, C. elok, C. laevigatus, C. papuensis and C. semenanjungensis; the presence of pre-cloacal pores distinguishes C. spinosus from C. darmandvillei, C. jellesmae, C. laevigatus, C. semenanjungensis, C. sermowaiensis and C. thirakhupti; the presence of a pre-cloacal groove distinguishes 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-cloacal and 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 distinguishes 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 of broad subdigital lamellae distinguish it from: C. tigroides and C. yoshii; the presence of more than 17 subdigital lamellae distinguishes it from: C. angularis; the presence of fewer than 22 subdigital lamellae distinguishes it from: C. cavernicolous, C. consobrinus, C. derongo, C. ingeri, C. irianjayensis, C. louisiadensis, C. marmoratus, C. matsuii, C. mimikianus, C. noveaguinea, C. philippinicus, C. sermowaiensis, and C. yoshii; and the presence of enlarged scales in the porebearing femoral series distinguishes it from: C. angularis, C. annulatus, C. aurensis, C. cavernicolous, C. elok, C. ingeri, C. intermedius, C. irianjayensis, C. jellesmae, C. laevigatus, C. lateralis, C. malayanus, C. matsuii, C. papuensis, C. peguensis, C. philippinicus, C. pubisulcus, C. quadrivirgatus, C. semenanjungensis, C. sermowaiensis, C. sumonthai, C.

spinosus aaroni agusanensis angularis annulatus aurensis baluensis brevipalmatus chanhomeae cavernicolous consobrinus darmandvillei derongo elok ingeri intermedius irianjayensis irregularis jarujini jellesmae laevigatus lateralis loriae louisiadensis malayanus marmoratus matsuii mimikianus novaeguineae papuensis pequensis philippinicus pubisulcus

Taxon

SVL

70–83 70–86.5 70–103 ? 45–70 92–95 72–86 64–73 79 64–81 97–121 85 105–112 56–68 65–76 57–81 147–163 56–71 70 63–75 43 85 ? 110–122 70–73 76 105 95 115–129 61–65 85 74–92 59–74

1

1 1 1 ? 1 0 1 1 1 1 1 1 1 0 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1

2

1 1 1 ? ? 0 1 1 1 0 1 0 1 1 1 1 1 1 0 1 ? 1 1 1 1 1 1 1 1 1 1 1 0

3

1 1 1 ? ? 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 1 1 1 1 1 1 1 1

4

1 1 1 ? ? 0 1 0 ? 1 1 1 1 0 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1

5

1 1 1 0 1 0 1 1 1 1 1

1 1 0 1 0 0 1 0

1 1 1 ? ? 0 0 1 1 1 1 1

6

40–43 34–40 46–51 40–42 50–60 45–51 40–45 35–45 36–38 51–58 58–65 36–40 46–48 44 40–43 40–50 36–41 41–46 28 32–58 30 60–64 44–49 30–38 58–62 40–50 51 34–48 35–46 ? 29–38 36–42 43–55

7

0 0 0 1 0 1 1 0 1 0 1 1 1 0 1 1 1 0 1 0 0 0 0 1 1 0 0 1 0 ? 1 0 0

8

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

9

18–21 21–28 23–28 15, 16 18–23 18–23 21–23 16–19 21–23 22–26 23–28 17 24–26 18, 19 23–27 21 31–35 19, 20 22 19–25 14 21, 22 20–24 23–31 21–23 23, 24 22 22–28 28–33 ? 16–18 22–24 17–22

0 ? 0 ? ? 0 1 1 ? 0 0 1 1 0 0 1 1 0 1 0 0 ? 1 1 0 1 1 ? 1 1 ? 0 0

10

0 1 1 1 0 0 1 1 0 0 1 ? 1 0 0 1 1 1 1 0 0 ? 1 1 1 1 0 1 1 1 0 0 0

11

12

4–7 11–20 3–14 0 0 0 6–9 6–7 32–34 0 1–6 18, 19 0 0 0 0 0 7–8 19 0 0 0 53–70 36–64 0 3–10 0 10–12 38–42 0 0 0 0

1 0 0 0 1 1 0 0 0 1 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 1 0 1 1

13

1 1 1 1 1 1 1 1 ? 1 1 ? 1 1 1 1 1 1 0 0 0 1 1 1 1 1 0 1 1 1 1 1 1

14

15

8–12 5–8 7–11 3 4–7 7 9–10 9–10 32–34 4 9–10 0 0 8 8 8 7–16 8 19 0 0 13 53–70 36–64 8–10 12–16 7 7–14 38–42 8, 9 7–9 8–10 7–9

0 0 0 1 0 0 0 0 1 0 0 0 ? 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 1 0 0 0 0

16

0 0 0 0 1 1 1 0 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 0 0

17

1 1 0 0 1 1 0 0 1 1 1 0 0 0 0 0 1 1 1 0 0 0 0 1 1 0 0 1 1 1 1 1 0

18

0 0 1 1 1 0 1 1 0 0 0 1 1 1 1 1 0 0 0 1 1 1 1 0 0 1 1 0 0 1 0 0 1

19

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

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C

56 K L 8H 6 C E C H J J C

5 A 9B C 4C D 1D E F 1C 1 1G 1H I C C J C

n

TABLE 2.—Diagnosis table of characters used to distinguish Cyrtodactylus spinosus from other Cyrtodactylus species. Indications of character states in the table are presence of a character state (1), absence of a character state (0), and data missing from the literature (?). Characters are notated as 1—tuberculation moderate to strong, 2—tubercles on forelimbs, 3—tubercles on hind limbs, 4—tubercles on head and/or occiput, 5—tubercles on at least 1/3 of tail, 6—number of ventral scales, 7—enlarged median subcaudals, 8—proximal subdigital lamellae broad, 9—number of subdigital lamellae on toe IV, 10—contact of posterior thigh scales abrupt, 11—enlarged femoral scales, 12—number of femoral pores, 13—pre-cloacal groove, 14—enlarged pre-cloacal scales, 15—number of pre-cloacal pores, 16—pre-cloacal and femoral pores/scales continuous, 17—reticulate pattern on head, 18—body banded, 19—body blotched, 20—body striped. Where derived from the literature, references are abbreviated by letters as follows: A—Gu¨nther and Ro¨sler (2002), B—Brown and Alcala (1978), C—Youmans and Grismer (2006), D—Grismer (2005), E—Taylor (1963), F—Bauer et al. (2003), G—de Rooij (1915), H—Brown and Parker (1973), I—Dring (1979), J—Brongersma (1934), K—Darevsky (1964), L—Werner (1896), M—Bauer et al. (2002), N—Smith (1925), O—Das and Lim (2000).

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0 1 ?1 0 0 0 0 0 0 0 0 0 0 0 ? 1 1 1 0 1 0 0 0 1 0 1 0 1 F O C

M N

C C

pulchellus quadrivirgatus redimiculus semenanjungensis seribuatensis sermowaiensis sumonthai sworderi thirakhupti tigroides tiomanensis yoshii

C C

115 51–71 78 59–69 75 68–93.5 71 63–77 72–80 83 84 75–96

1 1 ? 1 1 1 1 1 1 1 1 1

1 1 ? 1 1 1 1 1 1 1 1 1

1 1 ? 1 1 1 1 1 1 1 1 1

1 1 ? 1 1 1 1 1 1 ? 1 1

1 1 ? ? 1 0 0

33–35 34–42 ? 48–53 28–39 30–38 33–36 42–48 37–40 34 36–40 50–58

1 0 1 0 0 0 1 0 1 0 0 0

1 1 ? 1 1 1 1 1 1 0 1 0

19, 20 19, 20 20–24 17–21 19–22 22 18 20 20, 21 19–23 20–22 25–30

1 0 ? 1 1 0 ? 1 1 ? 1 0

1 1 ? 0 1 0 0 1 1 0 1 0

14–18 0 8–9 0 42–45 0 0 0 0 5–7 0 0

1 0 1 1 0 0 0 0 0 0 1 0

1 1 1 1 1 0 1 1 1 1 1 0

6–8 0–4 5–8 0 42–45 0 2 5, 6 0 8–9 3–5 8–12

1 0 ? 0 1 0 0 0 1 0 0 0

0 0 ? 0 0 0 0 0 1 0 0 0

1 0 ? 1 0 0 1 0 1 1 1 0

20 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 SVL n Taxon

TABLE 2.—Continued.

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sworderi, C. thirakhupti, C. tiomanensis and C. yoshii (Table 2). DISCUSSION We consider herpetological diversity on the island of Sulawesi to be underestimated owing to a lack of attention by taxonomists and comprehensive survey work (Brown et al., 2000; Evans et al., 2003a,b; Iskandar and Tjan, 1996; McGuire et al., 2007). Recent biotic survey and inventor efforts have found new species on Sulawesi and adjacent islands including two new species of flying lizard, genus Draco (McGuire et al., 2007) and additional species that are in need of description. Genetic analyses of some groups: Limnonectes (Evans et al., 2003a), Bufo celebensis (Evans et al., 2003b) and Lamprolepis (unpublished data) have revealed cryptic lineages. Exploration of the northern, central and southwestern regions of the island have resulted in the discovery of two new species of Cyrtodactylus—a new species described by Hayden et al. (2008) and C. spinosus. We suspect that additional species of Cyrtodactylus await discovery and several taxa currently masquerade under the widespread species C. jellesmae. Among SE Asian members of the genus, Cyrtodactylus spinosus is morphologically distinct, owing to the presence of spines, unique scalation, and distinct color pattern; it is consequently unlikely to be confused with any other species. Indeed, this species is so unique we cannot speculate on its closest relatives within the genus. Discovery of this new species within LoreLindu National Park further emphasizes the need for conservation efforts targeting this major center of SE Asian biodiversity. Despite the park’s having been designated as a protected area, farmers in the area continue to encroach the park’s boundary, using slashand-burn methods to clear land for agriculture. Both small-scale timber poaching and large-scale logging operations continue to degrade the park, especially along its unguarded southern boundary (Bickford et al., 2007). Improved security of the park boundary is needed to protect this stronghold of biodiversity that likely houses a large diversity of endemic species yet to be discovered.

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Acknowledgments.—We thank N. Andayani (Univ. of Indonesia; Wildlife Conservation Society), J. Supriatna (Univ. of Indonesia; Conservation International), Mumpuni (Museum Zoologicum Bogoriense, LIPI), and A. Riyanto (Museum Zoologicum Bogoriense, LIPI) for their assistance in obtaining permits to work in Indonesia, as well as with other logistical issues. We thank the following government officials who provided research and export permits, as well as specimen loans: A. Budiman (formerly of LIPI), S. Prijono (former Director of Museum Zoologicum Bogoriense). For assistance and companionship in the field, we are grateful to F. Chain, D. Halliwel, A. Rosyid, Shobi, and Ted Townsend (plus others). For discussion and/or comments on the manuscript, we thank L. Trueb. This work was supported by the National Science Foundation (DEB-BSI 0328700).

LITERATURE CITED BATUWITA, S., AND M. M. BAHIR. 2005. Description of five new species of Cyrtodactylus (Reptilia: Gekkonidae) from Sri Lanka. Raffles Bulletin of Zoology Suppl. 12:351–380. BAUER, A. M. 2002. Two new species of Cyrtodactylus (Squamata: Gekkonidae) from Myanmar. Proceedings of the California Academy of Sciences 53:75–88. BAUER, A. M. 2003. Description of seven new Cyrtodactylus (Squamata: Gekkonidae) with a key to the species of Myanmay. Proceedings of the California Academy of Sciences 54:463–498. BAUER, A. M., AND K. HENLE. 1994. Familia Gekkonidae (Reptilia, Sauria). Part 1. Australia and Oceania. Das Tierreich 109. BAUER, A. M., O. S. G. PAUWELS, AND L. CHANHOME. 2002. A new species of cave-dwelling Cyrtodactylus (Squamata: Gekkonidae) from Thailand. Natural History Journal of the Chulaongkorn University 2:19–29. BAUER, A. M., M. SUMONTHA, AND O. S. G. PAUWELS. 2003. Two new species of Cyrtodactylus (Reptilia: Squamata: Gekkonidae) from Thailand. Zootaxa 376:1–18. BICKFORD, D. P., J. SUPRIATNA, Y. ANDAYANI, D. ISKANDAR, B. J. EVANS, R. M. BROWN, T. TOWNSEND, UMILAELA, D. AZHARI, AND J. A. MCGUIRE. 2007. Indonesia’s protected areas need more protection - suggestions from island examples. Pp. 53–77. In N. S. Sodhi, G. Acciaioli, M. Erb, and A. K.-J. Tan (Eds.), Biodiversity and Human Livelihoods in Protected Areas: Case Studies from the Malay Archipelago. Cambridge University Press, Cambridge, U.K. BOULENGER, G. A. 1897. A catalogue of the reptiles and batrachians of Celebes, with special reference to the collections made by Drs. P. & F. Sarasin in 1893–1896. Proceedings of the Zoological Society of London 1897:193–237. BRONGERSMA, L. D. 1934. Contributions to Indo-Australian Herpetology. Zoologische Mededeelingen 17: 161–251. BROWN, R. M., AND A. C. DIESMOS. 2002. Application of lineage-based species concepts to oceanic island frog populations: the effects of differing taxonomic philosophies on the estimation of Philippine biodiversity. Silliman Journal 42:133–162. BROWN, R. M., AND S. I. GUTTMAN. 2002. Systematic evolution in Rana signata complex of Philippine and

[Vol. 64, No. 2

Bornean stream frogs: reconsideration of Huxley’s modification of Wallace’s Line at the Oriental-Australian faunal zone interface. Biological Journal of the Linnean Society 76:393–461. BROWN, R. M., J. SUPRIATNA, AND H. OTA. 2000. Discovery of a new species of Luperosaurus (Squamata; Gekkonidae) from Sulawesi, with a phylogenetic analysis of the genus and comments on the status of L. serraticaudus. Copeia 2000:191–209. BROWN, W. C., AND A. C. ALCALA. 1978. Philippine Lizards of the Family Gekkonidae. Silliman University Press, Dumaguete City, Philippines. BROWN, W. C., AND F. PARKER. 1973. A new species of Cyrtodactylus (Gekkonidae) from New Guinea with a key to species from the island. Brevoria 417. DAREVSKY, I. S. 1964. Two new species of gekkonid lizards from the Komodo island in Lesser Sundas Archipelago. Zoologischer Anzeiger 173:169–174. DAS, I., AND L. J. LIM. 2000. A new species of Cyrtodactylus (Sauria: Gekkonidae) from Pulau Tioman, Malaysia. The Raffles Bulletin of Zoology 48:223–231. DAVID, P., A. TEYNIE´, AND A. OHLER. 2004. A new species of Cyrtodactylus Gray, 1827 (Reptilia: Squamata: Gekkonidae) from southern Laos. Raffles Bulletin of Zoology 52:621–627. DE ROOIJ, N. 1915. The reptiles of the Indo-Australian Archipelago. I Lacertilia, Chelonia, Emydosauria. E. J. Brill, Leiden, The Netherlands. DE QUEIROZ, K. 1998. The General Lineage Concepts of species, species criteria, and the process of speciation. Pp. 57–75. In D. J. Howard and S. H. Berlocher (Eds.), Endless forms: species and speciation. Oxford University Press, New York, New York, U.S.A. DE QUEIROZ, K. 1999. The General Lineage Concept of species and the defining properties of the species category. Pp. 49–89. In R. A. Wilson (Ed.), Species: new interdisciplinary essays. Massachusetts Institute of Technology Press, Cambridge, Massachusetts, U.S.A. DRING, J. C. M. 1979. Amphibians and reptiles from northern Trengganu, Malaysia, with descriptions of two new geckos: Cnemaspis and Cyrtodactylus. Bulletin of the British Museum (Natural History), Zoology 34:181–241. EVANS, B. J., R. M. BROWN, J. A. MCGUIRE, J. SUPRIATNA, N. ANDAYANI, A. DIESMOS, D. ISKANDAR, D. J. MELNICK, AND D. C. CANNATELLA. 2003a. Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the Asian and Australian faunal zones. Systematic Biology 52:794–819. EVANS, B. J., J. SUPRIATNA, N. ANDAYANI, M. I. SETIADI, D. C. CANNATELLA, AND D. J. MELNICK. 2003b. Monkeys and toads define areas of endemism on Sulawesi. Evolution 57:1436–1443. GOLUBEV, M. L., AND N. N. SZCZERBAK. 1985. On the relationships of two Palearctic gecko genera: Tenuidactylus I Agamura (Reptila, Gekkonidae) [in Russian]. Pp. 61–62. In Problems of Herpetology, Sixth AllUnion Herpetological Conference Abstracts, Tashkent. Nauka, Leningrad. GRISMER, L. L. 2005. New species of bent-toed gecko (Cyrtodactylus Gray 1827) from Pulau Aur, Johor, West Malaysia. Journal of Herpetology 39:424–432.

June 2008]

HERPETOLOGICA

GRISMER, L. L., AND T. M. LEONG. 2005. New Species of Cyrtodactylus (Squamata: Gekkonidae) from southern Peninsular Malaysia. Journal of Herpetology 39: 584–591. GU¨NTHER, R., AND H. RO¨SLER. 2002. Eine neue Art der Gattung Cyrtodactylus Gray, 1827 aus dem Westen von Neuguinea (Reptilia: Sauria: Gekkonidae). Salamandra 38:195–212. HAYDEN, C. J., R. M. BROWN, G. GILLESPIE, M. I. SETIADI, UMILAELA, C. W. LINKEM, D. BICKFORD, A. RIYANTO, MUMPUNI, AND J. A. MCGUIRE. 2008. A new species of bent-toed gecko Cyrtodactylus Gray 1827 (Squamata: Gekkonidae) from the island of Sulawesi, Indonesia. Herpetologica 64. HEIDRICH, A., H. RO¨SLER, V. N. THANH, WOLFGANG BO¨HME, AND THOMAS ZIEGLER. 2007. Another new Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha - Ke Bang National Park, central Truong Son, Vietnam. Zootaxa 1445:35–48. ISKANDAR, D. T., AND K. N. TJAN. 1996. The amphibians and reptiles of Sulawesi, with notes on the distribution and chromosomal number of frogs. Pp. 39–46. In D. J. Kitchener and A. Suyanto (Eds.), Proceedings of the First International Conference on Eastern Indonesian-Australian Vertebrate Fauna. Manado, Indonesia. KLUGE, A. G. 1983. Cladistic relationships among gekkonid lizards. Copeia 1983:465–475. KLUGE, A. G. 1991. Checklist of gekkonoid lizards. Smithsonian Herpetology Information Resource 8:60–68. KLUGE, A. G. 1993. Gekkonoid Lizard Taxonomy. International Gecko Society, San Diego, California, U.S.A. KLUGE, A. G. 2001. Gekkotan Lizard Taxonomy. Hamadryad 26:1–209. MACEY, J. R., N. B. ANANJEVA, Y. WANG, AND T. J. PAPENFUSS. 2000. Phylogenetic relationships among Asian Gekkonid lizards formerly of the genus Cyrtodactylus based on cladistic analyses of allozymic data: monophyly of Cyrtopodion and Mediodactylus. Journal of Herpetology 34:258–265. MCGUIRE, J. A., R. M. BROWN, MUMPUNI, A. RIYANTO, AND N. ANDAYANI. 2007. The flying lizards of the Draco lineatus group (Squamata: Iguania: Agamidae): A taxonomic revision with descriptions of two new species. Herpetological Monographs 21:180–213. ORLOV, N. L., N. Q. TRUONG, R. A. NAZAROV, N. B. ANANJEVA, AND N. N. SANG. 2007. A new species of the genus Cyrtodactylus Gray, 1827 and redescription of Cyrtodactylus paradoxus (Darevsky et Szczerbak, 1997) [Squamata: Sauria: Gekkonidae] from south Vietnam. Russian Journal of Herpetology 14:145–152. PAUWELS, O. S. G., A. M. BAUER, M. SUMONTHA, AND L. CHANHOME. 2004. Cyrtodactylus thirakhupti (Squamata: Gekkonidae), a new cave-dwelling gecko from southern Thailand. Zootaxa 772:1–11. QUANG, H. X., N. L. ORLOV, N. B. ANANJEVA, A. G. JOHNS, H. N. THAO, AND D. Q. VINH. 2007. Description of a new species of the genus Cyrtodactylus Gray, 1827 (Squamata: Sauria: Gekkonidae) from the karst of north central Vietnam. Russian Journal of Herpetology 14:98–106.

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SMITH, M. A. 1925. A new ground-gecko (Gymnodactylus) from the Malay Peninsula. Royal Asiatic Society, Journal of the Malayan Branch, Singapore 3:87. SZCZERBAK, N. N., AND M. L. GOLUBEV. 1984. On generic assignment of the Palearctic Cyrtodactylus lizard species (Reptilia, Gekkonidae) [in Russian]. Vestnik Zoologii 1984:50–56. SZCZERBAK, N. N., AND M. L. GOLUBEV. 1986. Gecko Fauna of the USSR and Contiguous Regions [in Russian]. Naukova Dumka, Kiev, Ukraine. TAYLOR, E. H. 1963. Lizards of Thailand. The University of Kansas Science Bulletin 55:687–1077. WERNER, F. 1896. Zweiter Beitrag zur Herpetologie der Indo-Orientalischen Region. Verhandlungen der Kaiserlich-Ko¨niglichen zoologisch-botanischen Gesellschaft in Wien 46:6–24. WILEY, E. O. 1978. The evolutionary species concept reconsidered. Systematic Zoology 21:17–26. YOUMANS, T. M., AND L. L. GRISMER. 2006. A new species of Cyrtodactylus (Reptilia: Squamata: Gekkonidae) from the Seribuat Archipelago, west Malaysia. Herpetological Natural History 10:61–70. ZIEGLER, T., H. RO¨SLER, H. W. HERRMAN, AND V. N. THANH. 2002. Cyrtodactylus phongnhakebangensis sp. n., ein neuer Bogenfingergecko aus dem annamitischen Karstwaldassiv, Vietnam. Herpetofauna 24:11–25.

APPENDIX Specimens Examined Codes for institutions are CAS, California Academy of Sciences; KU, University of Kansas Natural History Museum; LSUMZ, Louisiana State University Museum of Zoology; MVZ, Museum of Vertebrate Zoology, University of California, Berkeley; MZB, Museum Zoologicum Bogoriense, Bogor Indonesia; NMPNG, National Museum of Papua New Guinea; UPNG, University of Papua New Guinea; CCA, Christopher C. Austin; RMB, Rafe M. Brown; JAM, Jimmy A. McGuire. Field collection numbers are in parentheses. Cyrtodactylus agusanensis: PHILIPPINES: MINDANAO: Province of Davao del Norte: 70 km S Bislig: LSUMZ 41601, 41602 (males), LSUMZ 41603 (female), LSUMZ 41604, 41605 (males); DINAGAT: Province of Surigao del Norte: Municipality of Loreto: KU 305564–5; SAMAR: Province of Eastern Samar: Municipality of Taft: Barangay San Rafael: KU 305569. Cyrtodactylus annulatus: PHILIPPINES: 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). Cyrtodactylus cavernicolus: EAST MALAYSIA: SARAWAK: Niah Cave: CAS 23726 (male). Cyrtodactylus consobrinus: EAST MALAYSIA: SARAWAK: Bintulu District: Sungei Seran: CAS 105993 (male); Kapit District: Sungai Mengiong: MVZ 11782 (male); BRUNEI: Temburang District: Sungai Belalang: LSUMZ 55833 (male); INDONESIA: PROPINSI SUMATERA UTARA: Bukit Lawang: Bahorak: MZB 4355 (male), 4356 (juvenile). Cyrtodactylus darmandvillei: INDONESIA: PROPINSI NUSA TENGGARA BARAT: Pulau Lombok.LSUMZ 81732 (JAM 3176) (male). Cyrtodactylus derongo: PAPUA NEW GUINEA: WESTERN PROVINCE: Derongo: NMPNG R23452 (female). Cyrtodactylus

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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); Poso Kabupaten: LSUMZ 8402 (male); Luwuk Kabupaten: LSUMZ 8405 (male), 8404, 8406 (females). Cyrtodactylus loriae: PAPUA NEW GUINEA: CHIMBU PROVINCE: Karimui: CAS 117964 (male), CAS 118018 (female), NMPNG R23625, R8347, R8348; PAPUA NEW GUINEA: UPNG 5687 (male); WESTERN HIGHLANDS PROVINCE: Kaironk: UPNG 0976 (female); MOROBE PROVINCE: Wau.: NMPNG 24730. Cyrtodactylus louisiadensis: PAPUA NEW 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,

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Taman Nasional Bentuang Karimun: MZB 3920 (female). Cyrtodactylus marmoratus: INDONESIA: JAVA: Djurung Sriti: 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 novaeguineae: PAPUA NEW GUINEA: SANDAUN PROVINCE: Utai: CCA 3142 (female), 3415 (juvenile); Bewani Station: CCA 3674 (male). Cyrtodactylus quadrivirgatus cf.: INDONESIA: SUMATRA: Propinsi Bengkulu: 46 km East of Bengkulu:.MVZ 239338 (male), 239578 (female). Cyrtodactylus philippinicus: PHILIPPINES: LUZON: Camarines del Sur Province: Municipality of Naga: KU 305571; NEGROS: Negros Oriental Province: Municipality of Dumagete, Baragay Valencia: KU 305572; LUBANG: Occidental Mindoro Province: Municipality of Lubang, Barangay Vigo: KU 303846, 303844. Cyrtodactylus sermowaiensis: PAPUA NEW GUINEA: SANDAUN PROVINCE: 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: NMPNG R24803–24806; EAST SEPIK PROVINCE: Maprik: NMPNG R22796; SANDAUN PROVINCE: Idam River: UPNG 3914– 3916.