Zootaxa 2488: 22–38 (2010) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition)

A new cryptic of Gray, 1831 (: ) from and

PERRY L. WOOD JR1,6, L. LEE GRISMER2, JESSE L. GRISMER1, THY NEANG3, THOU CHAV4, & JERMEY HOLDEN5 1Department of Biology, Villanova University, Villanova, PA 19085, USA. E-mail: [email protected], [email protected] 2Department of Biology, La Sierra University, Riverside, CA 92515-824, USA E-mail: [email protected] 3General Department of Nature Conservation and Protection (DNCP), Ministry of Environment, Phnom Penh, Cambodia. E-mail: [email protected] 4Department of Environment, Pursat Penh, Cambodia 567 High Street Meppershall, Beds, SG17 5LX, UK. E-mail: [email protected] 6Corresponding author. E-mail: [email protected]

Abstract

A new species of Acanthosaura (Gray 1831), A. cardamomensis sp. nov. from eastern Thailand and western Cambodia is described based on having a combination of unique morphological characteristics. It most closely resembles A. crucigera but differs in having significantly longer postorbital spines, occipital spines, nuchal spines, and dorsal spines; more scales between the fifth canthals; longer maximum dorsal scale length; longer maximum nuchal scale length; a significantly smaller diastema between the nuchal and dorsal crest spines; and significantly more supralabials. Mitochondrial DNA was used to calculate percent sequence divergence between samples of A. crucigera from southern and western Thailand and samples of A. cardamomensis sp. nov. from eastern Thailand and western Cambodia. The uncorrected pairwise sequence divergence between populations of A. cardamomensis ranged from 1.2– 3.1% and the range between populations of A. crucigera was 3.8%. The percent sequence divergence between A. cardamomensis sp. nov. and A. crucigera ranged between 10.0–14.6%.

Key words: Acanthosaura, armata, cardamomensis, crucigera, Cardamom Mountains

Introduction

Species of the Southeast Asian agamid Acanthosaura (Gray 1831) are diurnal, arboreal, insectivores ranging in elevation from sea level to 1450 m a.s.l. (Ananjeva 2008). Currently, the genus contains eight species ranging from northeastern India, east through Myanmar, Thailand, Cambodia, , , southern (including island), and southward through the Malay Peninsula and adjacent archipelagos (Grismer et al. 2006) to Sumatra, and the Anambas and Natunas Archipelagos (Grismer et al. 2008a,b; Leong et al. 2002; Taylor 1963; Manthey & Grossmann 1997; Manthey 2008). Recent molecular investigations of the relationships within Acanthosaura have established that some of the wide-ranging species were composed of multiple species (Kalybina-Hauf et al. 2004; Ananjeva et al. 2008). is one of these wide-ranging species, which extends from Myanmar east through Thailand, Cambodia, and southward to Northern Peninsular (Boulenger 1912; Grismer et al. 2006b; Manthey & Grossmann 1997; Manthey 2008; Pauwels et al. 2003; Taylor 1963). Wood et al. (2009) demonstrated that the montane population from Bukit Larut, in Peninsular Malaysia referred to as of A. crucigera (Boulenger 1912; Manthey 2008) was a different species, A. bintangensis. Within Cambodia, there are three species of Acanthosaura: A. capra (Günther 1861), A. coronata (Günther 1861) and A. crucigera (Boulenger 1885). and A. coronata are found in hilly

22 Accepted by S. Carranza: 20 Apr. 2010; published: 31 May 2010 eastern Cambodia (Stuart et al. 2006) and A. crucigera occurs in the Cardamom Mountains in western Cambodia (Grismer et al. 2008; Stuart & Emmett 2006). Boulenger (1885) considered a male specimen collected by Henri Mouhot in the mid nineteenth century from “Chartaboum” (= Chanthaburi Province) from the Thai Cardamom Mountains (BMNH 61.4.12.43) southeastern Thailand, as A. armata. Smith (1935) was the first to report A. crucigera from the southeastern region of the Cardamom Mountains and subsequent surveys throughout the Cardamom Mountains reported additional populations (Stuart & Emmett 2006; Daltry & Chheang 2000; Grismer et al. 2007, 2008a,b; Daltry & Traehdt 2003; Chuaynken et al. 2004). Stuart and Emmett (2006) commented on the differences in Boulenger’s and Smith’s taxonomic opinions and determined that they all resembled the Cambodian specimens of A. crucigera. Genetic and morphological analyses of these from eastern Thailand and Cambodia, previously ascribed to A. crucigera and A. armata have revealed they cannot be ascribed to any known species of Acanthosaura and are described herein as new.

Materials and methods

Specimens were collected by hand or blow pipe during all hours of the day from Phnom Aural (4-10 August 2005), Phnom Samkos (4-13 August 2006), and Phnom Dalai (4-10 July 2009). Photographs were taken to document color pattern in life prior to euthanization. Liver samples taken were stored in 100% ethanol and lizards were fixed in 10% formalin and later transferred to 70% ethanol. Fifty-two morphological characters were analyzed and taken from museum specimens (Appendix) and the literature (Hardwicke & Gray 1827; Gray 1831, 1845; Orlov et al. 2006; Günther 1861, 1864; Boulenger 1885, 1912; Smith 1935; Taylor 1963; Manthey & Grossmann 1997; Stuart & Emmett 2006; Stuart et al. 2006; Ananjeva et al. 2008; Wood et al. 2009) from all currently recognized species of Acanthosaura. Measurements were taken on the type series to the nearest 0.1 mm with Mitutoyo digital calipers under a Nikon SMZ 1000 dissecting microscope. All measurements were made on the left side of the body where applicable (fide Wood et al. 2009). Characters examined were: SVL—snout vent length, measured from the tip of the snout to the tip of the vent; TL—tail length, measured from the posterior margin of the vent to the tip of the tail; HL—head length, measured from posterior edge of the rectis of the jaw to the tip of the snout; HW—maximum head width, the width at the level of the tympanum; HD—maximum head height, measured across the parietal region; SL—snout length, measured from the anterior edge of the orbit to the tip of the snout; MW—mental width; MH—mental height; RW—rostral width; RH—rostral height; ORBIT—eye diameter, measured from the posterior to the anterior edge of the eye; TD—tympanum diameter, measured horizontally from the anterior to the posterior border of the tympanum; OS—length of the occipital spine, measured from the base to the tip; SUPRAL—number of supralabials; INFRAL—number of infralabials; DIASN—number of scales in the vertebral crest scale diastema counted from the posterior end of the nuchal crest to the anterior end of the dorsal crest; VENT— number of ventral scales counted at the midline from the anterior edge of the shoulders to the edge of the vent; FI—number of subdigital lamellae on the fourth finger; TO—subdigital lamellae on the fourth toe; NSSOS, number of scales surrounding the occipital spine; CS—number of canthus-rostralis-supracilliary scales, counted from the nasal scale to the posterior end of the ridge at the posterior margin of the orbit; NS—number of scales between the nasals; PM—number of postmental scales bordering the mental; TN—scales absent on tympanum (0) or present (1); YAS— presence (1) or absence (0) of a Y-shaped arrangement of enlarged scales on the snout; ND— presence (1) or absence (0) of a black, diamond shaped, nuchal collar; BEP—presence or absence of a black eye patch; ESBO, presence or absence of elliptical scales below the orbit (Table 1). Institutional abbreviations are BMNH, Natural History Museum, London, England; CUMZR, Chulalongkorn University Museum of Zoological Records, Bangkok, Thailand; THNHM Thailand Natural History Museum, Pathum Thani, Thailand; FMNH, Field Museum of Natural History, Chicago, Illinois, USA; LSUHC, La Sierra University Herpetological Collection, La Sierra University, Riverside, California, USA; UKMHC, Universiti Kebangsaan Malaysia Herpetological Collection, Bangi, Selangor, West Malaysia and ZRC, Zoological Reference Collection at the Raffles Museum of Biodiversity at the National University of Singapore.

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 23 Statistical analyses were used to test for significant differences in means. All data were analyzed using PASW statistics 17.0 (SPSS Inc., Chicago, IL). Kolmogorov-Smirnov and Shapiro-Wilk tests were used to test for normal distributions between the variables. When distributions were normal a two-sample t-test was used to test for significant differences between the means of different groups. Levene’s test for equal variances was also employed. When distributions were not normal, the nonparametric Mann-Whitney U test was used to test for significant differences between different means. Molecular data was collected from 12 specimens of Acanthosaura crucigera from Thailand and Cambodia to determine the percent sequence divergence between the different populations. For all the taxa used in the genetic study GenBank accession numbers can be found in the (Appendix). Genomic DNA was extracted from liver or muscle tissue that were preserved in 95–100% ethanol following Qiagen DNeasy tissue kit (Valencia, CA, USA). We used two primers, METF6 (L4437a; 5′–AAG CTT TCG GGC CCA TAC C– 3′) from Macey et al. (1997) and a primer new to this paper ACANTHND2.833.R1 (5′– AGG GAG GTT ATT GTT GCT AG–3′) to amplify a 698 bp fragment of the NADH dehydrogenase subunit 2 gene. The PCR protocol followed Jackman et al. (2008). The PCR products were purified using AMPure magnetic bead solution (Agentcourt Bioscience, Beverly, MA, USA) and were sequenced with Purified PCR products and were then sequenced using the Applied Biosystems BigDyeTM Terminator v3.1 Cycle Sequencing ready reaction kit or with the GE Healthcare DYEnamicTM ET Dye Terminator Kit (GE Healthcare, Piscataway, NJ, USA). Sequencing reactions were purified using CleanSeq magnetic bead solution (Agentcourt Bioscience, Beverly, MA, USA) and were analyzed with an ABI 3700 automated sequencer. Complementary strands were used to insure accuracy. The 698 bp sequences were edited using Geneious (2006) and were then aligned by eye. Sequences were then incorporated into MacClade (Maddison & Maddison 1992) and translated into amino acids to check the reading frame of the sequences and to check for premature stop codons. Uncorrected pairwise sequence divergence was calculated using PAUP*4.0b10 (Swofford 2002).

Results

An independent sample t-test and the nonparametric Mann-Whitney U tests indicate that there are significant differences between the means of selected characters in Acanthosaura crucigera from Thailand and A. crucigera from the Cardamom Mountains. Seven characters were significantly different; two meristic and five morphometric (Table 1, 2). Four of these characters were normally distributed and were analyzed using independent sample t-tests (PS/HL, OS/HL, NS/HL, NCS). The remaining three characters with significantly non-normal distributions were significantly different between A. crucigera and A. crucigera from the Cardamom Mountains (DS/HL, DIA/SVL, SUPRAL). An independent sample t-test and the nonparametric Mann-Whitney U tests indicate that there are significant differences between and A. crucigera from the Cardamom Mountains. Eleven characters were significantly different: eight meristic and three morphometric (Table 3, 4). Six of these characters were normally distributed and were analyzed using independent sample t-tests (PS/HL, OS/HL, NS/HL, DIASN, VENT, NSSOS). The remaining five with significantly non-normal distributions differ significantly between A. armata and A. crucigera from the Cardamom Mountains (SUPRAL, INFRAL, CS, SDLT, SDLF). The uncorrected sequence divergence within A. crucigera from eastern Thailand and the Cambodian Cardamom mountains populations ranged from 1.2–3.1%. The interspecifc sequence divergence between A. crucigera from Botpyinn Township, Pakchan Reserve Forest, Main Ma The camp, Kawthaung District, Tanintharyi Division, Myanmar, (CAS 229582) and A. crucigera from Kaeng Krachan National Park, Phetchaburi Province, Thailand (CUMZR 2008.05.26.1) 265 km to the southeast was 3.8%. The percent sequence divergence between populations of A. cf. crucigera from the Cardamom Mountains and A. crucigera from western Thailand (390 km to the west) and southern Myanmar (530 km to the southwest) was surprisingly high at 10.0–14.6%. The high percent sequence divergence along with morphological differences strongly indicates that these populations are not exchanging genes. Acanthosaura from eastern Thailand and

24 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL. western Cambodian represents genetically and morphological distinct species separate from A. crucigera in western Thailand and southeastern Myanmar and is described below.

FIGURE 1. The distribution of Acanthosaura crucigera, A. cf. crucigera, and A. cardamomensis in Cambodia, Thailand, and Myanmar (Daltry & Chheang 2000; Daltry & Traeholt 2003; Grismer et al. 2007, 2008a,b; Manthey 2008; Stuart & Emmet 2006; Stuart et al. 2006). The green shape depicts the approximate outline of the Thai and Cambodian Cardamom Mountains. The green triangle indicates the type locality of Acanthosaura crucigera (Tavoy, Burma). Acanthosaura cf. crucigera extends southward to the Malaysian border including Pulau Langkawi.

Systematics

Acanthosaura cardamomensis sp. nov. Cardamom Mountain Horned Agamid Figures 2–3

Acanthosaura armata Boulenger, G. A. 1885: 301. Gonocephalus armatus crucigerus Smith, M. A. 1935:160–161. Acanthosaura crucigera Daltry, J. C. & Cheang, D. 2000: 100. Acanthosaura crucigera Daltry, J. C. & Traeholt, C. 2003: 99–100. Acanthosaura crucigera Stuart, B. L. & Emmett, D. A. 2006: 12–13. Acanthosaura cf. crucigera Grismer, L. L., Chav T., Neang T., Wood Jr., P. L., Grismer, J. L., Youmans, T. M., Ponce, A., Daltry, J. C. & Kaiser, H. 2007: 221–222. Acanthosaura sp. Grismer, L. L., Neang T., Chav T., Wood Jr., P. L., Oaks, J. R., Holden, J., Grismer, J. L., Szutz, T. R. & Youmans, T. M. 2008: 165.

Holotype. Adult female (LSUHC 7447) collected on 5 August 2005 by Jesse L. Grismer, L. Lee Grismer & Perry L. Wood Jr., at 1400 h at 1,121 m a.s.l. from Camp 2, Phnom Aural Kampong Speu Province, Cambodia (12°01’N, 104°09’E).

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 25 FIGURE 2. Adult female holotype (LSUHC 7447) of Acanthosaura cardamomensis from Kampong, Phnom Aural, Cambodia (12°01’N, 104°08’E) at 1400 m a.s.l. (A) In life. (B) Lateral view of the scalation on the left side of the body. (C) Lateral view of the head. (D) Dorsal view.

TABLE 1. Results of an independent sample t-test for Acanthosaura crucigera from western Thailand and Myanmar and Acanthosaura cf. crucigera from eastern Thailand and Cambodia.

A. crucigera A. cf. crucigera Sample size Mean ± SD Sample Size Mean ± SD Test statistic P-value PS/HL n=8 0.18 ± 0.10 n=29 0.33 ± 0.07 t=-4.2 p<0.001 OS/HL n=8 0.19 ± 0.13 n=27 0.33 ± 0.08 t=-3.57 p<0.002 NS/HL n=8 0.23 ± 0.82 n=28 0.41 ± 0.09 t=-4.69 p<0.001 NCS n=8 10 ± 1 n=34 12 ± 2 t=-4.07 p<0.001

TABLE 2. Results of a nonparametric Mann-Whitney U test for Acanthosaura crucigera from western Thailand and Myanmar and Acanthosaura cf. crucigera from eastern Thailand and Cambodia. A. crucigera A. cf. crucigera Sample Size Sample Size Test statistic P-value DS n=8 n=33 z=-3.40 p<0.002 NS n=8 n=33 z=-3.18 p<0.002 DS/HL n=5 n=31 z=-4.00 p<0.001 DIA/SVL n=5 n=32 z= -2.49 p<0.010 SUPRA n=5 n=34 z=-3.13 p<0.015

26 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL. TABLE 3. Results of an independent sample t-test for Acanthosaura armata from Peninsular Malaysia and Singapore and Acanthosaura cf. crucigera from eastern Thailand and Cambodia. A. armata A. cf. crucigera Sample size Mean ± SD Sample Size Mean ± SD Test statistic P-value PS/HL n=13 0.43 ± 0.08 n=29 0.32 ± 0.07 t=-4.01 p<0.001 OS/HL n=13 0.23 ± 0.06 n=27 0.32 ± 0.07 t=3.75 p<0.002 NS/HL n=13 0.32 ± 0.07 n=28 0.40 ± 0.09 t=2.81 p<0.009 DIASN n=14 5.21 ± 2.19 n=33 10.8 ± 2.55 t=7.10 p<0.001 VENT n=14 62.2 ± 5.01 n=33 56.2 ± 3.11 t=-8.41 p<0.001 NSSOS n=14 4.5 ± 0.51 n=32 4.96 ± 0.64 t=2.39 p<0.022

TABLE 4. Results of a nonparametric Mann-Whitney U test for Acanthosaura armata from Peninsular Malaysia and Singapore and Acanthosaura cf. crucigera from eastern Thailand and Cambodia. A. armata A. cf. crucigera Sample Size Sample Size Test statistic P-value SUPRAL n=14 n=34 z=-2.41 p<0.017 INFRAL n=14 n=34 z=-4.42 p<0.001 CS n=14 n=34 z=-2.71 p<0.008 SDLT n=14 n=34 z=-1.97 p<0.050 SDLF n=14 n=34 z=-3.00 p<0.004

FIGURE 3. Type series of Acanthosaura cardamomensis from left to right; female holotype (LSUHC 7447), adult male (LSUHC 7877), adult male (LSUHC 7406), adult female (FMNH 180309), female paratype (FMNH 263260), sub adult male (FMNH 180307), juvenile (LSUHC 7446).

Paratypes. LSUHC 7406; adult male from Camp 1, Phnom Aural, Kampong Speu Province, Cambodia (12°01’N, 104°08’E) by Jesse L. Grismer on 5 August 2005 at 549 m a.s.l. LSUHC 7877; adult male Camp 1, Pursat Province, Cambodia (12°01’N, 104°08’E) by L. Lee Grismer on 6 August 2006 at 501 m a.s.l. BMNH

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 27 1932.10.4.5; sub adult male from Bokor, Kamchay Mountains, Cambodia, by M.A. Smith 1935 at 915 m a.s.l. BMNH 1915.8.14.11; sub adult female from Koh Chang Island, South East Siam M. A. Smith and C. B. Kloss. BMNH 61.4.12.43; sub adult male from Chartaboum” (= Chantaburi Province) from the Thai Cardamom Mountains, H. Mouhot. FMNH 180309; adult female from Nakon Ratchasima Prov, Thailand, Khao Yai National Park by W. Ronald Heyer on 10 June 1969. FMNH 180307; sub adult male from Nakon Ratchasima Prov, Thailand, Amphoe Pak Thong Chai, Sakaerat Exp Station collected by W. Ronlad W. Ronald Heyer on 9 May 1969. LSUHC 7446; juvenile from Camp 2, Kampong Speu Province, Phnom Aural, Cambodia (12°01’N, 104°09’E) collected by L. Lee Grismer on 8 August 2005 above 1,400 m a.s.l. FMNH 263260; adult female from hill evergreen forest along Prek Kaoh River, Bokor National Park, Kampot Province, near 10°37’19”N; 104°02’52”E Cambodia, at 800-900 m a.s.l. collected by Bryan L. Stuart on 14 may 2000.For referred material see Appendix I.

FIGURE 4. Paratype specimens from the BMNH. (A) Lateral view of the head of BMNH 1932.10.4.5. (B) Dorsal view of BMNH 1932.10.4.5. (C) Lateral view of the head of BMNH 1915.8.14.11. (D) Dorsal view of BMNH 1915.8.14.11. (E) Lateral view of the head of BMNH 61.4.12.43. (F) Dorsal view of BMNH 61.4.12.43.

Diagnosis. A large species (maximum SVL 149 mm in females, maximum SVL 133 mm in males) with a single long cylindrical spine above posterior margin of eye; large spine on occiput between tympanum and nuchal crest; tympanum naked, round; large rectangular rostral scale; dewlap present in males and females;

28 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL. females with dark coloration in the gular region; scales on flanks intermixed with randomly arranged clusters of larger keeled scales; nuchal crest present; 6–15 scales in nuchal diastema; vertebral crest composed of enlarged, pointed scales beginning at shoulder region continuing to ¾ down vertebral column then fading at base of tail; tail 120–160% of SVL; black collar and black eye patch present; posterior 2/3 of tail black to dark greenish; yellow to greenish spots on dorsum; dorsum light tan to dark green (Fig. 2; Table 5 compares these characters across all species of Acanthosaura).

FIGURE 5. Color pattern variation within Acanthosaura cardamomensis. (A) Female holotype. (B) Adult male from Phnom Aural. (C) Juvenile from Phnom Aural, Cambodia. (D) Adult male from Phnom Samkos, Cambodia. (E) Adult female from Khao Khitchakut National Park, Chanthaburi Province, Thailand (F) Sub adult male from Phnom Dalai, Cambodia.

Description of the holotype. (Fig. 2). Adult female, SVL 149.4 mm, TL 174.3 mm. Head length (HL 42.4 mm), head somewhat narrow (HW/SVL 17%), moderately tall (HD/HL 51%), triangular in dorsal and lateral profile; moderate snout length (SL/HL 44% mm), rostrum wide (RW/RH 219%), steeply sloping anteriorly; canthus rostralis prominent, forming a large projecting shelf extending above eye, composed of 13 enlarged scales; shelf terminates with a notch anterior to postorbital spine; rostral large in size, rectangular,

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 29 bordered laterally by first supralabials and posteriorly by four smaller scales; nasals crater-like, surrounded by one prenasal, four postnasal, and three small subnasal scales; eight scales between nasal scales; supranasals somewhat elongate; enlarged scales above orbit weakly keeled; three rows of moderately keeled scales below orbit extending from anterior margin of eye to middle of eye; eye moderate (ORBIT/HL 17%), orbit large (ORBIT/HL 25%); interorbital, frontal, and prefrontal scales keeled and smaller than enlarged row of keeled scales below the orbit; four enlarged, keeled, azygous prefrontal scales arranged in a Y-shaped pattern; scales of occiput flat to weakly keeled; parietal eyespot surrounded by a larger row of scales; large cylindrical epidermal spine above posterior margin of eye, surrounded by four lancolate scales; occipital spine surrounded by four enlarged keeled scales; single row of five enlarged, keeled suborbital scales extending from below posterior margin of eye to anterior margin of tympanum, decreasing in size posteriorly, terminal scale spine-like; elongate cylindrical epidermal spine on lateral margin of nape surrounded by a rosette of four smaller spines; tympanum naked, round, surrounded by small conical scales; 13 rectangular supralabials decreasing in size posteriorly; mental square above becoming triangular below, larger than adjacent infralabials; two postmentals similar in size, four scales contacting mental; chinshields enlarged, extending posteriorly to angle of jaw, separated from infralabials by one scale row anteriorly and three at angle of jaw; 12 rectangular infralabials, scales in center of series largest; gulars sharply keeled and spinose with a larger midventral row; extensible dewlap present; nuchal crest bearing a diastema of 11 scales at base of nape; nuchal crest composed of eight elongate, lancolate scales bordered on each side by two rows of enlarged, flat spinose scales; dorsal body crest present, extending from posterior margin of diastema onto base of tail, vertebral crest composed of enlarged epidermal, laterally compressed, spinose scales, bordered by two rows of slightly smaller paravertebral spinose scales; vertebral crest tapers in height three-fourth the way down vertebral column then fades at base of tail.

FIGURE 6. Morphological differentiation between adult females of (A) Acanthosaura crucigera (LSUHC 9462) from Sai Yok, Kanchantaburi, Thailand and (B) Acanthosaura cardamomensis from (LSUHC 7447) Phnom Aural, Kampong, Kampong Speu Province, Cambodia.

Body robust, triangular in cross-section; dorsal scales small with medially projecting spines; larger randomly arranged scales among dorsals; scales of pectoral region and abdomen larger than dorsals, keeled, spinose, arranged in semi-transverse rows; enlarged keeled scales anterior to vent; limbs relatively long; dorsal and ventral scales of forelimbs keeled, spinose, and nearly same size; five digits on manus; subdigital scales keeled; scales of hind limbs keeled and spinose; postfemoral scales small, interspersed with larger spinose scales; five digits on pes; subdigital scales keeled; 17 subdigital lamellae under fourth finger; 22 subdigital lamellae under fourth toe; tail 117% of SVL, covered with keeled spinose scales; keels on subcaudals posteriorly directed, same size as keels of other caudal scales; base of tail in adult males nearly twice as wide as in female.

30 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL. TABLE 5. All measurement (in mm) for all currently recognized species of Acanthosaura. PS–postorbital spine length, measured from the base to the tip of the spine; PS/HL–postorbital spine length, measured from the base to the tip of the spine divided by the head length; NS–the maximum length of the largest spine in the nuchal crest measured from the base to the tip; NS/HL–the maximum length of the largest spine in the nuchal crest measured from the base to the tip divided by the head length; NSL–longest nuchal scale, measured at the base; TD/HD–tympanum diameter divided by head depth; DS–maximum length of longest spine in the dorsal crest body measured from the base to the tip; DS/HL–maximum length of longest spine on the body measured from the base to the tip divided by the head length; OS–length of the occipital spine, measured from the base to the tip; OS/HL–length of the occipital spine, measured from the base to the tip divided by the head length; DSL–longest dorsal scale, measured at the base below the dorsal crest; WNC–maximum width of the spines in the nuchal crest, measured at the base; WDS–maximum width of the largest dorsal scale below the dorsal crest, measured at the base; DIAS–length of the diastema, measured from the posterior end of the nuchal crest to the anterior end of the nuchal crest; FOREL–forelimb length, measured from axilla to the proximal edge of the palmar region; HINDL–hindlimb length, measured from groin to the proximal edge plantar region; RS–number of scales bordering the rostral scale; NCS–number of scales between the fifth canthals; NSCSL–number of scales from the fifth canthal to the fifth supralabial; NR–number of scales between the nasals and the rostrals; NSSLC–number of scales between the seventh supralabial and the sixth canthal; LKP–presence or absence of light knee patch; GP–size of gular pouch scored as absent, small, medium or large; OF–presence or absence of oblique fold anterior to the fore limb insertions. For additional character abbreviations see materials and methods. / indicates missing data.

sp. nov. A. cardamomensis A. armata A. bintangensis A. capra coronata A. A.crucigera A. nataliae A. lepidogaster A. titiwangsaensis n=34 n=14 n=3 n=3 n=3 n=5 n=1 n=3 n=3 SVL 82–149 72.4–138 83.9–142 94–137.9 66–86.1 91.2–127 106.7– 76.45– 91.8–118.4 158 101.1 TL 103–188 96.6–190 112.8–206 133.6– 86.3–105 150.8–164 171–287 130.6– 136–174 182.1 144.1 HL 16.3–42.4 6.6–33.7 16.9–25.4 16.3–38.9 14.4–16.3 17.3–23.3 25.2–43.6 18.9–29.7 20–24.3 HW 16.4–27.7 15.3–23 17.5–23.4 16.8–27 13.6–17.5 15.6–21.4 20.2–27.8 13.40–19.1 17.5–23.4 HD 12.6–21.7 12.2–18.9 15.0–19.2 14.8–24.3 11.9–16.8 14.4–21 16.9–24.9 12–12.5 15.7–20.2 SL 8.6–18.7 6.3–16.6 7.9–11.3 7.6–16.6 6.9–8.4 7.9–12 12–19.9 9.3–10.2 9.7–12.5 ORBIT 6.0–12.7 5.4–12.2 8.4–12.6 7.6–11.6 6.9–7.5 5.5–10.8 7.2–10.9 4.7–9.1 9.8–13.2 TD 2.5–5.8 2.4–5.2 2.5–3.0 3.4–5.2 1.7–2.8 2.6–3.4 3.9–7 2.2–3.0 2.7–4 TD/HD 0.20–0.27 0.19–0.28 0.16 0.21–0.23 0.14–0.17 0.16–0.18 0.23–0.28 0.18–0.24 0.17–0.20 TN 0 0 0 0 0 0 0 0–1 0 PS 3.2–12.7 4.9–9.9 1.9–4.2 5.2–10.2 absent 2.5–7.8 7.7–17.8 2.2–2.5 3.3–4.4 PS/HL 0.14–0.45 0.22–0.56 0.07–0.19 0.36 0 0.14–0.33 0.36 0.10–0.11 0.14–0.18 NS 3.8–17.4 5.5–11.2 1.3–4.7 4.2–14.7 0 3.7–8.9 8.5–23.8 3.3–3.4 2.7–4.4 NS/HL 0.17–0.66 0.22–0.51 0.17–0.21 0.42–0.43 0 0.21–0.38 0.58 0.15 0.11–0.18 NSL 1.5–6.3 1.4–5.4 1.5–2.3 4.4–4.7 0 1.7–2.7 3.4 1.0–1.3 1.6–2.4 DS 2–14.2 4.9–11.3 1.8–2.2 3.5–6.8 absent 1.7–5.53 6–17.7 1.5–2.7 1.7–2.1 DS/HL 0.14–0.45 0.20–0.52 0.08–0.09 0.16–0.17 0 0.09–0.24 0.44 0.07–0.12 0.07–0.09 DSL 1.4–5.5 0.74–3.6 1.2–2.6 1.8–1.9 absent 1.3–1.9 2 1–2 1.5–1.9 WNC 1.8–4.2 1–2.2 1.6–2.1 2.3–4.1 0 1.8–3.4 3.1–4.8 1.5–1.5 1.4–1.6 WDS 1.3–3.3 1.3–2.3 1.7–2.4 2.2–4.3 absent 1.2–1.9 2.2–4.5 1.5–2.0 1.6–2.3 DIAS 2.7–8.3 1.2–6.8 5–7.9 2–6.7 absent 4.9–8.4 2.5 6.3–6.3 5.1–7.6 DIASN 6–15 1–8 11–15 4–7 absent 12–25 10 10–12 10–13

DIAS/ 0.03–0.07 0.01–0.06 0.043– 0.049 absent 0.048– 0.036 0.078– 0.049–0.067 SVL 0.065 0.083 0.082

continued next page

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 31 TABLE 5. (continued)

sp. nov. A. cardamomensis A. armata A. bintangensis A. capra coronata A. crucigera A. A. nataliae A. lepidogaster A. titiwangsaensis n=34 n=14 n=3 n=3 n=3 n=5 n=1 n=3 n=3 FOREL 31.7–56.8 33.7–48.9 33.9–61.5 54.2–83.8 30.2–35.3 35.2–49.8 60–85 33.0–37.1 38–51.7

HINDL 42–77.1 39–69.6 43.3–68.6 78.5–107.2 38.4–47.8 52.5–65 85–129.7 49.4–50.4 48.5–65.6

SUPRAL 11–15 10–14 12 10 12–13 10–12 11 10–13 12–13 INFRAL 10–14 12–15 11–12 12–13 11–13 10 11–12 9–13 11–12 VENT 50–65 51–68 51–55 55–66 53–58 55–61 64–71 55–61 47–57 FI 15–20 13–17 23 16–17 13–14 16–20 16–21 17–19 20–21 TO 20–25 19–26 26–28 22–24 17–19 23–26 20–27 23 23–27 TL/SVL 1.2–1.6 1.17–1.58 1.3–1.4 1.2–1.5 0.06–1.01 1.4–1.8 1.2–1.5 1.6–1.9 1.1–1.5

OS 4.1–13.3 4–9.4 1.2–2.6 absent 0 2.4–4.9 absent 3.2–3.4 1.8–2.3 OS/HL 0.24–0.56 0.16–0.38 0.1–0.11 0 0 0.13–0.5 0 0.14–0.15 0.09–0.1

NSSOS 4–6 4–6 6–7 absent 4–5 4–6 absent 5 4–5

CS 11–16 11–15 14–15 12–14 12–15 12–13 13 10–12 14–15 RW 1.7–4.7 1.8–4.5 3.6–5.3 4.2–4.6 0.8–0.9 2.7–4.0 6.13 2.8–3.0 3.6–5.2 RH 1.1–2.2 0.93–1.8 1.7–2.0 1.8–2.3 0.5–0.8 1.3–2.0 2.58 1.4–1.5 1.4–1.8 RS 5–9 5–9 7–9 7–8 5 7–9 6 5–9 9 NS 7–10 7–10 8 9 7–9 7–9 8 7–8 8 NCS 9–17 10–17 10–11 9 8–11 9–12 14 7–10 11–12 NSCSL 7–12 6–14 7–8 7–8 5–6 7–11 8 8 9–11

NR 1–2 1–2 1 1–2 3–4 1 1 1 1–2 NSSLC 10–19 10–22 9–12 9–11 6–11 9–14 16 10 11–14

MW 0.2–2.1 0.86–2.0 1.3–1.8 1.9–2.2 0.6–1.5 1–1.5 2.87 1.2–1.3 1.4–2.0 MH 0.9–2.0 0.84–2.24 1.4–2.1 1.7–2.2 1.3–1.6 1.1–1.7 2.03 1.2–1.3 1.4–2.4 PM 4–5 3–6 4–5 4 4–5 4 4 5 5 YAS 0–1 0–1 1 1 0–1 0–1 1 1 1 ND 1 0–1 1 1 0 1 0 1 1 LKP 1 1 0 1 1 1 0 1 0 BEP 1 0 1 1 0 1 1 0,1 1 ESBO 0 0 1 0 0 0 0 0 0 GP 1–4 1 3–4 3–4 0 1–2 4 1 2–4 OF 1 1 1 1 1 1 1 1 1

Coloration in life. Dorsal surface of head, body and limbs greenish-yellow, sides and top of the head lighter than body; black eye patch extending from posterior margin of nostrils through orbit and tympanum meeting dark nuchal marking on nape; lips cream colored, mottled in black on anterior labials and decreasing in mottling posteriorly; iris orangish brown; black collar complete dorsally, fading towards nape; gular region nearly uniformly black with a cream medial stripe on dewlap, extending form gular region to anterior edge of

32 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL. dewlap; body marbled with greenish black; marbled base pattern bearing randomly distributed yellow markings extending on to base of tail, where they fade; limbs greenish-yellow with scales outlined in black fading distally; tail faintly banded anteriorly, becoming nearly solid black posteriorly; undersides of limbs and belly creamed colored (Fig. 2). Variation. (Figs. 3–5). The paratypes resemble the holotype in all aspects except LSUHC 7877 (male) differs from the holotype by lacking green in the dorsal pattern, the marbling is larger and more uniform and it lacks the dark colored gular region. LSUHC 7406 (male) differs from the holotype in having a cream colored body with a faint marbled pattern bearing small randomly distributed yellow markings and lacking a dark colored gular region. FMNH 180309 (female) differs from the holotype in having a more distinct modeled pattern on the dorsum, more prominent banding on the tail and having a darker gular region. FMNH 263260 (female) has larger randomly distributed markings in the dorsal pattern. FMNH 180307 (male) differs from the holotype by having a more prominent marbled dorsal pattern and lacking dark coloration in the gular region. LSUHC 7446 (juvenile) differs from the holotype by lacking a distinct dorsal color pattern and coloration in the gular region. BMNH 1932.10.4.5 (male) differs from the holotype in having a more prominent dorsal pattern with more marbled markings, lacking the dark coloration in the gular region. BMNH 1915.8.14.11 (female) differs from the holotype in having larger marbled patterns on the dorsum and in having less coloration in the gular region. BMNH 61.4.12.43 (male) differs from the hollow type in having a much lighter dorsal pattern and by lacking coloration in the gular region. The black eye patch and dorsal pattern is extremely faded. Scalation and morphometric variation is presented in Table 6. Distribution. Acanthosaura cardamomensis occurs in the Cambodian and Thai Cardamom Mountains, with populations in Cambodia at Bokor National Park, Kampot Province; Phnom Samkos, Pursat Province; Phnom Dalai, Pursat Province; Phnom Aural, Kampong Speu Province to Thmai Baing Koh kang province. In Thailand this species is known from and Koh Chang Island, Trat Province; Khao Soi Dao District, Chantaburi Province; Khao Kitchakut National Park, Chantaburi Province; Amphoe Pak Thong Chai, Sakaerat, Nakhon Ratchasima Province; Khao Yai National Park, Nakhon Ratchasima Province, (Fig. 1). Natural history. This species can be found both during the day and at night. Individuals have been found on various substrates such as ferns approximately 0.5 meters above the ground, leaf litter, on sapling trees sleeping at night 2 m above the forest floor, and sleeping on vines 0.75 m up near the streams edge. Juveniles have been seen running across the leaf litter, found under pieces of bark in the leaf litter, and on small logs low to the ground. The majority of the juveniles were found close to the ground. When approached or provoked during the day some lizards will climb up the trees while others may drop to the ground and seek refuge under fallen or within hollow logs. Acanthosaura cardamomensis ranges from 400-1400 m a.s.l. The holotype is a gravid female carrying 18 eggs, collected during August 2005. A gravid female (FMNH 180309) collected on 10 July 1969 contained 23 eggs. We hypothesize that this species is breeding year round based on a number of juveniles collected at various times of the year: FMNH 180308, 10 May 1969; FMNH 263335, 30 January 2004; LSUHC 7394-95, 6 August 2005. Etymology. The specific epithet cardamomensis is in reference to the type locality, the Cardamom Mountains. Comparisons to other species. Acanthosaura cardamomensis most closely resembles A. crucigera but can be differentiated from it by having significantly longer spines in the nuchal crest to head length ratio (NS/ HL, 1.7–66% vs. 2.0–38%, n=33; p< 0.008), longer postorbital spine to the head length ratio (PS/HL, 14–45% vs. 14–33%, n=34; p< 0.039), longer dorsal spine to head length ratio (DS/HL, 12–49% vs. 9–24%, n=36; p< 0.001) than A. crucigera. Acanthosaura cardamomensis has a significantly smaller diastema to snout vent length ratio than A. crucigera (DIASVL, 3–7% vs. 5–8%, n=37; p< 0.014). Acanthosaura cardamomensis has significantly more infralabials (INFRAL, 10-14 vs. 10, n=39; p< 0.001), more canthus-rostralis-supracilliary scales (CS, 11–17 vs. 12–13, n=39; p< 0.039), more supralabials (SUPRAL, 11–15) 10–12, n= 39; p< 0.01) than A. crucigera. Acanthosaura cardamomensis has significantly fewer scales in the diastema (DIASN, 6–15 vs.12–25, n=38; p< 0.001) and subdigital lamellae beneath the fourth toe compared to A. crucigera (TO, 20– 25 vs. 23-26, n=39; p< 0.010). These data are summarized in Table 1–5.

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 33 TABLE 6. Selected measurements (in mm) for the type series of Acanthosaura cardamomensis. For character abbreviations see materials and methods and Table 5.

Sex LSUHC LSUHC LSUHC FMNH FMNH FMNH LSUHC BMNH BMNH BMNH 7447 7877 7406 180309 263260 180307 7446 1932.10.4. 1915.8.14.1 61.4.12.43 Holotype Paratype Paratype Paratype Paratype Paratype Paratype 5 Paratype 1 Paratype Paratype Female Male Male Female Female Male Juvenile Male Female Male SVL 149.4 133.1 125.5 136 130.6 105 92.6 118 99 102 TL 174.3 213.5 188.7 186 173.6 153 134.1 174 116 164 HL 42.4 26.1 36.8 25.6 25.4 20.3 29.5 22.8 21.8 21.3 HW 26.1 25.7 23.5 26.7 22.2 20.4 17.5 24.1 23 18.4 HD 21.7 23.4 18.4 20.9 18.5 17.1 15 20.5 16.7 15.3 SL 18.7 16.2 15 13.1 15.2 10.1 12.8 10.2 9 9.6 ORBIT 14.5 13.9 11.3 11.4 10.5 10.6 9.1 11 10.9 10.1 TD 5.8 5 5.6 3.6 4.8 4 3.9 5 4.1 3.9 TN 1 1 1 1 1 1 1 1 1 1 PS 12.7 11.8 11.1 8.5 6.8 4.5 9.6 10.2 3.2 5.1 PS/HL 0.3 0.45 0.3 0.33 0.27 0.22 0.32 0.44 0.14 0.24 NS 16.2 12.6 17 9 9.1 8 10.2 9.8 3.9 7.1 NS/HL 0.38 0.48 0.46 0.35 0.35 0.39 0.35 0.42 0.17 0.33 NSL 5.4 5.9 6.3 4 3.2 3.5 2.9 3.8 3 3.9 DS 18.1 10.4 14.2 7 5.9 5.3 8.9 6.2 6.3 5.6 DS/HL 0.43 0.4 0.38 0.27 0.23 0.26 0.31 0.27 0.28 0.26 DSL 2 5.5 2 3.1 4 3.3 2.3 2.8 2.4 3.4 WN 2.3 2.7 2.74 3.5 3.2 3 2.31 4.2 1.6 1.6 WDS 3.3 2 3.2 2.3 2.4 2.2 2 2.1 1.3 5.6 DIAS 8.3 5.4 5.3 4.6 5 4.1 3.8 8 4.2 3.3 DIASN 11 10 14 6 11 10 11 11 12 9 DIAS/SVL 0.06 0.04 0.042 0.033 0.038 0.039 0.028 0.067 0.042 0.031 FOREL 56.2 56.8 48.8 49.6 49.9 43.3 40.7 43 29 46 HINDL 76.8 77.1 63.4 68.3 65.8 56.6 55.4 59 42 60 SUPRAL 13 13 14 13 13 11 13 12 12 13 INFRAL 12 11 13 11 11 11 11 11 12 13 VENT 60 57 65 52 55 57 53 58 57 58 FI 17 17 17 15 15 17 17 17 17 16 TO 22 22 23 23 22 24 21 22 23 20 TL/SVL 1.2 1.6 1.5 1.4 1.3 1.5 1.4 1.5 1.17 1.6 OS 12.1 10.7 11.7 8.4 9.2 6.7 7.6 - 4 6.6 OS/HL 0.29 0.4 0.32 0.33 0.36 0.33 0.26 - 0.18 0.31 NSSOS 4 5 4 4 5 6 4 - 4 4 CS 13 14 14 14 13 14 11 16 13 15 RW 4.6 3.5 3.8 3.8 4.1 3.4 3.4 3.9 3.2 3.3 RH 2.1 2.1 1.9 1.9 1.8 1.4 1.4 2 1.4 1.8 RS 6 8 8 8 6 8 9 8 8 7 NS 8 8 8 7 7 8 8 8 8 8 NCS 14 9 13 9 12 10 15 10 14 10 NSCSL 10 7 9 8 9 8 11 7 7 7 continued next page

34 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL. TABLE 6. (continued) Sex LSUHC LSUHC LSUHC FMNH FMNH FMNH LSUHC BMNH BMNH BMNH 7447 7877 7406 180309 263260 180307 7446 1932.10.4. 1915.8.14.1 61.4.12.43 Holotype Paratype Paratype Paratype Paratype Paratype Paratype 5 Paratype 1 Paratype Paratype Female Male Male Female Female Male Juvenile Male Female Male NR 2 1 2 1 1 1 1 1 1 2 NSSLC 13 10 11 13 12 12 16 11 15 11 MW 1.8 0.9 1.5 1.6 1.9 1.4 1.2 1.6 1.7 1.8 MH 1.6 1.3 1.2 1.6 1.6 1.4 1.2 1.2 1.6 2 PM 4 4 5 4 4 4 4 4 4 4 YAS 1 1 1 1 1 1 1 0 0 0 ND 1 1 0 1 1 1 1 1 1 1 LKP 1 1 1 1 1 1 1 1 1 1 BEP 1 1 1 1 1 1 1 1 1 1 ESBO 0 0 0 0 0 0 0 0 0 0 GP 3 4 4 2 2 2 2 4 2 2 OF 1 1 1 1 1 1 1 1 1 1

Acanthosaura cardamomensis differs from A. armata by having significantly larger occipital spine to head length ratio (OS/HL, 14–45% vs. 22–56%, n=42; p< 0.002), larger nuchal scale length to head length ratio (NS/HL, 17–66% vs. 22–51%, n=41; p< 0.009 Acanthosaura cardamomensis has a significantly smaller postorbital spine to head length ratio (PS/HL, 14–45% vs. 22–56%, n=42; p< 0.001) than A. crucigera. Acanthosaura cardamomensis has significantly more scales between the diastema (DIASN, 6–15 vs. 1–8, n=47; p< 0.001), more supralabials (SUPRAL 11–15 vs. 10–14, n=48; p< 0.017), more canthus-rostralis- supracilliary scales (CS, 11–17 vs. 11–15, n=48; p< 0.008), more subdigital lamellae beneath the fourth finger (FI, 15-20 vs. 14-17, n=48; p< 0.003), and more scales bordering the occipital spine (NSSOS, 4–6 vs. 4–5, n=46; p< 0.022) than A. crucigera. Acanthosaura cardamomensis has significantly fewer infralabial scales (INFRAL, 10–14 vs. 12–15, n=48; p< 0.001), fewer scales in between the nasals (NS, 6–10 vs. 7–10, n=48; p< 0.022), fewer ventral scales (VENT, 50–65 vs. 51–68, n=47; p< 0.001), and fewer subdigital lamellae beneath the fourth toe (TO, 20–25 vs. 19–26, n=48; p< 0.05) than A. crucigera. Acanthosaura cardamomensis has a black eye patch, which is absent in A. armata. Acanthosaura cardamomensis differs from A. bintangensis by having a larger dorsal spine to the head length ratio (DS/HL 14–45% vs. 8–9%), fewer subdigital lamellae beneath the fourth finger (FI, 15–20 vs. 23), a larger occipital spine to head length ratio (OS/HL, 24–56% vs. 10–11%), and by having a light knee patch and lacking an elliptical scale below the orbit. Acanthosaura cardamomensis differs from A. capra by having more supralabial scales (SUPRAL, 11–15 vs. 10), by having an occipital spine verses its absence in A. capra, and by not having uniform dorsal scales as in A. capra. Acanthosaura cardamomensis differs from A. coronata by having a dorsal crest, postorbital spines, nuchal spines, dorsal spines, occipital spines, nuchal diamond, black eye patch, gular pouch and a diastema. Acanthosaura cardamomensis can further be differentiated from A. coronata by having more subdigital lamellae beneath the fourth finger (FI, 15–20 vs. 13–14), more subdigital lamellae beneath the fourth toe (TO, 20–25 vs. 17–19), more scales between the fifth canthal scale to the fifth supralabial (NSCSL, 7–12 vs. 5–6), and having fewer scales between the nasals and the rostrals (NR, 1–2 vs. 3–4). Acanthosaura cardamomensis differs from A. nataliae by having a longer diastema (DIAS, 2.7–8.3 mm vs. 2.5 mm), an occipital spine, black nuchal diamond, and light knee patches. Acanthosaura cardamomensis differs from A. lepidogaster by having a larger postorbital spines to head length ratio (PS/HL, 14–45% vs. 10–11%), by having a larger nuchal scale to the head length ratio (NS/HL, 17–66% vs. 15) having a larger dorsal spine to head length ratio (DS/HL, 14–45% vs. 7–12%) and by having a larger occipital spine to head length ratio (OS/HL, 24–56% vs. 14–15%).

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 35 Acanthosaura cardamomensis differs from A. titiwangsaensis by having larger dorsal spine to head length ratio (DS/HL, 14–45% vs. 7–9%), having a larger occipital spine divided by the head length ratio (OS/HL, 24–56% vs. 9–10%) and by having a light knee patch present.

Discussion

The taxonomic diversity within the genus Acanthosaura is well underestimated (Ananjeva et al. 2008). Within the past five years three new species have been described and one synonym has been resurrected (Wood et al. 2009; Orlov et al. 2006; Stuart et al. 2006). The Acanthosaura crucigera group is wide ranging and its morphological variation is conserved. It is not surprising to find cryptic diversity within the A. crucigera complex. A thorough and careful examination of museum specimens within this complex is needed to fully understand the cryptic diversity within the group. A robust phylogenetic analysis of all extant species of the genus Acanthosaura is needed to provide a working taxonomic framework for the genus. From the robust phylogenetic tree we can further understand the species level relationships of the group and it can subsequently be used to help delineate species within the group as a whole (Wood et al. in prep).

Acknowledgements

We wish to thank Fauna and Flora International and the General Department of Administration for Nature Conservation and Protection of the Ministry of the Environment, Cambodia, for facilitating our fieldwork. Exportation permits were granted by His Execllebcy, Chay Samith, a delegate of the Royal Government of Cambodia in charge as General Director. We would like to thank Timothy M. Youmans, Armando Ponce, Thomas R. Szutz and all of the villagers for assistance in the field. We owe thanks to the curators K. K. P. Lim (ZRC), H. Voris and A. Resetar (FMNH), Kumthorn Thirakhupt (CUHC), Colin McCarthy (BMNH), and Tanya Chan-ard (THNHM) for their generosity in loaning specimens. For and advice and help with statistical questions we thank Joshua LaPergola. We thank Aaron M. Bauer for critical comments on the manuscript. This research was supported in part by a grant to LLG from the National Geographic Society (8487-08) and the College of Arts and Sciences, La Sierra University. We would like to thank the Villanova University’s Department of Biology for funding to support lab work.

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36 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL. from Southwestern Cambodia. Journal of Herpetology. Grismer, L.L., Youmans, T.M., Wood, Jr., P.L., Ponce, A., Johnson, R., Wright, S.B. & Norsham, S.Y. (2006) Checklist of the herpetofauna of Pulau Langkawi with taxonomic comments. Hamadryad, 29, 15–32. Grismer, L.L., Chav T., Neang T., Wood Jr, P.L., Grismer, J.L., Youmans, T.M., Ponce, A., Daltry, J.C. & Kaiser, H. (2007) The herpetofauna of the Phnom Aural Wildlife Sanctuary and checklist of the herpetofauna of the Cardamom Mountains, Cambodia. Hamadryad, 31, 216–241. Grismer, L.L., Neang, T., Chav, T., Wood, Jr, P. L., Oaks, J.R., Holden, J., Grismer, J.L., Szutz, T.R. & Youmans, T.M. (2008a) Additional amphibians and reptiles from the Phnom Samkos Wildlife Sanctuary in the northwestern Cardamom Mountains, Cambodia, with comments on their and the discovery of three new species. The Raffles Bulletin of Zoology, 56, 161–175. Grismer, L.L., Neang, T., Chav, T. & Grismer, J.L. (2008b) Checklist of the amphibians and reptiles of the Cardamom region of southwestern Cambodia. Cambodian Journal of Natural History 2008 (1) 12–28. Günther, A.C.L.G. (1861) Second list of Siamese reptiles. Proceedings of the Zoological Society of London, 1861(8), 135–140. Günther, A.C.L.G. (1864) The Reptiles of British India. Ray Society, London, xxviii, 452, 26 lithographed plates, 19 text figs. Hardwicke, T. & Gray, E.J. (1828) Synopsis of the species of saurian reptiles collected in India. Zoological Journal, 3: 213. Jackman, T.R., Bauer, M.A., Greenbaum, E., Glaw, F. & Vences, M. (2008) Molecular phylogenetic relationships among species of the Malagasy-Comoran gecko genus Paroedura (Squamata: Gekkonidae). Molecular Phylogenetics and Evolution 46, 74–81. Kalyabina-Hauf, S., Ananjeva, N.B., Joger, U., Lenk, P., Murphy, R.W., Stuart, B.L., Orlov, N.L., Ho, C.T. & Wink, M. (2004) Molecular phylogeny of the genus Acanthosaura (Agamidae). Current Herpetology. 23, (1), 7–16. Leong, T.M., Grismer, L.L. & Mumpuni (2002) Preliminary checklists of the herpetofauna of the Anambas and Natuna Islands (South China Sea). Hamadryad, 27(2), 165–174. Macey, J.R., Larson, A., Ananjeva, N.B., Fang, Z. & Papenfuss, T.J. (1997) Two novel gene orders and the role of light- strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular Biology and Evolution 14, 91–104. Maddison, W.P. & Maddison, D.R. (1992) MacClade version 3: Analysis of Phylogeny and Character Evolution. Sinauer Associates, Sunderland, Massachusetts. Orlov, N.L., Truong, N.Q. & Nguyen, V.S. (2006) A New Acanthosaura allied to A. capra Günther, 1861 (Agamidae, Sauria) from Central Vietnam and Southern Laos. Russian Journal of Herpetology 13(1), 61–76. Pauwels, O.S.G., David, P., Chucheep, C. & Kumthorn, T. (2003) Reptiles of Phetchaburi Province, western Thailand: a list of species, with natural history notes, and a discussion on the biogeography at the Isthmus of Kra. The Natural History Journal of Chulalongkorn University, 3(1), 23–53. Smith, M.A. (1935) The fauna of British India, Including Ceylon and Burma. Reptilia and Amphibia, Vol II. Sauria. Taylor and Francis, London xiii + 440 pp., 1 pl. Stuart, B.L. & Emmett, D.A. (2006) A collection of amphibians and reptiles from the Cardamom Mountains, Southwestern Cambodia. Fieldiana, No. 109, 1–17. Stuart, B.L., Sok, K. & Neang, T. (2006) A collection of amphibians and reptiles from hilly eastern Cambodia. The Raffles Bulletin of Zoology, 54, 129–155. Swofford, D.L. (2002) PAUP* Phylogenetic Analysis Using Parsimony (* and other Methods). Sinauer Associates, Sunderland, Massachusetts. Taylor, E.H. (1963) The lizards of Thailand. Kansas University Science Bulletin, 46(5), 687–1077. Manthey, U. (2008) Agamid Lizards of Southern Asia, 1, (Terralog 7a). Edition Chimaira/ Verlag ACS GmbH, Frankfurt/M./Rodgau. Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Südostasiens. Natur und Tier-Verlag, Münster, 512 pp. Wood, P.L. Jr., Grismer, J.L., Grismer, L.L., Norhayati, A., Onn, C.K. & Bauer, A.M. (2009) Two new montane species of Acanthosaura Gray, 1831 (Squamata: Agamidae) from Peninsular Malaysia. Zootaxa, 2012, 28–46.

A NEW SPECIES OF ACANTHOSAURA Zootaxa 2488 © 2010 Magnolia Press · 37 Appendix

Acanthosaura armata Hardwicke & Gray, 1827. UKMHC 0636, Gunung Panti, Johor, West Malaysia; ZRC 2.310, Tebing Tinggi, Kelantan, West Malaysia. BMNH 1946.8.13.88, 1946.8.13.85, Types “Singapore”. LSUHC 5079, Gubir, Kedah, West Malaysia. LSUHC 6707, 6723-25, 6767, Pulau Pinang, Penang, West Malaysia. LSUHC 5668- 69, 5676, 5679, Temengor Forest Reserve, Perak, West Malaysia. Acanthosaura bintangensis Wood et al. 2008. LSUHC 9045, Bukit Larut, Perak, Peninsular Malaysia; ZRC 2.309 Bukit Larut, Perak, Peninsular Malaysia. FMNH 143140 Bukit Larut, Tuping, Perak, Peninsular Malaysia. Acanthosaura cardamomensis sp. nov. BMNH 1932.10.4.5, Bokor, Kamchay Mountains, Cambodia. FMNH 263226, 263261-62, 263227, Bokor National Park, Kampot province, Cambodia. LSUHC 7877-78, Camp 1, Pursat Province, Cambodia. BMNH 61.4.12.43, Chantaboum. THNHM 13719, Khao Soi Dao, Chanthaburi, Thailand. FMNH 1800309, Khao Yai National Park, Nakon Ratchasima Province, Thailand. BMNH 1915.8.14.11, Koh Chang Island, South East Siam. LSUHC 7337, 7393-95, 7406, 7446-47, 7451, 7457, 7466, Phnom Aural, Kampong, Speu Province, Cambodia. FMNH 180310-15, Sakaerat, Amphoe Pak Thong Chai, Nakon Ratchasima Province, Thailand. FMNH 180304-08, Sakaerat Express Station, Amphoe Pak Thong Chai, Nakon Ratchasima Prov, Thailand. FMNH 263335, Thmar Baing District, Koh Kong Province, Cambodia. Acanthosaura capra Günther, 1861. BMNH 1946.8.13.90, 1946.8.13.83, lectotype and paralectotype Chantaboum. Günther, 1861. FMNH 253409-10, Ankhe District, Gia-Lai province, Vietnam. BMNH 1946.8.13.81-82 Types Chantaboum. Acanthosaura crucigera Boulenger, 1885. CAS 243923, along gas pipeline, SE of Khotama Camp, Yephyu Township, Dawei District, Tanintharyi Division, Myanmar. BMNH 1946.8.19.64, 1946.8.13.84, Types Tavoy, Burma. BMNH 87.11.2.4-5 Tavoy, Tennaserm, Burma. CUMZR 2008.05.26.1, Kaeng Krachan National Park, Phetchaburi Province, Thailand. Acanthosaura lepidogaster Cuvier, 1829. LSUHC 4111, 4139-40, Diao Luo Shan, Hainan Island, China. Acanthosaura nataliae Orlov, Nguyen & Nguyen, 2006. LSUHC 8514, Vietnam. Acanthosaura titiwangsaensis Wood et al. 2008. LSUHC 9109, 9094, Bishop’s Trail, Fraser’s Hill, , Peninsular Malaysia. ZRC 2.5994 , Pahang, Telom Valley, Peninsular Malaysia.

Genetic material Acanthosaura crucigera Boulenger, 1885. CUMZR 2008.05.26.1, Kaeng Krachan National Park, Phetchaburi Province, Thailand (GenBank HM 143889); CAS 229582 Botpyinn Township, Pakchan Reserve Forest, Main Ma The camp, Kawthaung District, Tanintharyi Division, Myanmar (GenBank GU817389). Acanthosaura cardamomensis Wood, Grismer, Grismer, Neang, Chav Holden, 2010. FMNH 263335, Thmar Baing District, Koh Kong Province, Cambodia (GenBank GU817391). FMNH 263262, 263225-27, 263261, Kampot District, Kampot Province, Cambodia (GenBank GU817393, GU817397-99, GU817400). LSUHC 7337, 7393 Phnom Aural, Kampong, Kampong Speu Province, Cambodia (GenBank GU817392, GU817394). LSUHC 7877- 78, Camp 1, Pursat Province, Cambodia (GenBank GU817395-96).

38 · Zootaxa 2488 © 2010 Magnolia Press WOOD JR. ET AL.