Acanthosaura phuketensis (Squamata: Agamidae), a ...

Zootaxa 4020 (3): 473–494 www.mapress.com /zootaxa / Copyright © 2015 Magnolia Press

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http://dx.doi.org/10.11646/zootaxa.4020.3.4 http://zoobank.org/urn:lsid:zoobank.org:pub:893ACB72-A8EF-4330-BA59-5D88F46B88D0

Acanthosaura phuketensis (Squamata: Agamidae), a new long-horned tree agamid from southwestern Thailand OLIVIER S. G. PAUWELS1, MONTRI SUMONTHA2, KIRATI KUNYA3, AWAT NITIKUL4, PHAMON SAMPHANTHAMIT5, PERRY L. WOOD, JR. 6 & LEE L. GRISMER7 1

Département des Vertébrés Récents, Institut Royal des Sciences naturelles de Belgique, Rue Vautier 29, B-1000 Brussels, Belgium. E-mail: [email protected] 2 Ranong Marine Fisheries Station, 157 Saphanpla Rd., Paknam, Muang, Ranong, Thailand. E-mail: [email protected] 3 Nakhonratchasima Zoo, ZPO, 111 M. 1, Ratchasima – Pak Tongchai Rd., Chaimongkol, Muang Nakonrajsima, Thailand. E-mail: [email protected] 4 Khao Phra Thaeo Non-hunting Area, Thalang, Phuket, Thailand 5 Gibbon Rehabilitation Project, Ban Bangrong, Thalang, Phuket, Thailand 6 Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA. E-mail: [email protected] 7 Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California, 92515-8247 USA. E-mail: [email protected]

Abstract We describe a new lowland forest-dwelling species of the genus Acanthosaura from Phuket Island and the Phuket mountain range in southwestern Thailand. Acanthosaura phuketensis sp. nov., the 11th species in the genus, seems most closely related to A. crucigera from Myanmar and western Thailand and A. cardamomensis from the Cardamom Mountains, but can be differentiated from them by a combination of morphological and coloration characteristics. This new discovery stresses the importance of preserving the last forest patches remaining on Phuket Island, home to three other squamate endemics. Key words: Thai-Malay Peninsula, Phuket mountain range, Phuket Island, Khao Phra Thaeo Non-hunting Area, Khao Lak–Lam Ru National Park

Introduction The genus Acanthosaura Gray includes ten currently recognized species, and globally extends from northeastern India, east to southern China including Hainan Island, and south, through the Indochinese and Thai-Malay peninsulas, to Sumatra, and the Anambas and Natunas Archipelagos (Wood et al. 2010; Ananjeva et al. 2011). Phylogenetic studies (Kalyabina-Hauf et al. 2004, Ananjeva et al. 2008) showed that the genus was in need of revision as it included several undescribed and cryptic species as revealed by molecular data. Until the present Century, the most widespread species, Acanthosaura crucigera Boulenger, 1885, whose type locality is Tavoy in Myanmar, was believed to extend from Myanmar east through Thailand, Cambodia, and southward to Peninsular Malaysia. However, populations from eastern Thailand and Cambodia have been shown to belong to a distinct species, A. cardamomensis Wood, Grismer, Grismer, Neang, Chav & Holden, 2010 (Wood et al. 2010; Pauwels & Iskandar 2010). In addition, Wood et al. (2009) and Grismer (2011) have indicated that populations from southern Thailand and Peninsular Malaysia might actually belong to five species, all related to, but distinct from, A. crucigera. So far only two of them were named: A. bintangensis Wood, Grismer, Grismer, Norhayati, Chan & Bauer, 2009 and A. titiwangsaensis Wood, Grismer, Grismer, Norhayati, Chan & Bauer, 2009. Geographically situated south of the distribution of populations regarded as belonging to true A. crucigera and west of these five species mentioned above, is a population with extremely long cephalic horns and dorsal spines found in Phuket Island, southwestern Thailand, that has been referred in the literature as A. cf. crucigera

Accepted by S. Carranza: 3 Sept. 2015; published: 23 Sept. 2015

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(Sumontha et al. 2012, 2015). Its taxonomic status is particularly confusing in the literature. Grossmann & Tillack (2000) mentioned that the fact that this population has so long horns and spines might be the reason why it was mentioned by Boettger (1887) as A. armata (Hardwicke & Gray, 1827), a species whose type locality is Singapore. Grossmann & Tillack (2000) recognized that this population is differentiable from all other populations of Acanthosaura crucigera. Leong et al. (2003), probably unaware of Boettger’s record of A. armata for Phuket, announced a first record of A. armata for the island, representing “the second species of Acanthosaura known to occur on the island, apart from A. crucigera Boulenger, 1885 (Grossmann and Tillack, 2000)”, adding to the confusion. In order to clarify the taxonomic status of the Phuket Island Acanthosaura population, we collected a series of voucher specimens from forested areas on the island, and compared them with other species and populations. Morphological and coloration data show that this population is clearly distinct from all other named species, and we consequently describe and name it hereafter.

Material and methods The type series was collected by hand and preserved in 70° ethanol. Fourty-three meristic and mensural characters were noted on each specimen of the type-series (see Table 1). Measurements were taken to the nearest 0.1 mm with a Mitutoyo digital caliper. Paired measurements were made on the left side, as was done in the recent revisions in the Acanthosaura crucigera species group (Wood et al., 2009, 2010; Ananjeva et al., 2011). Paired meristic characters are given left/right. The list and methodology of measurements and meristic counts follow Wood et al. (2010): BEP: presence (1) or absence (0) of a black eye patch; CS: number of canthus rostralis - supraciliary scales, counted from the nasal scale to the posterior end of the ridge at the posterior margin of the orbit; DIAS: length of the diastema, measured from the posterior end of the nuchal crest to the anterior end of the dorsal crest; 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; DS: maximum length of the largest spine in the dorsal crest measured from the base to the tip; ESBO: presence (1) or absence (0) of elliptical scales below the orbit; EYE: eye diameter, measured from the posterior to the anterior edge of the eye; FI: number of subdigital lamellae on the fourth finger; FOREL: forelimb length, measured from axilla to the proximal edge of the palmar region; GP: size of gular pouch scored as absent, small, medium or large; HD: maximum head height, measured across the parietal region; HINDL: hindlimb length, measured from groin to the proximal edge of the plantar region; HL: head length, measured from posterior edge of the rectis of the jaw to the tip of the snout; HW: head width, maximum head width, the width at the level of the tympanum; INFRAL: number of infralabials; LKP: presence (1) or absence (0) of light knee patch; MH: mental height; MW: mental width; NCS: number of scales between the fifth canthals; ND: presence (1) or absence (0) of a black, diamond shaped, nuchal collar; NR: number of scales between the nasals and the rostrals; NS: number of scales between the nasals; NSCSL: number of scales from the fifth canthal to the fifth supralabial; NSL: maximum length of the largest spine in the nuchal crest measured from the base to the tip; NSSLC: number of scales between the seventh supralabial and the sixth canthal; NSSOS: number of scales surrounding the occipital spine; OF: presence (1) or absence (0) of oblique fold anterior to the fore limb insertions; ORBIT: orbit diameter, measured from the posterior to the anterior edge of the orbit; OS: length of the occipital spine, measured from the base to the tip; PM: number of scales bordering the mental (wrongly defined by Wood et al. 2010 as the ‘number of postmental scales bordering the mental’ while they actually counted the total number of scales bordering the mental as shown in their Table 5); PS: postorbital spine length, measured from the base to the tip of the spine; RH: rostral height; RS: number of scales bordering the rostral scale; RW: rostral width; SL: snout length, measured from the anterior edge of the orbit to the tip of the snout; SUPRAL: number of supralabials; SVL: snout-vent length, measured from the tip of the snout to the tip of the vent; TaL: tail length, measured from the posterior margin of the vent to the tip of the tail; TBW: tail base width, maximum width at tail base (in addition to characters abreviations listed in Wood et al. 2010); TD: tympanum diameter, measured horizontally from the anterior to the posterior border of the tympanum; TN: scales absent on tympanum (0) or present (1); TO: subdigital lamellae on the fourth toe; VENT: number of ventral scales counted at the midline from the anterior edge of the shoulders to the edge of the vent; WNC: maximum width of the spines in the nuchal crest, measured at the base; YAS: presence (1) or absence (0) of a Y-shaped arrangement of enlarged scales on the snout. We compared our new material with museum specimens of all currently recognized species of Acanthosaura but A. brachypoda Ananjeva, Orlov, Nguyen & Ryabov, 2011;

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see Appendix. Institutional abbreviations: BMNH, Natural History Museum, London, United Kingdom; CUMZ R, Chulalongkorn University Museum of Zoology, Reptile Collection, Bangkok, Thailand; IRSNB, Institut Royal des Sciences Naturelles de Belgique, Brussels; MNHN, Muséum National d’Histoire Naturelle, Paris; PSUZC-RT, Prince of Songkhla University Zoological Collection, Reptile Section, Songkhla, Thailand; QSMI, Queen Saovabha Memorial Institute, Thai Red Cross Society, Bangkok, Thailand; RFD, Royal Forest Department, Bangkok, Thailand; THNHM, Natural History Museum, National Science Museum, Technopolis, Pathum Thani, Thailand.

Systematics Acanthosaura phuketensis sp. nov. Figures 1–12 Acanthosaura armata: Boettger 1887: 38. Acanthosaura cf. crucigera: Chan-ard et al. 1999: 90, left. Acanthosaura crucigera: Grossmann & Tillack 2000: 30–31, 36, 38. Acanthosaura crucigera: Pauwels et al. 2000: 132–133. Acanthosaura crucigera: Pauwels et al. 2002: 27. Acanthosaura crucigera: Manthey 2008: 24: RA00044-4, RA00046-3; 25: RA00047-4. Acanthosaura cf. crucigera: Sumontha et al. 2012: 69. Acanthosaura cf. crucigera: Sumontha et al. 2015: 110.

Holotype. Ethanol-preserved whole adult male individual, THNHM 22542 (formerly Montri Sumontha’s field number MS 560) from Ban Bangrong (coordinates UTM UPS ca. 47 p0433130 0888891 = 08o02.465N 98o23.588E; altitude ca. 41 m asl), Thalang District, Phuket Island, Phuket Province, southwestern Thailand. Collected by Awat Nitikul on 2 June 2008. Paratypes. Five ethanol-preserved whole individuals, one adult male (hemipenes everted) IRSNB 2681 (formerly IRSNB 15141) from Phang-Nga Wildlife Breeding Station, Muang District, Phang-Nga Province, southwestern Thailand, collected by O. S. G. Pauwels on 26 July 1998, and two adult females PSUZC-RT 2012.5 and THNHM 22543 (formerly MS 559 and MS 225 respectively) and two subadult males, PSUZC-RT 2012.6 and QSMI 1166 (formerly MS 558 and MS 557 respectively) from the same locality, collecting date and collector as for the holotype. Diagnosis. A medium-sized species (maximum SVL 123.5 mm) with a single long cylindrical spine above posterior margin of eye; large spine on occiput between tympanum and nuchal crest; tympanum naked, roundish, large; moderately developed gular pouch, black in females; scales on flanks randomly intermixed with medium and large scales; nuchal crest present and strongly developed; 12-17 scales in the diastema between the nuchal and vertebral crests; vertebral crest strongly developed, composed of enlarged, pointed scales beginning at shoulder region and continuing till first 6th of tail while regularly decreasing in size; tail 1.4–1.7 times SVL; black collar present; black eye patch present, extending posteriorly to reach the nuchal crest. Description of the holotype. Adult male. SVL 123.5 mm. TaL 205.6 mm, tail complete. Head length 31.4 mm; head moderately long (HL/SVL 25%), somewhat narrow (HW/SVL 18%), not tall (HD/HL 0.59), triangular in dorsal and lateral profile. Snout short (SL/HL 35%); interorbital and frontal regions and rostrum wide (RW/RH 238%), steepy sloping anteriorly. Canthus rostralis prominent, forming a large projecting shelf extending above eye, composed of 14/14 enlarged scales; shelf terminates with a notch anterior to the postorbital spine. Rostral moderate in size, rectangular, bordered laterally by first supralabials and posteriorly by five smaller scales; nasal concave, surrounded by 1/1 prenasal anteriorly and 3/3 postnasals posteriorly; seven scales between nasal scales; supranasals somewhat elongate; enlarged scales above orbit weakly keeled; no enlarged row of keeled scales below orbit. Eye relatively large (EYE/HL 20%), orbit very large (ORBIT/HL 33%). Prefrontal and frontal scales slightly keeled and larger than scales between orbit and supralabials; scales on occiput weakly keeled; no enlarged parietal eyespot. Extremely elongate epidermal spine above posterior margin of eye, posteriorly recurved, surrounded by 6/ 5 enlarged scales. Suborbitals small, slightly keeled, extending by a row of 6/6 enlarged keeled scales from below posterior margin of eye to anterior margin of tympanum, increasing in size posteriorly. Elongate epidermal spine on

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FIGURE 1a. Lateral view of preserved Acanthosaura phuketensis sp. nov. adult male holotype. Photo by O. S. G. Pauwels. FIGURE 1b. Ventral view of preserved Acanthosaura phuketensis sp. nov. adult male holotype. Photo by O. S. G. Pauwels.

lateral margin of nape, anteriorly recurved (the left one broken near its base and healed), surrounded by a rosette of 5/5 much shorter spiny scales, themselves surrounded by a row of pointed scales. Tympanum exposed, oblong, about as wide as tall, surrounded by small conical scales. SL 11/11, rectangular, slightly decreasing in size posteriorly; mental squarish above, becoming triangular below, larger than 1st pair of IL; 2 postmentals of comparable size, widely in contact behind mental; 4 scales contacting the mental; chinshields enlarged, extending posteriorly past angle of jaw, separated from infralabials by one scale row anteriorly and three at angle of jaw; IL 12/12, rectangular, scales in center of series largest; gulars sharply keeled and spinose with 3 enlarged midventral rows. Dewlap very slightly extensible, gular pouch moderate. Nuchal crest composed of 7 very elongate, lanceolate scales bordered on each side by 2–3 rows of enlarged, spinose scales; nuchal crest followed by a diastema of 17 scales at base of nape. Dorsal body crest very developed, extending from posterior margin of diastema onto first sixth of tail; vertebral crest composed of enlarged, epidermal, laterally compressed, spinose scales, bordered by a


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single row of smaller paravertebral spinose scales; vertebral crest tapers slightly to base of tail, then fades progressively. Body moderately developed, triangular in cross-section. Dorsal scales small, medium-sized and large mixed without regular pattern, with keels projecting posteriorwards; scales of pectoral region and abdomen larger than dorsals, keeled, more or less arranged in transverse rows; keeled scales anterior to vent not enlarged. Limbs relatively long; dorsal and ventral scales of forelimbs keeled, spinose, about the same size. Five digits on manus; subdigital scales keeled, subdigital lamellae under 4th finger 17/16. Scales of hind limbs keeled and spinose; postfemoral scales small, interspersed with larger spinose scales. Five digits on pes; subdigital scales keeled, subdigital lamellae under 4th toe 22/23. Tail length 1.7 times SVL, tail covered with keeled spinose scales, keels on subcaudals directed posteriorly; subcaudals much longer than supracaudals; base of tail 14.3 mm wide.

FIGURE 2. Left head side of preserved adult male paratype Acanthosaura phuketensis sp. nov. IRSNB 2681. Photo by O. S. G. Pauwels.

Variation. Morphometrical and meristic data for the type series are shown in Table 1. All type specimens show 2 postmentals. They also all show a nuchal crest composed of 7 lanceolate scales, although the size of these scales is much larger in the holotype/adult males than in the adult female or subadults. The throat of the two adult females is blackish, while it is beige with black longitudinal reticulations in the adult male holotype. Distribution. Within Phuket Island, Acanthosaura phuketensis sp. nov. was recorded so far only from the type locality and nearby in Khao Phra Thaeo Non-hunting Area (Fig. 13–14), but it probably occurs in other remaining forested areas of the island. Chan-ard et al. (1999: 90) illustrated two individuals, including an adult male, that they identified as Acanthosaura cf. crucigera, from Khao Lak National Park in Phang-Nga Province. That same adult male and an adult female were illustrated by Grossmann & Tillack (2000: 31) under Acanthosaura crucigera. One of the individuals already illustrated by Chan-ard et al. (1999: 90) plus an additional individual from Khao Lak were illustrated by Manthey (2008: 24). Khao Lak park is situated along the sea on the mainland, at ca. 65 km north of the type locality of Acanthosaura phuketensis sp. nov. Based on the coloration and scalation characteristics visible on these photographs, these individuals are referrable to Acanthosaura phuketensis sp. nov. The phylogeny presented by Kalyabina-Hauf et al. (2004) and Ananjeva et al. (2008) groups three A. crucigera samples from



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Takua Pa District with a sample (IRSNB 15141, here renumbered IRSNB 2681 and designated paratype of A. phuketensis sp. nov.) from ‘Thai Muang’ (in fact Muang) District, all from Phang-Nga Province. On the map these authors provided, the Takua Pa locality is erroneously shown as if it was located on Phuket Island. Manthey (2008: 25) also showed the photograph of an adult male from Khao Sok in Surat Thani Province. The mention of A. cf. crucigera from the type locality of Cyrtodactylus ranongensis Sumontha, Pauwels, Panitvong, Kunya & Grismer, 2015 by Sumontha et al. (2015) in Suk Samran District in Ranong Province is referrable to A. phuketensis sp. nov. The known localities of Acanthosaura phuketensis sp. nov., spreading over Phuket, Phang-Nga, Surat Thani and Ranong provinces, are all located on the Phuket Range, a 200 km long continuation of the Tenasserim Range. The range of the new species is surrounded by other populations of the crucigera group with shorter horns, as is well illustrated through the photographs from various populations and localities presented by Manthey (2008).

FIGURE 3a. Dorsal view of preserved Acanthosaura phuketensis sp. nov. adult female paratype PSUZC-RT 2012.5. Photo by O. S. G. Pauwels. FIGURE 3b. Ventral view of preserved Acanthosaura phuketensis sp. nov. adult female paratype PSUZC-RT 2012.5. Photo by O. S. G. Pauwels.

Natural history. Acanthosaura phuketensis sp. nov. is primarily an arboreal species, as shown on published in situ photographs (Chan-ard et al., 1999; Grossmann & Tillack, 2000; Manthey 2008) and according to our personal field observations. The Phuket type specimens were collected at night while they were asleep on large trees in mature secondary forest. However this species also regularly descends on the ground. Pauwels et al. (2000) found by day an individual under a stone in the dry bed of a stream, a few meters from another individual found active on the ground, in mature secondary forest. This diurnal species can be locally abundant; many individuals were


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FIGURE 4a. Dorsal view of preserved Acanthosaura phuketensis sp. nov. adult female paratype THNHM 22543. Photo by O. S. G. Pauwels. FIGURE 4b. Ventral view of preserved Acanthosaura phuketensis sp. nov. adult female paratype THNHM 22543. Photo by O. S. G. Pauwels.

observed in mature secondary forest at the type locality, in primary forest in Khao Phra Thaeo Non-hunting Area, and in primary and mature secondary forest in Phang-Nga Wildlife Breeding Station. At the type locality it was observed by us (AN, KK, MS, OSGP, PS) in syntopy with 23 species of squamates, namely Cnemaspis phuketensis Das & Leong and C. vandeventeri Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, Cyrtodactylus brevipalmatus (Smith), C. oldhami (Theobald), C. phuketensis Sumontha, Pauwels, Kunya, Nitikul, Samphanthamit & Grismer, 2012 (Gekkonidae), Bronchocela cristatella (Kuhl), Calotes emma Gray (Agamidae), Ahaetulla fasciolata (Fischer) and A. prasina (Boie), Boiga cyanea (Duméril, Bibron & Duméril) and B. cynodon (Boie), Dendrelaphis caudolineatus (Gray), D. cyanochloris (Wall), D. formosus (Boie), D. haasi Van Rooijen & Vogel and D. striatus (Cohn), Ptyas carinata (Günther) and P. fusca (Günther) (Colubridae), Aplopeltura boa A NEW TREE AGAMID FROM PHUKET


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(Boie), Pareas carinatus (Boie) (Pareatidae), Python brongersmai Stull (Pythonidae), Trimeresurus phuketensis Sumontha, Kunya, Pauwels, Nitikul & Punnadee, 2011, Tropidolaemus wagleri Wagler (Viperidae), and with four species of anuran amphibians, Ingerana tasanae (Smith), Limnonectes blythii (Boulenger) (Dicroglossidae), Hylarana eschatia (Inger, Stuart & Iskandar) (Ranidae) and Nyctixalus pictus (Peters) (Rhacophoridae). In Suk Samran District in Ranong Province it has been observed in secondary forest at the proximity of Bronchocela cristatella, Calotes emma (Agamidae), Cyrtodactylus brevipalmatus and C. ranongensis, Gehyra mutilata (Wiegmann), Gekko gecko (Linnaeus), Hemidactylus frenatus (Schlegel), H. garnotii Duméril & Bibron and H. platyurus (Schneider), Ptychozoon lionotum Annandale (Gekkonidae), Ahaetulla mycterizans (Linnaeus) and A. prasina, Boiga dendrophila (Boie) and B. drapiezii (Boie) (Colubridae), Rhabdophis nigrocinctus (Blyth), Xenochrophis trianguligerus (Boie) (Natricidae), Pareas carinatus (Pareatidae) and Malayopython reticulatus (Schneider) (Sumontha et al., 2015). The species seems to be strictly sylvicolous, and was never found in highly degraded secondary forest nor in plantations. Figure 9 shows a subadult male Acanthosaura phuketensis sp. nov. eating a large leaf-colored grasshopper (Orthoptera). In captivity, A. phuketensis sp. nov. accepts meal worms, crickets, woodlice and earthworms (Pauwels et al., 2000 and unpubl. obs.). The abdominal cavity of the paratype IRSNB 2681 contained numerous nematodes.

FIGURE 5a. Dorsal view of preserved Acanthosaura phuketensis sp. nov. subadult male paratype QSMI 1166. Photo by O. S. G. Pauwels. FIGURE 5b. Ventral view of preserved Acanthosaura phuketensis sp. nov. subadult male paratype QSMI 1166. Photo by O. S. G. Pauwels.


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FIGURE 6a. Dorsal view of preserved Acanthosaura phuketensis sp. nov. subadult male paratype PSUZC-RT 2012.6. Photo by O. S. G. Pauwels. FIGURE 6b. Ventral view of preserved Acanthosaura phuketensis sp. nov. subadult male paratype PSUZC-RT 2012.6. Photo by O. S. G. Pauwels.

Etymology. The specific epithet is an adjective in reference to Phuket Island and Phuket Range, on which the type locality lies. We suggest the following common names: Kingkakhaownaam Phuket (Thai), Phuket Horned Tree Agamid (English), Acanthosaure de Phuket (French), Phuket-Nackenstachler (German), Phuketstekelnekagame (Dutch). Comparisons to other species. Table 2 shows a comparison of morphometric and meristic data for all currently recognized species of Acanthosaura and A. phuketensis sp. nov. It is mostly based on the interspecific comparison table provided by Wood et al. (2010, Table 5) and additional data from Hallermann (2000; only data for lectotype A. fruhstorferi (Werner) included), Ananjeva et al. (2011) and Bobrov (2013). However, it is to be noted that Wood et al. (2010) wrongly defined the character ‘ORBIT’ used in their comparison table, as the ‘eye diameter, measured from the posterior to the anterior edge of the eye’, while they actually meant and measured the orbit diameter, not the eye diameter. About the same table presented by Wood et al. (2010), it should also be noted that there was a lapsus calami regarding the ratio TaL/SVL for A. coronata, erroneously indicated as varying from ‘0.06–1.01’, while the corrected variation has been used in our Table 2. A close re-examination of the material regarded by Wood et al. (2010) to represent A. crucigera led to separate populations from western Thailand and at



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FIGURE 7. Live adult male Acanthosaura phuketensis sp. nov. in situ at the type locality of Cyrtodactylus ranongensis in Suk Samran District, Ranong Province, southwestern Thailand. Photo by M. Sumontha.

and near its type locality in Myanmar from those from the Kra Isthmus area based on coloration and morphometrical characteristics, as well as on molecular data (Wood et al., in prep.). Only the northern populations are actually referrable to A. crucigera, showing, similarly to the type material, a relatively low nuchal crest, a low dorsal crest, and no contact between the black eye patch and the nuchal crest in both sexes, among other characteristics. The morphological variation for A. crucigera that was presented by Wood et al. (2010) in their interspecific comparison table has consequently been reviewed to include only what we regard here as true A. crucigera (see Appendix: Material examined). Acanthosaura phuketensis sp. nov. can be differentiated from A. armata by its higher maximal length of the postorbital spines (11.8 vs. 9.9 mm), its much higher number of scales in the diastema between the nuchal and the dorsal crests (12–17 vs. 1–8), a lower number of infralabials (10–12 vs. 12–15) and the presence of a black eye patch (vs. absence). It can be distinguished from A. bintangensis by having a larger tympanum (TD/HD 0.22–0.33 vs. 0.16), much longer postorbital spines (PS 4.6–11.8 vs. 1.9–4.2 mm, and PS/HL 0.23–0.38 vs. 0.07–0.19), longer spines in the nuchal crest (NSL/HL 0.21–0.39 vs. 0.17–0.21), longer spines in the dorsal crest (DS 2.3–8.3 vs. 1.8–2.2 mm, and DS/HL 0.11–0.26 vs. 0.08–0.09), more ventral scales (57–67 vs. 51–55), less subdigital lamellae on the fourth finger (15–17 vs. 23), less subdigital lamellae on the fourth toe (21–24 vs. 26–28), much longer occipital spines (OS 2.6–9.5 vs. 1.2–2.6 mm, and OS/HL 0.13–0.30 vs. 0.10–0.11), less scales surrounding the occipital spine (4–5 vs. 6–7), more scales between the fifth canthals (12–13 vs. 10–11), presence of a light knee patch (vs. absence), less developed gular pouch (GP 0–2 vs. 3–4) and absence of an enlarged row of keeled scales below orbit (vs. presence). From A. brachypoda, A. phuketensis sp. nov. can be differentiated by a larger tympanum (TD/HD 0.22–0.33 vs. 0.21), much longer postorbital spines (PS/HL 0.27– 0.28 in the two adult female paratypes of A. phuketensis sp. nov. vs. 0.11 in the adult female holotype of A. brachypoda), longer spines in the nuchal crest (NSL/HL 0.23 and 0.30 in the two adult female paratypes of A. phuketensis sp. nov. vs. 0.16 in the adult female holotype of A. brachypoda), much longer spines in the dorsal crest (DS/HL 0.11 and 0.14 in the two adult female paratypes of A. phuketensis sp. nov. vs. 0.06), a distinctly higher


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number of scales in the diastema between the nuchal and the dorsal crests (12–17 vs. 7), less subdigital lamellae on the fourth finger (15–17 vs. 18), a much longer occipital spine (OS/HL 0.19 and 0.24 in the two adult female paratypes of A. phuketensis sp. nov. vs. 0.03), and less scales between the nasals (NS 7–8 vs. 9). The character DIAS is given both as 4.5 and 1.9 mm in the original description of A. brachypoda, which is based on a single specimen, so this character cannot be used here for comparison. The methodology provided for measuring FOREL and HINDL in the original description of A. brachypoda is insufficiently described to allow verifying that these measurements were taken according to our methodology, so we could not use these characters here; CS, NCS, NR, NSCSL, NSSLC and NSSOS were not provided in the original description of A. brachypoda. A. phuketensis sp. nov. can be distinguished from A. capra (Günther, 1861) based on a generally larger tympanum (TD/HD 0.22–0.33 vs. 0.21–0.23), shorter spines in the nuchal crest (NSL/HL 0.21–0.39 vs. 0.42–0.43), slightly higher maximal length of dorsal crest spines (8.3 vs. 6.8 mm), much higher number of scales in the diastema between nuchal and dorsal crests (12–17 vs. 4–7), presence of occipital spines (vs. absence), lower number of scales between the nasals (7–8 vs. 9), higher number of scales between the fifth canthals (12–13 vs. 9), generally more scales between the seventh supralabial and the sixth canthal (11–14 vs. 9–11) and a notably less developed gular pouch (GP 0–2 vs. 3– 4). From A. cardamomensis, it can be separated based on a much lower maximal length of dorsal crest spines (8.3 vs. 14.2 mm, with a sudden decrease in size before dorsum mid-length in male A. phuketensis sp. nov., posterior to dorsum mid-length in male A. cardamomensis), a generally higher number of scales in the diastema separating the nuchal and the dorsal crests (12–17 vs 6–15), a lower maximal length of occipital spines (9.5 vs. 13.3 mm, and OS/ HL 0.13–0.30 vs. 0.24–0.56), a generally less-developed gular pouch (GP 0–2 vs. 1–4); another character to separate them is based on coloration: in male A. phuketensis sp. nov. the black eye patch extends posteriorly to reach the nuchal crest, while it never does in A. cardamomensis (see species’ description and photographs in Wood et al. 2010 and all photographs of A. cf. crucigera from Cardamom Mountains [= A. cardamomensis] in Manthey 2008). Acanthosaura phuketensis sp. nov. is distinguishable from A. coronata (Günther, 1861) based on a larger tympanum (0.22–0.33 vs. 0.14–0.17 mm), presence of postorbital spines, occipital spines, of nuchal and dorsal crests (vs. absence), higher number of subdigital lamellae on the fourth finger (15–17 vs. 13–14), higher number of subdigital lamellae on the fourth toe (21–24 vs. 17–19), longer tail (TaL/SVL 1.4–1.7 vs. 0.6–1.0), higher number of scales between the fifth canthals (12–13 vs. 8–11), higher number of scales from the fifth canthal to the fifth supralabial (8–10 vs. 5–6), lower number of scales between the nasals and the rostrals (1–2 vs. 3–4), generally higher number of scales between the seventh supralabial and the sixth canthal (11–14 vs. 6–11), presence of a black, diamond shaped, nuchal collar (vs. absence) and presence of a black eye patch (vs. absence). From A. crucigera, it is distinguished by a larger tympanum (TD/HD 0.22–0.33 vs. 0.14–0.21), a higher maximal length of postorbital spines (11.8 vs. 7.8 mm), a distinctly higher maximal length of occipital spines (9.5 vs. 4.9 mm), a higher maximal length of nuchal crest spines (12.2 vs. 8.9 mm), a higher maximal length of dorsal crest spines (8.3 vs. 5.5 mm); although several morphometrical characters of both species are overlapping (see Table 2), an easy character to separate them is based on coloration: in male A. phuketensis sp. nov. the black eye patch extends posteriorly to reach the nuchal crest, while it never does in A. crucigera (see species’ original description by Boulenger 1885). From A. lepidogaster (Cuvier, 1829), A. phuketensis sp. nov. can be separated based on its much longer postorbital spines (4.6–11.8 vs. 1.5–2.5 mm, PS/HL 0.23–0.38 vs. 0.06–0.11), much higher maximal length of nuchal crest spines (4.1–12.2 vs. 2.9–3.4 mm, NSL/HL 0.21–0.39 vs. 0.12–0.15) and of occipital spines (9.5 vs. 3.4 mm), its much higher maximal length of dorsal crest spines (8.3 vs. 2.7 mm, DS/HL 0.11–0.26 vs. 0.07–0.12), its generally lower number of subdigital lamellae on the 4th finger (15–17 vs. 17–19), higher number of scales between the fifth canthals (12–13 vs. 7–10), higher number of scales between the seventh supralabial and the sixth canthal (11–14 vs. 10) and lower number of scales in contact with the mental (4 vs. 5). A. phuketensis sp. nov. can be separated from A. nataliae Orlov, Nguyen & Nguyen, 2006 by its much lower maximal length of postorbital spines (11.8 vs. 17.8 mm), nuchal crest spines (12.2 vs. 23.8 mm, NSL/HL 0.21–0.39 vs. 0.58) and dorsal crest spines (8.3 vs. 17.7 mm, DS/HL 0.11–0.26 vs. 0.44), a longer diastema between nuchal and dorsal crests (3.6–7.6 vs. 2.5 mm, DIAS/SVL 0.05–0.08 vs. 0.04) with more scales (12–17 vs. 10), presence of occipital spines (vs. absence), lower number of scales between the fifth canthals (12–13 vs. 14), lower number of scales between the seventh supralabial and the sixth canthal (11–14 vs. 16), presence of a black, diamond shaped, nuchal collar (vs. absence), presence of light knee patch (vs. absence) and much lesser development of gular pouch (GP 0–2 vs. 4). From A. titiwangsaensis, it can be distinguished by its larger tympanum (TD/HD 0.22–0.33 vs. 0.17–0.20), its much longer postorbital spines (4.6–11.8 vs. 3.3–4.4 mm, PS/HL 0.23–0.38 vs. 0.14–0.18), its much longer



483

occipital spines (2.6–9.5 vs. 1.8–2.3 mm, OS/HL 0.13–0.30 vs. 0.09–0.10), its much higher maximal length of nuchal crest spines (12.2 vs. 4.4 mm, NSL/HL 0.21–0.39 vs. 0.11–0.18), its much longer dorsal crest spines (2.3– 8.3 vs. 1.7–2.1 mm, DS/HL 0.11–0.26 vs. 0.07–0.09), its higher number of ventrals (57–67 vs. 47–57), its lower number of subdigital lamellae on 4th finger (15–17 vs. 20–21), lower number of scales bordering the rostral scale (5–8 vs. 9), lower number of scales bordering the mental (4 vs. 5), presence of light knee patch (vs. absence) and lesser development of gular pouch (0–2 vs. 2–4). TABLE 1. Morphometrical (in mm) and meristic data for the type series of Acanthosaura phuketensis sp. nov. For character abbreviations see material and methods. Paired meristic characters are given left/right. NA = not applicable. Holotype THNHM 22542 Adult male

Paratype Paratype IRSNB 2681 PSUZC-RT Adult male 2012.5 Adult female

Paratype THNHM 22543 Adult female

Paratype QSMI 1166 Subadult male

Paratype PSUZC-RT 2012.6 Subadult male

SVL

123.5

96.3

118.4

100.8

69.2

74.2

TaL

205.6

>101.2

189.8

145.0

107.0

113.2

TaL/SVL

1.7

NA

1.6

1.4

1.5

1.5

TBW

14.3

12.1

14.5

9.2

5.5

5.4

HL

31.4

26.9

29.6

27.4

19.7

23.0

HW

22.8

18.0

21.2

19.1

14.4

14.7

HD

18.6

14.8

18.6

16.3

10.9

12.3

SL

11.0

10.2

10.1

9.3

6.8

8.3

ORBIT

10.5

9.7

11.2

9.0

6.6

7.3

EYE

6.3

6.5

4.7

4.0

3.7

3.7

TD

4.7

3.2

4.3

3.6

3.5

4.0

TD/HD

0.25

0.22

0.23

0.22

0.32

0.33

TN

0

0

0

0

0

0

PS

11.8

9.1

8.0

7.7

4.6

6.5

PS/HL

0.38

0.34

0.27

0.28

0.23

0.28

NSL

12.2

9.1

6.8

8.1

4.1

6.5

NSL/HL

0.39

0.34

0.23

0.30

0.21

0.28

DS

8.3

6.5

3.3

3.9

2.3

2.5

DS/HL

0.26

0.24

0.11

0.14

0.12

0.11

WNC

2.9

2.4

2.4

2.0

1.4

1.7

DIAS

7.6

4.5

6.5

7.6

3.6

3.8

DIAS/SVL

0.06

0.05

0.05

0.08

0.05

0.05

DIASN

17

13

12

13

15

14

FOREL

38.8

33.5

42.9

33.9

22.3

27.5

HINDL

60.3

50.4

60.6

49.1

38.2

38.6

SUPRAL

11/11

11/11

12/11

10/11

12/11

10/12

INFRAL

12/12

11/11

12/12

10/10

11/12

11/11

VENT

66

57

59

57

67

58

FI

17/16

16/17

16/17

15/15

17/17

17/17

TO

22/23

22/23

23/24

21/21

22/23

23/23

OS

9.5

7.3

5.6

6.7

2.6

5.1

OS/HL

0.30

0.27

0.19

0.24

0.13

0.22

NSSOS

5/5

4/4

4/5

4/4

4/5

5/5

CS

14/14

13/13

11/12

10/11

12/13

12/12

......continued on the next page


PAUWELS ET AL.

TABLE 1. (Continued) Holotype THNHM 22542 Adult male

Paratype Paratype IRSNB 2681 PSUZC-RT Adult male 2012.5 Adult female

Paratype THNHM 22543 Adult female

Paratype QSMI 1166 Subadult male

Paratype PSUZC-RT 2012.6 Subadult male

RW

3.8

2.4

3.8

3.5

2.3

2.3

RH

1.6

1.3

1.4

1.7

1.1

1.4

RS

7

5

8

6

7

5

NS

7

8

7

7

8

8

NCS

13

13

12

12

13

12

NSCSL

9/9

9/10

10/9

8/9

9/8

9/9

NR

1/1

2/2

1/1

1/1

1/1

2/2

NSSLC

12/12

11/13

11/11

14/11

11/12

11/12

MW

1.1

1.3

1.4

1.4

0.5

0.9

MH

1.1

1.2

1.6

0.9

0.6

1.0

PM

4

4

4

4

4

4

YAS

1

1, not very distinct



0

0

ND

1

1

1

1

1

1

LKP

1

1

1

1

1

1

BEP

1

1

1

1

1

1

ESBO

0

0

0

0

0

0

GP

2

1

1

0

0

0

OF

1

1

1

1

1

1

FIGURE 8. Live adult male Acanthosaura phuketensis sp. nov., asleep on a branch in situ at Ban Bangrong, Phuket Island, southwestern Thailand. Photo by P. Samphanthamit.



485

FIGURE 9. Live subadult male Acanthosaura phuketensis sp. nov., eating a grasshopper on a tree in situ at Ban Bangrong, Phuket Island, southwestern Thailand. Photo by P. Samphanthamit.


PAUWELS ET AL.

FIGURE 10. Live adult female Acanthosaura phuketensis sp. nov. in situ at Ban Bangrong, Phuket Island, southwestern Thailand. Photo by P. Samphanthamit.



487

FIGURE 11. Live subadult female Acanthosaura phuketensis sp. nov. in situ at Ban Bangrong, Phuket Island, southwestern Thailand. Photo by P. Samphanthamit.


PAUWELS ET AL.

FIGURE 12. Live juvenile Acanthosaura phuketensis sp. nov. in situ at Ban Bangrong, Phuket Island, southwestern Thailand. Photo by P. Samphanthamit.

FIGURE 13. Map of Thailand showing the type locality of Acanthosaura phuketensis sp. nov. (black dot). Courtesy of W. Sodob.



489

FIGURE 14. Biotope of Acanthosaura phuketensis sp. nov. in Khao Phra Thaeo Non-hunting Area, Phuket Island, southwestern Thailand. Photo by O. S. G. Pauwels.

Discussion Our comparisons show that Acanthosaura phuketensis sp. nov. is clearly distinct from all the currently recognized species of Acanthosaura, and that it is part of the A. crucigera group, in which it seems most closely related to A. crucigera and the geographically distant A. cardamomensis. Its description is just one more step in the taxonomic review of the A. crucigera group, which is already known to include several additional undescribed species in the Thai-Malay Peninsula (Wood et al. 2010, Grismer 2011, Wood et al. in prep.). Similarly, the lepidogaster species group appears to comprise several undescribed species (Ananjeva et al. 2008), and the total number of recognized Acanthosaura species will keep increasing (Sumontha et al. in prep., Wood et al. in prep.). The main conservation threat for A. phuketensis sp. nov. is probably forest logging and degradation. Fortunately, it has already been reported from several protected areas, i.e., Khao Phra Thaeo Non-hunting Area in Phuket Province, Khao Lak - Lam Ru National Park and Phang-Nga Wildlife Breeding Station in Phang-Nga Province, and Khao Sok National Park in Surat Thani Province. It will probably be recorded from Kaeng Krung National Park, Khlong Phanom National Park and Khlong Yan Wildlife Sanctuary in Surat Thani Province, Khlong Nakha Wildlife Sanctuary in Ranong Province, and Si Phang Nga National Park and Ton Pariwat Wildlife Sanctuary in Phang-Nga Province, all offering forested areas and lying within the Phuket Range. The last forested areas on Phuket Island are currently subject to strong pressure because of unregulated infrastructure development and land use (Boupun & Wongsai 2013). The high reptile diversity of Phuket Island (Frith 1978, Pauwels & Bauer 2001, Sumontha et al. 2002a-b, Leong et al. 2003, this work), including three endemic forest dwelling species (Cnemaspis phuketensis Das & Leong, 2004, Cyrtodactylus phuketensis Sumontha et al. and Trimeresurus phuketensis Sumontha et al.), stresses the importance of preserving the last patches of primary and mature


PAUWELS ET AL.



491

1.4–2.9

3.6–7.6

12–17

0.05–0.08

22.3–42.9

38.2–60.6

10–12

10–12

WNC

DIAS

DIASN

DIAS/SVL

FOREL

HINDL

SUPRAL

INFRAL

0.11–0.26

0.23–0.38

PS/HL

2.3–8.3

4.6–11.8

PS

DS/HL

0

TN

DS

0.22–0.33

TD/HD

0.21–0.39

3.5–4.7

TD

NSL/HL

6.6–11.2

ORBIT

4.1–12.2

6.8–11.0

SL

NSL

14.4–22.8

10.9–18.6

19.7–31.4

HL

HD

107.0–205.6

HW

69.2–123.5

TaL

A. phuketensis sp. nov.

SVL

A. armata

12–15

10–14

39.0–69.6

33.7–48.9

0.01–0.06

1–8

1.2–6.8

1.0–2.2

0.20–0.52

4.9–11.3

0.22–0.51

5.5–11.2

0.22–0.56

4.9–9.9

0

0.19–0.28

2.4–5.2

5.4–12.2

6.3–16.6

12.2–18.9

15.3–23.0

6.6–33.7

96.6–190

72.4–138.0

83.9–142.0

A. bintangensis 11–12

12

43.3–68.6

33.9–61.5

0.04–0.07

11–15

5.0–7.9

1.6–2.1

0.08–0.09

1.8–2.2

0.17–0.21

1.3–4.7

0.07–0.19

1.9–4.2

0

0.16

2.5–3.0

8.4–12.6

7.9–11.3

15.0–19.2

17.5–23.4

16.9–25.4

112.8–206.0

117.0

A. brachypoda 11

12–13

?

?

?

7

?

1.6

0.06

1.9

0.16

4.7

0.11

3.2

0

0.21

3.6

8.3

12.2

17.2

20.6

30.3

185.4

94.0–137.9

A. capra 12–13

10

78.5–107.2

54.2–83.8

0.05

4–7

2.0–6.7

2.3–4.1

0.16–0.17

3.5–6.8

0.42–0.43

4.2–14.7

0.36

5.2–10.2

0

0.21–0.23

3.4–5.2

7.6–11.6

7.6–16.6

14.8–24.3

16.8–27.0

16.3–38.9

133.6–182.1

82–149

A. cardamomensis 10–14

11–15

42.0–77.1

31.7–56.8

0.03–0.07

6–15

2.7–8.3

1.8–4.2

0.14–0.45

2.0–14.2

0.17–0.66

3.8–17.4

0.14–0.45

3.2–12.7

0

0.20–0.27

2.5–5.8

6.0–12.7

8.6–18.7

12.6–21.7

16.4–27.7

16.3–42.4

103–188

66.0–86.1

A. coronata 11–13

12–13

38.4–47.8

30.2–35.3

Absent

Absent

Absent

0

0

Absent

0

0

0

Absent

0

0.14–0.17

1.7–2.8

6.9–7.5

6.9–8.4

11.9–16.8

13.6–17.5

14.4–16.3

86.3–105.0

92.2–127.0

A. crucigera 10–12

10–13

48.8–65.0

35.6–49.8

0.04–0.08

9–25

4.9–8.4

1.3–3.4

0.09–0.24

2.0–5.5

0.14–0.38

3.1–8.9

0.09–0.33

1.9–7.8

0

0.14–0.21

2.5–3.9

8.9–10.8

8.7–12.1

15.7–22.5

16.0–22.3

18.7–23.6

130.0–174.0

76.5–101.1

A. lepidogaster 9–13

10–13

49.4–50.4

33.0–37.1

0.08

10–12

6.3

1.5

0.07–0.12

1.5–2.7

0.12–0.15

2.9–3.4

0.06–0.11

1.5–2.5

0–1

0.18–0.24

2.2–3.0

4.7–9.1

9.3–10.2

12.0–12.5

13.4–19.1

18.9–29.7

130.6–144.1

91.8–118.4

11–12

12–13

48.5–65.6

38.0–51.7

0.05–0.07

10–13

5.1–7.6

1.4–1.6

0.07–0.09

1.7–2.1

0.11–0.18

2.7–4.4

0.14–0.18

3.3–4.4

0

0.17–0.20

2.7–4.0

9.8–13.2

9.7–12.5

15.7–20.2

17.5–23.4

20.0–24.3

136.0–174.0

……continued on the next page

11–12

11

85.0–129.7

60.0–85.0

0.04

10

2.5

3.1–4.8

0.44

6.0–17.7

0.58

8.5–23.8

0.36

7.7–17.8

0

0.23–0.28

3.9–7.0

7.2–10.9

12.0–19.9

16.9–24.9

20.2–27.8

25.2–43.6

171.0–287.0

106.7–158.0

A. nataliae

TABLE 2. Comparison of morphometrical (in mm) and meristic data for all currently recognized species of Acanthosaura and A. phuketensis sp. nov. For the ‘‘?’’ see the comparison section.

A. titiwangsaensis


PAUWELS ET AL.

2.6–9.5

0.13–0.30

OS

OS/HL

11–14

NR

NSSLC

0–1

1

1

1

0

0–2

1

YAS

ND

LKP

BEP

ESBO

GP

OF

4

1–2

NSCSL

PM

8–10

NCS

0.5–1.4

12–13

NS

0.6–1.6

7–8

RS

MH

5–8

RH

MW

2.3–3.8

1.1–1.7

RW

4–5

1.4–1.7

TaL/SVL

10–14

21–24

TO

CS

15–17

FI

NSSOS

57–67

VENT

A. phuketensis sp. nov.

TABLE 2. (Continued)

A. armata

1

1

0

0

1

0–1

0–1

3–6

0.8–2.2

0.9–2.0

10–22

1–2

6–14

10–17

7–10

5–9

0.9–1.8

1.8–4.5

11–15

4–6

0.16–0.38

4.0–9.4

1.2–1.6

19–26

13–17

51–68

A. bintangensis 1

3–4

1

1

0

1

1

4–5

1.4–2.1

1.3–1.8

9–12

1

7–8

10–11

8

7–9

1.7–2.0

3.6–5.3

14–15

6–7

0.10–0.11

1.2–2.6

1.3–1.4

26–28

23

51–55

A. brachypoda 1

0

0

1

1

1

1

4

2.1

2.9

?

?

?

?

9

7

2.3

3.5

?

?

0.03

1.0

1.58

24

18

63

A. capra 1

3–4

0

1

1

1

1

4

1.7–2.2

1.9–2.2

9–11

1–2

7–8

9

9

7–8

1.8–2.3

4.2–4.6

12–14

Absent

0

Absent

1.2–1.5

22–24

16–17

55–66

A. cardamomensis 1

1–4

0

1

1

1

0–1

4–5

0.9–2.0

0.2–2.1

10–19

1–2

7–12

9–17

7–10

5–9

1.1–2.2

1.7–4.7

11–16

4–6

0.24–0.56

4.1–13.3

1.2–1.6

20–25

15–20

50–65

A. coronata 1

0

0

0

1

0

0–1

4–5

1.3–1.6

0.6–1.5

6–11

3–4

5–6

8–11

7–9

5

0.5–0.8

0.8–0.9

12–15

4–5

0

0

0.6–1.0

17–19

13–14

53–58

A. crucigera 1

1–2

0

1

1

1

1

4

1.1–1.7

1.0–1.5

10–14

1–2

7–11

9–12

7–9

7–9

1.3–2.0

2.7–4.0

12–15

4–6

0.11–0.50

2.5–4.9

1.1–1.8

21–26

16–18

55–63

A. lepidogaster 1

0–1

0

0–1

1

1

1

5

1.2–1.3

1.2–1.3

10

1–2

8

7–10

7–8

5–9

1.4–1.5

2.8–3.0

10–12

5

0.14–0.15

3.2–3.4

1.6–1.9

22–23

17–19

55–61

A. nataliae 1

4

0

1

0

0

1

4

2.0

2.9

16

1

8

14

8

6

2.6

6.1

13

Absent

0

Absent

1.2–1.5

20–27

16–21

64–71

1

2–4

0

1

0

1

1

5

1.4–2.4

1.4–2.0

11–14

1–2

9–11

11–12

8

9

1.4–1.8

3.6–5.2

14–15

4–5

0.09–0.10

1.8–2.3

1.1–1.5

23–27

20–21

47–57

A. titiwangsaensis

secondary forest remaining on the island. Our research team has intentionally given a specific epithet relating to Phuket Island to each of the three squamate species we described from the island in order to draw the attention of the scientific community and Government stakeholders to this need to locally improve current conservation efforts and land use planning.

Acknowledgements We are grateful to Tanya Chan-ard (THNHM, Pathum Thani), Lawan Chanhome (QSMI, Bangkok), Georges L. Lenglet and Sébastien Bruaux (IRSNB, Brussels), Colin McCarthy (BMNH, London), Kumthorn Thirakhupt (Chulalongkorn University, Bangkok) and Sansareeya Wangkulangkul (PSU, Songkhla) for giving access to the collections under their care, to Natalia Ananjeva (Russian Academy of Sciences, St. Petersburg) for providing useful information and to Wachira Sodob for elaborating the map. We thank the Royal Forest Department (Chaweewan Hutachareon, Sanal Lienphayboon, Wirot Pimanrotchanakoon and Schwann Tunhikorn) and the National Research Council (Tuenchai Niyamangkoon and Warapan Wicharn) of Thailand for research and collecting permits for the Phang-Nga specimens, and Ong-Arj Laohawat, former director of Phang-Nga Wildlife Breeding Station, for working facilities.

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APPENDIX. Comparative material examined. Acanthosaura armata: see Wood et al. (2009: 46; 2010: 38) and Grismer (2011). A. bintangensis: see Wood et al. (2009: type series; 2010: 38) and Grismer (2011). A. cardamomensis: see Wood et al. (2010: 38). A. capra: see Wood et al. (2010: 38). A. coronata: see Wood et al. (2009: 46; 2010: 38). A. crucigera: BMNH 87.11.2.4–5 (latter specimen with incomplete tail, not used in the measurements and ratio involving tail length in Table 2), BMNH 1946.8.13.84, BMNH 1946.8.19.64, ‘Tavoy’, Myanmar; CUMZ R 2009.10.22.1–3, ‘Sai Yok National Park, Kanchanaburi Province, Thailand’. A. cf. crucigera: see Grismer (2011: 122). A. lepidogaster: see Wood et al. (2009: 46; 2010: 38). A. nataliae: see Wood et al. (2009: 46). A. phuketensis sp. nov. (non type material): MNHN 1998.0528, ‘Phang-Nga Wildlife Breeding Station, Muang District, PhangNga Province, Thailand’; MNHN 1999.7697 & RFD (Field Nr. P145), ‘Tham Nam Pud, Muang District, Phang-Nga Province, Thailand’. A. titiwangsaensis: see Wood et al. (2009: type series; 2010: 38); Grismer (2011).


PAUWELS ET AL.