Three New Species of Pseudocalotes Fitzinger (Squamata: Agamidae) from the Sky Island Archipelago of Peninsular Malaysia

Home , Bronchocela, Pseudocalotes

Zootaxa 4136 (3): 461–490 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2016 Magnolia Press ISSN 1175-5334 (online edition) http://doi.org/10.11646/zootaxa.4136.3.3 http://zoobank.org/urn:lsid:zoobank.org:pub:3A3A379B-5552-44C9-B35B-B26ECB1005EE Dragons in the mist: three new species of Pseudocalotes Fitzinger (Squamata: Agamidae) from the sky island archipelago of Peninsular Malaysia

L. LEE GRISMER1, EVAN S. H. QUAH2, PERRY L. WOOD, JR.3, SHAHRUL ANUAR2, ABDUL MUIN2, HAYDEN R. DAVIS1, MATTHEW L. MURDOCH1, JESSE L. GRISMER4, MICHAEL COTA5 & ANTHONY J. COBOS1 1Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515 USA. E-mail: [email protected] 2School of Biological Sciences, Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Penang, Malaysia. E-mail: [email protected]; ; E-mail: [email protected]; E-mail: [email protected] 3Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA. E-mail: [email protected] 4Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045-7651, USA. E-mail: [email protected] 5Natural History Museum, National Science Museum, Thailand, Technopolis, Khlong 5, Khlong Luang, Pathum Thani 12120 Thai- land. E-mail: [email protected]

Abstract

An integrative taxonomic analysis is used to delimit and describe three new species of Pseudocalotoes from the sky island archipelago of the Banjaran (=mountain range) Titiwangsa of Peninsular Malaysia. Pseudocalotes drogon sp. nov., from Fraser’s Hill, Pahang is basal to the sister species P. larutensis from Bukit Larut, Perak in the Banjaran Bintang and the new species P. rhaegal sp. nov. from Cameron Highlands, Pahang. Pseudocalotes drogon sp. nov. is differentiated from all other species of Psuedocalotes by having the combination of a flat rostrum; seven postrostrals; an interparietal; 11 circumorbitals; five canthals; 7–10 superciliaries; one scale between the rostral and nasal; nine supralabials; eight infralabials; 10 postnasal-suborbital scales; four postmentals; five or six sublabials; five or six chinshields; 47 smooth, wide, gular scales; weak transverse gular and antehumeral folds; two enlarged scales between the ear and eye; enlarged upper and lower posttemporals; a single enlarged supratympanic; no enlarged postrictals; three large scales bordering the dorsal margin of the ear opening; large pretympanic scales; eight scales in the nuchal crest not separated by a gap; enlarged vertebral scales extending to the tip of the tail; keeled and non-plate-like scales on flanks; 51 midbody scales; midventrals smaller than dorsals; 19 subdigital lamellae on the fourth finger; 23 subdigital lamellae on the fourth toe; preaxial scales on third toe enlarged and spinose; subdigital lamellae not unicarinate; HW/HL 0.52; HL/SVL 0.31; no elbow or knee patches; and a male dewlap color of lime-green bearing a central yellow spot. Pseudocalotes rhaegal sp. nov. is differentiated from all other Psuedocalotes by having the combination of a convex rostrum; 6–8 postrostrals; an interparietal; nine or 10 circumorbitals; five canthals; 7–10 superciliaries; one or two scales between the rostral and nasal scales; eight or nine supralabials; seven or eight infralabials; 11 or 12 postnasal-suborbital scales; four postmentals; four or five chinshields; 40–45 smooth, wide, gular scales; no transverse gular fold; a weak antehumeral fold; three or four enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; no large scales bordering the upper margin of the ear opening or in the pretympanic region; 6–8 enlarged nuchal crest scales not separated by a gap; enlarged vertebral scales extending to the base of the tail; weakly keeled, non-plate-like scales on the flanks; 52– 58 midbody scales; midventrals smaller than dorsals; 19–21 subdigital lamellae on the fourth finger; 22–26 subdigital lamellae on the fourth toe; preaxial scales on the third enlarged and rounded; subdigital lamellae not unicarinate; HW/HL 0.50–0.54; HL/SVL 0.28–0.30; no elbow or knee patches; and female dewlap color yellow bearing a purple base. The analyses also indicated that the new species, P. v i s e r i on sp. nov. from Genting Highlands, Pahang in the southern section of the Banjaran Titiwangsa is the sister species of P. flavigula from Cameron Highlands 121 km to the north and can be separated from all other species of Psuedocalotes by having the combination of three postrostrals; 10 circumorbitals; four or five canthals; 5–7 superciliaries; rostral and nasals in contact; supralabials contacting the nasal; six or seven supralabials; six or seven infralabials; two or three postmentals; 47 or 48 smooth, flat, gular scales; three chinshields; weak transverse gular and antehumeral folds; two enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; 7–9 nuchal crest scales lacking gaps and not extending beyond midbody; weakly keeled and plate-like scales on the flanks; 35–38 midbody scales; ventrals smaller than dorsals;

Accepted by S. Carranza: 19 May 2016; published: 7 Jul. 2016 461 22 or 23 subdigital lamellae on the fourth finger; 26 or 27 subdigital lamellae on the fourth toe; preaxial scales on the third toe not modified; subdigital scales not unicarinate; HW/HL 0.62; no white marking below the eye; dewlap in males yellow; and no elbow or knee patches. Pseudocalotes rhaegal sp. nov. most likely occurs in syntopy with P. flavigula in Ta- nah Rata at Cameron Highlands and its discovery adds to a growing body of literature detailing the recent descriptions of several new, upland, closely related, sympatric species in Peninsular Malaysia. Another new population referred to here as Pseudocalotes sp. nov. from the Hala-Bala Wildlife Sanctuary, Betong District, Yala Province, Thailand is discussed. The discovery and description of these three new Pseudocalotes from the upland regions of Peninsular Malaysia continues to underscore the remarkably high herpetological diversity and ecological complexity in this sky island archipelago that is still underestimated, unappreciated, and unprotected.

Key words: Peninsular Malaysia, Thailand, Agamidae, Pseudocalotes, new species, uplands

Introduction

Globally, upland ecosystems in tropical rainforests are renown for their high levels of endemism (Blackburn & Measey, 2009; Bell et al., 2010, Grismer, 2011; Grismer et al., 2010a,b; Tolley et al., 2011). This has become patently clear in the sky island archipelago of Peninsular Malaysia that remarkably has yielded 33 new, co- distributed species of amphibians and reptiles across five different mountain systems in only the last nine years of exploration (Chan et al. 2009, 2014; Grismer 2006a,b, 2007, 2008; Grismer et al. 2004, 2008, 2009a,b, 2010a,b, 2011, 2012, 2013a,b, 2014a,b, 2015a,b; Grismer & Quah 2015; Loredo et al. 2013; Matsui et al., 2009, 2014; Wood et al. 2008, 2009) with at least 13 more species awaiting description (Grismer et. al. in prep.; Quah et al. in prep.). Joining this growing list of newly discovered montane taxa is the genus Pseudocalotes whose species content is increased here with the discovery of three new populations from the Banjaran (=mountain range) Titiwangsa. Pseudocalotes comprises a morphologically diverse group of arboreal agamid lizards ranging from the normal to the bizarre in appearance (Manthey 2012). The 21 species of this genus currently recognized generally inhabit upland areas in tropical regions and collectively extend from India to southern China and thence southward to Sumatra and Java (Hallerman et al. 2010). Although the latest phylogeny thus far indicates this genus is monophyletic (J. Grismer et al. 2016), the broader interspecific relationships have yet to be the focus of a phylogenetic systematic analysis. Currently there are four cloud forest species of Pseudocalotes known from the Thai-Malay Peninsula that represent two distinct morphological groups (Grismer 2011). Pseudocalotes khaonanensis Chan-ard, Cota, Makchai, & Laoteow from the Nakhon Si Thammarat Mountains in southern Thailand and P. larutensis Hallermann & McGuire from the Banjaran Bintang near the west coast in northern Peninsular Malaysia have small, uniformly keeled dorsal and flank scales and a relatively narrow head with an elongate snout. Whereas P. flavigula (Smith) from the centrally located Banjaran Titiwangsa and P. dringi Hallermann & Böhme from the eastern Banjaran Timur in Peninsular Malaysia have large, plate-like, flank scales (much more pronounced in P. flavigula) and a wider head and shorter snout (Fig. 1). Our ongoing fieldwork in the sky-island archipelago of Peninsular Malaysia has resulted in the discovery of three new populations of Pseudocalotes and the collection of additional specimens from the type localities of P. flavigula and P. larutensis (Appendix). Two of the three new populations come from Cameron Highlands and Fraser’s Hill, Pahang (Fig. 2) and bear the P. larutensis-khaonanensis morphology as well as having relatively short tails, a naked tympanum, prominent nuchal crests that do not extend onto the body, and no elongate postorbital and nuchal spines—characters that separate them from all other agamid genera from the Thai-Malaya Peninsula and align them with the genus Pseudocalotes (Grismer 2011). The third population from Genting Highlands, Pahang resembles P. flavigula in body shape and general coloration but differs discretely from it in squamation. We have obtained a series of close-up color photographs of a fourth specimen of Pseudocalotes from the mountains in the Hala-Bala Wildlife Sanctuary, Betong District, Yala Province, Thailand along the Thai-Malay border that, like the specimens from Genting Highlands, appears superficially similar to P. flavigula but differs from it in squamation. Molecular analyses indicate that the three new populations from Peninsular Malaysia do not form a monophyletic group and morphological and color pattern analyses indicate that each new population is discretely diagnosable from each other and all other Pseudocalotes. The intent of this paper is to describe these three new species. We want to be clear that the phylogenetic component of this study is not intended to be inclusive of all or even most of the species of Pseudocalotes and was

462 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. constructed only from tissue samples on hand and augmented by sequences available on GenBank. The phylogeny generated herein is simply a tool to aid in establishing the specific identity of each new population, the potential relationships of the Malaysian species to one another, and to establish the percent sequence divergence among them.

FIGURE 1. Different ecomorphs of Pseudocalotes on the Thai-Malay Peninsula. Upper: adult male P. larutensis (LSUDPC 6205) from Bukit Larut, Perak, Peninsular Malaysia. Lower: adult male P. flavigula (LSUDPC 9698) from Tanah Rata, Cameron Highlands, Pahang, Peninsular Malaysia. Photos by L. Lee Grismer.

Materials and method

Molecular analysis. Ingroup sampling consisted of 16 individuals of eight species of Pseudocalotes (Table 1). Outgroups used to root the tree were Japalura slendida Barbour & Dunn, Acanthosaura cardamomensis Wood,

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 463 Grismer, Grismer, Neang, Chav, & Holden, and A. crucigera Boulenger based on the relationships of J. Grismer et al. (2016). Sequence data were generated for the three new populations of Pseudocalotes from Peninsular Malaysia as well as additional specimens of P. flavigula from Cameron Highlands, Pahang, P. larutensis from Bukit Larut, Perak, and P. fl ow eri from the Bokor Plateau, Cambodia. Sequences for all other species and the outgroups were downloaded from GenBank (Table 1). All new sequences are deposited in GenBank (Table 1).

TABLE 1. Specimens used for the molecular analyses of this study with species localities and GenBank accession numbers for ND2 and tRNAs. Voucher number Species Locality GenBank # LSUHC 7393 Acanthosaura cardamomensis Cardamom Mountains, Cambodia GU817394 CAS 229582 Acanthosaura crucigera unknown GU817389 LSUMZ 81212 Japalura splendida unknown AF288230 MVZ 224106 Pseudocalotes brevipes unknown AF128502 LSUHC 12223 Pseudocalotes drogon sp. nov. Fraser’s Hill, Malaysia KX258211 LSUHC 12420 Pseudocalotes flavigula Cameron Highlands, Malaysia KX258212 LSUHC 12580 Pseudocalotes flavigula Cameron Highlands, Malaysia KX258213 LSUHC 12581 Pseudocalotes flavigula Cameron Highlands, Malaysia KX258214 LSUHC 8572 Pseudocalotes floweri Bokor Plateau, Cambodia KX258215 CAS 207492 Pseudocalotes kakhienensis unknown GQ502784 LSUHC 10285 Pseudocalotes larutensis Bukit Larut, Malaysia KX258216 LSUHC 11289 Pseudocalotes larutensis Bukit Larut, Malaysia KX258217 LSUHC 12434 Pseudocalotes larutensis Bukit Larut, Malaysia KX258218 LSUHC 9052 Pseudocalotes larutensis Bukit Larut, Malaysia KX258219 TNHC 58040 Pseudocalotes larutensis Bukit Larut, Malaysia AF128503 LSUHC 12000 Pseudocalotes rhaegal sp. nov. Cameron Highlands, Malaysia KX258220 LSUHC 12179 Pseudocalotes rhaegal sp. nov. Cameron Highlands, Malaysia KX258221 LSUHC 12114 Pseudocalotes viserion sp. nov. Genting Highlands, Malaysia KX258222 LSUHC 12227 Pseudocalotes viserion sp. nov. Genting Highlands, Malaysia KX258223

Genomic DNA was isolated from liver or skeletal muscle samples stored in 95% ethanol using the Qiagen DNeasyTM tissue kit (Valencia, CA, USA). ND2 was amplified using a double-stranded Polymerase Chain Reaction (PCR) under the following conditions: 1.0 µl (~10–30 µg of DNA) genomic DNA, 1.0 µl (10 µM) light strand primer, 1.0 µl (10µM) heavy strand primer, 1µl (10 µM) dinucleotide pairs, 2.0 µl (1.5 mM) 5x buffer, 1.0 µl (1.5 mM) MgCl 10x buffer, 2.5 ul Taq polymerase, and 7.56 µl ultra-pure H2O (Table 2). PCR reactions were executed on an Eppendorf Mastercycler gradient thermal cycler under the following conditions: initial denaturation at 95°C for 2 min, followed by a second denaturation at 95°C for 35 s, annealing at 50°C for 35 s, followed by a cycle extension at 72°C for 35 s, for 31 cycles. All PCR products were visualized on a 1 % agarose gel electrophoresis. Successful PCR products were vacuum purified using MANU 30 PCR plates (Millipore) and purified products were resuspended in ultra-pure water. Purified PCR products were sequenced using the ABI Big-Dye Terminator v3.1 Cycle Sequencing Kit in an ABI GeneAmp PCR 9700 thermal cycler. Cycle sequencing reactions were purified with Sephadex G-50 Fine (GE Healthcare) and sequenced on an ABI 3730xl DNA Analyzer at the BYU DNA Sequencing center. Primers used for amplification and sequencing are presented in Table 2. Sequences were analyzed from both the 3' and the 5' ends separately to confirm congruence between the reads. Both the forward and the reverse sequences were uploaded and edited in GeneiousTM version v5.5.6 (Drummond et al. 2011). The protein-coding region of the ND2 sequence was aligned by eye. Mesquite v3.02 (Maddison & Maddison 2015) was used to calculate the correct amino acid reading frame and to confirm the lack of premature stop codons. Pairwise sequence divergence were calculated in MEGA v6.06 (Tamaura et al. 2013).

464 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. FIGURE 2. Distribution of the species of Pseudocalotes on the Thai-Malay Peninsula. Starred circles are type localities. Pseudocalotes khaonanensis from Kaho Nan National Park, Nakkon Si Thammarat Province, Thailand. Pseudocalotes sp. nov. from the Hala-Bala Wildlife Sanctuary, Yala Province, Thailand. Pseudocalotes dringi from the type locality at Gunung Tahan, Pahang and Gunung Lawit, Terengganu, Peninsular Malaysia. Pseudocalotes larutensis from Bukit Larut, Perak, Peninsular Malaysia. Pseudocalotes rhaegal sp. nov. from Robinson Falls, Cameron Highlands, Pahang, Peninsular Malaysia. Pseudocalotes drogon sp. nov. from Fraser’s Hill, Pahang, Peninsular Malaysia. Pseudocalotes flavigula from Gunung Brinchang, Cameron Highlands, Pahang, Peninsular Malaysia. Pseudocalotes viserion sp. nov. from Genting Highlands, Pahang, Peninsular Malaysia.

TABLE 2. Primers used for PCR amplification and sequencing. L4437b 5’-AAGCAGTTGGGCCCATACC-3’ Macey et al. (1999) H5934 5’- AGRGTGCCAATGTCTTTGTGRTT-3’ Macey et al. (1999)

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 465 For the phylogenetic analyses we applied two model-based methods, Maximum Likelihood and Bayesian Inference. The Bayesian Information Criterion (BIC) as implemented in IQ-TREE (Nguyen et al. 2015) calculated TPM3+G4, HKY+G4, and TPM3u+G4 to be the best-fit models of evolution for the first, second, and third codon positions, respectively. Maximum Likelihood analysis was performed using RAxML HPC v7.5.4 (Stamatakis 2008) via the command line, with an initial search of 200 replicates for the best tree. Nodal support was calculated with 1000 bootstrap pseudoreplicates via the rapid hill-climbing algorithm (Stamatakis 2008). Nodes that had bootstrap values (ML) of 95 and above were considered significantly supported. The Bayesian analysis was carried out in MrBayes v3.2 (Ronquist et al. 2012) via command line. Two simultaneous runs were performed with eight chains per run, seven hot and one cold following default priors. The analysis was run for 5,000,000 generations and sampled every 5000 generations from the Markov Chain Monte Carlo (MCMC). The analysis was halted after the average standard deviation split frequency was below 0.01. Conservatively, the first 25% of the trees from each run were discarded as burnin. A consensus tree was computed from the two parallel runs using the sumt function in MrBayes v3.2 (Ronquist et al. 2012). Nodes that had posterior probabilities (BI) of 0.95 or above were considered significantly supported (Wilcox et al. 2002). Morphological analysis. Color notes were taken from digital images of living specimens. The following measurements were taken with Mitutoyo dial calipers to the nearest 0.1 mm under a Nikon SMZ 1500 dissecting microscope on the left side of the body where appropriate: head length (HL), measured from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the widest part of the temporal region; snout-vent length (SVL), taken from the tip of snout to the vent; and tail length (TL), taken from the vent to the tip of the tail. Scale counts taken include postrostrals (numbers of scales contacting the rostral scale not including the first supralabials); postnasal-suborbital (number of scales extending from the nasal scale to below the posterior margin of the orbit [Fig. 3]); canthals (number of enlarged scales between the postnasal and the anteriormost superciliary [Fig. 3]); circumorbitals (enlarged scales forming a semicircular arc extending from the anterior margin of the orbit [where they are small] and grading into larger scales that terminate at the posterior margin of the orbit Fig. 4]); superciliaries (number of flat, elongate, imbricating scales immediately above the orbit); supralabials (number of scales extending from the rostral to just beyond the rictus [see Harvey et al. 2015:Fig.1); infralabials (number of scales extending from the mental to the scales positioned beneath the last supralabial [see Harvey et al. 2015:Fig.1]); postmentals (number of scales, excluding the first infralabials, in contact with the mental); chinshields (number of enlarged scales between the mental that extend posteriorly along the margin of the mandible medial to the infralabials and terminate with the scale anterior to the posteriormost scale in the series that is equal in size to the adjacent medial scale [Fig.4]); gular scales (number of scales between an imaginary transverse line level with the prebrachial margin and the postmentals or infralabials if the latter are in contact); nuchal crest (number of elongate, lancolate, vertebral scales on the nape of the neck); midbody scales (number of contiguous scales counted around the body equidistant between the forelimb and hind limb insertions); fourth finger lamellae (number of transverse, subdigital lamellae on the fourth finger counted from the base of the finger where it contacts the body of the manus to the claw [including the claw sheath]); and fourth toe lamellae (number of transverse, subdigital lamellae on the fourth toe counted from the base of the toe where it contacts the body of the pes to the claw [including the claw sheath]). The remaining scale characters scored were the presence or absence of an interparietal scale; the presence or absence of a scale(s) separating the rostral and first supralabial; the presence or absence of transverse gular and antehumeral folds (Figs. 3,4); condition of gular scales (flat, wide, keeled, smooth, acuminate); number of enlarged scales between the eye and the ear (Fig. 3); the presence or absence of enlarged crest scales on the body; the presence or absence and extent of large vertebral scales on the body and tail; the number and position of enlarged posttemporals and supratympanics (Figs. 3,4); size of scales bordering dorsal margin of the ear opening; size of scales in pretympanic region (pretympanics); the presence or absence of gap in the nuchal crest scale series; the size and condition of the flank scales (plate-like, small, keeled, smooth); number of keels on subdigital lamelae; relative size of ventral and dorsal scales; and the condition of preaxial lamellae of the third toe (Fig. 5). Color pattern among Pseudoclaotes varies tremendously both within and between species (Manthey 2010) making it difficult to quantify particular components as diagnostic characters for comparisons across all species. However, dewlap color pattern is less variable and species specific. Dewlaps of males are generally more colorful than that of females, but in many species, females bear the same color characteristics as the males but to a lesser extent. Therefore, in species where males are unknown, the female color pattern is noted and used as a proxy for that of the male. Because sample sizes were small for many of the species

466 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. in this analysis—including the new species being described—the utility of some of the characters purported to be diagnostic may break down with increases in samples sizes. Conversely, new diagnostic characters may also be discovered. Therefore, all taxonomic decisions are based on an integrative interpretation of both the morphological, color pattern, and molecular data. The above morphological and color pattern characters are scored across all species where possible in Table 3.

FIGURE 3. Lateral view of the head of Pseudocalotes viserion sp. nov. (LSUDPC 9909) from Genting Highlands, Pahang, Peninsular Malaysia illustrating descriptive scale characteristics. Photograph by Kurt Orion.

Although the majority of the character states listed in Table 3 come from Harvey et al. (2015:230), their data matrix is augmented here by filling in missing character states for some species; the modification of various counts for Pseudocalotes flavigula (Smith), P. floweri (Boulenger), P. kingdonwardi (Smith), and P. kakhienensis (Anderson) based on the examination of additional material; and the addition to the matrix of P. andamanensis (Boulenger), P. austeniana (Annandale), P. khaonanensis (Chan-ard, Cota, Makchai, & Laoteow, P. larutensis Hallermann & McGuire, P. saravacensis Inger & Steubing, and P. ziegleri Hallermann, Truong, Orlov, & Ananjeva. The additional specimens examined are listed in the Appendix. Literature sources for additional character states are Anderson (1879), Annandale (1908), Chan-nard et al. (2008); Boulenger (1891), Harikrishnan & Vasdevan (2013), Hallermann, & Böhme (2000), Hallermann et al. (2010), Inger & Steubing (1994), and

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 467 Mahony (2010). LSUHC refers to the La Sierra University Herpetological Collection, La Sierra University, Riverside, California, USA; ZRC refers to the Zoological Reference Collection at the Lee Kong Chian Natural History Museum, National University of Singapore, Singapore; CAS refers to the California Academy of Sciences, San Francisco, California, USA; TNHC refers to the Texas Natural History Collection, Texas Memorial Museum, Austin, TX, USA; THNHM refers to the Thailand Natural History Museum, National Science Museum, Khlong Luang, Pathum Thani, Bangkok; and LSUDPC refers to the La Sierra University Digital Photo Collection.

FIGURE 4. Left: dorsal view of head of Psedocalotes flavigula (LSUDPC 9698) from Tanah Rata, Cameron Highlands, Pahang, Peninsular Malaysia illustrating descriptive scale characteristics. Right: Ventral view of gular region of Psedocalotes rhaegal sp. nov. (LSUHC 12178) from Robinson Falls, Cameron Highlands, Pahang, Peninsular Malaysia illustrating descriptive scale characteristics. Photos by L. Lee Grismer.

Results

Although the monophyly of Pseudocalotes is not supported by the ML analysis (ML 65) it is supported by the Bayesian analysis (BI 0.95). However, there is no significant support for any of the deeper nodes in the tree (Fig. 6). The analyses do indicate that the Peninsular Malaysian species bearing different body scale and snout morphologies are likely to not form a monophyletic group but are only distantly related to one another. Support (1.00/88) does exists for a clade where P. drogon sp. nov. is basal to the sister species (1.00/80) P. rhaegal sp. nov. and P. larutensis and where P. flavigula and P. viserion sp. nov. are sister species (1.00/100)—although the latter pair have a 22% uncorrected pairwise sequence divergence between them (Fig. 6). The morphological and color pattern analyses support the molecular analyses in that P. drogon, sp. nov., P. rhaegal sp. nov., and P. viserion sp. nov. are discretely diagnosable from one another and from all other species of Pseudocalotes (Tables 3,4,5). Additionally, the morphological and color pattern analyses indicate that although P. flavigula and P. viserion sp. nov. are more similar to each other than they are to any other species in Peninsular Malaysia, they are discretely diagnosable from one another as well as differing from all other species of Pseudocalotes in having greatly enlarged, plate-like flank scales. Given the molecular, morphological, and color pattern data above, the three new populations are described below.

73.7 73.7 – 12 12 62 9 3 9 – – – – – poliani poliani 8 7 or Variable 10 7 6 or 5 No 1 9 8 or 10 9 or 2 1 or / Smooth Flat Yes Yes 1 4 3 or Yes No 7 4 3 or No Yes No 54 Yes 18 23 Serrate Fringe No 62.2 No / 46 46 44 28 30 67 ……continued on the next page next on the ……continued 6 7 10 9 – 10 – – – – – – – – – flavigula 4 No 11 10 or 5 4 or 5 No 1 6 7 4 3 or 40 Smooth Flat Weak Weak 3 2 or 2 No No 5 no No weak Yes 41 Yes 22 26 No No 54 No Yellow

81.5 81.5 – 48 48 66 22 27 7 12 7 7 8 3 13 3 – – – – – – – – – – – – kakhienensis 5 Yes 9 4 5 No/Yes 2 8 7 or 6 1 42 Smooth Flat Yes Yes 3 2 or 1 Yes Yes/no 7 0 No Yes No 59 Yes 20 23 Yes Variable 68.9 No Longitudinal stripes 83.3 83.3

– 54 54 24 27 10 6 6 8 8 3 11 2 – – – – – – – – – – – kingdonwardi kingdonwardi 4 3 or Yes 7 4 4 No/yes 2 1 or 6 6 3 2 or 36 Smooth Flat No Yes 1 3 2 or Yes No 8 0 No Yes No 42 Yes 20 23 Basally Variable 68.4 No No pigmented patch 76.8 76.8

– 12 12 55 28 32 – – – – 7 13 7 8 2 10 4 8 – – – – – – – – guttalineatus guttalineatus 5 No 9 5 5 No 0 10 9 or 7 2 / Keeled Flat No No 3 2 or 1 Yes Yes 10 4 Yes Yes No 45 Yes 22 26 Basally No 65.5 No blue Grayish lines

76.2 76.2 – 65 65 29 30 6 13 7 2 10 9 5 4 13 7 – – – – – – – – – – – – – cybelidermus 4 Yes 9 6 5 or 4 Yes/no 0 7 7 2 / Keeled Flat No No 1 2 1 or Yes Yes 9 4 Yes Yes No 51 No 22 24 Basally No 60.5 No Light green with cyan center Pseudocalotes. 7 – rhammanotus rhammanotus 5 Yes 10 9 or 6 5 No 2 9 8 5 4 or / Keeled Flat No Yes 2 1 Yes No 8 7 Yes Yes No 51 Yes 21 24 No No 64.5 No Brown with centerWhite

69.0 69.0 – 40 40 26 32 7 13 8 11 11 5 11 – – – – – – – – – – tympanistriga 4 No 9 6 5 or 6 No 2 1 or 9 9 2 / Keeled Flat No Yes 4 3 or 1 Yes No 4 3 2 or No No no 46 Yes 22 26 No No 60.8 No Same color as venter

Distribution of characters of Distribution states across the species of

TABLE 3. Postrostrals Interparietal Cicrcumorbitals Canthals Superciliaries Scale between rostral and nasal nasal Supralabials contacting Supralabials Infralabials Postmentals Gulars smoothGulars or keeled granular orGulars flat gular Transverse fold fold Antehumeral Enlarged between scales andeye ear Enlarged posttemporals Enlarged supratympanic Enlarged postrictals Nuchal crest crest in nuchal Gap Enlarged crest scales midbody beyond keeled scales Flank plate-like scales Flank midbody around Scales thanMidventrals smaller dorsals lamellae finger 4th 4th toe lamellae Preaxial lamellae on toemodified III Unicarinate lamellae beneath distal phalanges HW/HL Elbow & knee patches color Male dewlap

75 75 – khaonanensis khaonanensis / / / 6 8 Yes yes 8 9 / Acuminate Acuminate Flat No Yes 1 1 No No 9 No No Weak No 72 No 21 27 No No / No purple ……continued on the next page next on the ……continued austeniana austeniana 4 3 or No / 3 / No Yes 6 5 or 8 7 or 2 / Smooth Flat No Yes 4 Yes No No 7 No yes Weak No 51 / 19 31 / / / No on No color patch throat 12 12 14 62 30 6 5 – – – – – – andamanensis andamanensis 4 Yes 11 5 4 or 8 7 or Yes Yes 10 9 or 10 2 / Keeled Flat No Yes 3 yes No No 11 No No No No 57 Yes / 27 Yes No 0.59 / Yellow/White 52 52 – dringi dringi 5 / / 5 / No Yes 8 8 7 or 2 / Smooth Flat No No 2 yes No No 6 No No Weak No 48 Yes 20 26 No No 0.51 No Purple 73.6 73.6 – 76 76 21 27 – – – 7 13 7 7 10 10 4 13 – – – – – – – – brevipes 5 Variable 9 5 5 Variable 1 0 or 6 7 2 1 or / Keeled Acuminate Yes Yes 1 2 1 or Yes/no No 7 / No Yes No 66 No 16 20 Serrate Fringe No 68.1 Yes Purple 70.3 70.3 – 74 74 25 – – 7 9 4 10 – – – – microlepis 7 6 or Variable 10 5 7 6 or No 1 8 7 or 7 1 / Keeled Flat Yes No 1 1 No No 8 / No Weak No 71 No 19 18 or 21 Serrate Fringe No 62.2 No Brown 12 12 22 6 10 11 4 – – – – – – floweri 4 Yes 10 6 5 or 8 Yes/no 2 1 or 8 9 2 1 or 42 Smooth flat Yes Yes 2 1 No No 9 3 No Yes No 44 Yes 21 24, 25 Serrate Fringe No 67.5 No Purple

(Continued)

TABLE 3. Postrostrals Interparietal Circumorbitals Canthals Superciliaries Scale(s) between rostral and nasal Supralabials contacting nasal Supralabials Infralabials Postmentals Gulars smoothGulars or keeled granular orGulars flat gular Transverse fold fold Antehumeral Enlarged between scales andeye ear Enlarged posttemporals Enlarged supratympanics Enlarged postrictals Nuchal crest crest nuchal in Gap Enlarged crest scales midbody beyond keeled scales Flank plate-like scales Flank midbody around Scales thanVentrals dorsals smaller lamellae finger 4th 4th toe lamellae Preaxial lamellae on toemodified III Unicarinate lamellae beneath distal phalanges HW/lHL Elbow & knee patches color Male dewlap

470 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. sp. nov. sp. nov. 38 38 7 9 – – – viserion 3 No/yes 10 5 4 or 5 no yes 7 6 or 7 6 or 3 2 or 48 47 or Smooth Flat Weak Weak 2 1 No No 7 No No Weak Yes 35 Yes 23 22 or 27 26 or No No 0.62 No Yellow sp. nov. 0.54 0.54 – 45 45 58 21 26 8 10 8 – – – – – – – rhaegal 6 Yes 10 9 or 5 7 Yes Yes 9 8 or 8 7 or 4 40 Smooth Flat No Weak 4 3 or 1 No No 6 No Yes Weak No 52 Yes 19 22 Enlarged and rounded No 0.50 No purple w/ Cyan center

sp. nov. 10 – drogon drogon 7 Yes 11 5 7 Yes Yes 9 8 4 47 Smooth Flat Weak Weak 2 1 no No 8 No yes Keeled No 51 Yes 19 23 Enlarged and spinose No 0.52 No yellow Lime-green w/ center 0.65 0.65 – 14 14 69 55 20 25 7 6 8 10 10 9 – – – – – – – – – – – larutensis larutensis 5 Yes 11 4 6 Yes Yes 8 8 4 3 or 55 Weak Flat No Yes 3 2 or 1 No No 5 No No Weak No 52 Yes 17 22 No No 0.50 No Yellow/Purple 64 64 – 7 11 11 – – – ziegleri 7 / / 5 / No Yes 8 7 3 2 or Smooth Flat No / 2 no / No 7 6 or No No Smooth No 57 Yes / 24 Yes No 0.54 No Blue saravacensis / / / 4 / Yes Yes 7 7 2 Keeled Flat No No 3 2 yes No 7 No No Weak No 68 / 16 20 / No 0.59 No specklesBlack

(Continued) (Continued)

TABLE 3. Postrostrals Interparietal Cicrcumorbitals Canthals Supraciliaries Scale(s) between rostral and nasal Superlabials contacting nasal Supralabials Infralabials Postmentals Gulars smoothGulars or keeled granular orGulars flat gular Transverse fold fold Antehumeral Enlarged between scales andeye ear Enlarged posttemporals Enlarged posttympanics Enlarged postrictals Nuchal crest crest in nuchal Gap Enlarged crest scales midbody beyond keeled scales Flank plate-like scales Flank midbody around Scales thanVentrals smaller dorsals lamellae finger 4th 4th toe lamellae Preaxial lamellae on toemodified III Unicarinate lamellae beneath distal phalanges HW/lHL Elbow & knee patches color Male dewlap

Pseudocalotes drogon sp. nov. Drogon’s False Garden Lizard Figs. 5,6,7

Holotype. Adult male LSUHC 12223 collected on 24 March 2015 by L. Lee Grismer, Evan S. H. Quah, Perry L. Wood, Jr., Hayden R. Davis, Matthew L. Murdoch, Brando R. Burch, and Anthony J. Cobos at 2030 hrs 1 km south of Air Terjun Jeriau, Fraser’s Hill, Pahang, Peninsular Malaysia (3° 43.283 N 101° 43.035 E; 1066 m in elevation). Diagnosis. Pseudocalotes drogon sp. nov. is differentiated from all other Psuedocalotes by having the combination of a flat rostrum; seven postrostrals; an interparietal; 11 circumorbitals; five canthals; 7–10 superciliaries; one scale between the rostral and nasal; nine supralabials; eight infralabials; 10 postnasal-suborbital scales; four postmentals; five or six sublabials; five or six chinshields; 47 smooth, wide, gular scales; weak transverse gular and antehumeral folds; two enlarged scales between the ear and eye; enlarged upper and lower posttemporals; a single enlarged supratympanic; no enlarged postrictals; three large scales bordering the dorsal margin of the ear opening; large pretympanic scales; eight scales in the nuchal crest not separated by a gap; enlarged vertebral scales extending to the tip of the tail; keeled and non-plate-like scales on flanks; 51 midbody scales; midventrals smaller than dorsals; 19 subdigital scales on the fourth finger; 23 subdigital scales on the fourth toe; preaxial scales on third toe enlarged and spinose; subdigital scales not unicarinate; HW/HL 0.52; HL/SVL 0.31; no elbow or knee patches; and a male dewlap color of lime-green bearing a central yellow spot. These characters or a subset of them are scored across all species in Table 3. Description of holotype. Adult male SVL 91.4 mm; head relatively large with a somewhat elongate flat snout, subtriangular in lateral and dorsal profile; HW/HL 0.52; HL/SVL 0.31; interorbital region flat; rostrum sloped anteriorly, canthus rostralis sharp, composed of five large canthal scales; supraorbital scales slightly enlarged, keeled; a single scale separating rostral and nasal scale; single row of 11 enlarged, circumorbital scales bordering medial margins of supraorbital region, scales largest posteriorly; eight, flat, imbricate superciliary scales; scales of orbit (surrounding eye) granular; short series of two enlarged, acuminate, postorbital scales extending to above anterior margin of tympanum; dorsal, head scales acuminate; rostral low, rectangular, bordered laterally by first supralabials, posteriorly by seven smaller scales; external nares set in slightly elevated single, rectangular, large, nasal scale; five enlarged, acuminate, median scales in prefrontal region forming a Y-shaped series with two larger, keeled scales on each side of midline; parietal, occipital, and upper postorbital region covered with slightly enlarged, elevated, acuminate to keeled scales; an enlarged parietal scale on each side of midline bordered by a clump of four, contiguous, enlarged, acuminate, occipital scales; interparietal small, lacking eyespot; temporal scales unequal with small scales intermixed with enlarged, weakly keeled scales; single, raised, enlarged, keeled scale in temporal region followed by a single, enlarged, spinose, posttemporal scale; tympanum naked; auditory meatus bodered dorsally by three large supratympanic scales; nine (R,L) supralabials; 10 postnasal-suborbital scales; mental triangular, larger than adjacent infralabials; two larger postmentals, separated medially and forming first of a series of five(R) or six(L) enlarged chinshields that are separated from infralabials by one anteriorly and two posteriorly rows of smaller scales; eight (R,L) infralabials; gular scales flat, wide, directed posteromedially, 47 at midline; dewlap small; and transverse gular and antehumeral folds weak; nuchal crest composed of eight, large, contiguous, lanceolate scales beginning on occiput, reaching maximum height on nape, tapering to a low, vertebral crest on body composing a single row of slightly enlarged keeled scales extending to base of tail; body laterally compressed; dorsal body and flank scales weakly keeled, subimbricate, rectangular, arranged in indistinct, transverse, and posteroventrally oriented rows; flank scales larger than dorsal body scales; ventrals same size as dorsals; 51 scales around midbody; scales of pectoral region, belly, and precloacal regions strongly keeled; limbs slender, covered dorsally with keeled scales; five digits on pes and manus; hind limbs larger, slightly more robust than forelimbs; subdigital lamellae of toes I, II, IV, and V bicarinate; preaxial lamellae on toe III enlarged and spinose (Fig. 5); 19 lamellae beneath fourth finger; 23 lamellae beneath fourth toe; tail laterally compressed, slightly swollen at base, covered with keeled scales, vertebral row forming a serrate, dorsal ridge, 1.89 times SVL. Coloration in life (Fig. 7). Dark phase—Dorsal ground color of head, body, limbs, and tail very dark-grey; labials cyan; row of white scales between supralabials and orbit; scattered white spots on anterior portion of flanks and sides of neck; four wide, diffuse, darker, transverse, dorsal bands between limb insertions extending to ventral

472 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. edge of flanks; one band in pelvic region; nine bands on tail; limbs generally immaculate, dark-grey; groups of white scales in lateral margins of gular region; anterior gular region dull-white bearing dark-brown oblique lines; dewlap dingy lime-green with a dingy yellow center; ventral surfaces, beige, immaculate. Light phase—Dorsal body pattern same as dark phase but ground color a much lighter grey; dark dorsal bands prominent against dull- white interspace; cyan labials faded; gular region whitish bearing light-brown, oblique lines; dewlap lime-green with yellow center, dewlap scales bearing brownish centers; venter whitish, immaculate. Distribution. Pseudocalotes drogon sp. nov. is known only from the type locality 1 km south of Air Terjun Jeriau, Fraser’s Hill, Pahang Peninsular Malaysia (Fig. 2). It is expected however to be found further south to at least Genting Highlands. Etymology. The specific epithet drogon refers to this species’ resemblance in form and color to the dark dragon, Drogon—one of three dragons born in the Dothraki Sea and commanded by Daenerys Targaryen—the Mother of Dragons—in George R. R. Martin’s fictional work Game of Thrones.

FIGURE 5. Lateral view of preaxial scales on the third toe of the left hind limb in P. drogon sp. nov. (LSUHC 12223), P. rhaegal sp. nov. (LSUHC 12178), and P. flavigula (LSUHC 12420) showing the degrees of modification. Photos by L. Lee Grismer.

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 473 FIGURE 6. Maximum Likelihood tree topology showing the relationships of the new species Pseudocalotes drogon sp. nov., P. rhaegal sp. nov., and P. viserion sp. nov. to selected other species of Pseudocalotes. Numbers at the nodes are Bayesian posterior probabilities (BI)/Maximum Likelihood (ML) bootstrap values.

Natural history. Pseudocalotes drogon sp. nov. was collected at night at 2000 hrs while sleeping on a thin, horizontal branch of small tree in the vicinity of a small stream in hill dipterocarp forest (Fig. 7). The adult male was in the light color phase at the time of collection but became much darker during the day (Fig. 7). Comparisons. Pseudocalotes drogon sp. nov. can be differentiated from all other species of Pseudocalotes by having a lime-green dewlap with a yellow center and the unique combination of numerous other characteristics (Table 3). With the exception of Pseudocalotes flavigula and P. viserion sp. nov., P. drogon sp. nov. most closely resembles the other geographically proximate Thai-Malay Peninsula species P. khaononensis, P. larutensis, P. rhaegal sp. nov., and P. dri ngi . Pseudocalotes drogon sp. nov. can be differentiated from the former two species by a number of characteristics (Table 3), the most notable of which is the lack of enlarged, plate-like scales on the flanks (compare Fig. 7 with Figs. 10,12). From P. khaononensis, P. drogon sp. nov. can be differentiated by having a maximum SVL of 91.4 mm versus 104.5 mm; five as opposed to six canthals; eight as opposed to nine infralabials; smooth as opposed to acuminate gular scales; 51 as opposed to 72–75 scales around midbody; ventral scales being smaller as opposed to being larger than dorsal scales; 23 as opposed to 27 subdigital lamellae on the fourth toe; having as opposed to lacking enlarged, spinose, preaxial lamellae on the third toe; and males having a lime-green dewlap with a yellow center versus a purple dewlap (Fig. 8). From P. larutensis, P. drogon sp. nov. can be differentiated by having a maximum SVL of 91.4 mm versus 81.0 mm; 11 versus 11–14 circumorbitals; smooth as opposed to weakly keeled gular scales; males having a lime-green dewlap with a yellow center versus a yellow dewlap with a purple, horizontal, centrally positioned marking (Fig. 8). From P. dringi, P. drogon sp. nov. is separated by having a maximum SVL of 91.4 mm versus 70.3 mm; seven versus five postrostrals; the rostral and

474 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. nasal scale not contacting as opposed to contacting; four as opposed to two postmentals; a weak, antehumeral fold as opposed to lacking a fold; having as opposed to lacking enlarged, spinose, preaxial lamellae on the third toe; and males having a lime-green dewlap with a yellow center versus a purple dewlap (Fig. 8). From P. rhaegal, P. drogon sp. nov. can be differentiated by having 11 as opposed nine or 10 circumorbitals; 10 as opposed to 11 or 12 postnasal-suborbitals; 47 as opposed to 40–45 gular scales; a row of two versus three or four enlarged scales between the ear and eye; large as opposed to small pretympanic scales; a flat versus a convex rostrum; having as opposed to lacking enlarged vertebral scales on the tail; and 51 versus 52–58 midbody scales; enlarged, spinose, preaxial scales on the third toe as opposed to enlarged, rounded preaxial scales on the third toe; longer head (HL/ SVL = 0.31 versus 0.28–0.30); and a row of white scales extending from the nasal scale to just beyond the posterior border of the orbit as opposed to a suborbital white patch (Table 4). Differences from other species are listed in Table 3.

TABLE 4. Diagnostic character states of the type series of Pseudocalotes drogon sp. nov. and P. rhaegal sp. nov. Paired values are listed as left and right. P. drogon sp. nov. P. rhaegal sp. nov. LSUHC 12223 LSUHC 12000 LSUHC 12178 LSUHC 12179 Holotype Paratype Holotype Paratype Sex Male Female Female Female Postrostrals 7 8 8 6 Interparietal yes yes yes yes Circumorbitals 11 10 9 10 Canthals 5 5 5 5 Superciliaries 8 10 8 7 Scale between rostral and nasal 1 2 1,2 1 Rostrum Flat Convex Convex Convex Supralabial-nasal contact Yes Yes Yes Yes Supralabials 9 9 9 8 Infralabials 8 8 7 8 Postnasal-suborbitals 10 11,11 12,11 12,11 Postmentals 4 4 4 4 Chinshields 5,6 5,6 5,4 Damaged Gulars 47 42 40 45 Gulars wide or granular Wide Wide Wide Wide Gulars smooth or keeled Smooth Smooth Smooth Smooth Transverse gular fold Weak No No No Antehumeral fold Weak Weak Weak Weak Enlarged scales between eye and ear 2 3 3,4 4 Enlarged posttemporals 1 1 1 1 Posttympanics modified 0 0 0 0 Enlarged supratympanics 1 1 1 1 Pretympanic scales large Yes No No No Enlarged postrictals 0 0 0 0 Rostrum Flat Convex Convex Convex Nuchal crest 8 7 6 8 Gap in crest No No No No Nuchal crest scales on body No No No No ...... continued on the next page

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 475 TABLE 4. (Continued) P. drogon sp. nov. P. rhaegal sp. nov. LSUHC 12223 LSUHC 12000 LSUHC 12178 LSUHC 12179 Holotype Paratype Holotype Paratype Enlarged vertebral scales on tail Yes No No No Flank scales keeled Weak Weak Weak Weak Flank scales plate-like No No No No Midbody scales 51 54 52 58 Midventrals smaller than dorsals Same size Same size Yes Yes 4th finger lamellae 19 19 20 21 4th toe lamellae 23 22 25 26 Preaxial lamellae modified on toe III Enlarged/spinose Enlarged/rounded Enlarged/rounded Enlarged/rounded Unicarinate lamellae on toes No No No No Suborbital white patch No Yes Yes Yes Broken white line extending from nares Yes No No No to beyond posterior margin of eye HW/HL 0.52 0.54 0.54 0.50 Body bands extend to base of flanks Yes No No No Male dewlap color Lime-green with Cyan w/ purple Cyan w/ purple Cyan w/ purple yellow center center (female) center (female) center (female) HL/SVL 0.31 0.29 0.30 0.28 SVL 91.4 75.1 83.7 85.2 TL 173 123 140 140 TL/SVL 1.89 1.63 1.67 1.64

FIGURE 7. Type specimen of Pseudocalotes drogon sp. nov. from Fraser’s Hill, Pahang, Peninsular Malaysia. Upper left: adult male holotype (LSUHC 12223) in dark phase. Lower left: holotype in light phase. Upper right: dewlap color of holotype. Lower right: microhabitat at the type locality. Photos by L. Grismer.

476 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. FIGURE 8. Dewlap coloration in male Pseudocalotes drogon sp. nov. (LSUDPC 9497; photograph by L. Lee Grismer), P. larutensis (LSUDPC 9400; photograph by L. Lee Grismer), P. khaonanensis (LSUDPC 9910; photograph by Micheal Cota), and female P. rhaegal sp. nov. (LSUDPC 9488; photograph by Mohd. A. Muin).

Pseudocalotes rhaegal sp. nov. Rhaegal’s False Garden Lizard Figs. 5,6,8,9

Holotype. Adult female LSUHC 12178 collected on 18 March 2015 by L. Lee Grismer, Evan S. H. Quah, Shahrul Anuar, Mohd A. Muin, Perry L. Wood, Jr., Hayden R. Davis, Matthew L. Murdoch, Brandon R. Burch, and Anthony J. Cobos at 2030 hrs at Robinson Falls, Cameron Highlands, Pahang, Peninsular Malaysia (04° 43.283 N 101° 23.129 E; 1411 m in elevation). Paratypes. Adult female LSUHC 12179 bears the same locality collecting data as the holotype. Adult female LSUHC 12000 bears the same collecting locality and collectors but was collected on 4 September 2014. Diagnosis. Pseudocalotes rhaegal sp. nov. is differentiated from all other Psuedocalotes by having the combination of a convex rostrum; 6–8 postrostrals; an interparietal; nine or 10 circumorbitals; five canthals; 7–10 superciliaries; one or two scales between the rostral and nasal scales; eight or nine supralabials; seven or eight infralabials; 11 or 12 postnasal-suborbital scales; four postmentals; 4–6 chinshields; 40–45 smooth, wide, gular scales; no transverse gular fold; weak antehumeral fold; three or four enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; no large scales bordering the upper margin of the ear opening or in the pretympanic region; 6–8 enlarged nuchal crest scales not separated by a gap; enlarged vertebral scales extending to base of tail; weakly keeled, non-plate-like scales on flanks; 52–58 scales around midbody; midventrals smaller than dorsals; 19–21 subdigital lamellae on fourth finger; 22–26 subdigital lamellae on fourth toe; preaxial scales on third toe enlarged and rounded; subdigital lamellae not unicarinate; HW/HL 0.50–0.54; HL/SVL 0.28–0.30; no elbow or knee patches; and female dewlap bearing a purple base. These characters or a subset of them are scored across all species in Table 3.

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 477 Description of holotype. Adult female SVL 83.7 mm; head relatively large subtriangular in lateral and dorsal profile; HW/HL 0.54; HL/SVL 0.30; interorbital region flat; rostrum convex, sloped anteriorly, canthus rostralis sharp, composed of five large canthal scales; supraorbital scales slightly enlarged, keeled; single row of nine or 10 enlarged, circumorbital scales bordering medial margins of supraorbital region; eight flat, imbricate to subimbrictae, superciliary scales; scales of orbit (surrounding eye) granular; short row of three (R) four (L) enlarged, acuminate, postorbital scales extend to above anterior margin of tympanum; dorsal, head scales acuminate; rostral low, rectangular, bordered laterally by first supralabials, posteriorly by eight smaller scales; external nares in slightly elevated, rectangular, large, nasal scale; seven enlarged, acuminate, median scales in prefrontal region forming a Y-shaped series; parietal and occipital regions covered with enlarged, elevated, acuminate to keeled scales; an enlarged parietal scale on each side of midline; interparietals small, irregularly shaped, centralmost bearing a small eyespot; temporal scales unequal with small scales intermixed with enlarged, weakly keeled scales; one, enlarged, keeled, scale in temporal region followed by a single, enlarged, spinose, posttemporal scale; tympanum naked, not bordered dorsally by large supratympanic scales; nine (R,L) supralabials; 12 (R), 11 (L) large, postnasal-suborbital scales; mental triangular, larger than adjacent infralabials; four larger postmentals, separated medially by two small scales and forming first of a series of five (R) four (L) enlarged chinshields separated from infralabials by one anteriorly and two posteriorly rows of smaller scales; seven (R,L) infralabials; gular scales wide, smooth, directed posteromedially, 40 at midline; dewlap small; transverse gular and antehumeral folds weak.

FIGURE 9. Type series of Psedocalotes rhaegal sp. nov. from Robinson Falls, Cameron Highlands, Pahang, Peninsular Malaysia and habitat at the type locality. Upper left: LSUHC 12179, photograph by L. Lee Grismer. Upper right: LSUHC 12000, photograph by L. Lee Grismer. Lower left: LSUHC 12178, photograph by Mohd A. Muin. Lower right: Montane forest microhabitat at Robinson Falls, Cameron Highlands, Pahang.

Nuchal crest composed of six, large, contiguous, lanceolate scales beginning on occiput, reaching maximum height on nape, tapering to a low, vertebral crest on body composing a single row of slightly enlarged, keeled vertebral scales extending to base of tail; body laterally compressed; dorsal body and flank scales weakly keeled, subimbricate, rectangular, arranged in indistinct, transverse, and posteroventrally oriented rows; flank scales larger than dorsal body scales; ventrals same size as dorsals; 52 scales around midbody; scales of pectoral region, belly, and precloacal regions strongly keeled; limbs slender, covered dorsally with keeled scales; five digits on pes and

478 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. manus; hind limbs larger, slightly more robust than forelimbs; subdigital lamellae of toes I, II, IV, and V bicarinate; preaxial lamellae on toe III enlarged and rounded (Fig. 5); 20 lamellae beneath fourth finger; 25 lamellae beneath fourth toe; and tail laterally compressed, slightly swollen at base, covered with keeled scales but lacking an enlarged vertebral row, 1.67 times SVL. Coloration in life (Fig. 9). Dorsal ground color of head, body, and limbs, brown, mottled with shades of slightly darker brown; linear patch of light-colored suborbital scales beneath eye; four very faint, diffuse, dark bands between forelimb and hind limb insertions not extending to base of flanks; lighter interspace between bands mottled; scattered greenish scales in postorbital, temporal, lower flank, and pelvic regions; scales of forearms greenish with dark bands; seven dark caudal bands anteriorly separated by dull-green interspaces, dark bands fade on posterior one-third of tail; gular region beige with irregularly shaped, brown, thin, submandibular bands; gular pouch small, base of pouch cyan with purple center. Variation (Fig. 9). LSUHC 12179 very closely approximates the holotype in overall body coloration and pattern, except for being lighter overall. LSUHC 12000 bears the same general color pattern as the holotype and LSUHC 12178 but differs greatly from them in being overall more green in coloration and having four dark, diffuse, dorsal bands between the forelimb and hind limb insertions that are more prominent. Additionally, the tail is distinctly banded for nearly its entire length. Variation in squamation is presented in Table 4. Distribution. Pseudocalotes rhaegal sp. nov. is known only from Robinson Falls, Cameron Highlands, Pahang Peninsular Malaysia (Fig. 2) but is expected to range more widely along the Banjaran Titiwangsa. Etymology. The specific epithet rhaegal refers to this species’ resemblance in form and color to the greenish dragon, Rhaegal—one of three dragons born in the Dothraki Sea and commanded by Daenerys Targaryen—the Mother of Dragons—in George R. R. Martin’s fictional work Game of Thrones. Natural history. All Pseudocalotes rhaegal sp. nov. were collected at night between at 2000 and 2300 hrs while sleeping on thin, horizontal branches of small trees in the vicinity of a river flowing through the hill dipterocarp forest at Robinson Falls (Fig. 9). All were gravid females collected during mid-March or mid- September, indicating that this species may breed year-round. Comparisons. Pseudocalotes rhaegal sp. nov. can be differentiated from all other species of Pseudocalotes by having a cyan dewlap with a purple center (females) and the unique combination of numerous other characteristics (Table 3). With the exception of Pseudocalotes flavigula and P. viserion sp. nov., P. rhaegal sp. nov. most closely resembles the other geographically proximate Thai-Malay Peninsula species P. khaononensis, P. larutensis, P. drogon sp. nov., and P. dringi. Pseudocalotes rhaegal sp. nov. can be differentiated from P. flavigula and P. viserion sp. nov., by several characteristics the most notable of which is the lack of enlarged, plate-like scales on the flanks (Figs. 9,10). From Pseudocalotes khaononensis, P. rhaegal sp. nov. can be differentiated by having a maximum SVL of 85.2 mm versus 104.5 mm; five as opposed to six canthals; seven or eight as opposed to nine infralabials; smooth as opposed to acuminate gular scales; three or four versus one enlarged scale between the eye and the ear; enlarged vertebral scales extending beyond midbody as opposed to not extending beyond midbody; 52–58 as opposed to 72–75 scales around midbody; ventral scales being smaller as opposed to being larger than dorsal scales; 22–26 as opposed to 27 subdigital lamellae on the fourth toe; having as opposed to lacking enlarged, rounded, preaxial lamellae on the third toe; and having a cyan dewlap with a purple center (female) versus a purple dewlap in males (Fig. 8). From P. larutensis, P. rhaegal sp. nov. can be differentiated by having nine or 10 versus 11–14 circumorbitals; 40–45 versus 55–69 gulars; smooth as opposed to weakly keeled gular scales; enlarged vertebral scales extending beyond midbody as opposed to not extending beyond midbody; preaxial scales on third toe bearing enlarged, rounded, scales versus being unmodified; having a cyan dewlap with a purple tip (females) versus a yellow dewlap with a purple, horizontal, centrally positioned marking (Fig. 8). From P. dringi, P. rhaegal sp. nov. is separated by having a maximum SVL of 85.2 mm versus 70.3 mm; 6–8 versus five postrostrals; four as opposed to two postmentals; a weak, antehumeral fold as opposed to lacking a fold; having three or four versus two enlarged scales between eye and ear; enlarged vertebral scales extend beyond midbody as opposed to not extending beyond midbody; having as opposed to lacking enlarged, spinose, preaxial lamellae on the third toe; 52–58 scales around midbody versus 48–52; 23 versus 26 fourth toe subdigital lamellae; and having a cyan dewlap with a purple tip (females) versus a purple dewlap in males (Fig. 8). From P. drogon sp. nov., P. rhaegal sp. nov. can be differentiated by having nine or 10 as opposed to 11 circumorbitals; 11 or 12 as opposed to 10 postnasal- suborbitals; 40–45 as opposed to 47 gular scales; a row of three or four versus two enlarged scales between the ear and eye; lacking as opposed to having three large supratympanic scales; pretympanic scales small as opposed to

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 479 large; a convex versus a flat rostrum; lacking versus having enlarged vertebral scales on the tail; 52–58 versus 51 midbody scales; having enlarged, rounded, preaxial scales on the third toe as opposed to enlarged preaxial scales on the third toe being spinose; shorter snout (HL/SVL = 0.28–0.30 versus 0.31); and having a white patch of suborbital scales as opposed to lacking a white patch of suborbital scales (Table 4). Differences from other species are listed in Table 3.

Pseudocalotes viserion sp. nov. Viserion’s False Garden Lizard Figs. 3,6,10

Holotype. Adult female LSUHC 12227 collected on 26 March 2015 by Evan S. H. Quah at 1000 hrs crossing the radar tower road at Ulu Kali at Genting Highlands, Pahang, Peninsular Malaysia (03° 26.166 N 101° 47.021 E; 1754 m in elevation). Paratype. Adult male LSUHC 12141 found dead on the radar tower road at Ulu Kali at Genting Highlands, Pahang, Peninsular Malaysia (03° 26.12 N 101° 47.345 E; 1750 m in elevation) by L. Lee Grismer, Perry L. Wood, Jr., Hayden R. Davis, Matthew L. Murdoch, Brandon R. Burch, and Anthony J. Cobos. Diagnosis. Pseudocalotes viserion sp. nov. can be separated from all other species of Psuedocalotes by having a combination of three postrostrals; 10 circumorbitals; four or five canthals; 5–7 superciliaries; rostral and nasal in contact; supralabials contacting nasal; six or seven supralabials; seven or eight infralabials; two or three postmentals; three enlarged chinshields; 47 or 48 smooth, flat, gular scales; weak transverse gular and antehumeral folds; two enlarged scales between the ear and eye; an enlarged upper and lower posttemporal; an enlarged supratympanic; no enlarged postrictals; 7–9 nuchal crest scales lacking gaps and not extending beyond midbody; weakly keeled and plate-like scales on flanks; 35–38 scales around midbody; ventrals smaller than dorsals; 22 or 23 subdigital lamellae on fourth finger; 26 or 27 subdigital lamellae on fourth toe; preaxial scales on third not modified; subdigital lamellae not unicarinate; HW/HL 0.62; no white marking below the eye; dewlap in males yellow; and no elbow or knee patches. These characters or a subset of them are scored across all species in Table 3. Description of holotype. Adult female SVL 71 mm; head relatively large, triangular in lateral and dorsal profile; HW/HL 0.62; HL/SVL 0.30; interorbital and frontal regions flat; snout not elongate but convex, sloped anteriorly, canthus rostralis sharp, composed of four scales; supraorbital scales enlarged, keeled; single row of 10 enlarged, circumorbital scales bordering medial margins of supraorbital region, posteriormost of series enlarged and tuberculate; five, flat, imbricate superciliary scales; scales of orbit (surrounding eye) granular; three enlarged postorbital scales, posteriormost scale acuminate, keeled; dorsal, head scales weakly acuminate; rostral low, rectangular, bordered laterally by first supralabials, dorsolaterally by nasal scale, and posteriorly by three smaller scales; external nares in slightly elevated single, rectangular, large, nasal scale; three enlarged, acuminate, median scales in prefrontal region; parietal, occipital, and upper postorbital region covered with slightly enlarged, irregularly shaped, weakly acuminate scales; an enlarged, keeled parietal scale on right side of midline followed by a large, raised occipital scale; enlarged parietal on left side of midline broken up and adjacent occipital scale not as large or raised as the corresponding scale on right; interparietal small, bearing eyespot; temporal scales irregular in size; single enlarged, raised postorbital scale; two, raised, enlarged, keeled, posttemporal scale; tympanum naked, scales on dorsal margin not enlarged; six (R,L) supralabials; nine (R,L) postnasal-supraorbital scales; mental triangular; two large postmentals in medial point contact forming first in a series of three enlarged chinshields separated from infralabials by three anteriorly and 10 posteriorly rows of smaller scales; 7 (R), 8(L) infralabials; gular scales smooth, flat, directed posteromedially, 47 at midline; dewlap small; transverse gular and antehumeral folds weak. Seven, large, lanceolate nuchal scales beginning on occiput, reaching maximum height on nape, tapering to a low, vertebral crest on body composed of a single row of keeled, slightly enlarged scales that extend onto base of tail where they are largest; body round in cross-section; dorsal body and flank scales weakly keeled, subimbricate, rectangular, arranged in indistinct, transverse rows; flank scales larger than dorsal body scales, plate-like; ventrals smaller than dorsals; scales of pectoral region, belly, and precloacal regions strongly keeled; limbs moderate in stature, covered dorsally with keeled scales; five digits on pes and manus; hind limbs larger, more robust than forelimbs; subdigital lamellae of toes I, II, IV, and V bicarinate; 22 lamellae beneath fourth finger; 27 lamellae

480 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. beneath fourth toe; preaxial scales of third toe unmodified; tail laterally compressed, not swollen at base, covered with keeled scales, 1.68 times SVL.

FIGURE 10. Pseudocalotes viserion sp. nov. from the radar tower road, Genting Highlands, Pahang, Peninsular Malaysia. Left column: adult females LSUHC 12227 (upper, photograph by L. Lee Grismer), LSUDPC 9907 in light phase (middle, photo by Kurt Orion), and LSUDPC 9908 in dark phase (lower, photo by Kurt Orion). Right column (photographs by L. Lee Grismer): adult female LSUHC 9951 (upper), adult male LSUHC 9484 (middle and lower).

Coloration in life (Fig. 10). Dorsal ground color of body and tail dull yellow; labials, lateral regions of throat, proximal region of brachia and anterior portion of flanks white; ground color of top of head light-green, randomly mottled with small, irregularly shaped, dark markings; four very faint, wide, diffuse, dark vertebral patches between forelimb and hind limb insertions extending onto dorsal margins of flanks; ground color of forelimbs yellow-green, mottled with darkly edged scales; ground color of hind limbs dingy green, limbs faintly banded; wide, diffuse faint bands on anterior region of tail fading posteriorly; gular region beige, bearing a series of faint, diffuse, obliquely oriented, brown stripes in lateral margins; gular pouch yellow; subcaudal region gray; rest of venter beige, generally immaculate; bottoms of hands and feet slightly darker. Variation (Fig. 10). The male paratype (LSUHC 12141) resembles the female holotype (LSUHC 12227) in aspects of color pattern but the overall ground color is brown, not dull-yellow. It also has a greatly swollen tail base. Differences in scalation are presented in Table 5. Distribution. Pseudocalotes viserion sp. nov. is known only from the radar tower road at Ulu Kali at Genting Highlands, Pahang Peninsular Malaysia (Fig. 2).

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 481 TABLE 5. Diagnostic morphological and color pattern characters separating Psedocalotes flavigula and P. viserion sp. nov.

LSUHC LSUHC LSUHC LSUHC LSUHC ZRC FMNH 12114 12227 12420 12580 12581 2.5162 143903 paratype holotype viserion sp. nov. flavigula Postrostrals 3 3 5 6 4 7 / Interparietal No Yes No No No No No Cicrcumorbitals 10 10 10 10 10 10 10 Canthals 5 4 4 4 5 4 5 Superciliaries 7 5 5 6 7 7 7 Scale between rostral and nasal No No No Yes No No / Supralabials contacting nasal Yes Yes Yes Yes Yes Yes Yes Supralabials 7 6 6 7 6 6 7 Infralabials 7 7,8 7 7 6 7 8 postnasal-suborbitals 8 9 9 8 8 / 9 Postmentals 2 3 3 3 4 3 3 Chinshields 3 3 4 5 6 5 6 Gulars 48 47 46 40 40 / 42 Gulars smooth or keeled Smooth Smooth Smooth Weak Smooth Smooth Smooth Gulars granular or flat Flat Flat Flat Flat Flat Flat Flat Transverse gular fold Weak Weak Weak Weak Weak Weak Weak Antehumeral fold Weak Weak Weak Weak Weak Weak Weak Enlarged scales between 2 3 2 3 2 3 3 eye and ear Enlarged posttemporals 2 2 2 2 2 2 2 Enlarged supratympanics 1 1 1 1 1 1 1 Enlarged postrictals No No No No No No No Nuchal crest 9 7 7 9 8 6 10 Gap in crest No No No No No No No Enlarged crest scales No No No No No No No Beyond midbody Flank scales keeled Weak Weak Weak Weak Weak Weak Weak Flank scales plate-like Yes Yes Yes Yes Yes Yes Yes Scales around midbody 35 38 41 44 41 / 44 Midventrals smaller than Yes Yes Yes Yes Yes Yes Yes dorsals 4th finger lamellae 23 22 27 25 26 24,28 22 4th toe lamellae 26 27 30 27 30 / 28 Preaxial lamellae No No No No No No No modified on toe III HL/SVL / 0.3 0.31 0.3 0.29 / 0.1 TL/SVL 1.91 1.68 1.91 1.77 1.58 / 1.79 Male dewlap color Yellow / Yellow / / / Yellow Sex Male Female Male Female Female Female Male SVL 79717976778073

482 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. FIGURE 11. Microhabitat of Pseudocalotes viserion sp. nov. along the radar tower road at Ulu Kali at Genting Higlands, Panhang, Peninsular Malaysia. Photograph by L. Lee Grismer.

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 483 FIGURE 12. Pseudocalotes flavigula from Cameron Highlands, Pahang, Peninsular Malaysia. Upper left: Juvenile from Tanah Rata (LSUDPC 9902, photo by Zhaharil Dzulkafly). Middle left; Adult female from Brinchang Trail (LSUDPC 4350, photo by Leong Tzi Ming). Lower left and right: Adult male from Tanah Rata (LSUHC 12420, photo by L. Lee Grismer). Upper right and middle, respectively: Adult females from Tanah Rata (LSUHC 12581) and Brinchang Trail (LSUHC 12580); photos by Evan S. H. Quah.

Etymology. The specific epithet viserion refers to this species’ resemblance in form and color to the yellowish dragon, Viserion—one of three dragons born in the Dothraki Sea and commanded by Daenerys Targaryen—the Mother of Dragons—in George R. R. Martin’s fictional work Game of Thrones. Natural history. The holotype (LSUHC 12227) was collected during mid-morning at 1000 hrs under broken cloud cover as it was crossing the radar tower road at Ulu Kali at Genting Highlands. The paratype (LSUHC 12141) was found freshly killed on the same road at 1500 hrs. This short stretch of road runs along the crest of Ulu Kali and winds through a mossy cloud forest between 1,700 and 1,800 m in elevation (Fig. 11). The holotype was a gravid female that had just ovulated indicating that the reproductive season of this species extends through March. Comparisons. Pseudocalotes viserion sp. nov. is readily differentiated from all other species of Pseudocalotes except it sister species P. flavigula by having enlarged, plate-like scales on the flanks. It can be separated from P. flavigula by having larger, plate-like scales on the flanks (compare Figs. 10 and 12) as evidenced by having fewer midbody scale rows (35–38 versus 41–44); having more gular scales (47 or 48 versus 40–46); fewer subdigital lamellae on the fourth finger (22 or 23 versus 22–28); and fewer subdigital lamellae on the fourth toe (26 or 27

484 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. versus 26–30) (Table 5). There appears to be less yellow coloration in the gular region of P. f lavigul a in that the yellow does not extend onto the region of the throat anterior to the forelimb insertions as it does in P. viserion sp. nov. and the skin between the scales is white in P. flavigula as opposed to being yellow in P. viserion sp. nov. (Figs. 10,12). Additionally, these two species share a 22% uncorrected pairwise sequence divergence between them. Differences from other species are listed in Table 3.

Variation in Pseudocalotes flavigula

Smith (1924) described Pseudocalotes flavigula from a single specimen collected from the mossy forest along Brinchang Trail in Cameron Highlands (Fig. 13). Smith’s (1924) description was rather brief but was significantly augmented by Grismer (2011) with the examination of another specimen collected from Brinchang Trail (ZRC 2.5162) reported by (Leong 2001) and an additional specimen from Tanah Rata (FMNH 143903). Meristic variation among five of the six known specimens is presented in Table 5. The color pattern in P. flavigula is quite variable (Fig. 12). The dark, dorsal bands range from brown to black and from to distinct and diffuse to barely visible. The light-colored upper lip, lateral throat and proximal section of the brachium can range from immaculate white to lime-green and it may or may not extend onto the anterior portion of the flanks. There is considerable variation in overall ground color that ranges from dull-white to lime-green which we attribute to substrate matching (Fig. 12). We obtained a series of close-up photographs of a juvenile male Pseudocalotes (THNHM 25890) collected at 1400 m in elevation from a montane region in the Hala-Bala Wildlife Sanctuary, Betong District, Yala Province, Thailand along the Thai-Malay border. This habitat is contiguous with the Banjaran Titiwangsa and the data taken from the photographs—most notably the color pattern and the enlarged, plate-like flank scales (Fig. 14)—clearly align this specimen with the flavigula-viserion clade but until the specimen can be examined it will be recognized here as Pseudocalotes sp. nov.

FIGURE 13. Mossy forest habitat of Pseudocalotes flavigula at Brinchang Trail, Cameron Highlands, Pahang, Peninsular Malaysia. Photograph by L. Lee Grismer.

SKY ISLAND DRAGONS FROM PENINSULAR MALAYSIA Zootaxa 4136 (3) © 2016 Magnolia Press · 485 FIGURE 14. Pseudocalotes sp. nov., from the Hala-Bala Wildlife Sanctuary, Betong District, Yala Province, Thailand. Photograph by Sunchai Makchai.

Discussion

With phylogenies and phylogeographies for species across the sky island archipelago of Peninsular Malaysia beginning to accumulate, interesting biogeographic patterns are beginning to emerge. One such pattern involves lineages from Cameron Highlands in the central section of the Banjaran Titiwangsa being more closely related to lineages from Bukit Larut in the separate Banjaran Bintang, 77 km to the west across uninhabitable terrene, rather than being more closely related to lineages farther south at Fraser’s Hill or Genting Highlands in the same mountain range. This pattern is seen with Pseudocalotes rhaegal sp. nov., from Cameron Highlands being more closely related to P. larutensis from Bukit Larut than to P. drogon sp. nov. from Fraser’s Hill (Fig. 6). This same pattern occurs in the Slug Snake Asthenodipsas lasgalanensis (Loredo et al. 2013), certain species of Reed Snakes Macrocalamus (Quah et al. in prep.), and the Larut Skinks Larutia (Grismer et al. 2011). Another pattern seen is the deep genetic divergences between populations from Cameron Highlands and those from farther south at Fraser’s Hill or Genting Highlands. Such is the case for P. flavigula from Cameron Highlands and P. viserion sp. nov. from Genting Highlands; the Mourning Geckos Hemiphyllodactylus titiwangsaensis Zug from Cameron Highlands versus populations from Fraser’s Hill and Genting Highlands (Cobos et al. 2016); the Rock Geckos Cnemaspis temiah Grismer et al. from Cameron Highlands and Cnemaspis flavolineata (Nicholls) from Fraser’s Hill (Grismer et. al. 2014b); the Bent-toed Geckos Cyrtodactylus trilatofasciata Grismer, Wood, Jr., Quah, Anuar, Muin, Sumontha, Norhayati, Bauer, Wangkulangkul, Grismer, & Pauwels and C. sharkai Grismer, Wood, anuar, Quah, Muin, Mohamed, Onn, Sumarli, Loredo, & Heinz from Cameron Highlands and Merapoh, respectively, being distinct from Cyrtodactylus australotitiwangsaensis Grismer, Wood, Jr., Quah, Anuar, Muin, Sumontha, Norhayati, Bauer, Wangkulangkul, Grismer, & Pauwels from Frasers Hill and Genting Highlands (Grismer et al. 2014a); and multiple species of Reed Snakes in the genera Macrocalamus Günther and Calamaria Boie (Quah et al. in prep.).

486 · Zootaxa 4136 (3) © 2016 Magnolia Press GRISMER ET AL. As noted above, recent field work in the montane regions of Peninsular Malaysia has resulted in the discovery of 33 new species of amphibians and reptiles. Thus, discovering new species from such a bizarre group of montane lizards such as Pseudocalotes is exciting but not completely unexpected. The most remarkable aspect of these statistics is that these discoveries, for the most part, are not the result of searching previously unexplored regions or generated by the use of molecular analyses. Rather the majority of these species are newly discovered lineages that are easily diagnosed on the basis of morphology and color pattern and they come from areas that have been popular collecting localities for well over a century (e.g. Penang Hill, Bukit Larut, Cameron Highlands, Fraser’s Hill, Genting Highlands, etc.; see Grismer 2011 for a discussion). Few collections have been made in the largely unexplored Banjaran Timur of northeastern Peninsular Malaysia (see Grismer et al. 2010c and Sumarli et al. 2015 for discussions) and essentially nothing has been collected in the 150 km of the Banjaran Titiwangsa that extends south of Genting Highlands into the states of Negeri Sembilan and Johor. As we begin to focus our research on these southern and northeastern areas, we expect that a plethora of new species will be discovered. The obvious overarching message is that these upland regions serve as incredibly important, vast reservoirs of known and unrealized biodiversity whose content is being severely underestimated, unappreciated and unprotected (e.g. the rapid urbanization of Cameron Highlands and the proposed cable car up to Bukit Larut). If we want to implement any sort of sustainable management programs in order to conserve this biodiversity and these unique, fragile upland habitats, then field-based taxonomy not only must continue, it must be considered paramount.

Acknowledgements

We wish to thank Brandon R. Burch and Ruper G. Lewis for assistance in the field. Funding for LLG came from grants from the College of Arts and Sciences at La Sierra University and from a National Geographic Society Explorers Grant (9277-15). Shahrul Anuar was supported by Universiti Sains Malaysia and Ministry of Higher Education Research Grants. Field work for PLWJ was supported in part by a NSF grant EF-1241885 issued to Jack W. Sites Jr.

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APPENDIX

The following specimens were examined. Pseudocalotes floweri.—Cambodia; LSUHC 8673 Bokor Plateau, Kampot Province. Pseudocalotes flavigula.—West Malaysia, Pahang; LSUHC 12420, 12581 Cameron Highlands, Tanah Rata. FMNH 143903, ZRC 2.5162, LSUHC 12580 Cameron Highlands, Brinchang Trail. Pseudocalotes kakhienensis.—China; CAS 241969, 242381 Yunan Province. Pseudocalotes kingdonwardi.— China; CAS 242579 Yunan Province. Pseudocalotes larutensis.—West Malaysia, Perak, Bukit Larut LSUHC 9052, 9141–42, 10265, 10285, 11289, 12434.