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

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Three New Species of Pseudocalotes Fitzinger (Squamata: Agamidae) from the Sky Island Archipelago of Peninsular Malaysia 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 cir- cumorbitals; five canthals; 7–10 superciliaries; one scale between the rostral and nasal; nine supralabials; eight infralabi- als; 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 low- er 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 circu- morbitals; five canthals; 7–10 superciliaries; one or two scales between the rostral and nasal scales; eight or nine suprala- bials; 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 la- mellae 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 suprala- bials; six or seven infralabials; two or three postmentals; 47 or 48 smooth, flat, gular scales; three chinshields; weak trans- verse 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 yel- low; 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
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