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. viserion 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; 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 Tanah 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.
References
Anderson, J. (1879) Reptilia and amphibia. In: Anatomical and Zoological Researches: Comprising an Account of the Zoological Results of the Two Expeditions to Western Yunnan in 1868 and 1875. Volume I, 703–860, 969–975. Bernard Quarich, London, Vol. I, 705– 860,[1878]
Annandale, N. (1908) Description of a new Species of Lizard of the Genus Salea from Assam. Records of the Indian Museum, 2, 37–38.
Bell, R.C., Parra, J.L., Tonione, M., Hoskin, C.J., MacKenzie, J.B., Williams, S.E. & Moritz, C. (2010) Patterns of persistence and isolation indicate resilience to climate change in montane rainforest lizards. Molecular Ecology 19, 2531–455.
http://dx.doi.org/10.1111/j.1365-294x.2010.04676.xBlackburn, D.C. & Measey, G.J. (2009) Dispersal to or from an African biodiversity hotspot? Molecular Ecology, 18, 1904–1915.
http://dx.doi.org/10.1111/j.1365-294X.2009.04156.xBoulenger, G.A. (1891) On new or little known Indian and Malayan reptiles and batrachians. Annals and Magazine of Natural History, 8, 288–292.
http://dx.doi.org/10.1080/00222939109460437Chan, K.O., Grismer, L.L., Norhayati, A. & Belabut, D. (2009) A new species of Gastrophrynoides (Anura: Microhylidae): an addition to a previously monotypic genus and a new genus for Peninsular Malaysia. Zootaxa, 2124, 63–68.
Chan, K.O., Wood, P.L. Jr., Anuar, S., Muin, M.A., Quah, E.S.H., Sumarli, A.X. & Grismer, L.L. (2014) A new species of upland Stream Toad of the genus Ansonia Stoliczka (Aunra: Bufonidae) from northeastern Peninsular Malaysia. Zootaxa, 3764 (4), 427–440.
http://dx.doi.org/10.11646/zootaxa.3764.4.3Chan-ard, T., Cota, M., Mekchai, S. & Laoteow, S. (2008) A new species of the genus Pseudocalotes (Squamata: Agamidae) from peninsular Thailand. The Thailand Natural History Museum Journal, 3, 25–31.
Cobos, A., Grismer, L.L., Wood, P.L. Jr., Quah, E.S.H., Anuar, S. & Muin, M.A. (2016) Phylogenetic relationships of geckos of the Hemiphyllodactylus harterti group, a new species from Penang Island, Peninsular Malaysia, and a likely case of true cryptic speciation. Zootaxa, 4107 (3), 367–380.
http://dx.doi.org/10.11646/zootaxa.4107.3.5Drummond, A.J., Ashton, B., Buxton, S., Cheung, M., Cooper, A., Duran, C., Field, M., Heled, J., Kearse, M., Markowitz, S., Moir, R., Stones-Havas, S., Sturrock, S., Thierer, T. & Wilson, A. (2011) Geneious V5.6, Available from http://www.Geneious.Com (Accessed 5 Jul. 2016)
Grismer, J.L., Schulte II, J.A., Alexander, A., Wagner, P., Travers, S.L., Buehler, M.D., Welton, L.J. & Brown, R.M. (2016) The Eurasian Invasion: Phylogenomic data reveal multiple Southeast Asian origins for Indian Dragon Lizards. BMC Evolutionary Biology.
http://dx.doi.org/10.1186/s12862-016-0611-6Grismer, L.L. (2006a) A new species of Ansonia Stoliczka 1872 (Anura: Bufonidae) from Central Peninsular Malaysia and a revised taxonomy for Ansonia from the Malay Peninsula. Zootaxa, 1327, 1–21.
Grismer, L.L. (2006b) Two new species of skinks (Genus Sphenomorphus Fitzinger 1843) from the Seribuat Archipelago, West Malaysia. Herpetological Natural History, 9, 151–162.
Grismer, L.L. (2007) A new species of small montane forest floor skink (Genus Sphenomorphus Fitzinger 1843) from southern Peninsular Malaysia. Herpetologica, 63, 544–551.
http://dx.doi.org/10.1655/0018-0831(2007)63[544:ANSOSM]2.0.CO;2Grismer, L.L. (2008) A new species of insular skink (Genus Sphenomorphus Fitzinger 1843) from the Langkawi Archipelago, Kedah, West Malaysia with the first report of the herpetofauna of Pulau Singa Besar and an updated checklist of the herpetofauna of Pulau Langkawi. Zootaxa, 1691, 53–56.
Grismer, L.L. (2011) Lizards of Peninsular Malaysia, Singapore and Their Adjacent Archipelagos. Edition Chimaira, Frankfürt am Main. 728 pp.
Grismer, L.L. Anuar, S., Muin M.A., Quah, E.S.H. & Wood, P.L. Jr. (2013a) Phylogenetic relationships and description of a new upland species of Bent-toed Gecko (Cyrtodactylus Gray, 1827) of the C. sworderi complex from northeastern Peninsular Malaysia. Zootaxa, 3616 (3), 239–252.
http://dx.doi.org/10.11646/zootaxa.3616.3.2Grismer, L.L. & Chan, K.O. (2010a) Another new rock gecko (genus Cnemaspis Strauch 1887) from Pulau Langkawi, Kedah, Peninsular Malaysia. Zootaxa, 2419, 51–62.
Grismer, L.L., Chan, K.O., Grismer, J.L, Wood, P.L. Jr. & Norhayati, A. (2010c) A checklist of the herpetofauna of the Banjaran Bintang, Peninsular Malaysia. Russian Journal of Herpetology, 17, 147–160.
Grismer, L.L., Chan, K.O., Quah, E., Mohd, A.M., A.E. Savage, A.E., Grismer, J.L. Norhayati, A., Greer III1, L.F. & Remegio, A.-C. (2010b) Another new, diminutive Rock Gecko (Cnemaspis Strauch) from Peninsular Malaysia and a discussion of resource partitioning in sympatric species pairs. Zootaxa, 2569, 55–66.
Grismer, L.L., Grismer, J.L., Wood, P.L. Jr. & Chan, K.O. (2008) The distribution, taxonomy, and redescription of the geckos Cnemaspis affinis (Stoliczka 1887) and C. flavolineata (Nicholls 1949) with descriptions of a new montane species and two new lowland, karst-dwelling species from Peninsular Malaysia. Zootaxa, 1931, 1–24.
Grismer, L.L., Grismer, J.L. & Youmans, T.M. (2004) A new species of Leptolalax (Anura: Megophryidae) from Pulau Tioman, West Malaysia. Asiatic Herpetological Research, 10, 8–11.
Grismer, L.L., Norhayati, A. & Chan, K.O. (2009a) A new, diminutive, upland Sphenomorphus Fitzinger 1843 (Squamata; Scincidae) from the Belum-Temengor Forest Complex, Peninsular Malaysia. Zootaxa, 2312, 27–38.
Grismer, L.L., Norhayati, A., Chan, K.O., Belabut, D., Muin, M.A., Wood, P.L. Jr. & Grismer, J.L. (2009b) Two new diminutive species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from Peninsular Malaysia. Zootaxa, 2019, 40–56.
Grismer, L.L. & Quah, E.S.H. (2015) The Rediscovery of Sphenomorphus malayanus Doria, 1888 (Squamata: Scincidae) from the Titiwangsa Mountain Range of Peninsular Malaysia and its re-description as S. senja sp. nov. Zootaxa, 63–70.
http://dx.doi.org/10.11646/zootaxa.3931.1.4Grismer, L.L., Quah, E.S.H., Siler, C.D., Chan, K.O., Wood, P.L. Jr., Grismer, J.L., Anuar, S. & Norhayati, A. (2011) Peninsular Malaysia’s first limbless lizard: a new species of the genus Larutia (Böhme) from Pulau Pinang with a phylogeny of the genus. Zootaxa, 2799, 29–40.
Grismer, L.L., Wood, P.L. Jr., Anuar, S., Muin, M.A., Quah, E.S.H., McGuire, J.A., Brown, R.A., Ngo, V.T. & Pham, H. (2013b) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169, 849–880.
http://dx.doi.org/10.1111/zoj.12064Grismer, L.L. Wood, P.L. Jr., Anuar, S., Quah, E.S.H., Muin, M.A., Mohamed, M., Chan, K.O., Sumarli, A.X., Loredo, A.I. & Heinz, H.M. (2014a) The phylogenetic relationships of three new species of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae) from poorly explored regions in northeastern Peninsular Malaysia. Zootaxa, 3786 (3), 359–381.
http://dx.doi.org/10.11646/zootaxa.3786.3.6Grismer, LL., Wood, PL. Jr., Anuar, S., Quah, ESH., Muin, MA., Chan, KO., Sumarli, AX., & Loredo A.L. (2015a) Repeated evolution of sympatric, paleoendemic species in closely related, co-distributed lineages of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) across a sky-island archipelago in Peninsular Malaysia. Zoological Journal of the Linnean Society, 174, 859–876.
http://dx.doi.org/10.1111/zoj.12254Grismer, L.L., Wood, P.L. Jr., Anuar, S., Riyanto, A., Norhayati, A., Muin, M.A, Sumontha, M., Grismer, J.L., Chan, K.O., Quah, E.S.H. & Pauwels, O.S.G. (2014b) Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia. Zootaxa, 3880 (1), 1–147.
http://dx.doi.org/10.11646/zootaxa.3880.1.1Grismer, L.L., Wood, P.L. Jr., Lee, C.H., Quah, E.S.H., Anuar, S., Ngadi, E. & Sites, J.W. Jr. (2015b) An integrative taxonomic review of the agamid genus Bronchocela (Kuhl, 1820) from Peninsular Malaysia with descriptions of new montane and insular endemics. Zootaxa, 3948 (1), 1–23.
http://dx.doi.org/10.11646/zootaxa.3948.1.1Grismer, L.L., Wood, P.L. Jr., Quah, E.S.H., Anuar, S., Muin, M.A. Sumontha, M., Norhayati, A., Bauer, A.M., Wangkulangkul, S., Grismer, J.L. & Pauwels, O.S.G. (2012) A phylogeny and taxonomy of the Thai-Malay Peninsula Bent-toed Geckos of the Cyrtodactylus pulchellus complex (Squamata: Gekkonidae): combined morphological and molecular analyses with descriptions of seven new species. Zootaxa, 3520, 1–55.
Hallermann, J. & Böhme, W. (2000) A review of the genus Pseudocalotes (Squamata: Agamidae), with description of a new species from West Malaysia. Amphibia-Reptilia, 21, 193–210.
http://dx.doi.org/10.1163/156853800507372Hallermann, J.,Truong, N.Q., Orlov, N. & Ananjeva, N. (2010) A new species of the genus Pseudocalotes (Squamata: Agamidae) from Vietnam. Russian Journal of Herpetology, 17, 31–40.
Harikrishnan, S. & Vasudevan, K. (2013) Rediscovery of Calotes andamanensis Boulenger, 1891, and assessment of its generic allocation (Squamata: Sauria: Agamidae). Herpetozoa, 26, 3–13.
Harvey, M.B., Hamidy, A., Kurniawan, N., Shaney, K. & Smith, E.N. (2015) Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia. Zootaxa, 3841 (2), 211–238.
http://dx.doi.org/10.11646/zootaxa.3841.2.3Inger, R.F. & Stuebing, R.B. (1994) First record of the lizard genus Pseudocalotes (Lacertilia: Agamidae) in Borneo, with description of a new species. Raffles Bulletin of Zoology, 42, 961–965.
Leong, T.M. (2001) A first female topotype of the montane agamid Pseudocalotes flavigula (Smith, 1924) from Peninsular Malaysia, with description of coloration in life. Hamadryad, 26, 364–366.
Loredo, A.I., Wood, P.L. Jr., Quah, E.S.H., Anuar, S.H., Greer, L., Norhayati, A. & Grismer, L.L. (2013) Cryptic speciation within Asthenodipsas vertebralis (Boulenger, 1900) (Squamata: Pareatidae), the description of a new species from Peninsular Malaysia, and the resurrection of A. tropidonotus (Lidth de Jude, 1923) from Sumatra: an integrative taxonomic analysis. Zootaxa, 3664 (4), 505–524.
http://dx.doi.org/10.11646/zootaxa.3664.4.5Macey, J.R., Schulte, J.A., Larson, A., Tuniyev, B.S., Orlov, N. & Papenfuss, T.J. (1999) Molecular phylogenetics, tRNA evolution, and historical biogeography in anguid lizards and related taxonomic families. Molecular phylogenetics and evolution, 12 (3), 250–272.
http://dx.doi.org/10.1006/mpev.1999.0615Maddison, W. & Maddison, D. (2015) Mesquite: a modular system for evolutionary analysis. version 3.04. 2015. Avaliable from: http://mesquiteproject.org (Accessed 5 Jul. 2016)
Mahony, S. (2010) Systematic and taxonomic revaluation of four little known Asian agamid species, Calotes kingdonwardi Smith, 1935, Japalura kaulbacki Smith, 1937, Salea kakhienensis Anderson, 1879 and the monotypic genus Mictopholis Smith, 1935 (Reptilia: Agamidae). Zootaxa, 2514, 1–23.
Manthey, U. (2010) Terralog 7b. Agamid Lizards of Southern Asia/ Agamen des südlichen Asien. Draconinae 2 Leiolepinae. Chimaira, Frankfurt. 168pp Matsui, M., Belabut, D. & Norhayati, A. (2014) Two new fanged frogs from Peninsular Malaysia (Anura: Dicroglossidae). Zootaxa, 3881 (1), 75–93.
http://dx.doi.org/10.11646/zootaxa.3881.1.6Matsui, M., Belabut, D.M., Ahmad, N. & Yong, H.S. (2009) A new species of Leptolalax (Amphibia, Anura, Megophryidae) from Peninsular Malaysia. Zoological Science. Tokyo 26, 243–247.
http://dx.doi.org/10.2108/zsj.26.243Minh, B.Q., Nguyen, M.A.T., & von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30, 1188–1195.
http://dx.doi.org/10.1093/molbev/mst024Nguyen, L.-T., Schmidt, H.A., Haeseler, A. von. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32, 268–274.
http://dx.doi.org/10.1093/molbev/msu300
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) Mr. Bayes 3.2: Efficient Bayesian Phylogenetic Inference and Model Choice across a Large Model Space. Systematic Biology, 61, 539–542.
http://dx.doi.org/10.1093/sysbio/sys029Smith, M.A. (1924) Two new lizards and a new tree frog from the Malay Peninsula. Journal of the Federated States Museum. 11, 183–186.
Stamatakis, A. (2008) Raxml-Vi-Hpc: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics (Oxford), 22, 2688–2690.
http://dx.doi.org/10.1093/bioinformatics/btl446Sumarli, A.X., Grismer, L.L., Anuar, S., Muin, M.A. & Quah, E.S.H. (2015) First report on the amphibians and reptiles of a remote mountain, Gunung Tebu in northeastern Peninsular Malaysia. Checklist, 11, 1–32.
http://dx.doi.org/10.15560/11.4.1679Tamura, K., Stecher, G., Peterson, D., Filipski, A., & Kumar, S. (2013) MEGA6: molecular evolutionary genetics analysis version 6.0. Molecular biology and evolution, mst197.
http://dx.doi.org/10.1093/molbev/mst197Tolley, K.A., Colin, R.T., Measey, G.J., Menegon, M., Branch, W.R. & Matthee, C. (2011) Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hot spot. Journal of Biogeography.
http://dx.doi.org/10.1111/j.1365-2699.2011.02529.xWilcox, T.P., Zwickl, D.J., Heath, T.A. & Hillis, D.M. ( 2002) Phylogenetic relationships of the Dwarf Boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, 25, 361–371.
http://dx.doi.org/10.1016/S1055-7903(02)00244-0Wood, P.L. Jr., Grismer, J.L., Grismer, L.L., Norhayati, A., Chan, K.O. & Bauer, A.M. (2009) Two new montane species of Acanthosaura Gray, 1831 (Squamata: Agamidae) from Peninsular Malaysia. Zootaxa, 2012, 28–46.
Wood, P.L. Jr., Grismer, L.L., Norhayati A. & Juliana S. (2008) Two new species of torrent-dwelling toads Ansonia Stoliczka, 1870 (Anura: Bufonidae) from Peninsular Malaysia. Herpetologica, 64/3, 321–340.
http://dx.doi.org/10.1655/07-065.1