Abstract
Two new species of diminutive, sympatric, lowland, leaf-litter skinks of the genus Tytthoscincus Linkem, Diesmos & Brown from the Sekayu region of Hulu Terengganu, Terengganu State in northeastern Peninsular Malaysia are described on the basis genetic and morphological data. One of the new species, T. monticolus sp. nov., was collected in a hilly riparian area along Sungai (=river) Bubu and is most closely related to an undescribed species from the Tembak Reservoir area. The other, T. keciktuek sp. nov. collected along Sungai Peres, is most closely related to T. perhentianensis Grismer, Wood, & Grismer from Pulau (=island) Perhentian Besar. Sympatry and syntopy of multiple, specialized, unrelated, leaf-litter species of Tytthoscincus was previously only known from upland areas and these new species represent the first example of lowland of sympatry. More importantly, however, these endemic species add to a growing body of research and discoveries that continue to underscore the unrealized biodiversity of the riparian systems of Hulu Terengganu and the Sekayu region and their need for protection and continued study.
References
Brandley, M.C., Huelsenbeck, J.P. & Wiens, J.J. (2008) Rates and patterns in the evolution of snake-like body form in squamate reptiles: evidence for repeated re-evolution of lost digits and long-term persistence of intermediate body forms. Evolution, 62, 2042–2064.
https://doi.org/10.1111/j.1558-5646.2008.00430.xDrummon, A.J., Ashton, B., Buxton, S., Cheung, M., Cooper, A. & Duran C. (2011) Geneious V5.4. Available from http://www.geneious.com/ (Accessed 28 May 2018)
Greer, A.E. (1989) The Biology and Evolution of Australian lizards. Chipping Norton: Survey Beatty and Sons Pty. Ltd.
Grismer, L.L., Ismail, L.H.B, Awang, M.T., Rizal, S.A. & Ahmad, A.B. (2014) A new species of lowland skink (genus Lipinia Gray, 1845) from northeastern Peninsular Malaysia. Zootaxa, 3821 (4), 457–464.
http://dx.doi.org/10.11646/zootaxa.3821.4.4
Grismer, L.L., Muin, M.A., Wood, P.L., Anuar, S. & Linkem, C.W. (2016a) The transfer of two clades of Malaysian Sphenomorphus Fitzinger (Squamata: Scinidae) into the genus Tytthoscincus Linkem, Diesmos, & Brown and the description of a new Malaysian swamp-dwelling species. Zootaxa, 4092 (2), 231–242.
http://dx.doi.org/10.11646/zootaxa.4092.2.6Grismer, L.L., Wood, Jr., P.L., Lim, K.P.L. & Liang, L.J. (2017b) A new species of swamp-dwelling skink (Tytthoscincus) from Singapore and Peninsular Malaysia. Raffles Bulletin of Zoology, 65, 574–584.
Grismer, L.L., Wood, Jr., P.L., Quah, E.S.H., Aunar, S., Ngadi, E.B., Izam, N.A.M. & Ahmad, N. (2017a) Systematics, ecomorphology, cryptic speciation and biogeography of the lizard genus Tytthoscincus Linkem, Diesmos & Brown (Squamata: Scincidae) from the sky-island archipelago of Peninsular Malaysia. Zoological Journal of the Linnean Society.
https://doi.org/10.1093/zoolinnean/zlx067
Grismer, L.L., Wood, Jr., P.L., Syafiq, M.F., Badli-Sham, B.H., Rizal, S.A., Ahmad, A.B. & Quah, E.S.H. (2016b) On the taxonomy and phylogeny of the skinks Lipinia sekayuensis Grismer, Ismail, Awang, Rizal, & Ahmad and Lipinia surda Boulenger from Peninsular Malaysia. Zootaxa, 4147 (1), 59–66.
https://doi.org/10.11646/zootaxa.4147.1.3
Huelsenbeck, J. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755.
Huelsenbeck, J.P., Ronquist, F., Nielsen, R. & Bollback, J.P. (2001) Bayesian Inference of Phylogeny and Its Impact on Evolutionary Biology. Science, 294, 2310–2314. D
https://doi.org/10.1126/science.1065889Jombart, T., Devillard, S. & Balloux, F. (2010) Discriminant analysis of principal components: a new method for the analysis of genetically structured populations. BMC Genetics, 11, 94.
https://doi.org/10.1186/1471-2156-11-94
Kaiser, H.F. (1960) The application of electronic computers to factor analysis. Education and Psychological Measurement, 20, 141–151.
Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A., & Jermiin, L.S. (2017) ModelFinder: Fast Model Selection for Accurate Phylogenetic Estimates. Natural Methods, 14, 587–589.
https://doi.org/10.1038/nmeth.4285Karin, B.R., Das, I., & Bauer, A.M. (2016) Two new species of diminutive leaf-litter skinks (Squamata: Scincidae: Tytthoscincus) from Gunung Penrissen, Sarawak, Malaysia (northern Borneo). Zootaxa, 4093 (3), 407–423.
https://doi.org/10.11646/zootaxa.4093.3.7
Kumar, S., Stecher, G., & Tamura, K. (2016) MEGA7: Moleculat evolutionary genetics analaysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870-–1874.
https://doi.org/10.1093/molbev/msw054
Lee, M.S.Y., Skinner, A. & Camacho, A. (2013) The relationship between limb reduction, body elongation and geographical range in lizards (Lerista, Scincidae). Journal of Biogeography, 2013, 1–8.
https://doi.org/10.1111/jbi.12094Linkem, C.W. (2013) Molecular Phylogenetics and Biogeography of Sphenomorphini (Squamata: Scincidae). Unpublished PhD Thesis, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, Kansas.
Linkem, C.W., Diesmos, A.C. & Brown, R.M. (2011) Molecular systematics of the Philippine forest skinks (Squamata: Scincidae: Sphenomorphus): testing morphological hypotheses of interspecific relationships. Zoological Journal of the Linnean Society, 163, 217–1243.
https://doi.org/10.1111/j.1096-3642.2011.00747.x
Macey, J.R., Schulte, J.A., Larson, A., Tuniyev, B.S., Orlov, N. & Papenfuss, T.J. (1997) Molecular phylogenetics, tRNA evolution, and historical biogeography in anguid lizards and related taxonomic families. Molecular Phylogenetics and Evolution, 12, 250–272.
https://doi.org/10.1006/mpev.1999.0615
Maddison, W.P. & Maddison, D.R. (2015) Mesquite: a modular system for evolutionary analysis. Version 3.04. Available from: http://mesquiteproject.org (Accessed 28 May 2018)
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, LA.
Minh, Q., Nguyen, M.A.T. & von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30, 1188–1195.
https://doi.org/10.1093/molbev/mst024
Ng, P.K.L. & Ahmad, A.B. (2016) A new genus and new species for an unusual semi-terrestrial poyamid crab (Decopoda: Brachyura) with a bilobed mandibular palp from Peninsular Malaysia. Journal of Crustacean Biology, 36, 823–832.
https://doi.org/10.1163/1937240X-00002492
Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32, 268–274.
https://doi.org/10.1093/molbev/msu300
R Core Team. (2015) R: A language and environment for statistical computing. R Foundation for Statistical Computing. Vienna. Available from: http://www.R-project.org (Accessed 28 May 2018)
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
https://doi.org/10.1093/bioinformatics/btg180
Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer v1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (Accessed 28 May 2018)
Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 6.5 (16 August 2016). [Electronically accessible at http://www.asih.org/, American Society of Ichthyologists and Herpetologists, Washington, DC.]
Schmitz, A., Brandley, M.C., Mausfeld, P., Vences, M., Glaw, F., Nussbaum, R.A. & Reeder, T.W. (2005) Opening the black box: phylogenetics and morphological evolution of the Malagasy fossorial lizards of the subfamily “Scincinae”. Molecular Phylogenerics and Evolution, 34, 118–133.
https://doi.org/10.1016/j.ympev.2004.08.016
Siler, C.D. & Brown, R.M. (2011) Evidence for repeated acquisition and loss of complex body-form characters in an insular clade of Southeast Asian semi-fossorial skinks. Evolution, 65, 2641–2663.
https://doi.org/10.1111/j.1558-5646.2011.01315.xSumarli, A., Grismer, L.L., Wood, Jr., P.L., Ahmad, A.B., Rizal, S., Ismail, L., Izam, N.A.M., Ahmad, N. & Linkem, C. (2016) The first riparian skink (Genus: Sphenomorphus Strauch, 1887) from Peninsular Malaysia and its relationship to their Indochinese and Sundaic species. Zootaxa, 4173 (1), 29–44.
http://doi.org/10.11646/zootaxa.4173.1.3Taylor, E.H. (1935) A taxonomic study of the cosmopolitan lizards of the genus Eumeces with an account of the distribution and relationships of its species. University of Kansas Science Bulletin, 23, 1–643.
Wilcox, 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.
https://doi.org/10.1016/S1055-7903(02)00244-0
Wu, N.C., Alton, L.A., Clemente, C.J., Kearney, M..R. & White, CR. (2015) Morphology and burrowing energetics of semi-fossorial skinks (Liopholis spp.). Journal of Experimental Biology, 218, 2416–2426.
https://doi.org/0.1242/jeb.113803