Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia

Abstract

A well-supported and well-resolved phylogeny based on a concatenated data set from one mitochondrial and two nuclear genes, six morphological characters, and nine color pattern characters for 44 of the 50 species of the Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) is consistent with the previous taxonomy of Cnemaspis based solely on morphology and color pattern. Cnemaspis is partitioned into four major clades that collectively contain six species groups. The monophyly of all clades and species groups is strongly supported and they are parapatrically distributed across well-established, biogeographical regions ranging from southern Vietnam westward through southern Indochina, southward through the Thai-Malay Peninsula, then eastward to Borneo. Eight new species (Cnemaspis omari sp. nov. from the Thai-Malaysian border; C. temiah sp. nov. from Cameron Highlands, Pahang, Malaysia; C. stongensis sp. nov. from Gunung Stong, Kelantan, Malaysia; C. hangus sp. nov. from Bukit Hangus, Pahang, Malaysia; C. sundagekko sp. nov. from Pulau Siantan, Indonesia; C. peninsularis sp. nov. from southern Peninsular Malaysia and Singapore, and C. mumpuniae sp. nov. and C. sundainsula sp. nov. from Pulau Natuna Besar, Indonesia) are described based on morphology and color pattern and all but C. sundagekko sp. nov. are included in the phylogenetic analyses. Cnemaspis kendallii is polyphyletic and a composite of six species. An updated taxonomy consistent with the phylogeny is proposed for all 50 species and is based on 25 morphological and 53 color pattern characters scored across 594 specimens. Cladogenetic events and biogeographical relationships within Cnemaspis were likely influenced by this group's low vagility and the cyclical patterns of geographical and environmental changes in Sundaland over the last 25 million years and especially within the last 2.5 million years. The phylogeny indicates that nocturnality, diurnality, substrate preferences, and the presence of ocelli in the shoulder regions have evolved independently multiple times.

A new species of Rock Gecko of the genus Cnemaspis Strauch, 1887 (Squamata: Gekkonidae) from Belitung Island, Indonesia

“…The specimens were captured by hand, euthanized with 20% benzocaine diluted in ethanol, fixed with 10% buffered formalin, and stored in 70% ethanol. The following measurements followed Grismer et al (2014a) and were taken with Mitutoyo dial calipers to the nearest 0.1 mm, using an AmScope dissecting microscope when necessary. Measurements and meristic counts were made on the right side of each specimen (except for MZB.Lace.14075, for which lamellae counts were obtained from the left fingers on account of damage to the right hand).…”

“…We compared the specimens to specimens and descriptions of all congeners (Appendix 1). Some of the information on character states and their distribution in other Southeast Asian species were obtained from Grismer et al (2014a), Amarasinghe et al (2015), Iskandar et al (2017), and Kurita et al (2017). All type specimens were deposited in the Museum Zoologicum Bogoriense (MZB), Indonesian Institute of Sciences (LIPI), Cibinong, Bogor, Indonesia.…”

“…The new species is differentiated from all other species in the Southern Sunda clade (sensu Grismer et al 2014a) by having a unique combination of characters including: (1) a maximum SVL of 54.1 mm, (2) five or six postmental scales, (3) enlarged submetacarpal scales on the first finger, (4) enlarged submetatarsal scales on the first toe, (5) keeled ventral scales, (6) absence of precloacal pores, (7) absence of enlarged femoral scales, (8) absence of shield-like subtibial scales, (9) caudal tubercles encircling the tail, (10) an interrupted median row of enlarged keeled subcaudals, (11) presence of a distinct furrow on the lateral surface of the tail (12) 22-24 lamellae beneath fourth toe, and (13) two postcloacal tubercles on each side of the tail base.Key words: Cnemaspis, Belitung Island, Indonesia, new species …”

“…The genus is polyphyletic, with at least three divergent unrelated clades (Gamble et al 2012). In Southeast Asia, Cnemaspis ranges from southern Laos in the north, southward through Vietnam, Cambodia, and Thailand, and the Thai-Malay Peninsula, eastward to Borneo through the Seribuat, Anambas and Natuna archipelagoes, and southward to Sumatra and the Mentawai Archipelago (de Rooij, 1915;Grismer et al 2010Grismer et al , 2013Grismer et al , 2014aWood et al 2013;Amarasinghe et al 2015;Iskandar et al 2017). The genus Cnemaspis Strauch currently includes 135 recognized species (Uetz et al 2017), five of which are present on Borneo: C. kendallii (Gray, 1845), C. nigridia (Smith, 1925), C. dringi , C. paripari Grismer & Chan, 2009 and C. leucura Kurita, Nishikawa, Matsui & Hikida, 2017.…”

“…Several new species have been described from Indonesia in recent years (e.g. Grismer et al 2014a;Amarasinghe et al 2015;Iskandar et al 2017), bringing the total number of currently recognized species in the country to 18 (Uetz et al 2017).…”