Marta S. Grismer

Two new Bent-toed Geckos of the Cyrtodactylus pulchellus complex from Peninsular Malaysia and multiple instances of convergent adaptation to limestone forest ecosystems

A new species of limestone cave-adapted gecko of the Cyrtodactylus pulchellus complex, C. hidupselamanya sp. nov., is described from an isolated karst formation at Felda Chiku 7, Kelantan, Peninsular Malaysia. This formation is scheduled to be completely quarried for its mineral content. From what we know about the life history of C. hidupselamanya sp. nov., this will result in its extinction. A new limestone forest-adapted species, C. lenggongensis sp. nov., from the Lenggong Valley, Perak was previously considered to be conspecific with C. bintangrendah but a re-evaluation of morphological, color pattern, molecular, and habitat preference indicates that it too is a unique lineage worthy of specific recognition. Fortunately C. lenggongensis sp. nov. is not facing extinction because its habitat is protected by the UNESCO Archaeological Heritage of the Lenggong Valley due to the archaeological significance of that region. Both new species can be distinguished from all other species of Cyrtodactylus based on molecular evidence from the mitochondrial gene ND2 and its flanking tRNAs as well as having unique combinations of morphological and color pattern characteristics. Using a time-calibrated BEAST analysis we inferred that the evolution of a limestone habitat preference and its apparently attendant morphological and color pattern adaptations evolved independently at least four times in the C. pulchellus complex between 26.1 and 0.78 mya.

Three new karst-dwelling Cnemaspis Strauch, 1887 (Squamata; Gekkoniade) from Peninsular Thailand and the phylogenetic placement of C. punctatonuchalis and C. vandeventeri

Three new species of Rock Geckos Cnemaspis lineogularis sp. nov., C. phangngaensis sp. nov., and C. thachanaensis sp. nov. of the chanthaburiensis and siamensis groups are described from the Thai portion of the Thai-Malay Peninsula. These new species are distinguished from all other species in their two respective groups based on a unique combination of morphological characteristics, which is further supported by mitochondrial DNA (mtDNA) from the NADH dehydrogenase subunit 2 gene (ND2). Cnemaspis lineogularis sp. nov. is differentiated from all other species in the chanthaburiensis group by having a smaller maximum SVL 38 mm, 13 paravertebral tubercles, enlarged femoral scales, no caudal bands, and a 19.5–23.0% pairwise sequence divergence (ND2). Cnemaspis phangngaensis sp. nov. is differentiated from all other species in the siamensis group by having the unique combination of 10 infralabial scales, four continuous pore-bearing precloacal scales, paravertebral tubercles linearly arranged, lacking tubercles on the lower flanks, having ventrolateral caudal tubercles anteriorly present, caudal tubercles restricted to a single paraveterbral row on each side, a single median row of keeled subcaudals, and a 8.8–25.2% pairwise sequence divergence (ND2). Cnemaspis thachanaensis sp. nov. is distinguished from all other species in the siamensis group by having 10 or 11 supralabial scales 9–11 infralabial scales, paravertebral tubercles linearly arranged, ventrolateral caudal tubercles anteriorly, caudal tubercles restricted to a single paravertebral row on each side, a single median row of keeled subcaudal scales, lacking a single enlarged subcaudal scale row, lacking postcloaclal tubercles in males, the presence of an enlarged submetatarsal scale at the base if the 1st toe, and a 13.4–28.8% pairwise sequence divergence (ND2). The new phylogenetic analyses place C. punctatonuchalis and C. vandeventeri in the siamensis group with C. punctatonuchalis as the sister species to C. huaseesom and C. vandeventeri as the sister species to C. siamensis, corroborating previous hypotheses based on morphology. The discovery of three new karst-dwelling endemics brings the total number of nominal Thai Cnemaspis species to 15 and underscores the need for continued field research in poorly known areas of the Thai-Malay Peninsula, especially those that are threatened and often overlooked as biodiversity hot spots.

Phylogenetic taxonomy of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with descriptions of three new species from Myanmar

ABSTRACT A phylogenetic taxonomy of the gekkonid genus Hemiphyllodactylus based on molecular, morphological, and ecological data recovered 28 species, including three new species from the upland regions of the Shan Plateau in eastern Myanmar. Hemiphyllodactylus linnwayensis sp. nov. is a forest-adapted species that may also be a human commensal and H. montawaensis sp. nov. and H. tonywhitteni sp. nov. are karst forest-adapted species. The discovery of three new species from montane regions in eastern Myanmar extends the distribution of a larger monophyletic group of Hemiphyllodactylus westward to the eastern edge of the Ayeyrawady Basin through a series of semi-contiguous, parallel mountain ranges originating in western China and northern Thailand. The discovery of the karst forest-adapted H. montawaensis sp. nov. and H. tonywhitteni sp. nov. further emphasizes the unrealized herpetological diversity endemic to karst ecosystems and the need for increased field work throughout such habitats in South-East Asia. www.zoobank.org/urn:lsid:zoobank.org:pub:E42FA075-E8E0-4005-98AB-12E8D5F23A07

Taxonomy, phylogeny, and distribution of Bronchocela rayaensis (Squamata: Agamidae) on the Thai-Malay Peninsula.

An integrative taxonomic analysis used to identify a new population of Bronchocela from Phuket Island, Thailand indicates it is conspecific with B. rayaensis from the Langkawi Archipelago of northwestern Peninsular Malaysia. An additional specimen photographed from Khura Buri District, Phang-nga Province is also considered to be B. rayaensis and represents a northern range extension of 295 km from the Langkawi Archipelago.

Three more new species of Cyrtodactylus (Squamata: Gekkonidae) from the Salween Basin of eastern Myanmar underscore the urgent need for the conservation of karst habitats

ABSTRACT An integrative phylogenetic taxonomic analysis recovers three additional new species of karst-associated Cyrtodactylus Gray (Squamata: Gekkonidae) – C. bayinnyiensis sp. nov., C. chaunghanakwaensis sp. nov. and C. naungkayaingensis sp. nov. – from a narrow zone in the Salween Basin of Kayin and Mon states in eastern Myanmar from which nine new species were recently described. This degree of unprecedented diversity and site-specific endemism will no doubt continue to rise when at least 44 unsurveyed karstic habitat-islands in this same area are also explored. These data indicate that karst habitats not only serve as foci for speciation, but their rugged terrain spares them from agricultural development and, as such, they are the only habitats in the Salween Basin wherein much of the pre-agricultural herpetofauna can survive. This continues to underscore the fact that karst habitats in Myanmar harbour a significant portion of that country’s herpetofauna, some of which remains undescribed. Despite eastern Myanmar constituting some of the most extensive karstic regions in South-east Asia, they are the least legally protected, with only 1% of their terrain recognised as vulnerable. Until karst habitats in Myanmar are thoroughly investigated, a significant portion of this country’s herpetological diversity will remain underestimated and unprotected. Therefore, issues associated with karst conservation and management in Myanmar should be elevated to a new level of urgency.

A new Cyrtodactylus Gray, 1827 (Squamata, Gekkonidae) from the Shan Hills and the biogeography of Bent-toed Geckos from eastern Myanmar

A phylogenetic taxonomic analysis indicates that a newly discovered population of Cyrtodactylus from the vicinity of Ywangan Town in the Shan Hills, Shan State, Myanmar is a new species (C. ywanganensis sp. nov.) and the earliest diverging member of the linnwayensis group within the previously defined Indochinese clade. The DIVALIKE+J model of a BioGeoBEARS biogeographic analysis indicates that the Indochinese clade evolved in the Shan Hills and Salween Basin of eastern Myanmar and dispersed into Indochina on at least three separate occasions from 18.6-13.4 mya. Once there, uplift of the Tenasserim Mountains and Thai Highlands created the intermedius group, the oldhami group, and C. tigroides of western and southern Thailand which form sister lineages to the linnwayensis group, yathepyanensis group, and the sinyineensis group, respectively, of eastern Myanmar. Diverging lineages within the Indochinese clade highlight the importance of the Thai Highlands and Tenasserim Mountains in that groups evolution and speciation. The discovery of C. ywanganensis sp. nov. in karstic habitats in the Shan Hills continues to underscore the unrealized karst-associated herpetological diversity of this vast, relatively unexplored, upland region and the need for additional field studies.

A new species of Mud Snake (Serpentes, Homalopsidae, Gyiophis Murphy & Voris, 2014) from Myanmar with a first molecular phylogenetic assessment of the genus

A newly discovered species of homalopsid snake from the genus Gyiophis Murphy & Voris is described from the lowlands of Mawlamyine District in Mon state, southeastern Myanmar. Gyiophis salweenensis sp. nov. is presumed to be closely related to G. maculosa Blanford and G. vorisi Murphy based on the similarities in pholidosis and patterning but can be separated from G. maculosa by the shape of its first three dorsal scale rows that are square, ventral scale pattern that lacks a central spot, and a faint stripe on dorsal scale rows 1-4. It can be further distinguished from G. vorisi by its lower number of ventral scales (129 vs. 142-152), lower number of subcaudals (30/29 vs. 41-58), narrow rostral scale, and having more rows of spots on the dorsum (four vs. three). A preliminary molecular analysis using 1050 base pairs of cytochrome b (cytb) recovered G. salweenensis sp. nov. as the sister species to the Chinese Mud Snake (Myrrophis chinensis). G. maculosa and G. vorisi were unavailable for the analysis. The discovery of G. salweenensis sp. nov. highlights the need for more surveys into the herpetological diversity of eastern Myanmar which remains very much underestimated.

Two more new species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from the Shan Hills of eastern Myanmar (Burma)

An integrative phylogenetic analysis recovers two new species of the gekkonid genus Hemiphyllodactylus (Bleeker) from the Shan Hills of eastern Myanmar. Hemiphyllodactylus ywanganensis sp. nov. and H. uga sp. nov. are nested within the eastern Myanmar clade of a previous genus-wide phylogenetic analysis and form a more exclusive monophyletic group with H. linnwayensis. These species differ from each other and all other Hemiphyllodactylus in having unique combinations of character states involving postmental and subcaudal scale morphology; maximum SVL; digital formulae; numbers of chin scales, circumnasals, intersupranasals (=postrostrals), labials, longitudinally arranged dorsal and ventral scales, and pore-bearing femoroprecloacal scales; as well as subtle differences in coloration and pattern. The phylogenetic affinities of the eastern Myanmar clade are similar to those of an endemic clade of Cyrtodactylus from the Shan Hills in that both are more closely related to Indochinese taxa east of Myanmar as opposed to other Indo-Burmese species. The discovery of these new species underscores the underappreciated herpetological diversity of limestone ecosystems as well as the remote nature of the rugged uplands of the Shan Hills and emphasizes the need for continued field work in this region.

A new species of Crocodile Newt Tylototriton (Caudata: Salamandridae) from Shan State, Myanmar (Burma)

A phylogenetic taxonomic analysis of a monophyletic subgroup of the salamandrid genus Tylototriton revealed that a newly discovered population from Ngar Su Village, 1 km south of Ywangan, Shan State, Myanmar is a new species and most closely related to T. shanorum from approximately 80 km to the west in the vicinity of Taunggyi, Shan State. Tylototriton ngarsuensis sp. nov. differs from other closely related species of Tylototriton on basis of varying combinations of characteristics associated with it shorter head, larger size, rib nodule morphology, and overall drab, very dark, coloration, along with its genetic differentiation. Tylototriton ngarsuensis sp. nov. also appears to breed later in the year than most other species. Unfortunately, this species like many other Asian newts, is being harvested for the pet and medicinal trade and given its restricted distribution, this could pose a serious threat to its long-term survival.

A phylogenetic taxonomy of the Cyrtodactylus peguensis group (Reptilia: Squamata: Gekkonidae) with descriptions of two new species from Myanmar

A phylogenetic taxonomy of species in the Cyrtodactylus peguensis group from the Ayeyarwady Basin of Myanmar is constructed based on color pattern, morphology, and molecular systematic analyses using the mitochondrial gene NADH dehydrogenase subunit 2. Newly collected samples from the type locality of C. peguensis and other localities indicate that this clade is endemic to central Myanmar and contains at least seven species, four of which are undescribed. Three species, including C. peguensis occur in the low hills of the Bago Yoma Range within the central portion of the Ayeyarwady Basin. Two of these, C. myintkyawthurai sp. nov. from the northern and central Bago Yoma and C. meersi sp. nov. which is syntopic with C. peguensis in the southern Bago Yoma are described herein. As more lowland hilly areas bordering, and within the Ayeyarwady Basin are surveyed, more new species of this group are likely to be discovered. These discoveries continue the recent surge of descriptions of new species of Cyrtodactylus that are being discovered in Myanmar.