Filling the BINs of life: Report of an amphibian and reptile survey of the Tanintharyi (Tenasserim) Region of Myanmar, with DNA barcode data

Despite threats of species extinctions, taxonomic crises, and technological advances in genomics and natural history database informatics, we are still distant from cataloguing all of the species of life on earth. Amphibians and reptiles are no exceptions; in fact new species are described nearly every day and many species face possible extinction. The number of described species continues to climb as new areas of the world are explored and as species complexes are examined more thoroughly. The use of DNA barcoding provides a mechanism for rapidly estimating the number of species at a given site and has the potential to record all of the species of life on Earth. Though DNA barcoding has its caveats, it can be useful to estimate the number of species in a more systematic and efficient manner, to be followed in combination with more traditional, morphology-based identifications and species descriptions. Herein, we report the results of a voucher-based herpetological expedition to the Tanintharyi (Tenasserim) Region of Myanmar, enhanced with DNA barcode data. Our main surveys took place in the currently proposed Tanintharyi National Park. We combine our results with photographs and observational data from the Chaung-naukpyan forest reserve. Additionally, we provide the first checklist of amphibians and reptiles of the region, with species based on the literature and museum. Amphibians, anurans in particular, are one of the most poorly known groups of vertebrates in terms of taxonomy and the number of known species, particularly in Southeast Asia. Our rapid-assessment program combined with DNA barcoding and use of Barcode Index Numbers (BINs) of voucher specimens reveals the depth of taxonomic diversity in the southern Tanintharyi herpetofauna even though only a third of the potential amphibians and reptiles were seen. A total of 51 putative species (one caecilian, 25 frogs, 13 lizards, 10 snakes, and two turtles) were detected, several of which represent potentially undescribed species. Several of these species were detected by DNA barcode data alone. Furthermore, five species were recorded for the first time in Myanmar, two amphibians (Ichthyophis cf. kohtaoensis and Chalcorana eschatia) and three snakes (Ahaetulla mycterizans, Boiga dendrophila, and Boiga drapiezii)

COVID-19 and civil unrest undoing steady gains in karst conservation and herpetological research in Myanmar, and an impediment to progress

The COVID-19 pandemic and political turmoil in Myanmar has dealt a severe blow to the country’s progress in herpetological research and the protection of limestone habitats. Both afflictions have reversed much of the scientific and conservation gains made in the past decade, and continue to hinder exploratory surveys and continued monitoring of threatened karst ecosystems. There is an urgent need to resume field studies and conservation efforts as soon as possible and continue enhancing the capacity of local scientific and technical staff in Myanmar

Sex Chromosome Turnover in Bent-Toed Geckos (\u3cem\u3eCyrtodactylus\u3c/em\u3e)

Lizards and snakes (squamates) are known for their varied sex determining systems, and gecko lizards are especially diverse, having evolved sex chromosomes independently multiple times. While sex chromosomes frequently turnover among gecko genera, intrageneric turnovers are known only from Gekko and Hemidactylus. Here, we used RADseq to identify sex-specific markers in two species of Burmese bent-toed geckos. We uncovered XX/XY sex chromosomes in Cyrtodactylus chaunghanakwaensis and ZZ/ZW sex chromosomes in Cyrtodactylus pharbaungensis. This is the third instance of intrageneric turnover of sex chromosomes in geckos. Additionally, Cyrtodactylus are closely related to another genus with intrageneric turnover, Hemidactylus. Together, these data suggest that sex chromosome turnover may be common in this clade, setting them apart as exceptionally diverse in a group already known for diverse sex determination systems

Cyrtodactylus pyadalinensis Grismer & Wood & Thura & Win & Quah 2019, sp. nov.

<i>Cyrtodactylus pyadalinensis</i> sp. nov. <p>Panluang-Pyadalin Cave Bent-toed Gecko</p> <p>(Figs. 5, 6)</p> <p> <b>Holotype.</b> Subadult male CAS 226143 collected during the evening on 16 July 2002 by G.O.U. Wogan, R. S. Lucas, J. V. Vindum, Thin Thin, and A. K. Shein from Panluang-Pyadalin Cave Wildlife Sanctuary, Ywangan Township, Shan State, Myanmar (21.107000°N, 96.352111°E; 220 m in elevation).</p> <p> <b>Paratypes.</b> Subadult male CAS 226142 collected during the evening on 16 July 2002 by Htun Win from Panluang-Pyadalin Cave Wildlife Sanctuary, Ywangan Township, Shan State, Myanmar (21.115801°N, 96.360694°E; 346 m in elevation). Adult female LSUHC 13932 collected at 2100 hrs on 26 March 2018 by Perry L. Wood Jr., Nyo Min Htwe, and L. Lee Grismer from immediately outside the Pyadalin Cave, Panluang-Pyadalin Cave Wildlife Sanctuary, Ywangan Township, Taunggyi District, Shan State, Myanmar (21.13275°N, 96.34026°E; 306m in elevation.)</p> <p> <b>Additional material examined.</b> Hatchling LSUHC 13933 bearing the same locality and collection data as LSUHC 13932 except that it was collected by Nyo Min Htwe, Perry L. Wood Jr., and L. Lee Grismer.</p> <p> <b>Diagnosis.</b> <i>Cyrtodactylus pyadalinensis</i> <b>sp. nov.</b> differs from all other species in the <i>peguensis</i> group by having the unique combination of eight supralabials and 6–8 infralabials; 31–33 paravertebral tubercles; 19–21 longitudinal rows of body tubercles; 38–40 ventral scales; 16–18 subdigital lamellae on the fourth toe; 14 or 15 femoral pores in males; nine or 10 precloacal pores in males; two or three rows of enlarged, post-precloacal scales; top of head bearing dark blotches; 4–6 dark body bands; dark body bands lacking paravertebral elements; and maximum SVL of 72.1 mm (Table 4).</p> <p> <b>Description of holotype.</b> Subadult male, SVL 51.1 mm; head moderate in length (HL/SVL 0.28), wide (HW/ HL 0.61), somewhat flattened (HD/HL 0.37), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region concave, canthus rostralis rounded; snout elongate (ES/HL 0.39), rounded in dorsal profile; eye large (ED/HL 0.21); ear opening elliptical, moderate in size (EL/HL 0.09); eye to ear distance greater than diameter of eye; rostral rectangular, partially divided dorsally, bordered posteriorly by large left and right supranasals separated small internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal, posteriorly by three postnasals (upper largest), ventrally by first supralabial; eight (R,L) rectangular supralabials extending to below midpoint of eye; seven (R,L) infralabials tapering smoothly to below posterior margin of eye; scales of rostrum and lores flat, larger than granular scales on top of head and occiput; scales on top of head and occiput intermixed with slightly enlarged tubercles; dorsal supraciliaries not elongate or keeled; mental triangular, bordered laterally by first infralabials and posteriorly by large, left and right trapezoidal postmentals that contact medially for 70% of their length posterior to mental; one row of slightly large chinshields tapering posteriorly to fourth infralabial; and gular and throat scales small, granular, grading posteriorly into larger, flatter, smooth, subimbricate to imbricate, pectoral and ventral scales.</p> <p>Body relatively short (AG/SVL 0.44) with weak ventrolateral folds; dorsal scales small, interspersed with larger, semi-regularly arranged, moderately keeled tubercles; tubercles extend from occiput onto base of tail but no farther; tubercles on occiput and nape smaller than those on posterior portion of body; approximately 21 longitudinal rows of dorsal tubercles; 31 paravertebral tubercles; approximately 40 flat, imbricate, ventral scales larger than dorsal scales; nine pore-bearing precloacal scales; two rows of large post-precloacal scales; and no deep precloacal groove or depression.</p> <p>Forelimbs moderate in stature, relatively short (FL/SVL 0.15); flat scales of anterior margin of forearm larger than those on body, not interspersed with tubercles; palmar scales raised; digits relatively short, well-developed, inflected at basal, interphalangeal joints, slightly narrower distal to inflections; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.16), covered dorsally by granular scales interspersed with slightly larger, weakly keeled tubercles and anteriorly by large, flat, imbricate scales; ventral scales of femora flat, imbricate, larger than dorsals, lacking a distinct row of enlarged femoral scales; small postfemoral scales form an abrupt union with large, flat ventral scales of posteroventral margin of thigh; subtibial scales flat, imbricate; plantar scales slightly raised; digits relatively short, well- developed, inflected at basal, interphalangeal joints, slightly narrower distal to inflections; 18 subdigital lamellae (R,L) on fourth toe; claws well-developed, base of claw sheathed by a dorsal and ventral scale; two enlarged postcloacal tubercles at base of tail; and postcloacal scales flat.</p> <p>Tail original, 56.9 mm in length, 4.9 mm in width at base, tapering to a point; dorsal scales square and flat; transversely enlarged, median, subcaudal scales twice as wide as long, not extending onto lateral margin of tail in original section.</p> <p> <b>Coloration in life (Fig. 6).</b> Dorsal ground color of head body, limbs, and tail yellow; top of head bearing large, dark-brown, irregularly shaped, conjoined blotches edged in yellow; dark-brown, wide, nuchal loop extending from posterior margin of one eye, across occiput, to posterior margin of other eye; nape bearing a wide, dark-brown band edged in yellow; five wide, dark-brown body bands between limb insertions edged in yellow lacking paravertebral components, posterior three obliquely oriented; large, round, dark-brown markings between body bands; 6–8 smaller, diffuse brown blotches along lower margins of flanks; one dark-brown post-sacral band edged in yellow not bearing paravertebral elements; 12 dark-brown caudal bands wider than the 12 yellow caudal bands; dorsal portion of forelimbs darkly mottled to banded; dorsal portion of hind limbs bearing irregularly shaped, darkbrown blotches edged in yellow; and all ventral surfaces generally beige, immaculate.</p> <p> <b>Variation.</b> The paratypes generally approach the holotype in most aspects of coloration and pattern. The most notable difference is in the dorsal banding pattern where the paratypes have more transversely oriented dark, dorsal bands as opposed to the holotype whose bands are more obliquely oriented. In the paratype CAS 226142, the central band between the limb insertions is somewhat oval-shaped and bears a central light spot. The dark dorsal bands in the paratype LSUHC 13933 are considerably narrower than those of all the other specimens in the type series and the distal one-half of the tail is missing. The paratype LSUHC 13932 has a regenerated tail bearing a dark-beige ground color overlain with small, dark, irregularly shaped markings. Meristic differences among specimens of the type series are resented in Table 5.</p> <p> <i>……continued on the next page</i></p> <p> 22 <b>TABLE 5.</b> Meristic, mensural, and color pattern data for <i>Cyrtodactylus pyadalinensis</i> <b>sp. nov.</b> and <i>C. nyinyikyawi</i> <b>sp. nov.</b> / = data unobtainable.</p> <p> <i>……continued on the next page</i></p> <p> <b>Distribution.</b> <i>Cyrtodactylus pyadalinensis</i> <b>sp. nov.</b> is known only from the vicinity of the Panluang-Pyadalin Cave in the Panluang-Pyadalin Cave Wildlife Sanctuary, Ywangan Township, Taunggyi District, Shan State, Myanmar (Fig. 1).</p> <p> <b>Etymology.</b> The specific epithet, <i>pyadalinensis,</i> is a toponym referring to the type locality in the vicinity of the Pyadalin Cave.</p> <p> <b>Natural history.</b> The type series of <i>Cyrtodactylus pyadalinensis</i> <b>sp. nov.</b> were all collected in the vicinity of the Kinda Reservoir between the Panulaung River and the Pyadalin Cave. This area is within the foothills and rocky plain of the Nwalabo Mountain range on the western fringe of the Shan Plateau (Fig. 1) between 213 and 306 m in elevation. The habitat is composed of low-lying, highly eroded terrain and scree of the Nwalabo Mountains. It bears scattered karstic rocks and boulders surrounded by disturbed, drought-adapted, scrub Indiang Forest vegetation that is seasonally burned (Fig. 7). All specimens were found at night between 1900 and 2300 hrs among small rocks and leaf-leaf litter.</p> <p> <b>Comparisons.</b> <i>Cyrtodactylus pyadalinensis</i> <b>sp. nov.</b> descends from one of the deeper divergences of the <i>peguensis</i> group and the sister species to the clade (<i>C. niyniykyawi</i> <b>sp. nov.</b> (<i>C. peguensis</i> (<i>C. pyinyaungensis</i> and <i>C. myintkyawthurai</i>))) from which it differs by an uncorrected pairwise sequence divergence of 9.0–10.3%. From <i>C. meersi</i> and <i>C. annandalei</i> which occur outside this clade, it differs by 10.7–11.0% and 14.0–14.3%, respectively. It differs from all other species except <i>C. nyinyikyawi</i> <b>sp. nov.</b> in the dark-brown dorsal bands lacking as opposed to having paravertebral elements. It differs further from <i>C. annandalei</i> in that the top of the head is blotched as opposed to being unicolor. Differences from <i>C. nyinyikyawi</i> <b>sp. nov.</b> are presented in the Comparisons section above. Statistically significant mean differences in meristic characters among <i>C. pyadalinensis</i> <b>sp. nov.</b>, <i>C. myintkyawthurai,</i> and <i>C. pyinyaungensis</i> are presented in Tables 3 and 4.</p>Published as part of <i>Grismer, L. Lee, Wood, Perry L., Thura, Myint Kyaw, Win, Nay Myo & Quah, Evan S. H., 2019, Two more new species of the Cyrtodactylus peguensis group (Squamata: Gekkonidae) from the fringes of the Ayeyarwady Basin, Myanmar, pp. 274-294 in Zootaxa 4577 (2)</i> on pages 284-291, DOI: 10.11646/zootaxa.4577.2.3, <a href="http://zenodo.org/record/3993550">http://zenodo.org/record/3993550</a&gt

Two more new species of the Cyrtodactylus peguensis group (Squamata: Gekkonidae) from the fringes of the Ayeyarwady Basin, Myanmar

Grismer, L. Lee, Wood, Perry L., Thura, Myint Kyaw, Win, Nay Myo, Quah, Evan S. H. (2019): Two more new species of the Cyrtodactylus peguensis group (Squamata: Gekkonidae) from the fringes of the Ayeyarwady Basin, Myanmar. Zootaxa 4577 (2): 274-294, DOI: https://doi.org/10.11646/zootaxa.4577.2.

Cyrtodactylus nyinyikyawi Grismer & Wood & Thura & Win & Quah 2019, sp. nov.

<i>Cyrtodactylus nyinyikyawi</i> sp. nov. <p>Shwe Settaw Bent-toed Gecko</p> <p>(Fig. 4)</p> <p> <b>Holotype.</b> Adult female CAS 226139 collected on 14 September 2002 at 1030 hrs by Thin Thin, Kyi Soe Lwin, and Hla Tun from Shwe Settaw Wildlife Sanctuary, Min Bu Township, Magway Region, Myanmar (20.05972°N, 94.59611°E; 137 m in elevation).</p> <p> <b>Diagnosis.</b> <i>Cyrtodactylus nyinyikyawi</i> sp. nov. differs from all other species in the <i>peguensis</i> group by having the unique combination of nine supralabials; eight infralabials; 35 paravertebral tubercles; 20 longitudinal rows of body tubercles; 35 ventral scales; 19 subdigital lamellae on the fourth toe; four rows of enlarged, post-precloacal scales; keeled, conical, body tubercles; top of head bearing dark blotches; five dark, body bands; dark body bands lacking paravertebral elements; and maximum SVL of 64.5 mm (Table 3).</p> <p> <b>Description of holotype.</b> Adult female, SVL 64.5 mm; head moderate in length (HL/SVL 0.25), wide (HW/ HL 0.62), somewhat flattened (HD/HL 0.42), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region concave, canthus rostralis rounded; snout elongate (ES/HL 0.40), rounded in dorsal profile; eye large (ED/HL 0.20); ear opening elliptical, moderate in size (EL/HL 0.09); eye to ear distance greater than diameter of eye; rostral rectangular, partially divided dorsally, bordered posteriorly by large left and right supranasals separated small internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal, posteriorly by three postnasals (upper largest), ventrally by first supralabial; nine (R) supralabials extending to below midpoint of eye ball; eight rectangular infralabials tapering smoothly to below posterior margin of eye ball; scales of rostrum and lores flat, larger than granular scales on top of head and occiput; scales on top of head and occiput intermixed with slightly enlarged tubercles; dorsal supraciliaries not elongate or keeled; mental triangular, bordered laterally by first infralabials and posteriorly by large, left and right trapezoidal postmentals that contact medially for 70% of their length posterior to mental; one row of slightly enlarged chinshields tapering posteriorly to fourth infralabial; and gular and throat scales small, granular, grading posteriorly into larger, flatter, smooth, subimbricate to imbricate, pectoral and ventral scales.</p> <p>Body relatively short (AG/SVL 0.52) with weak ventrolateral folds; dorsal scales small, interspersed with larger, semi-regularly arranged, weakly keeled tubercles; tubercles extend from occiput onto base of tail but no farther; tubercles on occiput and nape smaller than those on posterior portion of body; approximately 20 longitudinal rows of dorsal tubercles; approximately 35 paravertebral tubercles; 35 flat, imbricate, ventral scales larger than dorsal scales; seven dimpled, precloacal scales; and four rows of enlarged post-precloacal scales.</p> <p>Forelimbs moderate in stature, relatively short (FL/SVL 0.13); flat scales of anterior margin of forearm larger than those on body, not interspersed with tubercles; palmar scales raised; digits relatively short, well-developed, inflected at basal, interphalangeal joints, slightly narrower distal to inflections; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.17), covered dorsally by granular scales interspersed with slightly larger, weakly keeled tubercles and anteriorly by large, flat, imbricate scales; ventral scales of femora flat, imbricate, larger than dorsals, lacking a distinct row of enlarged femoral scales; small postfemoral scales form an abrupt union with larger, flat ventral scales of posteroventral margin of thigh; subtibial scales flat, imbricate; plantar scales slightly raised; digits relatively short, welldeveloped, inflected at basal, interphalangeal joints, slightly narrower distal to inflections; 19 subdigital lamellae (R,L) on fourth toe; claws well-developed, base of claw sheathed by a dorsal and ventral scale; two enlarged postcloacal tubercles at base of tail; postcloacal scales flat.</p> <p>Proximal 14.4 mm of tail original, posterior 32.4 mm regenerated, 5.4 mm in width at base, tapering to a point; dorsal scales of original of tail granular rapidly becoming flatter posteriorly; dorsal scales of regenerated tail large, flat, semi-regular in arrangement; and transversely enlarged, median, subcaudal scales twice as wide as long, not extending onto lateral margin of tail in original section.</p> <p> <b>Coloration in life (Fig. 4).</b> Dorsal ground color of head body, limbs, and tail yellow; top of head bearing large, dark-brown, irregularly shaped blotches edged in yellow; dark-brown, wide, nuchal loop extending from posterior margin of one eye, across occiput, to posterior margin of other eye; nape bearing a wide, dark-brown band edged in yellow; four wide, dark-brown body bands between limb insertions edged in yellow lacking distinct, paravertebral components; large, round, dark-brown markings between body bands two and three and three and four; seven or eight smaller, somewhat diffuse brown blotches along lower margins of flanks; one wide, dark-brown post-sacral band edged in yellow not bearing paravertebral sections; one dark-brown caudal band on original portion of tail; regenerated portion of tail light-colored bearing diffuse, randomly arranged, dark markings; dorsal portion of forelimbs darkly mottled to banded; dorsal portion of hind limbs bearing irregularly shaped, dark-brown blotches edged in yellow. All ventral surfaces generally beige, immaculate.</p> <p> <b>Distribution.</b> <i>Cyrtodactylus nyinyikyawi</i> <b>sp. nov.</b> is known only from the type locality of the Shwe Settaw Wildlife Sanctuary, Min Bu Township, Magway Region, Myanmar (Fig. 1).</p> <p> <b>Etymology.</b> The specific epithet, <i>nyinyikyawi</i> is a patronym honoring Nyi Nyi Kyaw the Director General of the Forestry Department for his contributions to conservation efforts in Myanmar in general and to our work in particular.</p> <p> <b>Natural History.</b> The holotype CAS 226139 is a gravid female collected on the ground in secondary dry deciduous hardwood forest at 1030 hrs along the edges of a small seasonal lake. Being gravid with two eggs indicates that the monsoon month of September falls within this species’ reproductive season.</p> <p> <b>Comparisons</b> <i>Cyrtodactylus nyinyikyawi</i> <b>sp. nov.</b> is the sister species of a clade that includes the sister species <i>C. peguensis</i> and <i>C. pyinaungensis</i> and <i>C. myinykyawthurai</i> (Fig. 2). It differs from <i>C. peguensis</i> by a 9.3% uncorrected pairwise sequence divergence, from <i>C. myintkyawthurai</i> by an 11.3–12.3% sequence divergence, and from <i>C. pyinaungensis</i> by a sequence divergence of 10.0–10.3%. Outside this clade, <i>C. nyinyikyawi</i> <b>sp. nov.</b> differs from <i>C. meersi</i> by a sequence divergence of 10%, from the Panluang-Pyadalin Cave population if differs by 9.3– 9.7%, and from <i>C. annandalei</i> it differs by 10.3%. It differs from all species of the <i>peguensis</i> group by having a higher number of paravertebral tubercles (35 vs. 25–33, collectively) and differs from all other species except the Panluang-Pyadalin Cave population by the dark dorsal bands lacking distinct, paravertebral elements as opposed to having them (Figs. 4, 5). It differs further from <i>C. annandalei</i> in that the top of the head is blotched as opposed to being unicolor It differs further from the Panluang-Pyadalin Cave population in having 35 as opposed to 38–40 ventral scale rows. Other differences separating <i>C. nyinyikyawi</i> <b>sp. nov.</b> form other <i>peguensis</i> group species are listed in Tables 2 and 3.</p> <p> continued. <i>myintkyawthurai</i>, <i>C. pyinyaungensis</i>, and <i>C. pyadalinensis</i> <b>sp. nov.</b></p> <p> <b>Remarks.</b> Some (<i>i.e.</i> Dayrat 2005; Thomas Hbrek, <i>in litt,</i> 2018) have grave concerns about descriptions of new species based on only a single specimen, and posit that this should ‘never’ be done because such a description cannot take into account intraspecific variation that could potentially preclude its specific recognition. The myopic nature of this opinion notwithstanding, it is not only incorrect philosophically—as the ontological existence of a species is independent of its diagnosis (Frost & Kluge 1994)—it is counterproductive in reality. Additionally, such tactics would impede biodiversity studies in general and taxonomy in particular. Estimates have shown that 19% of all new vertebrate species described between 2000 and 2010 were based on a single specimen (Lim <i>et al.</i> 2012) and that number is likely to have increased in the last seven years—an indication that often, this is the logical first step in constructing species delimitation hypotheses (<i>i.e.</i> integrative taxonomies). Furthermore, with well-supported phylogenetic data such as that herein indicating that the specimen in question is not nested within or a sister species to any other species and shares a 9.3–12.3% uncorrected pairwise sequence divergence from its closest relatives, renders any morphological arguments to the contrary moot—regardless of these arguments’ erroneous conflation and confusion of ontology and epistemology. However, in this case, <i>Cyrtodactylus nyinyikyawi</i> <b>sp. nov.</b> has widely differing morphological and color pattern characters that at this point, distinguish it from all other species in the <i>peguensis</i> group, thus further eclipsing assumptions that it may be conspecific with something else. We <i>are</i> concerned about describing a new species based on a single specimen but only because the diagnosis is incomplete, not because it has anything to do with the reality of the specimen representing a distinct, independently evolving lineage based on the molecular evidence. Given the general ongoing biodiversity crisis throughout Southeast Asia, we felt it prudent to describe this species for potential protective status rather than delay its publication for the sake of a better diagnosis.</p> <p> The weak part of recognizing this specimen as a distinct species is not the incomplete diagnosis, but that the species was initially delimited on the basis of a single-locus mtDNA phylogeny. It is well-documented that mtDNA phylogenies can reveal significant structure in a data set by recovering sequentially nested monophyletic groups even though within that same data set, nuclear genes can indicate significant gene flow among these groups (<i>e.g.</i> Shaw 2002; Fisher-Reid & Wiens,2011; Toews & Brelsford 2012), thus precluding their species status. This weakens any hypothesis of specific identity based <i>solely</i> on mtDNA data. Nonetheless, given the current data available concerning its phylogenetic relationships and the discrete morphological and color pattern differences separating <i>C. nyinyikyawi</i> <b>sp. nov.</b> from its congeners in the <i>peguensis</i> group, we regard its specific identity as a robust, testable hypothesis.</p>Published as part of <i>Grismer, L. Lee, Wood, Perry L., Thura, Myint Kyaw, Win, Nay Myo & Quah, Evan S. H., 2019, Two more new species of the Cyrtodactylus peguensis group (Squamata: Gekkonidae) from the fringes of the Ayeyarwady Basin, Myanmar, pp. 274-294 in Zootaxa 4577 (2)</i> on pages 280-283, DOI: 10.11646/zootaxa.4577.2.3, <a href="http://zenodo.org/record/3993550">http://zenodo.org/record/3993550</a&gt

Discovery of the westernmost population of the genus Ansonia Stoliczka (Anura, Bufonidae) with the description of a new species from the Shan Plateau of eastern Myanmar

Quah, Evan S. H., Grismer, L. Lee, Wood, Perry L., Thura, Myint Kyaw, Oaks, Jamie R., Lin, Aung (2019): Discovery of the westernmost population of the genus Ansonia Stoliczka (Anura, Bufonidae) with the description of a new species from the Shan Plateau of eastern Myanmar. Zootaxa 4656 (3): 545-571, DOI: https://doi.org/10.11646/zootaxa.4656.3.1

Four new Burmese species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from distantly related parapatric clades from the Shan Plateau and Salween Basin

Grismer, L. Lee, Wood, Perry L., Quah, Evan S. H., Thura, Myint Kyaw, Oaks, Jamie R., Lin, Aung (2020): Four new Burmese species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from distantly related parapatric clades from the Shan Plateau and Salween Basin. Zootaxa 4758 (1): 45-82, DOI: https://doi.org/10.11646/zootaxa.4758.1.

Hemiphyllodactylus ngwelwini Grismer & Wood & Quah & Thura & Oaks & Lin 2020, sp. nov.

<i>Hemiphyllodactylus ngwelwini</i> sp. nov. Ngwe Lwin’s Slender Gecko <p>(Figs. 4, 5)</p> <p> <b>Holotype.</b> Adult male (LSUHC 14473) collected on 2 August 2019 at 1955 hrs by L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, and Aung Lin at the Thayeumin Cave, State, Myanmar (20.72288°N 96.58994°E WGS; 1057 m in elevation).</p> <p> <b>Paratypes</b>. Females and juvenile LSUHC 14474–76 bear the same collection data as the holotype. Females LSUHC 14328–29 and male 14330 were collected from Pwe Hla Village, Shan State (20.84125°N 96.69030°E WGS; 1416 m in elevation) by L. Lee Grismer, Perry L. Wood, Jr, Evan S. H. Quah, Myint Kyaw Thura, Jamie R. Oaks, and Aung Lin on 14 November 2018 and female LSUHC 14489 bears the same collecting locality but was collected by L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, and Aung Lin on 3 August 2019. Female LSUHC 14326 and male 14327 from the Myintmahati Cave, Shan State (20.59082°N 96.61198°E WGS; 1326 m in elevation) were collected by L. Lee Grismer, Perry L. Wood, Jr, Evan S. H. Quah, Myint Kyaw Thura, Jamie R. Oaks, and Aung Lin on 15 November 2018.</p> <p> <b>Diagnosis.</b> <i>Hemiphyllodactylus ngwelwini</i> <b>sp. nov.</b> can be separated from all other species of <i>Hemiphyllodactylus</i> by possessing the unique combination of having a maximum SVL of 40.2 mm; 9–13 chin scales; enlarged postmentals; five circumnasal scales; 1–3 intersupranasals (=postrostrals); 8–11 supralabials; 8–10 infralabials; 11–14 longitudinally arranged dorsal scales at midbody contained within one eye diameter and seven or eight ventral scales; four subdigital lamellae on the first finger and toe; 15–22 continuous, pore-bearing, femoroprecloacal scales in males; no plate-like subcaudal scales; adult females variably yellow; a dark postorbital stripe extending to at least base of neck; dorsolateral light-colored spots usually present on trunk; no dark, dorsolateral or ventrolateral stripe on trunk; dark zig-zag of paravertebral markings on trunk variable; light-colored postsacral marking variably bearing anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all Burmese species in Tables 3 and 6 and from all other species of <i>Hemiphyllodactylus</i> from southern China and western Thailand (clades 3 and 4 in Grismer <i>et al.</i> (2017: Table 3)).</p> <p> <b>Description of holotype.</b> Adult male, SVL 34.4 mm; head triangular in dorsal profile, depressed, distinct from neck; lores flat; rostrum moderate in length (SN/SVL 0.10); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernible; snout moderate, rounded in dorsal profile; eye large; ear opening elliptical, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supra- nasals; two differently sized intersupranasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); eight (R, L) rectangular supralabials tapering to below posterior margin of eye; 9, 10 (R, L) rectangular infralabials tapering to below posterior margin of eye; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials, posteriorly by two large nearly square postmentals; each postmental in contact with first infralabial, bordered laterally by single slightly enlarged sublabial; 10 chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate throat and even larger pectoral scales which grade into slightly larger, subimbricate ventrals.</p> <p>Body somewhat elongate (AG/SVL 0.49), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 12 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, eight ventral scales contained within one eye diameter; precloacal scales slightly larger than abdominal scales; pore-bearing precloacal scales continuous with pore-bearing femoral scales, totaling 21 pore-bearing femoroprecloacal scales; single enlarged tubercle on anterior margin of hemipenial swelling; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-3-3-3 (R, L); four transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with flat, juxtaposed scales dorsally and larger, flat subimbricate scales ventrally; plantar scales low, flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-3-3-3 (R, L); four transversely expanded lamellae on digit I; claws on digits II–V well-developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; caudal scales not occurring in whorls; dorsal caudal scales of original tail larger than dorsal body scales, flat, subcycloid, subimbricate; subcaudals slightly larger than dorsal caudals, not plate-like. Raw and ratiometric mensural data are presented in Table 7.</p> <p>....Continued next page</p> <p> <b>Coloration in life (Fig. 4).</b> All <i>Hemiphyllodactylus</i> are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below was taken when the holotype was photographed the morning after capture, approximately 12 hours following the time of collection while during its light-phase. Ground color of top of head, body, and limbs, gray and densely mottled with darker markings; top of head overlain with dark, semi-reticulate pattern; broad, dark, diffuse pre- and postorbital stripe extends from the external nares, through eye to just posterior of forelimb insertion on the body; pairs of diffuse, dark, paravertebral markings counter-shaded posteriorly with diffuse white spots extend from nape to base of tail transforming into a distinct, dark (nearly black), post-sacral band; immaculate, beige post-sacral marking immediately posterior to black post-sacral band not bearing light-colored, anteriorly projecting arms; dorsum and flanks faintly mottled with diffuse speckling; limbs bearing irregularly shaped, diffuse, dark markings; original tail bearing eight dark bands; gular region generally immaculate, except for darker lateral areas and faint stippling in scales; pigmentation density increases posteriorly with abdomen being generally gray; ground color of dorsal caudal region beige, bearing nine black diffuse bands not encircling tail; median subcaudal region faintly orange, generally immaculate.</p> <p> <b>Variation (Figs. 4, 5).</b> The color patterns of the paratypes generally match that of the holotype and no interpopulational differences were observed (Table 6). The dark, dorsal pattern of LSUHC 14326, 14328, 14330, 14489 is not as bold as that of the holotype. LSUHC 14476 is a juvenile with a broken tail. The tails of LSUHC 14326–28, 14330, and 14489 are regenerated and generally unicolor gray. Variation in scales counts, mensural data, and additional minor aspects in coloration are presented in Table 7.</p> <p> <b>Distribution.</b> <i>Hemiphyllodactylus ngwelwini</i> <b>sp. nov.</b> is known from three localities across a distance of approximately 29 km from Pwe Hla Village in the north to the Thayeumin and Myintmahati caves in the south, Shan State (Fig. 1).</p> <p> <b>Natural History.</b> All individuals from Pwe Hla Village were found on man-made structures in highly disturbed forest. LSUHC 14328–29 and LSUHC 14489 were collected on the walls of cement water tanks and LSUHC 14330 was collected from the underside of a wooden roof from a nearby rest shelter along the road. Both specimens from the Myintmahati Cave population (LSUHC 14326–27) were collected on cement structures immediately outside of a limestone cave in highly disturbed forest. LSUHC 14473 (the holotype) and LSUHC 14474–76 from the Thayeumin Cave population were found outside the limestone cave on corrugated tin shacks, cement buildings, and other man-made structures (Fig. 6) between a rice paddy and an isolated tract of highly disturbed forest.</p> <p> <b>Etymology.</b> The specific epithet recognizes and honors Mr. Ngwe Lwin, northern Program Manager of Fauna and Flora International in Myanmar. Mr. Ngwe Lwin has been supportive and invaluably instrumental in facilitating our field work in Myanmar since October of 2017.</p> <p> <b>Comparisons</b>. The molecular analyses indicate that <i>Hemiphyllodactylus ngwelwini</i> <b>sp. nov.</b> is a genetically distinct member of the north lineage composed of three, putatively, interbreeding populations being that the intrapopulational uncorrected pairwise sequence divergence across 29 km is only 1.0% and that individuals from the three populations are polyphyletic with respect to one another (Fig. 1, Table 8). <i>Hemiphyllodactylus ngwelwini</i> <b>sp. nov.</b> is the sister species to a clade composed of <i>H. ywanganensis</i> and the sister species <i>H. uga,</i> and <i>H. linnwayensis</i> (Fig. 1) from which it bears an uncorrected pairwise sequence divergence of 8.4% from <i>H. linnwayensis,</i> 9.0% from <i>H. uga,</i> and 8.5% from <i>H. ywanganensis</i> (Table 8). <i>H. ngwelwini</i> <b>sp. nov.</b> differs significantly from <i>H. linnwayensis, H. montawaensis,</i> and <i>H. tonywhitteni</i> in mean values of CS (10.8 vs 5.0, <i>p</i> = 7.42 -06; 10.8 vs 6.3, <i>p</i> = 4.93 -05; and 10.8 vs. 6.6, <i>p</i> = 5.14 -05; respectively; Table 3); differs significantly from <i>H. tonywhitteni</i> in mean values of DS (12.7 vs 14.8, <i>p</i> = 0.014; Table 3); and from <i>H. montawaensis</i> it differs significantly in adjusted mean values of HL (1.985 vs 2.852, <i>p</i> = 0.021; Fig. 2, Table 3). <i>Hemiphyllodactylus ngwelwini</i> <b>sp. nov.</b> differs from <i>H. uga</i> and <i>H. ywanganensis</i> (n = 2) by having four subdigital lamelae on the first finger as opposed to two or three in <i>H. uga</i> and three in <i>H. ywanganensis</i> and four subdigital lamelae on the first toe as opposed to two or three in the latter two species. However, the sample sizes of the latter two species (n =4 and n = 2, respectively) are so small that these values are likely to change with the addition of more samples. Owing to the high intraspecific variability of color pattern characters in <i>H. ngwelwini</i> <b>sp. nov.</b> (Figs. 4, 5), no interspecific differences between it and other members of the north lineage were found.</p>Published as part of <i>Grismer, L. Lee, Wood, Perry L., Quah, Evan S. H., Thura, Myint Kyaw, Oaks, Jamie R. & Lin, Aung, 2020, Four new Burmese species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from distantly related parapatric clades from the Shan Plateau and Salween Basin, pp. 45-82 in Zootaxa 4758 (1)</i> on pages 57-63, DOI: 10.11646/zootaxa.4758.1.2, <a href="http://zenodo.org/record/3730792">http://zenodo.org/record/3730792</a&gt

Hemiphyllodactylus kyaiktiyoensis Grismer & Wood & Quah & Thura & Oaks & Lin 2020, sp. nov.

<i>Hemiphyllodactylus kyaiktiyoensis</i> sp. nov. Kyaiktiyo Mountain Slender Gecko <p>(Figs. 7, 9)</p> <p> <b>Holotype.</b> Adult female (LSUHC 14032) collected on 4 November 2018 at 1900 hrs by Evan S. H. Quah, L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, Jamie R. Oaks, and Aung Lin along a trail leading down the northwest facing flank of the mountain from the Golden Rock Pagoda, Mon State, Myanmar (17.47840°N 97.10042°E WGS; 1057 m in elevation).</p> <p> <b>Paratypes</b>. Female paratypes (LSUHC 14030–31, 14033–34) bear the same collection data as the holotype.</p> <p> <b>Diagnosis.</b> <i>Hemiphyllodactylus kyaiktiyoensis</i> <b>sp. nov.</b> can be separated from all other species of <i>Hemiphyllodactylus</i> by possessing the unique combination of having a maximum SVL of 43.4 mm; 8–10 chin scales; enlarged postmentals; five circumnasal scales; 3–5 intersupranasals (=postrostrals); 7–10 supralabials; eight or nine infralabials; 12–16 longitudinally arranged dorsal scales at midbody contained within one eye diameter and eight or nine ventral scales; four subdigital lamellae on the first finger and four or five on first toe; no plate-like subcaudal scales; adult females not yellow; a dark postorbital stripe extending to at least base of neck; dorsolateral light-colored spots on trunk; no dark, dorsolateral stripe on trunk; faint, dark, ventrolateral stripe on trunk; no dark-colored pattern on trunk; wide, light-brown, nearly unicolor, vertebral region on trunk; beige, postsacral marking bearing anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all Burmese species in Tables 3 and 6 and from all other species of <i>Hemiphyllodactylus</i> from southern China and western Thailand (clades 3 and 4 in Grismer <i>et al.</i> [2017: Table 3]).</p> <p> <b>Description of holotype</b> Adult female SVL 43.4 mm; head triangular in dorsal profile, depressed, distinct from neck; lores flat; rostrum moderate in length (SN/SVL 0.11); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening elliptical, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supranasals; three equally sized intersupranasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); nine (R, L) rectangular supralabials tapering to below posterior margin of eye; 8 (R, L) rectangular infralabials tapering to below posterior margin of eye; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest, slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by two rectangular postmentals; each postmental in contact with first infralabial and bordered laterally by slightly smaller sublabial; 10 chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate throat and even larger pectoral scales which grade into slightly larger, subimbricate ventrals.</p> <p>Body somewhat elongate (AG/SVL 0.54), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 13 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, eight ventral scales contained within one eye diameter; precloacal scales larger than abdominal scales; no pore-bearing femoroprecloacal scales; single enlarged tubercle on lateral margin of tail base; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular, U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-3-4-3 (R, L); four transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with raised, juxtaposed scales dorsally and by larger, flat, subimbricate scales anteriorly and ventrally; plantar scales slightly raised, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular, and Ushaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-4-4-4 (R, L); five transversely expanded lamellae on digit I; claws on digits II–V well-developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; caudal scales occurring in whorls; dorsal caudal scales of original tail larger than dorsal body scales, flat, subcycloid, subimbricate; ventrolateral caudals enlarged, weakly flared giving a fringe-like appearance punctuated every five or six scales by even larger, raised scales; subcaudals flat slightly larger than dorsal caudals, not plate-like; posterior one-half of tail regenerated; dorsal caudals of regenerated portion smaller, more irregular in shape; other caudal scales generally same shape as those in original tail. Morphometric data are presented in Table 9.</p> <p> <b>Coloration in life (Fig. 7).</b> All <i>Hemiphyllodactylus</i> are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below is of that when the holotype was photographed the morning after capture, approximately 12 hours after the time of collection when in its light-phase. Ground color of top of head, body, limbs, and tail dull-yellow or straw; top of head overlain with a dense dark-brown mottling giving top of head a brownish appearance; brown, semi-reticulate pattern on top of head; broad, dark, diffuse pre- and postorbital stripe extends from external nares, through eye to forelimb, preorbital portion very faint; wide, nearly immaculate, light-brown stripe with crenulated margins extends from nape to base of tail, nearly covering entire dorsum; invaginations of crenulated margins contain a diffuse, light-colored spot that collectively appear as regularly spaced, dorsolateral spots along trunk; faint, dark, ventrolateral stripe on trunk; immaculate, beige, postsacral marking bears light-colored, anteriorly projecting arms; flanks faintly mottled with diffuse brown speckling; limbs bearing irregularly shaped, diffuse, brown bands and markings; base of toes bearing a faintly orange spot; gular region mottled with brown and faint stippling in scales; pigmentation density in scales decreases posteriorly with the abdomen being generally beige with faint stippling; original portion of tail bearing four faint, irregularly shaped, brown bands and heavily mottled interspaces; enlarged scales of ventrolateral fringe highlighted in white; subcaudal region densely stippled; regenerated portion of tail brownish with faint, dark markings.</p> <p> <b>Variation (Figs. 7, 9).</b> The color pattern of the paratypes (LSUHC 14030–31, 14033–34) generally match that of the holotype. The dark, dorsal patterns of LSUHC 14031 and 14033 are slightly bolder. LSUHC 14030 has a more lightly colored dorsal pattern overall and the dorsolateral trunk spots are more obvious. Variation in scales counts, mensural data, and additional minor aspects in coloration are presented in Table 9.</p> <p> <b>Distribution.</b> <i>Hemiphyllodactylus kyaiktiyoensis</i> <b>sp. nov.</b> is known only from the type locality on Kyaiktiyo Mountain in Mon State, Myanmar but is expected to occur in other, nearby upland areas (Fig. 1).</p> <p> <b>Natural History.</b> All individuals of the type series were found on the brick wall of a house at night beneath a neon light approximately 2.5 meters above the ground in disturbed hill forest (Fig. 10). No other geckos were found on the wall but <i>Hemidactylus garnotii, Hemidactylus brookii, Gehyra mutilata,</i> and <i>Cyrtodactylus aequalis</i> were abundant on all other nearby vegetation and human-made structures. Specimens LSUHC 14032–34 were gravid with two eggs indicating the month of November falls within the reproductive season of this species. Although no males were found, we cannot determine if this species is parthenogenetic or not.</p> <p> <b>Etymology.</b> The specific epithet is a toponym referring to the type locality of Kyaiktiyo Mountain, Mon State, Myanmar.</p> <p> <b>Comparisons</b>. The molecular analyses indicate that <i>Hemiphyllodactylus kyaiktiyoensis</i> <b>sp. nov.</b> is a genetically distinct member of the south lineage and is the sister species to <i>H. pinlaungensis</i> <b>sp. nov.</b> (Fig. 1) from which it bears an uncorrected pairwise sequence divergence of 9.5% (Table 10). <i>Hemiphyllodactylus kyaiktiyoensis</i> <b>sp. nov.</b> differs significantly from <i>H. pinlaungensis</i> <b>sp. nov.</b> in having a lower mean value of CS (9.2 vs 11.2, <i>p</i> = 0.014) and a lower adjusted mean value of SN (1.242 vs 1.363) <i>p</i> = 0.018, respectively; Table 6, Fig. 3). It differs further from <i>H. pinlaungensis</i> <b>sp. nov.</b> by having a significantly different centroid position based on the factor loadings of the PC1–3 (<i>p</i> = 0.002; Fig. 3). This latter metric can not be considered a diagnostic character but serves as a measure of the quantitative difference between these two species in multivariate space. Its differences from <i>H. zwegabinensis</i> <b>sp. nov.</b> are listed above in the comparisons section for that species.</p>Published as part of <i>Grismer, L. Lee, Wood, Perry L., Quah, Evan S. H., Thura, Myint Kyaw, Oaks, Jamie R. & Lin, Aung, 2020, Four new Burmese species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from distantly related parapatric clades from the Shan Plateau and Salween Basin, pp. 45-82 in Zootaxa 4758 (1)</i> on pages 68-70, DOI: 10.11646/zootaxa.4758.1.2, <a href="http://zenodo.org/record/3730792">http://zenodo.org/record/3730792</a&gt