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.
A new species of karst-adapted gekkonid lizard of the genus Cnemaspis Strauch is described from Gua Gunting and Gua Goyang in a karst region of Merapoh, Pahang, Peninsular Malaysia whose unique limestone formations are in immediate danger of being quarried. The new species differs from all other species of Cnemaspis based on its unique suite of morphological and color pattern characters. Its discovery underscores the unique biodiversity endemic to karst regions and adds to a growing list of karst-adapted reptiles from Peninsular Malaysia. We posit that new karst-adapted species endemic to limestone forests will continue to be discovered and these regions will harbor a significant percentage of Peninsular Malaysia's biodiversity and thusly should be conserved rather than quarried.
The fungal pathogen Batrachochytrium dendrobatidis (Bd) infects amphibians on every continent where they occur and is linked to the decline of over 200 amphibian species worldwide. At present, only three published Bd surveys exist for mainland Asia, and Bd has been detected in South Korea alone. In this article, we report the first survey for Bd in Peninsular Malaysia. We swabbed 127 individuals from the six amphibian families that occur on Peninsular Malaysia, including two orders, 27 genera, and 47 species. We detected Bd on 10 out of 127 individuals from four of five states and five of 11 localities, placing the 95% confidence interval for overall prevalence at 4-14%. We detected no variation in Bd prevalence among regions, elevations, or taxonomic groups. The infection intensity ranged from 1 to 157,000 genome equivalents. The presence of Bd infections in native species without clinical signs of disease suggests that Bd may be endemic to the region. Alternately, Bd may have been introduced from non-native amphibians because of the substantial amphibian food trade in Peninsular Malaysia. Under both scenarios, management efforts should be implemented to limit the spread of non-native Bd and protect the tremendous amphibian diversity in Peninsular Malaysia.
An integrative taxonomic analysis is used to identify and describe two new species of the agamid genus Bronchocela (Kuhl) from Peninsular Malaysia: an upland species B. shenlong sp. nov. from Bukit Larut, Perak in the Bintang Mountain Range and Parit Falls, Cameron Highlands, Pahang in the Titiwangsa Mountain Range and an insular species, B. rayaensis sp. nov., from Pulau Langkawi, Kedah off the northwest coast on the border with Thailand. Both species are diagnosed from each other and all other species of Bronchocela on the basis of body shape, scale morphology, and color pattern. The analysis also demonstrates the remarkable genetic similarity of B. cristatella (Kuhl) throughout 1120 km of its range from northern Peninsular Malaysia to western Borneo despite its highly variable coloration and pattern. The two new species are appended to a rapidly growing list of newly described lizard species (60 to date) from Peninsular Malaysia tallied within the last decade.
An integrative taxonomic analysis of the gekkonid lizard genus Hemiphyllodactylus was conducted using molecular, morphological, color pattern, and ecological data on a clade of 19 species, including two new species from karst landscapes in northern Laos and Vietnam. Hemiphyllodactylus houaphanensis sp. nov. (Laos) and H. vanhoensis sp. nov. (Vietnam) are karst-dwelling species whose discovery further emphasizes the unrealized herpetological diversity and site-specific endemism in karst ecosystems across Southeast Asia and the need for increased field work to protect these biodiverse landscapes.
During field work on Hon Son Island, we discovered a new species of the genus Cyrtodactylus inferred from genetic and morphological evidence. Phylogenetic analyses based on the mitochondrial gene ND2, delimit Cyrtodactylus borgattaorum sp. nov. as a new species in the C. intermedius group and the strongly supported sister species of C. phuquocensis in both Bayesian inference and Maximum Likelihood analyses. The new species is distinguished from all other members of the intermedius group by having the combination of 12 supralabials; 10 or 11 infralabials; 22 or 23 small, rounded, smooth, paravertebral tubercles; 12 rows of small, rounded, smooth, longitudinally arranged body tubercles; 35-39 ventrals; 8-10 expanded subdigital lamellae, 12 or 13 unexpanded subdigital lamellae, and 21 or 22 total subdigital lamellae on the fourth toe; 21 or 22 enlarged femorals; 8 or 9 enlarged precloacals; 8 or 9 precloacal pores in males; three rows of enlarged post-precloacals; postcloacal tubercles; enlarged femorals and enlarged precloacals not continuous; proximal femorals nearly same size of distal femorals; body tubercles smooth, greatly reduced; no pocketing between digits on the hands and feet; two dark-colored pigmented blotches on top of head; four dark-colored dorsal bands lacking lightened centers, thinner than light-colored interspaces between bands, bordered by prominent white tubercles; no dark body markings in light-colored interspaces between bands; limbs bearing white tubercles; and six dark-colored and light-colored caudal bands. Cyrtodactylus borgattaorum sp. nov. lives in syntopy with Cyrtodactylus eisenmanae which is the first record of syntopy between granite cave dwelling Cyrtodactylus.
A new species of gekkonid lizard is described from Phnom Khpoh, an isolated karstic hill within an extensive karstic archipelago in Battambang Province, western Cambodia. Phylogenetic analysis using a 1041 base pair fragment of the mitochondrial gene ND2 recovered Hemiphyllodactylus khpoh sp. nov. as the sister species of H. pardalis, found 335 km to the west in southern Thailand. Analyses of meristic, morphometric, and categorical characters of morphology and color pattern differentiated H. khpoh sp. nov. from both the morphologically distinct H. pardalis and the very similar Hemiphyllodactylus sp. from Phnom Kulen, approximately 130 km to the northeast in Siem Reap Province. The statistical morphological analyses and comparisons also showed that H. khpoh sp. nov. is well-differentiated from the recently described H. bokor and H. samkos from the nearby Cardamom Mountains as well from species in clade 6-a recently designated lineage from China, Laos, and Vietnam to which H. khpoh sp. nov. + H. pardalis compose the sister lineage. The discovery of H. khpoh sp. nov. highlights the rich but understudied biodiversity of karstic landscapes, emphasizing the need for continued research in the extensive unexplored karstic archipelago of western Cambodia. This region is poised to reveal unparalleled gekkonid diversity, comparable to that observed in the similarly sized karstic archipelago of the Salween Basin in southern Myanmar. Although karstic landscapes are proving to be some of the most biodiverse habitats on the planet, less than 1% of them are afforded any formal protection.
An integrative taxonomic analysis of the Ptychozoon lionotum group across its range in Indochina and Sundaland recovers P. lionotum sensu lato Annandale, 1905 as paraphyletic with respect to P. popaense Grismer, Wood, Thura, Grismer, Brown, Stuart, 2018a and composed of four allopatric, genetically divergent, ND2 mitochondrial lineages. Multivariate and univariate analyses of continuous and discrete morphological and color pattern characters statistically and discretely diagnose each lineage from one another and together, with maximum likelihood and Bayesian inference analyses, provide the foundation for the recognition of each lineage as a new species-hypotheses corroborated with a Generalized Mixed Yule Coalescent species delimitation analysis. Ptychozoon cicakterbang sp. nov. ranges throughout Peninsular Malaysia to Pulau Natuna Besar, Indonesia; P. kabkaebin sp. nov. is endemic to northern and central Laos; and P. tokehos sp. nov. ranges from southern Thailand south of the Isthmus of Kra northward to Chiang Mai, fringing the Chao Phraya Basin and ranging southward through Cambodia to southern Vietnam. Ptychozoon lionotum sensu stricto ranges from northwestern Laos through southern Myanmar to eastern India. The phylogeographic structure within each species varies considerably with P. lionotum s.s. showing no genetic divergence across its 1,100 km range compared to P. cicakterbang sp. nov. showing upwards of 8.2% sequence divergence between syntopic individuals. Significant phylogeographic structure exists within P. tokehos sp. nov. and increased sampling throughout Thailand may require additional taxonomic changes within this species.
The gekkonid genus Cyrtodactylus is a highly diverse group of lizards (280 + species), which covers an expansive geographic range. Although this genus has been the focus of many taxonomic and molecular systematic studies, species on the Southeast Asian island of Borneo have remained understudied, leading to an unclear evolutionary history with cascading effects on taxonomy and biogeographic inferences. We assembled the most comprehensive multilocus Bornean dataset (one mitochondrial and three nuclear loci) that included 129 novel sequences and representatives from each known Cyrtodactylus species on the island to validate taxonomic status, assess species diversity, and elucidate biogeographic patterns. Our results uncovered a high proportion of cryptic diversity and revealed numerous taxonomic complications, especially within the C. consobrinus, C. malayanus, and C. pubisulcus groups. Comparisons of pairwise genetic distances and a preliminary species delimitation analysis using the Automatic Barcode Gap Discovery (ABGD) method demonstrated that some wide-ranging species on Borneo likely comprise multiple distinct and deeply divergent lineages, each with more restricted distributional ranges. We also tested the prevailing biogeographic hypothesis of a single invasion from Borneo into the Philippines. Our analyses revealed that Philippine taxa were not monophyletic, but were likely derived from multiple separate invasions into the geopolitical areas comprising the Philippines. Although our investigation of Bornean Cyrtodactylus is the most comprehensive to-date, it highlights the need for expanded taxonomic sampling and suggests that our knowledge of the evolutionary history, systematics, and biogeography of Bornean Cyrtodactylus is far from complete.
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.
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.
An integrative taxonomic analysis of three newly discovered populations of the gekkonid genus Cyrtodactylus Gray from Merapoh, Pahang; Gunung Stong, Kelantan; and Gunung Tebu, Terengganu indicate they are part of the C. pulchellus complex and each is a new species and thusly named Cyrtodactylus sharkari sp. nov., C. jelawangensis sp. nov., and C. timur sp. nov., respectively. Each species bears a unique suite of morphological and color pattern characters separating them from each other and all other nominal species in the C. pulchellus complex. Their phylogenetic relationships to each other and other species in the C. pulchellus complex were unexpected in that they are not in accordance with the general distribution of the species in this complex, underscoring the intricate historical biogeography of the Thai-Malay Peninsula. These descriptions highlight our current lack of knowledge concerning the herpetological diversity and distribution of species in northeastern Peninsular Malaysia.
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.
Phylogenetic and morphological analyses delimit and diagnose, respectively, a new population of a karst-dwelling Cyrtodactylus from extreme northern Thailand. The new species, Cyrtodactylusphamiensis sp. nov., of the chauquangensis group inhabits karst caves and outcroppings and karst vegetation in the vicinity of Pha Mi Village in Chiang Rai Province, Thailand. Within the chauquangensis group, Cyrtodactylusphamiensis sp. nov. is the earliest diverging species of a strongly supported clade composed of the granite-dwelling C.doisuthep and the karst-dwelling sister species Cyrtodactylus sp. 6 and C.erythrops. The nearly continuous karstic habitat between the type locality of Cyrtodactylusphamiensis sp. nov. and its close relatives Cyrtodactylus sp. 6 and C.erythrops, extends for approximately 200 km along the border region of Thailand and the eastern limit of the Shan Plateau of Myanmar. Further exploration of this region, especially the entire eastern ~ 95% of the Shan Plateau, will undoubtably recover new populations whose species status will need evaluation. As in all other countries of Indochina and northern Sundaland, the continual discovery of new karst-dwelling populations of Cyrtodactylus shows no signs of tapering off, even in relatively well-collected areas. This only highlights the conservation priority that these unique karstic landscapes still lack on a large scale across all of Asia.
A new species of rock-dwelling Leiolepis is described from the Khorat Plateau in northeastern Thailand. Leiolepisglaurung sp. nov. can be differentiated from all other sexual species of Leiolepis by a combination of having a black gular region with a wide medial yellow stripe, a yellow ventrum with black mottling, bright red to orange subcaudal coloration, having reduced to no expandable flanks, and having only one black transverse bar on the flanks. This is the first rocky habitat-adapted Leiolepis. Leiolepisglaurung sp. nov. demonstrates numerous ecological adaptations to survive in these rocky habitats. Leiolepis are known for their expandable flanks with bright display colors, however Leiolepisglaurung sp. nov. has reduced or no ability to expand its flanks. We hypothesize this is an adaptation to reduce their body diameter to better fit into smaller rocky burrows unlike the larger and deeper burrows constructed in looser soils by other Leiolepis species. This discovery increases the number of Leiolepis species in Thailand to six, and worldwide to 11.
The gekkonid lizard Cyrtodactylus intermedius was formerly considered to be a single widespread species in hilly areas across eastern Thailand through southern Vietnam but has recently been partitioned into a complex of 12 nominal species across its range. A population belonging to the C. intermedius group was recently found in an isolated limestone karst block in Kampot Province in southwestern Cambodia, part of which lies within the recently designated Phnom Preah Kuhear Loung Natural Heritage Site. Comparisons of morphometric, meristic, qualitative morphological and color pattern data, as well as a molecular analysis using the mitochondrial ND2 gene, revealed that this population differs from all other named members of the C. intermedius group, and so is described here as a new species. The karst-dwelling C. regicavernicolus sp. nov. was recovered as the sister species to C. laangensis, the most geographically proximate member of the complex and one that is also restricted to a small limestone karst habitat.
A new insular species Cnemaspis bidongensis sp. nov. (Squamata: Gekkonidae), is described from Pulau Bidong, Terengganu, Peninsular Malaysia and bears a unique suite of morphological and color pattern characters that differentiate it from all other congeners. Cnemaspis bidongensis sp. nov. is the sister species to C. kendallii (Gray) and represents the fifth insular endemic species of Cnemaspis on archipelagos along the east coast of Peninsular Malaysia. This species survived massive deforestation of the small island of Bidong (260 ha) from the mid 1970s to the early 1990s when the island served as a Vietnamese refugee camp and harbored as many as 40,000 people at one time. We hypothesize that this species' generalized lifestyle contributed to its survival, allowing it to seek refuge in rocky microhabitats.
With 90 recognized species, kukri snakes in the genus Oligodon Fitzinger constitute the second largest snake radiation in the world. Oligodon species are collectively distributed across the Asian continent and possess several ecological and morphological attributes that are unique amongst other snakes. Despite their high levels of species richness, evolutionary relationships within Oligodon are poorly understood due to a limited number of samples and genetic markers available in earlier phylogenies. In this study, we assembled the largest molecular dataset of Oligodon to date, which we use to assess the systematics and biogeography of the entire genus. Based on a combination of maximum likelihood and Bayesian phylogenies using fragments of three mitochondrial genes (12 s, 16 s, CytB) and three nuclear genes (Rag1, C-mos, BDNF), we identify eight deeply divergent clades within Oligodon, of which only two correspond with species groupings that were recognized by previous morphological classifications. Four species delimitation methods employed on the mitochondrial portion of the dataset resulted in dramatically divergent estimations of molecular operational taxonomic units (mOTUs). When combined, all four methods support the existence of unrecognized species-level lineages, but also indicate that several other Oligodon species are poorly differentiated genetically and require additional integrative taxonomic research to properly resolve. Based on divergence dating, we demonstrate that Oligodon began to diversify during the early Neogene and hypothesize that the most recent common ancestor of the genus originated in mainland Southeast Asia. We conclude by recognizing eight phylogenetically defined species groups and identify sampling gaps that require further investigation once new data becomes available. This study contributes to a greater understanding of snake evolution on the Asian continent and acts as a baseline for future studies of this speciose genus.
Protected Areas (PAs) are the cornerstone of biodiversity conservation. Here, we collated distributional data for >14,000 (~70% of) species of amphibians and reptiles (herpetofauna) to perform a global assessment of the conservation effectiveness of PAs using species distribution models. Our analyses reveal that >91% of herpetofauna species are currently distributed in PAs, and that this proportion will remain unaltered under future climate change. Indeed, loss of species' distributional ranges will be lower inside PAs than outside them. Therefore, the proportion of effectively protected species is predicted to increase. However, over 7.8% of species currently occur outside PAs, and large spatial conservation gaps remain, mainly across tropical and subtropical moist broadleaf forests, and across non-high-income countries. We also predict that more than 300 amphibian and 500 reptile species may go extinct under climate change over the course of the ongoing century. Our study highlights the importance of PAs in providing herpetofauna with refuge from climate change, and suggests ways to optimize PAs to better conserve biodiversity worldwide.