Since the book-formatted, monographic treatment of the lizards of Peninsular Malaysia, Singapore, and their adjacent islands by Grismer (2011), 47 additional species have been added to that fauna bringing the total to 174. One species, Cyrtodactylus stresemanni, was removed from the lizard fauna based on its likely mislabeled type locality; 44 species were described as new-29 gekkonids, 11 scincids, five agamids, and one dibamid; and three others, Cnemaspis narathiwatensis, Cyrtodactylus brevipalmatus, and Scincella melanosticta were recently discovered in northern Peninsular Malaysia. Taxonomic changes for seven previously described species occurring after 2011 are noted and significant, new locality records are documented here for additional species. The intent of this review is to provide an up to date peer-reviewed checklist for the lizard species of this region prior to a pending updated book monograph.
The taxonomic status of Asthenodipsas vertebralis in Borneo has been plagued with uncertainty over the last eighty years. An examination of museum collections resulted in the discovery of a voucher specimen of A. vertebralis from Sarawak, East Malaysia that confirms the presence of the species in Borneo and a previously unrecognised species, Asthenodipsas ingeri sp. nov. from Mount Kinabalu, Sabah. Asthenodipsas ingeri sp. nov. can be differentiated from its congeners by the combination of an absence of preoculars and suboculars, 1st and 3rd pair of infralabials in contact, two pairs of posterior inframaxillaries, supralabials 3-5 (sometimes 3 4) in contact with orbit, 15/15/15 rows of dorsal scales, presence of sharp vertebral keel, and divided subcaudals. This study highlights the importance of careful re-examination of museum collections that could potentially harbour new species hiding in plain sight. These discoveries also add to the growing number of reptile and amphibian species being described from Borneo that shows no signs of abating but are already potentially threatened by the continued deforestation and developments on the island.
A reassessment of the taxonomy of Lygosoma bampfyldei based on morphology and color pattern indicates that it is a species complex containing L. bampfyldei Bartlett, 1895 from the Rajang River, Sarawak and Croker Range, Sabah in East Malaysia; L. peninsulare sp. nov. from Bukit Larut, Perak and 13.5 km east of Jeli, Kelantan, Peninsular Malaysia; L. kinabatanganensis sp. nov. from the Kinabatangan District, Deramakot camp (=Deramakot Sabah Forestry Department), Sabah, East Malaysia; and L. schneideri Werner, 1900 from Djapura, Indragiri, Sumatra, Indonesia-resurrected herein from the synonymy of L. bampfyldei. The new taxonomy aligns itself well with a growing body of literature demonstrating that semi-fossorial and fossorial Sundaic skinks are more diverse than previously considered.
Species in the genus Pseudocalotes are generally rare and known only from a few specimens. Pseudocalotes drogon was described based on a single male from Frasers Hill, Pahang, Peninsular Malaysia. During a recent field survey at Frasers Hill, a female specimen was collected and identified as P. drogon based on morphological characters and a molecular phylogenetic analysis. The morphological description of P. drogon is expanded here, based on the male holotype and the newly collected female specimen.
A new, diminutive species of Rock Gecko Cnemaspis tubaensis sp. nov. of the C. kumpoli group, is described from Tuba Island, Langkawi Archipelago, Kedah, Peninsular Malaysia and is differentiated from all other species in the kumpoli group by having a unique combination of morphological and color pattern characteristics, including a maximum SVL of 37.0 mm; 10 or 11 supralabials; eight or nine infralabials; 15-18 semi-linearly arranged paravertebral tubercles; lateral caudal furrow present; lateral caudal tubercles on the anterior portion of the tail; caudal tubercles not encircling tail; five or six precloacal pores; 28 or 29 subdigital lamellae on the fourth toe; smooth ventrals; smooth subcaudals with an enlarged median row of scales; subcaudal region light-grey and speckled with yellow; absence of light-colored ocelli on the shoulder; no yellow postscapular band; dorsum light-brown with sage-green blotches and black spots; flanks with scattered yellow spots; absence of black gular markings in both sexes; and 13.0-22.0% uncorrected pairwise sequence divergences in the NADH dehydrogenase subunit 2 gene (ND2). Cnemaspis tubaensis sp. nov. is the fourth species of Cnemaspis to be described from the Langkawi Archipelago and underscores the underestimated biodiversity of the islands which is in need of more thorough herpetological inventories.
A new species of Acroeimeria Paperna & Landsberg, 1989 is described from the spotted house gecko, Gekko monarchus (Schlegel) from Peninsular Malaysia. Oöcysts of Acroeimeria grismeri n. sp. are spheroidal to subspheroidal with a smooth bi-layered wall, measure on average 18.4 × 17.3 µm, and have a length/width (L/W) ratio of 1.1; a micropyle and an oöcyst residuum are absent but variable polar granule(s) are present, commonly in Brownian movement. Sporocysts are ellipsoidal and measure on average 8.6 × 6.7 µm, L/W 1.3; Stieda, sub-Stieda and para-Stieda bodies are absent. The sporocyst residuum is composed of numerous spheroidal granules in the center of the sporocyst. This is the initial species of coccidian reported from G. monarchus and one of the few reported from any reptile from Peninsular Malaysia.
The Asiatic coral snakes are basal in the phylogeny of coral snakes. Although envenoming by the Asiatic coral snakes is rarely fatal, little is known about their venom properties and variability from the American coral snakes. Integrating reverse-phase high performance liquid chromatography and nano-liquid chromatography-tandem mass spectrometry, we showed that the venom proteome of the Malaysian banded or striped coral snake (Calliophis intestinalis) was composed of mainly phospholipases A2 (PLA2, 43.4%) and three-finger toxins (3FTx, 20.1%). Within 3FTx, the cytotoxins or cardiotoxins (CTX) dominated while the neurotoxins' content was much lower. Its subproteomic details contrasted with the 3FTx profile of most Micrurus sp., illustrating a unique dichotomy of venom phenotype between the Old and the New World coral snakes. Calliophis intestinalis venom proteome was correlated with measured enzymatic activities, and in vivo it was myotoxic but non-lethal to mice, frogs and geckos at high doses (5-10 μg/g). The venom contains species-specific toxins with distinct sequences and antigenicity, and the antibodies raised against PLA2 and CTX of other elapids showed poor binding toward its venom antigens. The unique venom proteome of C. intestinalis unveiled a repertoire of novel toxins, and the toxicity test supported the need for post-bite monitoring of myotoxic complication. SIGNIFICANCE: Malaysian banded or striped coral snake (Calliophis intestinalis) has a cytotoxin (CTX)-predominating venom proteome, a characteristic shared by its congener, the Malayan blue coral snake (Calliophis bivirgata). With little neurotoxins (NTX), it illustrates a CTX/NTX dichotomy of venom phenotype between the Old World and the New World coral snakes. The low toxicity of the venom imply that C. intestinalis bite envenoming can be managed via symptomatic relief of the mild to moderate pain with appropriate analgesia. Systemically, the serum creatine kinase level of patients should be monitored serially for potential complication of myotoxicity. The distinct antigenicity of the venom proteins implies that the empirical use of heterologous antivenom is mostly inappropriate and not recommended.
The taxonomic position of the rare Selangor Mud Snake (Raclitia indica) Gray to other species of homalopsids has remained uncertain due to the scarcity of specimens in collections and the lack of genetic material for comparison. Here we report the first molecular phylogenetic examination of this species based on recently acquired material. The study recovered R. indica nested within the clade of advanced, fanged homalopsids and the sister species to Erpeton tentaculatus Lácèpede. We also present notes on variation observed in the new specimens as well as range extensions for the species.
A new species of limestone karst-adapted gecko of the Cyrtodactylus pulchellus complex, C. dayangbuntingensis sp. nov., is described from Dayang Bunting Island of the Langkawi Archipelago off the northwest coast of Peninsular Malaysia. It is the third species of the group to be described from the archipelago after C. langkawiensis and C. macrotuberculatus. The 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. This discovery underscores the need for continued surveys of the many islands in the archipelago to properly ascertain its true herpetological diversity.
A new species of Ansonia is described from the Shan Plateau of Myanmar based on an integrative taxonomic analysis that differentiates it from all other congeners. Molecular phylogenetic analyses based on the mitochondrial genes 12S and 16S rRNA and tRNA-val recover A. kyaiktiyoensis sp. nov. as the sister species to A. inthanon from Thailand but differs from it and other congeners by at least a 5.0% sequence divergence. It is further differentiated by the following combination of morphological characters: (1) maximum SVL 24 mm in males and females; (2) first finger shorter than second; (3) absence of interorbital and tarsal ridges; (4) presence of light-coloured interscapular spot; (5) presence of yellow rictal tubercle; (6) absence of wide, light-coloured patch below eye; (7) presence of large, discrete, bright-yellow submandibular spots along the underside of lower jaw; (8) iris yellow-gold; (9) presence of markings on the snout consisting of streaks below the eye to the lip, and on the canthus rostralis to the nostril; (10) dorsum grey-brown with orange-beige spots, a dark-brown X-shaped marking on the back surrounding the interscapular spot, and dark-coloured markings on rump; (11) fore- and hind limbs with orange-beige cross-bars; and (12) venter light-gray with yellow spotting, especially near flanks and underside of hind limbs. Ansonia kyaiktiyoensis sp. nov. is the westernmost known record for the genus and the only species west of the Salween Basin. Its discovery echoes the increasing number of herpetological discoveries being made in upland regions fringing the Ayeyarwady and Salween Basins.
A review of the taxonomic status of the Smooth Slug Snake (Asthenodipsas laevis) in Borneo resulted in the discovery of two previously unrecognised species from the highlands of Sabah, East Malaysia. Asthenodipsas jamilinaisi sp. nov. and A. stuebingi sp. nov. are presumed to be closely related to A. laevis based on similarities in pholidosis and patterning but can be separated from A. laevis by their dorsal scale rows of 15/15/15 vs 15/15/13 and the presence of a sharp vertebral keel. Asthenodipsas jamilinaisi sp. nov. can be further differentiated from A. stuebingi sp. nov. by the greatly enlarged size of the vertebral scales, higher number of ventrals in males (173-175 vs 165), higher number of subcaudals (53 vs 35-47) and colour pattern (dark overall with indistinct bands vs lighter head and body with clear, distinct bands and a dark neck patch). The discovery highlights the need for more careful examination of much of the herpetofauna of Borneo that still remains underestimated and understudied. There is an urgent need for continued surveys into its diversity and the collection of genetic material for integrated taxonomic revisions.
An integrative taxonomic analysis of Subdoluseps herberti from southern Thailand and Peninsular Malaysia and S. samajaya from Sarawak, East Malaysia (Borneo) recovers the former as paraphyletic with respect to the latter. The analyses recover the three southernmost populations of S. herberti in Peninsular Malaysia as conspecific and the sister lineage of S. samajaya, whereas S. herberti from Thailand and northern Peninsular Malaysia constitute the sister species to S. samajaya plus the southern three Peninsular Malaysian populations. As such, the southern populations are described herein as S. malayana sp. nov. and all three species are referred to as the S. herberti group. Clade boundaries and breaks within this group on the Thai-Malay Peninsula occurring at the Isthmus of Kra, across the Kangar-Pattani line, and between the Thai-Malay Peninsula and Borneo are consistent with phylogeographic patterns of other Sundaic taxa. The discovery of S. malayana sp. nov. continues to underscore the fact that, despite the well-studied nature of the lizard fauna of Peninsular Malaysia, much of it still remains unrealized and for conservation efforts to move forward, field research followed by expeditiously revised taxonomies must continue.
A reappraisal of the taxonomic status of the Dark-necked Slug Snake (Asthenodipsas malaccana Peters, 1864) across its range revealed that populations from Borneo are not conspecific with true A. malaccana from the Thai-Malay Peninsula and Sumatra, and is therefore described herein as new. Asthenodipsas borneensis sp. nov. can be distinguished from A. malaccana and other congeners by the absence of a preocular and suboculars, seven or eight supralabials with 3rd and 4th in contact with orbit, 4-7 infralabials with 2nd or 3rd pair in contact, two pairs of posterior inframaxillaries, 15/15/15 rows of dorsal scales, presence of sharp vertebral keel, divided subcaudals, maximum recorded SVL=441 mm, 166-179 ventrals, 35-48 subcaudals, head white to greyish brown and dorsum beige to orange-brown with a conspicuous dark-brown or black patch on the neck followed by multiple, narrow, vertical, dark bands along the rest of the body and tail. This discovery adds to a growing number of new slug snake species recently described from Southeast Asia and highlights the underestimated diversity in this family, especially in Borneo. Taxonomic revisions of the reptiles and amphibians of Borneo are still needed before the true diversity of the island and the relationships of the various taxa can be fully understood.
Accurately delimiting species boundaries is a nontrivial undertaking that can have significant effects on downstream inferences. We compared the efficacy of commonly used species delimitation methods (SDMs) and a population genomics approach based on genomewide single-nucleotide polymorphisms (SNPs) to assess lineage separation in the Malaysian Torrent Frog Complex currently recognized as a single species (Amolops larutensis). First, we used morphological, mitochondrial DNA and genomewide SNPs to identify putative species boundaries by implementing noncoalescent and coalescent-based SDMs (mPTP, iBPP, BFD*). We then tested the validity of putative boundaries by estimating spatiotemporal gene flow (fastsimcoal2, ABBA-BABA) to assess the extent of genetic isolation among putative species. Our results show that the A. larutensis complex runs the gamut of the speciation continuum from highly divergent, genetically isolated lineages (mean Fst = 0.9) to differentiating populations involving recent gene flow (mean Fst = 0.05; Nm > 5). As expected, SDMs were effective at delimiting divergent lineages in the absence of gene flow but overestimated species in the presence of marked population structure and gene flow. However, using a population genomics approach and the concept of species as separately evolving metapopulation lineages as the only necessary property of a species, we were able to objectively elucidate cryptic species boundaries in the presence of past and present gene flow. This study does not discount the utility of SDMs but highlights the danger of violating model assumptions and the importance of carefully considering methods that appropriately fit the diversification history of a particular system.
The gekkonid genus Cyrtodactylus is the third largest vertebrate genus on the planet with well over 300 species that range across at least eight biogeographic regions from South Asia to Melanesia. The ecological and morphological plasticity within the genus, has contributed to its ability to disperse across ephemeral seaways, river systems, basins, land bridges, and mountain ranges-followed by in situ diversification within specific geographic areas. Ancestral ranges were reconstructed on a mitochondrial phylogeny with 346 described and undescribed species from which it was inferred that Cyrtodactylus evolved in a proto-Himalaya region during the early Eocene. From there, it dispersed to what is currently Indoburma and Indochina during the mid-Eocene-the latter becoming the first major center of origin for the remainder of the genus that seeded dispersals to the Indian subcontinent, Papua, and Sundaland. Sundaland became a second major center of radiation during the Oligocene and gave rise to a large number of species that radiated further within Sundaland and dispersed to Wallacea, the Philippines, and back to Indochina. One Papuan lineage dispersed west to recolonize and radiate in Sundaland. Currently, Indochina and Sundaland still harbor the vast majority of species of Cyrtodactylus.
Understanding the processes that underpin adaptive evolutionary shifts within major taxonomic groups has long been a research directive among many evolutionary biologists. Such phenomena are best studied in large monophyletic groups that occupy a broad range of habitats where repeated exposure to novel ecological opportunities has happened independently over time in different lineages. The gekkonid genus Cyrtodactylus is just such a lineage with approximately 300 species that range from South Asia to Melanesia and occupy a vast array of habitats. Ancestral state reconstructions using a stochastic character mapping analysis of nine different habitat preferences were employed across a phylogeny composed of 76% of the known species of Cyrtodactylus. This was done in order to ascertain which habitat preference is the ancestral condition and from that condition, the transition frequency to more derived habitat preferences. The results indicate that a general habitat preference is the ancestral condition for Cyrtodactylus and the frequency of transitioning from a general habitat preference to anything more specialized occurs approximately four times more often than the reverse. Species showing extreme morphological and/or ecological specializations generally do not give rise to species bearing other habitat preferences. The evolution of different habitat preferences is generally restricted to clades that tend to occur in specific geographic regions. The largest radiations in the genus occur in rocky habitats (granite and karst), indicating that the transition from a general habitat preference to a granite or karst-dwelling life style may be ecologically uncomplicated. Two large, unrelated clades of karst-associated species are centered in northern Indochina and the largest clade of granite-associated species occurs on the Thai-Malay Peninsula. Smaller, independent radiations of clades bearing other habitat preferences occur throughout the tree and across the broad distribution of the genus. With the exception of a general habitat preference, the data show that karst-associated species far out-number all others (29.6% vs. 0.4%-10.2%, respectively) and the common reference to karstic regions as "imperiled arcs of biodiversity" is not only misleading but potentially dangerous. Karstic regions are not simply refugia harboring the remnants of local biodiversity but are foci of speciation that continue to generate the most speciose, independent, radiations across the genus. Unfortunately, karstic landscapes are some of the most imperiled and least protected habitats on the planet and these data continue to underscore the urgent need for their conservation.
The taxonomic status of Cyrtodactylus zebraicus in Peninsular Malaysia has been plagued with uncertainty over the last three decades owing to a lack of vouchered material. Recent collections confirmed the presence of this species in the northernmost state of Perlis and for the country. An expanded description of the newly collected Peninsular Malaysian specimens as well as the holotype of C. zebraicus is provided along with a comparison with other Cyrtodactylus species in the country. This study adds to the growing number of herpetofaunal species with Indo-Burmese affinities discovered in northern states of the country and the biogeographic importance of this region for the exchange of fauna and flora. The findings also highlight the importance of continued field work along the biogeographic interchange of the Banjaran Nakawan that separates southwestern Thailand from northwestern Peninsular Malaysia and the need to sample and collect voucher specimens that can be deposited in proper scientific collections for current and future research.
Limnonectes hascheanus and Limnonectes limborgi are two very similar-looking and closely related species and are sometimes referred to as the Limnonectes hascheanus-limborgi complex (Inger & Stuart 2010). Inger & Stuart (2010) tackled the systematics of the complex and confirmed the status of L. limborgi as a distinct species and not a junior synonym to L. hascheanus by providing molecular data and morphological characters of its distinctiveness. The geographic ranges of the two species were also reported to be distinct where L. hascheanus is mainly restricted to the southern part of the Thai-Malay Peninsula while L. limborgi is distributed from southern Myanmar north into northern Thailand and Laos before curving around into central Laos, northeastern Thailand, Cambodia, and southern Vietnam (Inger & Stuart 2010).
We use data sets from the Cyrtodactylus brevipalmatus group with limited genetic and morphological sampling to demonstrate that not accounting for sampling error may adversely influence decisions regarding species delimitation and diagnosis. Lack of geographic sampling between the endpoints of a species range may recover notable interpopulational genetic differentiation consistent with species-level differentiation. Additionally, small population sample sizes may fail recover statistically different diagnostic morphological differences. Combined, these types of sampling error can produce results seemingly consistent with the recognition of cryptic speciesgenetically delimited populations lacking diagnostic morphological characters. This is the current situation within some lineages of the C. brevipalmatus group whereas in others, sampling error is less problematic and does not jeopardize their taxonomy. We note the potential negative effects for comparative biology as a whole if sampling error is not taken into consideration prior to constructing taxonomies.
Liopeltis is a genus of poorly known, infrequently sampled species of colubrid snakes in tropical Asia. We collected a specimen of Liopeltis from Pulau Tioman, Peninsular Malaysia, that superficially resembled L. philippina, a rare species that is endemic to the Palawan Pleistocene Aggregate Island Complex, western Philippines. We analyzed morphological and mitochondrial DNA sequence data from the Pulau Tioman specimen and found distinct differences to L. philippina and all other congeners. On the basis of these corroborated lines of evidence, the Pulau Tioman specimen is described as a new species, L. tiomanica sp. nov. The new species occurs in sympatry with L. tricolor on Pulau Tioman, and our description of L. tiomanica sp. nov. brings the number of endemic amphibians and reptiles on Pulau Tioman to 12.