Amphibians of Sekayu lowland forest have been studied more than a decade, with discoveries of new records of species showing no sign of abating between the years 2003 to 2020, indicating the remarkably rich diversity of anurans in this forest. Despite ceaseless anthropogenic activities in this area, this study successfully recorded 52 species of amphibians from 32 genera in the lowland forest of Sekayu. The species composition consisted of a single species from the family Ichthyophiidae and 51 species of anurans of 31 genera and six families. The number of species recorded has steadily increased especially during more recent surveys from 2015 to 2020. This study augments the total number of amphibian species recorded from Hulu Terengganu by ten additional species, increasing the total to 70 species for the district.
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.
Two new species of diminutive, sympatric, lowland, leaf-litter skinks of the genus Tytthoscincus Linkem, Diesmos Brown from the Sekayu region of Hulu Terengganu, Terengganu State in northeastern Peninsular Malaysia are described on the basis genetic and morphological data. One of the new species, T. monticolus sp. nov., was collected in a hilly riparian area along Sungai (=river) Bubu and is most closely related to an undescribed species from the Tembak Reservoir area. The other, T. keciktuek sp. nov. collected along Sungai Peres, is most closely related to T. perhentianensis Grismer, Wood, Grismer from Pulau (=island) Perhentian Besar. Sympatry and syntopy of multiple, specialized, unrelated, leaf-litter species of Tytthoscincus was previously only known from upland areas and these new species represent the first example of lowland of sympatry. More importantly, however, these endemic species add to a growing body of research and discoveries that continue to underscore the unrealized biodiversity of the riparian systems of Hulu Terengganu and the Sekayu region and their need for protection and continued study.
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.
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.
Molecular phylogenetic analyses of the sister species Sphenomorphus stellatus and S. praesignis based on the mitochondrial genes 12S and 16S rRNA recover the former as paraphyletic with respect to the latter in that a specimen of S. stellatus from the type locality in Peninsular Malaysia is more closely related to S. praesignis than to Indochinese populations of S. stellatus. Furthermore, the phylogeny indicates that the Indochinese populations represent two species, thus resulting in four major lineages within this clade. These relationships are consistent with multivariate and univariate analyses of morphological and discrete color pattern data which statistically define and diagnose the four lineages and together with the molecular data, provide the foundation for robust, testable, species-level hypotheses. As such, S. stellatus is herein restricted to Peninsular Malaysia; S. annamiticus is resurrected for the circum-continental populations ranging through southeastern Thailand, southern Cambodia, and southern Vietnam; a new species-S. preylangensis sp. nov.-is described from an isolated mountain, Phnom Chi, from the Prey Lang Wildlife Sanctuary in central Cambodia; and the taxonomy of S. praesignis remains unchanged. The description of S. preylangensis sp. nov. underscores the necessity to conserve this remnant of lowland evergreen rainforest in the Prey Lang Wildlife Sanctuary.
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.
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 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.
A phylogenetic taxonomy of species in the Cyrtodactylus peguensis group from the Ayeyarwady Basin of Myanmar is constructed based on color pattern, morphology, and molecular systematic analyses using the mitochondrial gene NADH dehydrogenase subunit 2. Newly collected samples from the type locality of C. peguensis and other localities indicate that this clade is endemic to central Myanmar and contains at least seven species, four of which are undescribed. Three species, including C. peguensis occur in the low hills of the Bago Yoma Range within the central portion of the Ayeyarwady Basin. Two of these, C. myintkyawthurai sp. nov. from the northern and central Bago Yoma and C. meersi sp. nov. which is syntopic with C. peguensis in the southern Bago Yoma are described herein. As more lowland hilly areas bordering, and within the Ayeyarwady Basin are surveyed, more new species of this group are likely to be discovered. These discoveries continue the recent surge of descriptions of new species of Cyrtodactylus that are being discovered in Myanmar.
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.
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.
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.
Systematic assessments of species extinction risk at regular intervals are necessary for informing conservation action1,2. Ongoing developments in taxonomy, threatening processes and research further underscore the need for reassessment3,4. Here we report the findings of the second Global Amphibian Assessment, evaluating 8,011 species for the International Union for Conservation of Nature Red List of Threatened Species. We find that amphibians are the most threatened vertebrate class (40.7% of species are globally threatened). The updated Red List Index shows that the status of amphibians is deteriorating globally, particularly for salamanders and in the Neotropics. Disease and habitat loss drove 91% of status deteriorations between 1980 and 2004. Ongoing and projected climate change effects are now of increasing concern, driving 39% of status deteriorations since 2004, followed by habitat loss (37%). Although signs of species recoveries incentivize immediate conservation action, scaled-up investment is urgently needed to reverse the current trends.
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 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 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.
The family Xenophidiidae is an extremely rare and little-known family of snakes that contains only two species, Xenophidion acanthognathus Günther Manthey and X. schaeferi Günther Manthey that are each known only from their holotypes. We report on the rediscovery of X. schaeferi from two new locations in southern Peninsular Malaysia. The new specimens differ markedly from the holotype of X. schaeferi in colour pattern and are more akin to the patterning of X. acanthognathus from Borneo. However, molecular analyses of one of the new specimens reveal that it only differs from the holotype of X. schaeferi by a minimum sequence divergence of 0.27%. An expanded description of the species based on these specimens is provided and phenotypic variation within the species is discussed. We also report the first record of the genus Xenophidion from West Sumatra, Indonesia. These sensational discoveries continue to underscore the fact that southern Peninsular Malaysia and upland areas of Southeast Asia in general still harbour unrealized herpetological diversity and are still in need of field research-based taxonomic studies.