Cyrtodactylus metropolis sp. nov. from Batu Caves massif, Selangor, Peninsular Malaysia is differentiated from all congeners by having a unique suite of morphological and color pattern characteristics. Remarkably, this species has been overlooked despite a plethora of field studies at Batu Caves from 1898 to the present and no specimens had ever been examined until now. As with all other limestone forest-adapted Cyrtodactylus in Peninsular Malaysia, C. metropolis sp. nov. is not a cave-adapted species but is far more common on the exterior surfaces of the Batu Caves limestone massif and its surrounding limestone vegetation. We suggest that researchers devote time exploring the exterior surfaces of limestone massifs as well the interiors of their caves.
A new species of gekkonid, Hemiphyllodactylus chiangmaiensis sp. nov., from northwestern Thailand is separated from all other species of Hemiphyllodactylus by a set of features including: a maximum SVL of 41.2 mm; 8-12 chin scales extending transversely from unions of second and third infralabials and posterior margin of mental; lamellar formula on hand 3-3-3-3 or 3-4-3-3; lamellar formula on foot 3-3-3-3 or 3-4-4-4; continuous precloacal and femoral pores; a unique dorsal color pattern; and caecum and oviducts pigmented. These characters place this species in the speciose H. typus group. Hemiphyllodactylus chiangmaiensis sp. nov. fills a biogeographical hiatus in the distribution of this genus across northern Indochina.
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.
Sympatric species that initially overlap in resource use are expected to partition the environment in ways that will minimize interspecific competition. This shift in resource use can in turn prompt evolutionary changes in morphology. A classic example of habitat partitioning and morphological differentiation are the Caribbean Anolis lizards. Less well studied, but nevertheless striking analogues to the Anolis are the Southeast Asian Draco lizards. Draco and Anolis have evolved independently of each other for at least 80 million years. Their comparison subsequently offers a special opportunity to examine mechanisms of phenotypic differentiation between two ecologically diverse, but phylogenetically distinct groups. We tested whether Draco shared ecological axes of differentiation with Anolis (e.g., habitat use), whether this differentiation reflected interspecific competition, and to what extent adaptive change in morphology has occurred along these ecological axes. Using existing data on Anolis, we compared the habitat use and morphology of Draco in a field study of allopatric and sympatric species on the Malay Peninsula, Borneo and in the Philippines. Sympatric Draco lizards partitioned the environment along common resource axes to the Anolis lizards, especially in perch use. Furthermore, the morphology of Draco was correlated with perch use in the same way as it was in Anolis: species that used wider perches exhibited longer limb lengths. These results provide an important illustration of how interspecific competition can occur along common ecological axes in different animal groups, and how natural selection along these axes can generate the same type of adaptive change in morphology.
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.
Molecular and morphological analyses indicate that a new upland species of the Cyrtodactylus sworderi complex, C. tebuensis sp. nov. from Gunung Tebu, Terengganu, Malaysia is most closely related to C. sworderi and together they form the sister lineage to C. quadrivirgatus. Cyrtodactylus tebuensis sp. nov. is differentiated from all other species of Sundaland Cyrtodactylus on the basis of having the unique combination of large, conical, keeled body tubercles; tubercles present on top of head, occiput, nape, and limbs, and extending posteriorly beyond base of tail; 43-51 ventral scales; no transversely enlarged, median subcaudal scales; proximal, subdigital lamellae transversely expanded; 17-21 subdigital lamellae on fourth toe; an abrupt transition between posterior and ventral femoral scales; enlarged femoral scales; no femoral or precloacal pores; no precloacal groove; body bearing four wide, bold, dark brown stripes (lateral stripe on each flank and a pair of paravertebral stripes); and a pairwise sequence divergence of 13.0% from its closest relative C. sworderi based on the mitochondrial gene ND2. Cyrtodactylus tebuensis sp. nov. is the first endemic upland species of gekkonid from northeastern Peninsular Malaysia and underscores the necessity for additional field work in all upland systems.
An integrative taxonomic analysis based on additional specimens and color photographs of Lipinia sekayuensis and additional color photographs of L. surda from Pulau Tioman and the Gunung Panti Forest Reserve, Peninsular Malaysia confirm the previous hypotheses that L. sekayuensis is a valid species and is the sister species of L. surda. The two species share a 12.8% sequence divergence between them.
An integrative taxonomic analysis based on the mitochondrial gene ND2 and its flanking tRNAs, morphology, and color pattern indicates that a newly discovered gecko described herein as Hemiphyllodactylus cicak sp. nov. from Penang Hill on the Island of Penang, Peninsular Malaysia is a member of the H. harterti group. Hemiphyllodactylus cicak sp. nov. is most closely related to the clade composed of the sister species H. harterti from Bukit Larut, Perak in the Bintang Mountain Range and H. bintik from Gunung Tebu, Terengganu from the Timur Mountain Range. These three allopatric species form a monophyletic group that extends approximately 270 km across three isolated mountain ranges in northern Peninsular Malaysia. The molecular analysis also indicates that H. titiwangsaensis from the Titiwangsa Mountain Range is composed of three genetically distinct allopatric populations. The southern two populations from Fraser's Hill and Genting Highlands, Pahang have an uncorrected pairwise sequence divergence of 3.5% whereas these two populations have 12.4 and 12.8 % sequence divergences, respectively, from the northern population at Cameron Highlands, Pahang. Although the high sequence divergence clearly distinguishes the southern two populations from the former as a different species, all three populations are morphologically indistinguishable, leading to the hypothesis of a true, cryptic speciation event.
Recently discovered populations of skinks of the genus Sphenomorphus from central Peninsular Malaysia represent a new species, S. sungaicolus sp. nov., and the first riparian skink known from Peninsular Malaysia. Morphological analyses of an earlier specimen reported as S. tersus from the Forest Research Institute of Malaysia (FRIM), Selangor indicate that it too is the new riparian species S. sungaicolus sp. nov. Additionally, two specimens from the Tembat Forest Reserve, Hulu Terengganu, Kelantan and another from Ulu Gombak, Selangor have been diagnosed as new the species. The latter specimen remained unidentified in the Bernice Pauahi Bishop Museum, Honolulu, Hawaii since its collection in June 1962. Morphological and molecular analyses demonstrate that S. sungaicolus sp. nov. forms a clade with the Indochinese species S. maculatus, S. indicus, and S. tersus and is the sister species of the latter. Sphenomorphus sungaicolus sp. nov. can be differentiated from all other members of this clade by having a smaller SVL (66.5-89.6 mm); 39-44 midbody scale rows; 72-81 paravertebral scales; 74-86 ventral scales; a primitive plantar scale arrangement; and 20-22 scale rows around the tail at the position of the 10th subcaudal.
A new species of scincid lizard, Lipinia sekayuensis sp. nov. from Hutan Lipur Sekayu, Terengganu State in northeastern Peninsular Malaysia is most similar to L. surda (Boulenger) but differentiated from it and all other species of Lipinia by having the combination of an adult SVL of 42.3 mm; six supralabials; five infralabials; four supraoculars; prefrontals widely separated; two loreals; fused frontoparietals; lower eyelids bearing a large, transparent disc; 21 midbody scale rows; 56 paravertertebral scale rows; 65 ventral scale rows; enlarged, precloacal scales; 10 subdigital lamellae on the third finger; 11, 15, and seven lamellae on the third, fourth, and fifth toes, respectively; distal subdigital lamellae keeled; a median row of slightly enlarged, subcaudal scales present; a generally unicolor, dark-brown dorsum bearing nine very faint, diffuse, darker stripes; and an external ear opening replaced by a scaly, auditory depression.
A new species of Bent-toed Gecko Cyrtodactylus guakanthanensis sp. nov. of the C. sworderi complex is described from a limestone forest in Perak, Peninsular Malaysia whose karst formations at the type locality are within an active quarry. Cyrtodactylus guakanthanensis sp. nov. can be distinguished from all other Sundaland species by having the following suite of character states: adult SVL 77.7-82.2 mm; moderately sized, conical, weakly keeled, body tubercles; tubercles present on occiput, nape, and limbs, and extend posteriorly beyond base of tail; 37-44 ventral scales; no transversely enlarged, median, subcaudal scales; proximal subdigital lamellae transversely expanded; 19-21 subdigital lamellae on fourth toe; abrupt transition between posterior and ventral femoral scales; enlarged femoral scales; no femoral or precloacal pores; precloacal groove absent; wide, dark postorbital stripes from each eye extending posteriorly to the anterior margin of the shoulder region thence forming a transverse band across the anterior margin of the shoulder region; and body bearing five (rarely four) wide, bold, dark bands. Destruction of the karst microhabitat and surrounding limestone forest will extirpate this new species from the type locality and perhaps drive it to complete extinction given that it appears to be restricted to the particular microhabitat structure of the type locality and is not widely distributed throughout the karst formations. As with plants and invertebrates, limestone forests are proving to be significant areas of high herpetological endemism and should be afforded special conservation status rather than turned into cement.
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 new species of lowland karst dwelling Cnemaspis Strauch 1887, C. grismeri sp. nov. is described from the southeastern base of the Banjaran Bintang in northern Peninsular Malaysia. It is differentiated from its congeners by a unique combination of characters including size, coloration and scalation. Cnemapis grismeri sp. nov. is most closely related to C. mcguirei, an upland species endemic to the Banjaran Bintang. This phylogeographic pattern is also seen in the upland and lowland Banjaran Bintang species of Cyrtodactylus bintangtinggi and C. bintangrendah, respectively (Grismer et al. 2012). The discovery of yet another endemic gekkonid in the poorly explored karst regions of Peninsular Malaysia underscores the necessity for concentrated collecting efforts in these unique landscapes.
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.
We describe two new species of skinks from Gunung Penrissen, Sarawak, Malaysia, in northern Borneo, Tytthoscincus batupanggah sp. nov. and T. leproauricularis sp. nov. Morphological and molecular analyses both corroborate the two new species as unique compared to all other Tytthoscincus and additional Sphenomorphus that are candidates for taxonomic placement in the genus Tytthoscincus. Despite their phenotypic similarity and sympatric distribution, a molecular analysis shows that the new species are not sister taxa and exhibit a deep genetic divergence between each of their respective sister taxa. We discuss how historical climatic and geographic processes may have led to the co-distribution of two relatively distantly related phenotypically similar species. In light of these discoveries, we also emphasize the importance of conserving primary montane tropical rainforest for maintaining species diversity.
A new species of Bent-toed Gecko, Cyrtodactylus gunungsenyumensis sp. nov. of the sworderi complex, is described from Hutan Lipur Gunung Senyum, Pahang, Peninsular Malaysia and is differentiated from all other species in the sworderi complex by having a unique combination of characters including a maximum SVL of 74.7 mm; low, rounded, weakly keeled, body tubercles; 34-40 paravertebral tubercles; weak ventrolateral body fold lacking tubercles; 38-41 ventral scales; an abrupt transition between the posterior and ventral femoral scales; 20-23 subdigital lamellae on the fourth toe; enlarged femoral scales; no femoral or precloacal pores; no precloacal groove; wide caudal bands; and an evenly banded dorsal pattern. Cyrtodactylus gunungsenyumensis sp. nov. is a scansorial, karst forest-adapted specialist endemic to the karst ecosystem surrounding Gunung Senyum and occurs on the vertical walls of the limestone towers as well as the branches, trunks, and leaves of the vegetation in the associated karst forest. Cyrtodactylus gunungsenyumensis sp. nov. is the seventh species of karst forest-adapted Cyrtodactylus and the sixteenth endemic species of karst ecosystem reptile discovered in Peninsular Malaysia in the last seven years from only 12 different karst forests. This is a clear indication that many species remain to be discovered in the approximately 558 isolated karst ecosystems in Peninsular Malaysia not yet surveyed. These data continue to underscore the importance of karst ecosystems as reservoirs of biodiversity and microendemism and that they constitute an important component of Peninsular Malaysia's natural heritage and should be protected from the quarrying interests of foreign industrial companies.
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.