A new species of Himalayan skink of the genus Ablepharus (Squamata: Scincidae)

373 A new species of Himalayan skink of the genus Ablepharus © 2025 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Germany Open access at https://www.salamandra-journal.com 30 October 2025 ISSN 0036–3375 SALAMANDRA 61(4): 373–394 SALAMANDRA German Journal of Herpetology A new species of Himalayan skink of the genus Ablepharus (Squamata: Scincidae) Andrey M. Bragin1,2, Ht. Decemson2, Hmar Tlawmte Lalremsanga2, Zeeshan A. Mirza3 & Nikolay A. Poyarkov4 1 Joint Russian-Vietnamese Tropical Research and Technological Center, Nghia Do, Cau Giay, Hanoi, Vietnam 2 Developmental Biology and Herpetology Laboratory, Department of Zoology, Mizoram University, Aizawl, Mizoram, 796004, India 3 Max Planck Institute for Biology, Max-Planck-Ring 1, 72076 Tübingen, Germany 4 Department of Vertebrate Zoology, Biological Faculty, M. V. Lomonosov Moscow State University, Moscow 119234, Russia Corresponding author: Andrey M. Bragin, ORCID 0000-0002-3621-9763, e-mail: b[email protected] Manuscript received: 11 April 2025 Accepted: 18 August 2025 by Andreas Schmitz Abstract. Based on morphological and genetic evidence, we evaluated the taxonomic status of a deeply divergent highland population of Himalayan ablepharine skinks (genus Ablepharus Fitzinger in Lichtenstein, 1823) from Uttarakhand state in northern India. This lineage, here described as a new species, forms a well-supported clade of Ablepharus and differs from the morphologically similar species by a significant divergence in the 16S rRNA and cytochrome b mitochondrial DNA gene sequences. From the phylogenetic analysis of a 2,961 bp concatenated alignment of the ND2, cyt b, 16S, and 12S rRNA mitochondrial DNA genes and diagnostic morphological characters, we allocate the newly discovered population from Duggal Bittha village, Chopta region to the A. tragbulensis-A. ladacensis species complex (Clade 1 according to Bragin et al. 2024) and describe it as Ablepharus flammeus sp. n. Our discovery brings the number of Ablepharus species in India to seven and in the Himalayan region to eleven and emphasizes the incompleteness of knowledge on the herpetofaunal diversity of this highland region. In this study, we also compare known morphological data for ablepharine skinks from the Himalayan region and discuss the hidden cryptic diversity within this group of skinks. Key words. Sauria, Ablepharus, biodiversity, India, morphology, phylogenetics, taxonomy. Introduction The Karakorum, Himalaya, and Tibet mountain systems are inhabited by at least eight species of the genus Ablepharus: A. himalayanus Günther, 1864, A. sikimmensis Blyth, 1854, A. ladacensis Günther, 1864, A. tragbulensis Alcock, 1898, A. capitaneus Ouboter, 1986 (according to Eremchenko et al. 1998, Bragin et al. 2024), A. mahabharatus Eremchenko, Shah & Panfilov, 1998, A. nepalensis Eremchenko & Helfenberger, 1998 (Uetz et al. 2024). Three of the aforementioned species have wide ranges spanning vast mountainous areas in the east (A.himalayanus and A. ladacensis) and west (A. sikimmensis) of the region. As is often the case with small members of the family Scincidae, ablepharine skinks have similar external morphologies (Chapple et al. 2023). However, from the outset of research on this group (see Bragin et al. 2024), it was evident that the focal taxa are comprised of cryptic species. The taxonomic history of the species A. ladacensis and A. himalayanus is especially interesting. Günther (1864) described these morphologically distinct species as two independent species within the genus Eumeces Wiegmann, 1834. However, for unclear reasons, Ouboter (1986) considered them as part of a single species, Scincella ladacensis, and he relegated the latter to a subspecific status: Scincella ladacensis ladacensis and S. ladacensis himalayana. Ouboter (1986) further noted greater variability of morphological characters within the group. This later led to the reinstatement of the species status of Asymblepharus himalayanus (Eremchenko et al. 1998) and the description of two subspecies within A. ladacensis (Eremchenko 1992): Asymblepharus ladacensis ladacensis Eremchenko, 1992 from northwestern India and Asymblepharus ladacensis stimsoni Eremchenko, 1992 from Nepal. However, even in the latest, most complete morphological revision of the ablepharine skinks of Nepal (Eremchenko in Schleich & Kästle 2002), Eremchenko gives the following diag- 374 Andrey M. Bragin et al. nosis for the A. himalayanus species: “Very variable–from small, slim lizards, for example a population of Simla (NW India, Himachal Pradesh State), some populations in Kashmir; to big wide-headed and long-tailed lizards from Gulmarg (Gulmarg, 2,704 m), Lidder (Lidar) Valley and Marri, all in Kashmir”, with a wide range of not only metric but also meristic characters. Such a range of characters, wider than the morphological differences between recognized species, strongly hints at several hidden cryptic taxa within Ablepharus group. There have been no attempts to reconstruct the phylogeny of ablepharine skinks using molecular methods for a long time. However, the latest research by Bragin et al. (2024) proposes a molecular hypothesis of the relationships within ablepharine skinks based on a comprehensive sampling of taxa (encompassing 18 out of the 23 recognized ablepharine skink species), reconstructing four major clades inhabiting the Karakorum, Himalaya, and Tibet mountain systems, of which three belong to the genus Ablepharus, and the other one includes all known species of the genus Protoblepharus. Moreover, the analysis not only confirmed currently recognized species, but also reconstructed eight to ten additional, potentially new species-level lineages inside the Ablepharus group. These were found mainly within the broadly distributed and morphologically variable species A. himalayanus and A. ladacensis, which constitute cryptic species complexes. Thus, the obtained results of previous studies on this skink group show the underestimation of the taxonomic diversity of ablepharine skinks and help to outline a work plan for the future. During our recent field surveys in Uttarakhand state of India, we encountered an unusual population of ablepharine skinks superficially similar to A. himalayanus in overall morphological habitus and body coloration. Closer morphological examination of scalation, patterns of coloration, and morphometric characteristics of this population demonstrated clear morphological differences from other representatives of Ablepharus inhabiting the Himalayas. In the present paper, we demonstrate that these differences are concordant with a significant divergence between mtDNA gene sequences among this population. Subsequent phylogenetic analyses of four mtDNA genes (ND2, cyt b, 16S, and 12S rRNA) confirmed the placement of the population from the highlands of Chopta region within A.tragbulensis-A. ladacensis species complex (Clade 1 according to Bragin et al. 2024), where this population forms a deeply divergent lineage. Therefore, in the present paper, we describe the Ablepharus population from the highlands of Uttarakhand, India as a new species and discuss the taxonomy of Himalayan ablepharine skinks. Materials and methods Collection We used tissues from the herpetological collections of the Developmental Biology & Herpetology (DB&H) Laboratory of Mizoram University (MZMU, Aizawl, India) (summarized in Supplementary Table S1). For alcohol-preserved voucher specimens stored in museum collections, we removed a small sub-sample of muscle or liver, preserved it in 96% ethanol, and stored samples at -70 °C. We analyzed four tissue samples representing two newly discovered populations of Ablepharus inhabiting Uttarakhand state in northern India. In our analysis, we also included 281 GenBank sequences from ninety specimens of fifteen nominal Ablepharus species, six specimens of three nominal Protoblepharus species, and sixteen species of nonablepharine outgroups used for rooting the phylogenetic tree (Supplementary Table S1). The geographic location of sampled Ablepharus populations is presented in Figure 1. The collected specimens were subsequently deposited in the following herpetological collections: holotype deposited in the herpetological collection of the Zoological Survey of India, Kolkata (ZSI), and paratypes deposited in the herpetological collection of the Developmental Biology & Herpetology (DB&H) Laboratory of Mizoram University (MZMU, Aizawl, India). For all aspects of species concepts and delimitation, we follow the General Lineage Concept (sensu De Queiroz 2007), where a species represents a single evolutionarily independent lineage following a separate trajectory compared to its relatives. We specifically delimit evolutionary independence when lineages are reciprocally monophyletic, exhibit substantial genetic divergence, and are morphologically diagnosable. Laboratory procedures For molecular phylogenetic analyses, total genomic DNA was extracted from ethanol-preserved liver tissue using standard phenol-chloroform-proteinase K (final concentration 1 mg/ml) extraction procedures with consequent isopropanol precipitation (protocols followed by Russell & Sambrook 2001). For mtDNA, we amplified sequences covering the ND2 (mitochondrially encoded NADH dehydrogenase 2), cyt b (mitochondrially encoded cytochromeb), 16S rRNA, and 12S rRNA mtDNA genes to obtain a 2961 bp-length of continuous fragments of mtDNA. These fragments have also proven to be particularly useful in taxonomic studies of the ablepharine skinks and closely related taxa (Macey et al. 2006, Hedges 2014, Ankita et al. 2017, Che et al. 2020, Weng et al. 2021, Mirza et al. 2022, Bragin et al. 2024). In total, we amplified a 1,600bp-long fragment of the ND2 gene, 406 bp-long fragment of the cyt b gene, 550 bp-long fragment of the 16S rRNA gene, and 405 bp-long fragment of the 12S rRNA gene. Primers used in PCRs and sequencing were taken from the literature and are summarized in Supplementary Table S2. For primer pairs of the ND2 gene, we used the following PCR protocol: (1) initial denaturation step at 95 °C for 3 min; (2) 36 cycles of denaturation at 95 °C for 30 sec, annealing at 47 °C for 45 sec and extension at 72 °C for 1min; (3) final extension at 72 °C for 10 min; and (4) cool- 375 A new species of Himalayan skink of the genus Ablepharus ing step at 4 °C for storage. For primer pairs of the 16S rRNA gene: (1) initial denaturation step at 94 °C for 5 min; (2) 40 cycles of denaturation at 95 °C for 1 min, annealing at 47 °C for 1min and extension at 72 °C for 1 min; (3) final extension at 72 °C for 10 min; and (4) cooling step at 4 °C for storage. For primer pairs of the 12S rRNA gene: (1)initial denaturation step at 95 °C for 3 min; (2) 36 cycles of denaturation at 95 °C for 30 sec, annealing at 47 °C for 45sec and extension at 72 °C for 1 min; (3) final extension at 72°C for 10 min; and (4) cooling step at 4 °C for storage. For cyt b sequences we used a modified PCR protocol with a touchdown: (1) initial denaturation step at 95 °C for 5 min; (2)10 cycles of denaturation at 95 °C for 1 min, annealing for 1min with temperature decreasing from 50°C to 45°C (with cool-down at 0.5 °C per each cycle) and extension at 72 °C for 1 min; (3) 30 cycles of denaturation at 95 °C for 1min, annealing at 47 °C for 1 min and extension at 72 °C for 1 min; (4) final extension at 72 °C for 10min; and (5) cooling step at 4 °C for storage. PCRs were run on a Bio-Rad T100TM Thermal Cycler; sequence data collection and visualization were performed on an ABI 3730xl automated sequencer (Applied Biosystems, Foster City, CA, USA). PCR purification and cycle sequencing were done commercially through Evrogen (Moscow, Russia). All unique sequences were deposited in GenBank (see Supplementary Table S1). Genetic data analysis In addition to newly obtained sequences, we also used 281 DNA sequences of Scincidae from GenBank in our final alignments; sequences of Scincella cf. potanini, S. huanrenensis, S. modesta, S. vandenburghi, S. doriae, S. rupicola, S. melanosticta, S. reevesii, Sphenomorphus incognitus, S. indicus, Isopachys gyldenstolpei, Tropidophorus hangnam, Plestiodon chinensis, P. liui, P. elegans, and Ateuchosaurus chinensis were selected as outgroup taxa to help root Figure 1. Map showing the distribution of the studied Ablepharus populations’ locations. The shapes and colours of the figures correspond to those in the names of the samples in Figure 2. Conventions: a figure with a dot indicates the type localities of the corresponding species: pink circle – Ablepharus flammeus sp. n.; green hexagon – A. cf. tragbulensis, green square – A. ladacensis sp.5; green triangle – A. ladacensis sp. 1; green diamond – A. ladacensis sp. 2; green circle – Ablepharus sp. 1; green inverted triangle – A.ladacensis sp. 3; green star – A. ladacensis sp. 4; red square – A. himalayanus sp. 1; red triangle – A. himalayanus sp. 4; red circle – A. himalayanus sp. 3; red diamond – A. nepalensis; yellow square – Ablepharus sikimmensis; yellow circle – A. mahabharatus; blue circle – Protoblepharus sp.; blue triangle – P. apatani ; blue square – P. medogensis; blue diamond – P. nyingchiensis. Localities: 1– Pakistan, Kumrat; 2 – India and Pakistan, Kashmir, Tragbul Pass, about 50 km NW from Srinagar; 3 – India, Jammu and Kashmir, Gulmarg; 4 – India, Jammu and Kashmir, Sonamarg; 5 – India, Ladakh; 6 – India, Kashmir; 7 – India, Himachal Pradesh, Shimla; 8 – India, Uttarakhand, Garhval; 9 – India, Himachal Pradesh, Manali; 10 – India, Himachal Pradesh, Naggar; 11 – India, Himachal Pradesh, Ghiagi; 12 – India, Himachal Pradesh, Batsari; 13 – India, Himachal Pradesh, Shimla; 14 – India, Himachal Pradesh, Chail; 15 – India, Uttarakhand, Rajaji National Park; 16 – India, Uttarakhand, Ful Chatn; 17 – India, Himachal Pradesh, Naragan; 18 – India, Uttarakhand, Ful Chatn; 19 – India, Uttarakhand, Nanketal; 20 – India, Uttarakhand, Seven Lakes; 21 – India, Uttarakhand, Dhorali; 22 – India, Himachal Pradesh, Chamba; 23 – India, Uttarakhand, Dehradun; 24 – India, Uttarakhand, Farsula; 25 – Nepal, Western region, Piple; 26 – India, Uttarakhand, Chopta; 27 – India, Himachal Pradesh, Dkhankhar; 28 – India, Himachal Pradesh, Dkhankhar; 29 – India, Himachal Pradesh, Chitkul; 30 – India, Uttarakhand, Seven Lakes; 31 – India, Uttarakhand, Dhorali; 32 – China, Tibet, Baidengpo; 33 – China, Tibet, Xizang; 34 – Nepal, Western region, ChomrungKomrong Danda; 35 – Nepal, Western region, Damphus; 36 – Nepal, Central region, Gadawari; 37 – India, probably Sikkim; 38 – India, Sikkim; 39 – Nepal, Western region, Suikhet-Naudanda; 40 – Nepal, Western region, Pokhara, Phedi; 41 – Nepal, Central region, northern slopes of Mahabharat mountain range, along Bhainse-Pass Daman; 42 – Nepal, Central region, Gadawari; 43 – China, Tibet, Xigaze, Zhangmu (Khasa); 44 – India, probably Sikkim; 45 – India, Arunachal Pradesh, Talle; 46 – China, Tibet, Nyingchi; 47 – China, Tibet, Medog. 376 Andrey M. Bragin et al. our tree. Details on taxonomy, localities, GenBank accession numbers, and associated references for all examined specimens are summarized in Supplementary Table S1. Nucleotide sequences were initially aligned in MAFFT v6 (Katoh et al. 2002) with default parameters, subsequently checked by eye in BioEdit v7.2.5 (Hall 1999), and adjusted as needed. We reconstructed phylogenetic trees using a mtDNA concatenated dataset of ND2–cyt b–16S rRNA–12S rRNA fragments, used to assess species groups and estimate cryptic diversity within ablepharine skinks. The optimum partitioning schemes for alignments were identified with PartitionFinder 2.1.1 (Lanfear et al. 2012) using the greedy search algorithm under an AIC criterion. Phylogenies were hypothesized via maximum likelihood (ML) and Bayesian Inference (BI). We used IQTREE (Nguyen et al. 2015) to reconstruct ML phylogenies. Confidence in tree topology for ML analysis was assessed by 1,000 bootstrap replications for ML analysis (ML BS). Bayesian inference (BI) was performed in MrBayes v3.1.2 (Ronquist & Huelsenbeck 2003). Metropolis-coupled Markov chain Monte Carlo (MCMCMC) analyses were performed and run with one cold chain and three heated chains for one million generations and sampled every 1000 generations. Five independent MCMCMC run iterations were performed and 1,000 trees were discarded as burn-in. Nodal support was assessed by calculating posterior probabilities (BI PP). The best-fitting model for both BI and ML analyses of the ND2, 12S rRNA, and 16S rRNA gene suggested by the AIC was the GTR+I+G model. For cyt b gene, following the AIC, the GTR+I+G model was used for the first codon partition, the GTR+G for the second codon partition, and the SYM+I+G for the third codon partition. We a priori regarded tree nodes with BI PP values over 0.95 and ML BS values 75% or greater as sufficiently resolved, while BI PP values between 0.95 and 0.90 and ML BS values between 75% and 50% were regarded as tendencies. Lower values were regarded as indicating unresolved nodes (Huelsenbeck & Hillis 1993). Species delimitation Based on the Assemble Species by Automatic Partitioning (ASAP; Puillandre et al. 2021) species delimitation method, we examined putative species boundaries beyond those currently recognized by taxonomists. Barcode gaps, presumably occurring between intraand interspecific distances, were separately used to partition the 16S rRNA and cyt b datasets into species hypotheses (initial partition). Resulting inferences were then recursively applied to yield finer partitions (recursive partitions) until no further partitioning was possible. ASAP analysis was run on the 16S rRNA and cyt b datasets through a web-based interface (https://bioinfo.mnhn.fr/abi/public/asap/#) using default parameters (Simple Distance (p-distances)). In addition, both interand intraspecific uncorrected genetic p-distances were calculated using MEGA X for the 16S rRNA and cyt b datasets (Kumar et al. 2018). Morphological data analysis Measurements were acquired using digital calipers XPERT Xp150 to the nearest 0.1 mm. Measurements of type series were modified from Ouboter (1986), Das et al. (1998) and Grismer et al. (2019), including snout–vent length (SVL), from the tip of the rostral scale to the vent; tail length (TaL), from the vent to the tip of the tail; tail width (TaW), measured at the base of tail; axilla-groin length(AGD), measured from the posterior margin of the forelimb to the anterior margin of the hind limb; head length (HL1), from the tip of the snout to the angle of jaws from the lateral view; head length (HL2), from the tip of the snout to the posterior parietal from the dorsal view; interorbital distance (IOD), the distance between anterior edges of the orbits from the dorsal view; snout–ear length (SEL), from the tip of the snout to the anterior margin of the ear opening; head width (HW), the widest point of the head posterior to the eyes; head height (HH), the highest part of the head posterior to the eyes; body width (MBW), the widest point of the body closer to the hind limbs; internarial distance (IN), the distance between inner edges of nares from the dorsal view; eyeball diameter (ED), the anterior to the posterior margins of the eyeball; tympanum diameter (TD), measured as the longest length across the ear opening; rostral width (RW), the greatest distance between the lateral edges of the rostral scale; rostral height (RH), the greatest distance between the anterior and posterior edges of the rostral scale; snout length (SNL), from the tip of the snout to the anterior border of the orbit; eye– tympanum length (EED), from the posterior margin of the orbit to the anterior margin of the ear opening; eye–nostril length (END), from the margin anterior of the orbit to the posterior margin of the nares; snout–forelimb length (SNL), from the tip of the snout to the anterior margin of the forelimb insertion on the body; forelimb length (FLL), from the insertion point of the forelimb on the body to the tip of the fourth finger on an outstretched limb; hindlimb length (HLL), from the insertion point of the hindlimb on the body to the tip of the fourth toe on an outstretched limb; tibia length (TBL), measured along flexed tibia from the knee bend to the base of the pes; and length of the fourth toe of the hind limb (T4L). We also provide the following ratios: HL2/SVL; HW/SVL; HD/HL2; SNL/HW; ED/SNL; ED/HL2; SVL/BW; EED/END; SVL/HW; SVL/ HL2; SVL/FLL; SVL/HLL so as to compare our data with data in Ouboter (1986) and Das et al. (1998). The metric characteristics of juvenile specimens were not taken into account. Scale counts were made on the right or each side of the body under a Nikon SMZ460 stereo zoom microscope. Scale counts taken were the number of supraoculars contacted by the frontoparietal (F-SO); the degree of contact of prefrontals with each other, and with the frontal (PF); the number of nuchal scales on each side of the head (Nu) counted the number of enlarged scales (one and a half times larger than the dorsal scales), located immediately behind the parietals and interparietal and running 377 A new species of Himalayan skink of the genus Ablepharus along the dorsal surface of the neck; number of primary, secondary, and tertiary temporals (PT, ST, and TT, respectively); the character of overlapping of the temporal and parietal scales; number of supraciliary scales (SC), supraoculars (SO), loreals (AL, PL), preoculars (PR), presuboculars (PRS), postoculars (PO), postsuboculars (PSO), supralabials (SL) and infralabials (IF) on each side of the head; number of projecting ear lobules (PEL) counted as a number of large granules noticeably protruding above the edge of the ear opening; character of the scalation of the lower eyelid and transparent window; number of midbody scale rows (MBSR) counted as the number of longitudinal scale rows encircling the body at a point midway between the limb insertions; paravertebral scale rows (PVSR) counted as the number of scales in a line from parietals to a point on the dorsum opposite the vent; dorsal scales rows (DBR), counted between the dorsolateral dark stripes; ventral scale rows (VSR) counted as a row of scales between the postmental and the precloacal scales; number of enlarged precloacal scales (PrC); fourth toe and finger subdigital lamellas (TL4 and FL4, respectively); the degree of contact between adpressed forelimb and hindlimb (FH) and adpressed hind limbs and elbows (F-E); size and arrangement of rows of scales on the inner thighs (FemS); size (enlarged or not relative to adjacent scales) of subcaudal scales. The hemipenes of the holotype are described according to Vergilov et al. (2017) and Vergilov & Zlatkov (2024). Color pattern characters included the presence or absence of dark vertebral, dorsolateral, and lateral stripes (continuous or breaking into separate spots forming a dotted line); the presence or absence of white lateral stripes bordering the lower edge of the dark lateral stripe (or their breaking up into separate spots forming a dotted line); general background coloration of the dorsum and venter, and venter of males (or/and females) during the mating season; the presence or absence of a white stripe under the eye; coloration of the crown of the head; presence or absence of subcaudal banding; presence or absence of spots on the dorsal and along lateral sides of the body. Nomenclatural acts The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The LSID (Life Science Identifier) for this publication is: urn:lsid:zoobank. org:pub:145737E5-5D28-4AFF-A923-A3565253B3AD. The electronic edition of this work was published in a journal with an ISSN and has been archived and is available from the following digital repositories: www.zenodo.org and www.salamandra-journal.com. Results Sequence characteristics Our aligned matrix of all mtDNA data comprised 2.961bp (assembled from 1.600 bp fragment from ND2, 406 bp fragment from cyt b, 550 bp fragment from 16S rRNA, and 405 bp fragment from 12S rRNA). It included 92 samples, representing 17 species of Ablepharus (75% of the currently recognized species, including 86% of the species that inhabit the Himalayas and two newly described species), three species (of three currently recognized species) of the phylogenetically closely related genus Protoblepharus, and 16 samples from outgroup taxa (see Table S1). Information on fragment lengths and variability is summarized in Table S3. PartitionFinder 2.1.1 proposed the partition schemes and substitution models, summarized in Supplementary Table S3. We deposited the newly obtained sequences in GenBank under the accession numbers PV097228–PV097229 (cyt b), PV097230–PV097231 (ND2), PV105692–PV105692 (16S), and PV105694–PV105695 (12S) (see Table S1). Phylogenetic relationships BI and ML phylogenetic analyses resulted in almost identical topologies (Fig. 2). The topology of the matrilineal genealogy was largely consistent with the phylogeny of ablepharine skinks presented by Bragin et al. (2024) and recovered five main clades grouped into two major reciprocally monophyletic groups, corresponding to the genera Ablepharus and Protoblepharus (Fig. 2). Monophyly of the genus Ablepharus was poorly supported in BI analysis, but had significant support in ML analysis (0.6/97, hereafter node values given for BI PP/ML UFBS, respectively; Fig.2) as well as monophyly of the genus Protoblepharus (-/99); Ablepharus group included four strongly supported clades, three of which included species inhabiting the mountain systems of Tibet, the Himalayas, the Hindu Kush and the Karakoram (1) Clade 1 (0.9/82) included two subclades mainly corresponding to two nominal species of Ablepharus inhabiting the Western Himalaya region and the Karakoram Mountain system: A. tragbulensis and A. ladacensis. The first subclade (0.91/81) included three relatively deeply divergent mitochondrial lineages with all nodes strongly supported (1/100): two specimens of A. tragbulensis from Karakoram, Pakistan; four specimens of Ablepharus sp. 1 from the northwestern part of Uttarakhand, India; and, including two specimens of Ablepharus sp. n. from newly discovered population from Chopta in the central part of Uttarakhand, India. The second subclade (1/99) included five mitochondrial lineages, corresponding to A. ladacensis species complex. (2) Clade 2 (0.73/86) also included two subclades mainly corresponding to two nominal species of Ablepharus inhabiting the Western and Central Himalaya region: A.nepa lensis and A. himalayanus. The first subclade (1/100) included specimens of A. nepalensis from a type locality in 378 Andrey M. Bragin et al. Central Nepal, and the second subclade (0.51/95) included five mostly strongly supported mitochondrial lineages, corresponding to A. himalayanus species complex. (3) Clade 3 (0.99/98) joined two species of Ablepharus from Central and Eastern Himalayas regions: A. mahabharatus and A. sikimmensis. The population of Ablepharus Figure 2. Molecular phylogeny plotted on the Maximum-Likelihood consensus tree based on mtDNA sequences (ND2, cyt b, 16S rRNA and 12S rRNA genes) of Himalayan snake-eyed skinks and congeners. Support values on branches are posterior probabilities (PP) resulting from a separate partitioned Bayesian analysis followed by ML bootstraps. Colour bars on the right of each clade correspond to the results of species delimitation analyses based on ASAP. The numbers in brackets correspond to the sample data/locality number in Table 1 and on the map in Figure 1. The figures in sample names and colours of the clade’s selection correspond to those on the map in Figure 1. Inset photograph of Ablepharus flammeus (Uttarakhand, Chopta) by Andrey M. Bragin. 379 A new species of Himalayan skink of the genus Ablepharus sp. n. under study from Chopta, Uttarakhand, India, belongs to clade 1 (Fig. 2), where it is reconstructed as a member of a subclade also including A. tragbulensis from Pakistan and A. sp. 1 from northwestern Uttarakhand, India. However, the monophyly of this subclade appears to be insufficiently resolved and had strong support in BI-analysis but was poorly supported in ML-analysis (0.91/81). Three included lineages all had strong node support (1/100). Genetic distances The uncorrected p-distances for the 16S rRNA and cyt b gene fragments among examined members of the genus Ablepharus are presented in Table S4. For the 16S gene, interspecific distances among the members of the genus Ablepharus varied from p = 1.8% (between A. alaicus and A.eremchenkoi) to p = 13.2% (between A. ladacensis sp. 4 and A. anatolicus); for the cyt b gene, interspecific distances varied from p = 7.5% (between A. alaicus and A. eremchenkoi) to p = 24.7% (between A. deserti and A himalayanus sp. 2). The newly discovered Ablepharus sp. n. lineage from Chopta is highly divergent from other congeners with the minimal genetic distance of p = 5.5% (with A. sikimmensis sp. 2) and the maximal distance of p = 11.6% (with A.budaki) of sequence divergence in 16S rRNA mtDNA gene; and with the minimal genetic distance of p = 14.6% (with A.ladacensis sp. 3) and the maximal distance of p= 22.0% (with A. budaki) of sequence divergence in cyt b mtDNA gene. To assess the number of putative species-level lineages within the ablepharine skinks, we implemented the distance-based ASAP species delimitation analysis for the 16S rRNA and cyt b gene fragments. The result of ASAP analysis did not differ for the two mtDNA genes, resolved all the currently recognized species of the ablepharine skinks of the Himalayas region, and agreed well with recent ASAP results for this group (Bragin et al. 2024). ASAP analysis clearly recovered recently identified species-level lineage from the Chopta in Uttarakhand, India. Systematics Our molecular results are further corroborated by the morphological analysis, which recovered a number of important diagnostic characters that allow distinguishing the populations of Ablepharus sp. n. from the Chopta, Uttarakhand, India and all other congeners (summarized below). These results support our hypothesis that this recently discovered lineage of Ablepharus sp. n. from Uttarakhand, India represents a previously unknown species, which we formally describe below. Ablepharus flammeus sp. n. Figs 5–7 ZooBank LSID: urn:lsid:zoobank.org:act: A6EB3BF4-EFF6-4E93-B2C8-3AABCC188A09 Holotype: ZSI-R-29457 (field no. BAM-171) (Figs 3, 5, 6, 7), adult male, collected by Andrey M. Bragin on 29 October 2022 from Duggal Bittha, Chopta, Uttarakhand, India (30.48° N, 79.17° E) at an elevation of 2,520 m a.s.l. Figure 3. Holotype of Ablepharus flammeus sp. n. (ZSI-R-29457, male) in life. Photograph by Andrey M. Bragin. 380 Andrey M. Bragin et al. Paratypes: Six males, one female, and two juveniles. Males: MZMU3657, MZMU3652, MZMU3658, MZMU3659, and MZMU3653 (field no. BAM-170, BAM-172, BAM-173, BAM-174, and BAM-175, respectively) collected on 29 October 2022 from Duggal Bittha, Chopta, Uttarakhand, India (30.48° N, 79.17° E) at an elevation of 2,520 m a.s.l.; MZMU3660 (field no. BAM-176) collected on 29 October 2022 from the vicinity of Tungnath Temple, Chopta, Uttarakhand, India (30.48° N, 79.20° E) at an elevation of 3,305m a.s.l. Female: MZMU3656 (field no. BAM-169) collected on 29 October 2022 from Duggal Bittha, Chopta, Uttarakhand, India (30.48° N, 79.17° E) at an elevation of 2,520m a.s.l. Juveniles: MZMU3654 and MZMU3655 (field no. BAM-167 and BAM-168, respectively) collected on 29 October 2022 from Duggal Bittha, Chopta, Uttarakhand, India (30.48° N, 79.17° E) at an elevation of 2,520 m a.s.l. Andrey M. Bragin collected all the above specimens. Etymology: The specific epithet of the new species “flammeus” is an adjective in nominative singular, in masculine gender, formed from the Latin words “flamma”, meaning “fire”. The name of the new species reflects the fiery coloration of the neck, belly, inner thighs, forearms, and the ventral part of the tail in males and juveniles, as well as the intense yellow coloration of these body parts in females. Notably, these small ectothermic lizards thrive at high altitudes exceeding 3,000 meters above sea level, with some individuals observed basking near the edges of snowfields. The name also symbolizes the ardent disposition of these lizards and their resilient struggle for survival. Furthermore, the species’ habitat lies near the Tungnath Temple – the highest mountain temple dedicated to the god Shiva – which also serves as the type locality for the species. Fire, as one of Shiva’s symbols, represents both destruction and the fight for life, reinforcing the essence of the species’ name. We propose “Fire-bellied Ground Skink” as the English common name, “Ognennobryukhii gologlaz” (“Огненнобрюхий гологлаз”) as the Russian common name, and “Laiteldulsen” as the Mizo common name for this species. Diagnosis: Ablepharus flammeus is a medium-sized, robust ablepharine skink with (1) snout–vent length (SVL) 41.6–50.6 mm; (2) head length (HL2) 7.7–9.1 mm; (3) tail length (TaL) 52.6–68.3 mm; (4) head, body, and caudal scales smooth; (5) supranasals absent; (6) nasal semi-divided; (7)the lower eyelid is movable, covered with small arranged granules and with large transparent window; (8)tympanum rounded, deeply sunk, with one (70%), two (20%) or three (10%) projecting lobules on the anterior border; (9) prefrontals two, separated by frontonasal and frontal or, rarely, prefrontals touch at one point; (10)frontal elongated, in contact with 1st and 2nd supraoculars laterally, rarely in contact with 3rd supraoculars; (11) frontoparietals two; (12) interparietal large, diamond-shaped, slightly elongated posteriorly, in contact with frontoparietals anteriorly and parietals posteriorly; (13) parietals in contact posterior to interparietal, not divided; (14) supraoculars four; (15)supralabials seven; (16) infralabials six; (17) chin shields in four pairs; (18) eight nuchals in four pairs behind parietals; (19)followed by four transverse dorsal rows between dorsolateral dark stripes, all similar in size; (20)50–52 paravertebral scales rows; (21) 52–56 ventral longitudinal rows (including gulars); (22) 26–28 midbody scale rows; (23)two enlarged precloacals; (24) the central subcaudal row enlarged; (25) 12 subdigital lamellae beneath fourth finger; (26) 16 subdigital lamellae beneath fourth toe; (27)femoral scales large and oriented longitudinally; (28) dorsum brown, with absent or rarely dotted two dark vertebral stripes; (29) two dark dorsolateral stripes; (30)venter, lower flanks, ventral region of the neck, tail, and limbs of males and juveniles from orange to red, and yellow in females. Description of holotype: An adult male in a good state of preservation with a 25 mm long mid-ventral incision. The hemipenes of the specimen are fully everted to determine the sex of the specimen and for description. The specimen Figure 4. The type locality of Ablepharus flammeus sp. n. in the vicinity of Chopta, Uttarakhand, India: (A) Gentle slopes covered with multilayer high-stem polydominant deciduous forests formed by Quercus semecarpifolia, Q. floribunda, Acer acuminatum, Aesculus indica, Q. leucotrichophora and Alnus nepalensis at an altitude of 2520 m a.s.l.; (B) steep slopes covered with patches of low mixed forest formed by Abies pindrow, Quercus semecarpifolia, Q. floribunda, Rhododendron arboreum, etc. grading into subalpine and alpine meadows. Photographs by Andrey M. Bragin. 381 A new species of Himalayan skink of the genus Ablepharus is preserved in a linear manner; limbs are curved slightly inward and backward. Snout–vent length (SVL) 41.6 mm; tail length (TaL) 62.4mm; total length 104 mm; snout acute in dorsal and lateral aspects (EED/END ratio 1.57), slightly projecting beyond lower jaws; nostril with lateral orientation, slightly oval, posterior edge oriented diagonally upward, situated much closer to the snout tip than to the eye (END/SNL ratio 0.71); head robust, longer than wide (HL2/HW ratio 1.14). Body slender (SVL/MBW ratio 5.9); head not distinct from the neck, but cervical interception weakly expressed in front of the attachment point of the forelimbs. Forelimbs and hindlimbs are relatively short and stout (SVL/FLL ratio 3.67; SVL/HLL ratio 2.65), normally developed, each with five fingers, the tips of digits of forelimb and hindlimb not meeting when the limbs are adpressed against each other along the body axis, adpressed hind limbs not reaching elbows when the hind limbs are adpressed (Fig. 5); forearm very short (ForL/SVL ratio 0.08); tibia longer than forearm (TibL/SVL ratio 0.12); digits moderately long, slender and ending with distinctly visible, sharp, thin, slightly curved, and pointed claws (Figure 5). Subdigital lamellae smooth on manus and pes, entire; number of subdigital lamellae without taking into account claw sheath: right manus 6-10-12-12-8; right pes 4-9-12-16-10. Relative lengths of digits (measurements in parentheses in mm): left manus IV > III >V > II > I; left pes IV (3.4 mm) > III (3.1 mm) > V (2.8mm) > II (2.6 mm) > I (1.2 mm). Scales on dorsal and ventral surfaces of the limbs are smooth and imbricate, femoral scales are large and oriented longitudinally. Scales on the palmar and plantar regions much smaller than the associated digit lamellae and the scales on ventral surfaces of limbs; rounded or slightly elongated, smooth, and subimbricate. Other measurements and ratios of the holotype and paratypes are given in Tables 1, 2. Rostral wider (1.9 mm) than high (1.1 mm), in contact with first supralabials, nasals and frontonasal (Figs 5, 6). A pair of nasals not in contact with each other. Nasal posteriorly in contact with slightly smaller anterior loreal; laterally widely in contact with first supralabials. A large single frontonasal in broad contact with the rostral anteriorly and the frontal posteriorly; widely in contact with nasal, the anterior loreal, and prefrontal laterally. A pair of prefrontals separated by the frontal. Prefrontal anteriorly in contact with frontonasal, posteriorly in contact with frontal, first supraocular, and first supraciliary scale, laterally in contact with anterior and posterior loreals. Frontal large, nearly twice longer than wide, irregularly pentagonal with slightly rounded tapering posterior margin, contacting frontonasal and prefrontals anteriorly, in wide contact with first and second supraoculars laterally, in wide contact with frontoparietals posteriorly. Frontoparietals widely in contact with parietals and interparietal scale posteriorly, contacting frontal and second, third and fourth supraoculars anterolaterally. Interparietal rhomboid, posteriorly in contact with parietals; parietal eye indistinct. Parietals anteriorly in contact with frontoparietals, interparietal and fourth supraoculars; laterally touching upper secondary temporal, upper and lower pretemporals; in contact with two pairs of nuchals posteriorly. Parietal overlapping upper secondary temporal. Four supraoculars, the second being the largest, decreasing in size in both directions (Figs 5, 6). Scales on the lateral aspect of the head bear pits, throughout their surface and dispersedly placed. The density of the pits on the scales is rare on dorsal head scales and barely noticeable towards the posterior borders of parietals (Figs 5, 6). Nostril with lateral orientation slightly oval, posterior edge oriented diagonally upward, located closer to the posterior edge of nasal. Nasal semi-divided, the groove runs from the central posterior edge of the nostril to the posterior edge of the nasal. Postnasals absent. Anterior and posterior loreals roughly quadrangular, posterior loreal wider than anterior loreal; anterior loreal anteriorly in contact with nasal and frontonasal, posteriorly in contact with posterior loreal and prefrontal, laterally in contact with first and second supralabials; posterior loreal anteriorly in contact with anterior loreal and prefrontal, posteriorly in contact with first supraciliary scale and two preoculars, laterally in contact with second and third supralabials. Three preoculars, the upper one smaller than the lower one; one presubocular, slightly larger than the middle preocular (Figs5, 6). The upper eyelid bordered by five supraciliary scales and seven ciliary scales; and the lower eyelid with nine ciliary scales; the scales covering the lower eyelid are small, round, granule-like, spaced apart from each other; the lower eyelid bears oval, horizontally oriented, large transparent window. The upper edge of the transparent window is practically in contact with the lower edge of the ciliaries, almost half of the horizontal diameter of the orbit long (0.35 mm high, 0.59 mm wide, ED/transparent window wide ratio 2.29). Seven supralabials, gradually increasing in size, with the fifth and sixth supralabials being the largest; fifth supralabials forming the lower border of the eyelid. Two small equal-sized postoculars anteriorly in contact with the last supraciliary scale and in touch at one point with the last supraocular, posteriorly in contact with upper and lower pretemporals, laterally in contact with upper postsuboculars. Three postsuboculars, where the middle one is the largest, posteriorly in contact with lower pretemporal and primary temporal, laterally in contact with lower postoculars and fifth and sixth supralabials. One large rhomboid primary temporal anteriorly in contact with lower pretemporal, two postsuboculars and sixth supralabial, posteriorly in contact with upper and lower secondary temporals and seventh supralabial. Two secondary temporals, the upper one larger than the lower one. Parietal and lower secondary temporal overlap upper secondary temporal. Secondary temporals anteriorly in contact with parietal, lower pretemporal, primary temporal and seventh supralabial, posteriorly in contact with nuchal, upper and lower tertiary temporals and upper postsupralabial. Two tertiary temporals, approximately the same size. The lower one overlapping the upper one. Two, equal-sized, posteriorly elongated postsupralabials behind the seventh supralabial. One row of large scales between postsupralabials and the anterior edge of the ear opening. The ear opening is large, slightly oblate in front 388 Andrey M. Bragin et al. Table 3. Comparative characters between Ablepharus flammeus sp. n., A. tragbulensis, A. himalayanus, A. ladacensis, A. capitaneus, A. sikimmensis, A. nepalensis, A. mahabharatus. Abbreviations are listed in the Materials and methods. The data is presented in the following form: minimum value–maximum value (average ± standard deviation). n – sample size, n.a. – data not available. Morphological measurements given in millimeters. Elev. – Elevation in m a.s.l.; Rep. – Reproduction. A. flammeus sp. n. A. tragbulensis A. himalayanus A. ladacensis A. capitaneus A. sikimmensis A. nepalensis A. mahabharatus Reference This study Das et al. 1998 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Eremchenko et al. 1998 Eremchenko et al. 1998 n=10 n=5 n=71 n=101 n=9 n=141 n=5 n=2 Elev. 2520–3305 2740 2600–3100 2590–5490 1100–2100 1200–3200 1500–1580 1950–2300 Rep. none none viviparous– oviparous viviparous oviparous oviparous none none SVL 41.6–50.6 (45.5±7.4) 44.2–56.5 (49.72±2.07) 66.8–78.5 (68.2±10) 36–70.5 (45.8±9.5) *–53.7 (43.8±9.3) *–55.8 (39.1±8.7) 32.5–41 (35.3±3.58) 30–44 (37±9.9) TaL 52.6–68.3 (59.2±10.0) 69.5 n.a. n.a. n.a. n.a. 58–69 (63.5±7.78) – (48±0) TaW 3.7–5.3 (4.4±0.8) 4.5–4.9 (4.7±0.28) n.a. n.a. n.a. n.a. n.a. n.a. AGD 24.4–31.3 (27.6±4.8) n.a. n.a. n.a. n.a. n.a. 14–20 (17.14±2.78) 13–22 (17.5±6.36) HL1 5.1–8.2 (7.1±1.3) 6.2–8.7 (7.45±1.77) n.a. n.a. n.a. n.a. n.a. n.a. HL2 7.7–9.1 (8.4±0.9) n.a. n.a. n.a. n.a. n.a. 7.7–8.7 (7.98±0.41) 7.2–9.5 (8.35±1.63) IOD 3.2–4.3 (3.6±0.6) 3.8–4 (3.9±0.14) n.a. n.a. n.a. n.a. n.a. n.a. SEL 8.5–9.4 (9.0±1.1) n.a. n.a. n.a. n.a. n.a. n.a. n.a. HW 6.0–7.1 (6.5±0.8) 4.9–5.6 (5.25±0.49) n.a. n.a. n.a. n.a. 5.5–6.2 (5.9±0.25) 5–6 (5.5±0.71) HH 3.9–5.7 (4.8±0.8) 4.1–4.5 (4.3±0.28) n.a. n.a. n.a. n.a. n.a. n.a. MBW 6.3–8.3 (7.4±1.4) 5.7–7.5 (6.6±1.27) n.a. n.a. n.a. n.a. n.a. n.a. IND 1.2–2.5 (1.7±0.5) 1.8–2 (1.9±0.14) n.a. n.a. n.a. n.a. n.a. n.a. ED 1.4–2.1 (1.8±0.3) 2.2–2.5 (2.35±0.21) n.a. n.a. n.a. n.a. n.a. n.a. TD 0.7–1.2 (0.9±0.2) 0.8–0.8 (0.8±0) n.a. n.a. n.a. n.a. n.a. n.a. SNL 3.2–3.9 (3.6±0.5) 3.9–4.2 (4.05±0.21) n.a. n.a. n.a. n.a. n.a. n.a. SFIL 15.1–19.3 (17.3±2.6) n.a. n.a. n.a. n.a. n.a. n.a. n.a. FLL 10.5–12.2 (11.4±1.6) 12–13.3 (12.65±0.92) n.a. n.a. n.a. n.a. 9–11 (9.64±0.86) 9.8–11 (10.4±0.85) HLL 13.9–18.8 (16.1±2.9) 17.8–18.7 (18.25±0.64) n.a. n.a. n.a. n.a. 13–17 (14.38±1.52) 13.8–17.5 (15.65±2.62) TBL 3.1–4.8 (3.9±0.9) 5.8–6.3 (6.05±0.35) n.a. n.a. n.a. n.a. n.a. n.a. T4L 4.7–6.3 (5.4±0.9) n.a. n.a. n.a. n.a. n.a. n.a. n.a. EED 3.6–4.7 (4.0±0.6) 3.2–3.7 (3.45±0.35) n.a. n.a. n.a. n.a. n.a. n.a. END 2.2–2.9 (2.5±0.4) 2.6–2.7 (2.65±0.07) n.a. n.a. n.a. n.a. n.a. n.a. RW 1.4–2.1 (1.7±0.3) 1.7–1.7 (1.7±0) n.a. n.a. n.a. n.a. n.a. n.a. RD 1.0–2.1 (1.3±0.5) 0.9–1 (0.95±0.07) n.a. n.a. n.a. n.a. n.a. n.a. 389 A new species of Himalayan skink of the genus Ablepharus total number of species of the genus Ablepharus from 20 to 21; Ablepharus flammeus represents the seventh Ablepharus species known for India. Despite the recent significant progress, our knowledge of molecular phylogeny, classification, and distribution of Himalayan ablepharine skinks remains incomplete. There is an urgent need to clarify the conservation status of newly described populations. The new species Ablepharus flammeus may well represent an example of an endemic confined to a specific plant community of highmountain hard-leaved deciduous forests, where anthropogenic activities can have a destructive impact. The discovery of new ablepharine skinks in the Himalayan region was not unexpected. In recent years, skinks have been described with unprecedented frequency (Chapple et al. 2023). However, until the present study, starting from the end of the 20th century, the species diversity of the Himalayan representatives of the genus Ablepharus had remained unchanged (Eremchenko et al. 1998, Eremchenko in Schleich & Kästle 2002). The ability to at least quadruple the known diversity of ablepharine skinks has been demonstrated by earlier research (Bragin et al. 2024). However, the information presented here is still based on only a few trips to various locations inside this enigmatic mountain region. As research into the Himalayan Mountain subsystems continues, a far greater number of species could be expected. It is also crucial to conduct further research on the genetics, morphology, and ecology of the newly discovered and existing species. Because of our limited collection of genetic markers and the resulting sequence database for the populations under study, we are unable to fully explain the evolutionary history of ablepharine skinks. Identification of even known Ablepharus species by morphological characters is also significantly difficult. To date, there are only two identification keys, both incomplete, including either only representatives of the Ablepharus species to the west of the Himalayas (Eremchenko & Szczerbak 1986), or only the Ablepharus species living in the territory of Nepal (Eremchenko in Schleich & Kästle 2002). Both identification keys are incomplete and contain a number of highly variable morphological characters. Lastly, a reassessment of the current synonyms of the known species would be imperative. The available descriptions of ablepharine skink species are also quite fragmented in terms of the choice of methods for calculating meristic morphological characters as well as the choice of sets of these characters. For instance, it is necessary to recheck specimens of A. tragbulensis because it is doubtful according to Das et al. (1998) whether this species of ablepharine skink has two postmentals and eight supraand seven infralabial scales as opposed to all other species’ single, unpaired postmental and seven supraand six infralabials. Other discrepancies are possible. We reiterate the A. flammeus sp. n. A. tragbulensis A. himalayanus A. ladacensis A. capitaneus A. sikimmensis A. nepalensis A. mahabharatus Reference This study Das et al. 1998 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Eremchenko et al. 1998 Eremchenko et al. 1998 n=10 n=5 n=71 n=101 n=9 n=141 n=5 n=2 HL2/SVL 0.17–0.23 (0.19±0.02) 0.14–0.15 (0.15±0.01) n.a. n.a. n.a. n.a. n.a. n.a. HW/SVL 0.12–0.17 (0.14±0.01) 0.1–0.11 (0.1±0.01) n.a. n.a. n.a. n.a. n.a. n.a. HH/HL2 0.49–0.67 (0.57±0.07) 0.52–0.66 (0.59±0.1) n.a. n.a. n.a. n.a. n.a. n.a. SNL/HW 0.47–0.62 (0.55±0.04) 0.75–0.8 (0.77±0.03) n.a. n.a. n.a. n.a. n.a. n.a. ED/SNL 0.4–0.59 (0.49±0.07) 0.56–0.6 (0.58±0.02) n.a. n.a. n.a. n.a. n.a. n.a. ED/HL2 0.17–0.25 (0.21±0.03) 0.29–0.35 (0.32±0.05) n.a. n.a. n.a. n.a. n.a. n.a. SVL/ MBW 5.53–7.08 (6.17±0.48) 7.53–7.75 (7.64±0.16) n.a. n.a. n.a. n.a. n.a. n.a. EED/END 1.45–1.92 (1.62±0.14) 1.23–1.37 (1.3±0.1) n.a. n.a. n.a. n.a. n.a. n.a. SVL/HW 6.18–8.14 (7±0.65) 9.02–10.09 (9.55±0.76) *–* (6.9±0.7) *–* (6.7±0.5) *–* (8±0.7) *–* (6.7±0.6) 5.51–6.61 (6.02±0.41) 6–7.33 (6.67±0.94) SVL/HL2 4.94–6.04 (5.4±0.47) 6.49–7.13 (6.81±0.45) *–* (5.5±0.6) *–* (5.1±0.6) *–* (6.5±0.6) *–* (5.3±0.7) 4.17–4.71 (4.42±0. 24) 4.17–4.63 (4.4±0.33) SVL/FLL 3.7–4.5 (4±0.3) 3.68–4.25 (3.97±0.4) *–* (3.7±0.4) *–* (3.2±0.3) *–* (4.1±0.2) *–* (3.8±0.4) 3.61–3.73 (3.66±0.05) 3.06–4 (3.53±0.66) SVL/HLL 2.5–3.5 (2.9±0.3) 2.48–3.02 (2.75±0.38) *–* (2.7±0.3) *–* (2.5±0.2) *–* (2.7±0.2) *–* (2.7±0.3) 2.32–2.63 (2.46±0.11) 2.17–2.51 (2.34±0.24) Table 3 continued 390 Andrey M. Bragin et al. Table 4. Summary diagnostic characters among Ablepharus flammeus sp. n., A. tragbulensis, A. himalayanus, A. ladacensis, A. capitaneus, A. sikimmensis, A. nepalensis, A. mahabharatus. Abbreviations are listed in the Materials and methods. The data is presented in the following form: minimum value–maximum value (average ± standard deviation). n – sample size, n.a. – data not available. Data recorded through “/” indicate the value on the right/left side of the head or body. Elev. – Elevation in m a.s.l.; Rep. – Reproduction. A. flammeus sp. n. A. tragbulensis A. himalayanus A. ladacensis A. capitaneus A. sikimmensis A. nepalensis A. mahabharatus Reference This study Das et al. 1998 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Eremchenko et al. 1998 Eremchenko et al. 1998 n=10 n=5 n=71 n=101 n=9 n=141 n=5 n=2 Elev. 2520–3305 2740 2600–3100 2590–5490 1100–2100 1200–3200 1500–1580 1950–2300 Rep. None none viviparous– oviparous viviparous oviparous oviparous none none PF no, or rarely touch in one point n.a. n.a. no, or rarely touch in one point or wide contact n.a. n.a. no no F-SO 10%–1st, 2nd, 3rd/ 1st, 2nd, 3rd; 90%–1st, 2nd/ 1st, 2nd n.a. 64%–1st, 2nd, 3rd/ 1st, 2nd, 3rd; 36%–1st, 2nd/ 1st, 2nd 97%–1st, 2nd, 3rd/ 1st, 2nd, 3rd; 3%–1st, 2nd/ 1st, 2nd 1st, 2nd/ 1st, 2nd 1st, 2nd/ 1st, 2nd 1st, 2nd/ 1st, 2nd 50%–1st, 2nd, 3rd/ 1st, 2nd, 3rd; 50%–1st, 2nd/ 1st, 2nd Nu 4/4 (4±0) n.a. n.a. 3–4/3–4 (3.7±0.9) 3–4/3–4 (3.3±0.6) 3–4/3–4 (3.1±0.9) 3–4/3 (–) 3–4/4–4.5 (–) PR 2–3 (2.8±0.4) 2 (2±0) n.a. n.a. n.a. n.a. n.a. n.a. SC 5 (5±0) n.a. 3–7 (5.5±0.8) 5–8 (6.2±0.7) 5–7 (6.1±0.4) 4–8 (5.8±0.6) n.a. 7 (7±0) Cnot thickened not thickened 47% distinctly thickened; 32% slightly thickened distinctly thickened, the last but one ciliar is very narrow not thickened not thickened n.a. n.a. PO 2 (2±0) 3 (3±0) 3–5 (4.5±0.6) (PO + PSO) 3–5 (3.8±0.5) (PO + PSO) 5 (–) (PO + PSO) 4–6 (5±0.3) (PO + PSO) n.a. n.a. PSO 3 (3±0) 2 (2±0) 2 (2±0) SL 7 (7±0) 8 (8±00) 7 (–) 7 (–) 7 (–) 7 (–) 7 (7±0) 7 (7±0) Timp tympanum rounded, deeply sunk, with lobules on the anterior border tympanum large, slit-like, with lobules on the anterior border tympanum with lobules on the anterior border tympanum slit-shaped, deeply sunk, usually with some distinct projecting lobules tympanum slit-shaped, very small, without projecting lobules tympanum rounded and very small, deeply sunk, without projecting lobules tympanum is relatively large, not deeply sunk, without projecting lobules tympanum relatively large, not deeply sunk, without pro - jecting lobules PEL 1–3 (1.3±0.7) present – (2.7±1.1) – (3±1.1) – (0.9±1.5) n.a. n.a. n.a. IF 6 (6±0) 7 (7±0) 6 (–) 6 (–) 6 (–) 6 (–) n.a. n.a. PM 1 (1±0) 2 (2±0) n.a. n.a. n.a. n.a. n.a. n.a. MBSR 26–28 (27.3±1.1) 28–34 (31±4.24) 24–32 (27.8±1.9) 28–38 (34.1±1.7) 30–32 (–) 21–29 (24.57±1.5) 30–32 (31.4±0.89) 30 (30±0) PVSR 52–56 (54.3±1.1) n.a. 51–62 (57.8±2.8) 58–77 (66.3±4) 62–67 (63.8±1.7) 45–66 (53.9±4.2) n.a. n.a. DBR 4 n.a. 4 6 6 4 6 6 VSR 60–66 (62.3±2.1) 70–73 (71.5±2.12) 57–75 (65.2±4.5) 71–90 (80.2±4.5) 61–72 (65.4±3.2) 33–71 (41.08±4.3) 39–44 (42.2±1.92) 40–43 (41.5±2.12) PrC 2 2 2 2 2 2 2 2 FL4 12 (12±0) n.a. 9–16 (12.9±1.5) 13–18 (15.4±1.5) 10–12 (12±–) 9–13 (10.7±1) n.a. n.a. 391 A new species of Himalayan skink of the genus Ablepharus necessity of indicating an arbitrary large number of measurements rather than just proportions, ratio (Günther 1864) and ranges of values of metric or meristic characters for all specimens (Ouboter 1986, Das et al. 1998) without supplying data for each specimen, as this impedes future work and renders previously acquired data meaningless. In this paper we provide a template for interpreting the head scalation (see Fig. 6), for its further application to other taxa in ablepharine skinks. At this stage, until a thorough morphological examination of specimens representing the recently discovered genetic lineages of ablepharine skinks has been completed, it is difficult to say which morphological characters are best for identification and have minimal variability. Apparently, the most informative morphological characters for identifying known species are: number of midbody scale rows counted as the number of longitudinal scale rows encircling the body at a point midway between the limb insertions; paravertebral scale rows counted as the number of scales in a line from, but not including, the nuchal scales to a point on the dorsum opposite the vent; transverse dorsal scales rows, counted between the lateral dark bars; ventral scale rows counted as a row of scales between the postmental and the precloacal scales; lamellae under the fingers, the number of fourth toe and finger subdigital lamellae, preoculars, presuboculars, postoculars and postsuboculars, as well as the number and the character of the stripes and bands on the dorsal and lateral sides of body, and the nature of the orange or red coloring of lateral and ventral sides of body of males and females. High variable morphological characters include the degree of contact of prefrontals with each other and with the frontal; the total number of ciliaries and nuchal scales, and number of nuchal scales on each side of the head counted as the number of the scales between opposing upper secondary temporal on each side of the head that contact the parietals. Morphological characters that were considered utilizing various techniques and, presumably, data that are currently widely dispersed and inappropriate for identification: the number of nuchal scales (Ouboter 1986, Eremchenko & Szczerbak 1986, Eremchenko et al. 1998, Eremchenko in Schleich & Kästle 2002), the number of pairs of chin scales (Das et al. 1998), the number of scales on the lateral side of the body counted from acetabular to glenoid cavities (Günther 1864), and the exact number of separate preoculars and separate presuboculars (in the currently available descriptions we should focus on their total number – preoculars + presuboculars) (Ouboter 1986, Eremchenko in Schleich & Kästle 2002). Expansion of the sampling of known species is necessary to understand the ranges of morphological variability. Furthermore, there are currently no adult specimens available for A. mahabharatus. Numerous morphological characters that were thought to be unchangeable later proved to be quite variable, which led researchers to look for more variations in hemipenial, craniological, or osteological structures. For instance, only nine species of ablepharine skinks have comprehensive descriptions of their hemipenes (Eremchenko & Panfilov 1990, Vergilov et al. 2017, Vergilov & Zlatkov 2024). Even though individual Himalayan ablepharine skink representatives appear to be widely distributed, there is a lack of information regarding their ecology, which can potentially be species-specific and can be a major factor in the conservation of known biodiversity. Information on reproduction strategies is thus lacking for the great majority of Ablepharus and Protoblepharus species found in the Himalayan region. Of particular interA. flammeus sp. n. A. tragbulensis A. himalayanus A. ladacensis A. capitaneus A. sikimmensis A. nepalensis A. mahabharatus Reference This study Das et al. 1998 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Eremchenko et al. 1998 Eremchenko et al. 1998 n=10 n=5 n=71 n=101 n=9 n=141 n=5 n=2 TL4 16 (16±0) 21 (21±0) 14–22 (17.2±1.9) 17–24 (20.6±1.6) 15–17 (17 ±–) 12–18 (15.17±1.1) 15.5–17 (16±0.61) 16.5–17.5 (17±0.71) F-H no in adults; yes for juveniles yes n.a. n.a. n.a. n.a. n.a. n.a. F-E no in adults; no in juveniles n.a. n.a. n.a. n.a. n.a. n.a. n.a. FemS femoral scales large and oriented longitudinally n.a. femoral scales large and oriented longitudinally femoral scales large and oriented longitudinally; presence of granulular scales on the upper femoral scalation a group of femoral scales is oriented across the thigh; upper femoral scales large femoral scales large and oriented longitudinally a group of femoral scales is oriented across the thigh; upper femoral scalation small a group of femoral scales is oriented across the thigh; upper femoral scalation small Table 4 continued 392 Andrey M. Bragin et al. Table 5. Summary diagnostic coloration patterns among Ablepharus flammeus sp. n., A. tragbulensis, A. himalayanus, A. ladacensis, A. capitaneus, A. sikimmensis, A. nepalensis, A. mahabharatus. n – sample size, n.a. – data not available. A. flammeus sp. n. A. tragbulensis A. himalayanus A. ladacensis A. capitaneus A. sikimmensis A. nepalensis A. mahabharatus Reference This study Das et al. 1998 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Ouboter 1989, Eremchenko in Schleich & Kästle 2002 Eremchenko et al. 1998 Eremchenko et al. 1998 n=10 n=5 n=71 n=101 n=9 n=141 n=5 n=2 Elev. in m a.s.l. 2520–3305 2740 2600–3100 2590–5490 1100–2100 1200–3200 1500–1580 1950–2300 Dark continuous vertebral stripes absent or rarely dotted two dark dorsal stripes ten or eleven sharply defined, alternate dark-brown and grayishwhite stripes absent; rarely dotted stripe or band (specimens from the western region of Nepal) four diffuse dotted stripes absent rarely dotted dark vertebral stripe narrow dark brown vertebral line absent Continuous dorsolateral stripes black n.a. yellow-golden to greyish, rarely diffuse whitish whitish, wide, distinct rarely, golden absent diffuse greyish Dark continuous lateral band broad dark brown broad dark brown distinct dark dark brown bronze-brown distinct brown blackish brown wide uniform black White continuous lateral stripes, bordered the lower edge of dark lateral stripe clear narrow broken white with an irregular edge n.a. present narrow whitish present present; in both sexes becomes orange to red in mating season between arms and legs narrow white with an irregular edge straight narrow white Coloration of the dorsum light grayyellow-brown to pale brown n.a. bronze-brown brown to olive light brown to grayish brown bronze-brown light brown light brown Coloration of venter grayish to yellowish white n.a. greyish blue greenish to blue grayish to yellowish white white to greenish white, yellow or greyish grayish greenish n.a. coloration of males (or/and females*) during the mating season venter, lower flanks, neck and ventral region of the tail orange to red n.a. lower flanks and ventral region of the tail orange to red, sometimes venter also become orange to red n.a. orange venter orange venter (in both males and females) *; white lateral stripes in both sexes becomes orange to red in mating season between arms and legs* n.a. n.a. Spots on the dorsal side irregular dark or black spots n.a. black spots, sometimes associated with white n.a. small, irregularly located black spots irregular dark or black spots diffuse light and dark strokes diffuse small brown specks Spots along lateral sides small white dots on the lower edge of the lateral band, sometimes irregularly dotted with black cream spots light spots n.a. small, brown small white dots on the lower edge of the lateral band, sometimes irregularly dotted with black rare light specks absent Dark, transverse, subcaudal bars or subcaudal banding absent n.a. absent absent absent absent absent absent Coloration of the crown of the head several large dark spots dark brown mottling on a gray forehead n.a. with black, brown and whitish specks n.a. n.a. n.a. n.a. White stripe under the eye present n.a. present present absent dotted present present 393 A new species of Himalayan skink of the genus Ablepharus est is the pattern of distribution of ablepharine skink species and the relative sizes of their habitats. We observe a picture that species living predominantly at low altitudes in the pine forest belt have fairly extensive distribution ranges in which there is practically no intraspecific variability. Examples of such species are A. sikimmensis, which appears to range from Central Nepal east over 450 km to Sikkim in India (Fig. 1). Sub-specific structure within submontane species is weakly expressed (as in the case of A. sikimmensis, where species delimitation analyses tend to group reconstructed mtDNA lineages) (Fig. 2, Supplementary Table S4). Conversely, high-mountain species are examples of confusing species complexes with a high degree of genetic and morphological interspecific distances (Fig. 2, Supplementary Table S4). Furthermore, after careful examination, it is quite likely that each of these lineages will have a very small range; currently, this is supported by the species Ablepharus flammeus, A.nepalensis, and A. tragbulensis, which are only known from a few locations and are not found anywhere else. The low-elevation zones are a barrier for high-elevation endemics that are restricted in distribution in the sky islands. Numerous undescribed lineages of the A. ladacensis and A. himalayanus species complexes will probably exhibit a comparable distribution (Fig. 1). This distribution is likely since ancestral lineages that colonized the Himalayan foothills either remained in them or moved up, where conditions on individual peaks can significantly vary. In essence, individual peaks acted as “sky islands” well isolated by growing gorges and glaciers, where newly arrived lineages underwent a process of gradual speciation. Thus, on individual peaks in one region, we can find several species of one species complex. In contrast, in the same region, all lower latitudes will be inhabited by another but only one species (Fig. 1). It is necessary to understand that the peak values for genus Ablepharus are the maximum among skinks (Chapple et al. 2023), and with such a scatter of altitudes, the boundaries between ancestral lineages could largely serve as the boundaries of plant communities in altitudinal zones. Clade 3 (Fig. 2), which included mid-mountain species like A. sikimmensis and A. mahabharatus, is likely a better example of such speciation because A.mahabharatus, inhabiting altitudes between 1,950 to 2,300 m a.s.l., is found to be more genetically homogeneous, while A. sikimmensis that reach altitudes of up to 3,200 m a.s.l. tends to split into several poorly distinguishable mitochondrial lineages, where deeper lineages are found in the highlands, while outer ones inhabit foothill habitats. It is possible that individual ancestral lineages could have reached some areas of the mountain ranges independently, after the colonization of the lower heights by their relatives. This hypothesis seems more logical than the hypothesis of diversification of taxa as the Himalayas grew, given the dating of colonization by ablepharine skinks of the Sino-Japanese distribution range (Ghosh et al. 2023) and the association of most of the basal lineages of ablepharine skinks and their Lygosominae relatives with humid forests (Ouboter 1986, Eremchenko & Szczerbak 1986, Mirza et al. 2022), most of the flora of which consists of immigrated plant taxa that have evolved and diversified over millions of years following the Himalayan formation (Singh & Singh 1987, Pandit & Kumar 2013, Pandit et al. 2014, Manish & Pandit 2018). The data obtained in this research greatly complicates the hypothesis put forth by Ouboter (1986) and Eremchenko & Szczer bak (1986) regarding the invasion of the Himalayas by three lineages (corresponding to the ancestral lineages of A. sikimmensis, A. himalayanus, and A. ladacensis), and their subsequent expansion into the Eastern, Central, and Western Himalayan regions, respectively. We observed that far more intricate taxonomic diversity that has gone largely unreported, with a complicated web of habitats that include poorly understood sympatric zones and unexplored mechanisms of potential hybridization. More samples from still inadequately explored Himalayan regions (particularly from Ladakh and Sikkim in India, Bhutan, and Eastern and Western Nepal) and more genetic markers are needed to provide a more precise biogeographical scenario. Acknowledgements The first author deeply appreciates the friendship and unwavering support of his companions, Platon V. Yushchenko and Yulia D. Kozhevnikova, during the journey to the Himalayan peaks, without whom the new species would not have been discovered and described. AMB is also eternally grateful to Rajiv Bisht Puran Singh and his wonderful family, who provided shelter and all possible assistance and support during his stay in Uttarakhand. We thank Alexey V. Trofimets (member of MSU HerpLab) for his support and assistance. AMB and NAP are grateful to Andrei N. Kuznetsov (JVRTSTRC, Hanoi) for his long-standing support and encouragement. We thank Pratyush P. Mohapatra (ZSI, Kolkata) for his assistance with the registration of the holotype. ZAM acknowledges the Max Planck Society for support. We are grateful to the anonymous reviewers for their helpful comments, which enabled us to improve the previous version of the manuscript. This work was supported by the Russian Science Foundation to NAP (Grant No. 22-14-00037-P, molecular and data analysis). References Ankita, R., A. Bharti, V. Kumar & D. Kumar (2017): The preliminary molecular study of four skink species in Rajaji Tiger Reserve (RTR), Uttarakhand, using 12S rRNA mitochondrial locus. – Mitochondrial DNA Part B, 2: 495–499. Bragin, A. M., S. L. Litvinchuk, L. J. Borkin, D. A. Melnikov, D. V. Skorinov, D. Jablonski, R. Masroor, H. T. Lalremsanga, Z. A. Mirza, C. Dufresnes & N. A. Poyarkov (2024): Hidden on the roof of the world: mitochondrial data reveals exceptional genetic diversity of Himalayan ablepharine skinks (Reptilia: Scincidae). – Russian Journal of Herpetology, 31: 351–368. Chapple, D. G., A. Slavenko, R. Tingley, J. E. Farquhar, M. Camaiti, U. Roll & S. Meiri (2023): Built for success: distribution, morphology, ecology and life history of the world’s skinks. – Ecology and Evolution, 13: e10791. Che, J. K., F. Yan & Y. Zhang (2020): Amphibians and reptiles in Tibet – diversity and evolution. – Science Press, Beijing. Das, I., B. Dattagupta & N. Gayen (1998): Systematic status of Lygo soma himalayanum tragbulensis Alcock, “1897” 1898 (Sauria: 394 Andrey M. Bragin et al. Scincidae) collected by the Pamir boundary commission, 1885. – Russian Journal of Herpetology, 5: 147–150. De Queiroz, K. (2007): Species concepts and species delimitation. – Systematic Biology, 56: 879–886. Eremchenko, V. K. (1992): On systematics of Asymblepharus ladacensis (Günther, 1864) (Sauria: Scincidae, Lygosominae). – pp. 58–65 in: Eremchenko, V. K., A. M. Panfilov & E. I. Tsarinenko (eds): Research conspectus on cytogenetics and systematics of some Asian species of the Scincidae and Lacertidae. – Ilim Press, Bishkek. Eremchenko, V. K. (2002): Family Scincidae (Skinks). – pp. 723–768 in: Schleich, H. H. & W. Kästle (eds): Amphibians and Reptiles of Nepal. Biology, Systematics, Field Guide. – ARG Ganther Verlag KG, Ruggell. Eremchenko, V. K. & A. M. Panfilov (1990): Ablepharus darvazi sp. nov. – new species Ablepharus (Sauria, Scincidae) from Tadjikistan. – Izvestiya Akademii Nauk Kirgizskoi SSR, Khimiko-tekhnologicheskie biologicheskie nauki, 4: 56–63. Eremchenko, V. K. & N. N. Szczerbak (1986): Ablepharine lizards of the USSR and adjacent countries. – Frunze, Ylym Publ. Eremchenko, V. K., N. Helfenberger, K. B. Shah & A. M. Panfilov (1998): Two new species of skinks (Scincidae: Lygosominae) from Nepal. – Vestnik Natsionalnoi Akademii Nauk Kyrgyzskoi Respubliki, 4: 41–45. Ghosh, A., M. Sil, K. B. Ukuwela & A. Datta-Roy (2023): Independent origins or single dispersal? Phylogenetic study supports early Cenozoic origin of three endemic Indo-Sri Lankan Lygosomine skink genera. – Zoologica Scripta. Grismer, L. L., P. L. Wood, E. S. H. Quah, S. Anuar, N. A. Poyarkov, T. Neang, N. L. Orlov, P. Thammachoti & H. Seiha (2019): Integrative taxonomy of the Asian skinks Spheno mor phus stellatus (Boulenger, 1900) and S. praesignis (Boulenger, 1900), with the resurrection of S. annamiticus (Boettger, 1901) and the description of a new species from Cambodia. – Zootaxa, 4683: 381–411. Günther, A. C. L. G. (1864): The Reptiles of British India. – Ray Society, London. Hall, T. A. (1999): BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. – Nucleic Acids Symposium Series, 41: 95–98. Hedges, S. B. (2014): The high-level classification of skinks (Reptilia, Squamata, Scincomorpha). – Zootaxa, 3765: 317–338. Huelsenbeck, J. P. & D. M. Hillis (1993): Success of phylogenetic methods in the four-taxon case. – Systematic Biology, 42: 247– 264. IUCN (2025): The IUCN Red List of Threatened Species. Version 2024-2. – Available online at https://www.iucnredlist.org. Katoh, K., K. Misawa, K. Kuma & T. Miyata (2002): MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. – Nucleic Acids Research, 30: 3059–3066. Kumar, S., G. Stecher, M. Li, C. Knyaz & K. Tamura (2018): MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. – Molecular Biology and Evolution, 35: 1547–1549. Lanfear, R., B. Calcott, S. Y. W. Ho & S. Guindon (2012): PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. – Molecular Bio logy and Evolution, 29: 1695–1701. Macey, J. R., J. A. Schulte, J. L. Strasburg, J. A. Brisson, A. Larson, N. B. Ananjeva, Y. Wang, J. F. Parham & T. J. Papenfuss (2006): Assembly of the eastern North American herpetofauna: new evidence from lizards and frogs. – Biology Letters, 2: 388–392. Manish, K. & M. K. Pandit (2018): Geophysical upheavals and evolutionary diversification of plant species in the Himalaya. – PeerJ, 6: e5919. Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. da Fonseca & J. Kent (2000): Biodiversity hotspots for conservation priorities. – Nature, 403: 853–858. Mirza, Z. A., A. M. Bragin, H. Bhosale, G. G. Gowande, H. Patel & N. A. Poyarkov (2022): A new ancient lineage of ablepharine skinks (Sauria: Scincidae) from eastern Himalayas with notes on origin and systematics of the group. – PeerJ, 10: e12800. Nguyen, L. T., H. A. Schmidt, A. Von Haeseler & B. Q. Minh (2015): IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. – Molecular Bio logy and Evolution, 32: 268–274. Ouboter, P. E. (1986): A revision of the genus Scincella (Reptilia, Sauria, Scincidae) of Asia, with some notes on its evolution. – Zoo logische Verhandelingen, 229: 1–66. Pandit, M. K. & V. Kumar (2013): Land-use change and conservation challenges in the Indian Himalaya: Past, present, and future. – pp. 121–133 in: Sodhi, N. S., L. Gibson & P. H. Raven (eds): Conservation biology: Voices from the tropics: Wiley-Blackwell. Pandit, M. K., K. Manish & L. P. Koh (2014): Dancing on the roof of the world: ecological transformation of the Himalayan landscape. – BioScience, 64: 980–992. Puillandre, N., S. Brouillet & G. Achaz (2021): ASAP: assemble species by automatic partitioning. – Molecular Ecology Resources, 21: 609–620. Ronquist, F. & J. P. Huelsenbeck (2003): MrBayes 3: Bayesian phylogenetic inference under mixed models. – Bioinformatics, 19: 1572–1574. Russell, D. W. & J. Sambrook (2001): Molecular Cloning: A Laboratory Manual. – Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY. Singh, J. & S. Singh (1987): Forest vegetation of the Himalaya. – The Botanical Review, 53: 80–192. Uetz, P., P. Freed & J. Hošek (eds) (2025): The Reptile Database. – Available from: http://www.reptile-database.org, accessed 8 August 2025. Vergilov, V. S., B. P. Zlatkov & N. D. Tzankov (2017): Hemipenial differentiation in the closely related congeners Ablepharus kitaibelii (Bibron & Bory De Saint-Vincent, 1833), and Ablepharus budaki (Göçmen, Kumlutaş & Tosunoğlu, 1996) (Squamata: Sauria: Scincidae). – Herpetozoa, 30: 39–48. Vergilov, V. S. & B. P. Zlatkov (2024): Morphology of hemipenes and its taxonomic implication in the fused eyelids species of the genus Ablepharus (Squamata: Scincidae). – Journal of Comparative Zoology (Zoologischer Anzeiger), 312: 79–91. Weng, S. Y., S. Huang, L. Yang, L. F. Peng, C. Wei & J. C. Li (2021): The complete mitochondrial genome of Asymblepharus himalayanus (Scincidae; Sauria) and its phylogenetic position within Scincidae. – Mitochondrial DNA Part B, 6: 3031–3032. Supplementary data The following data are available online: Supplementary Table S1. Localities, voucher information, and GenBank accession numbers for all specimens used in molecular analyses in this study. Supplementary Table S2. Details of primers used in the study for PCR amplification and sequencing. Supplementary Table S3. Characteristics of analyzed mtDNA sequences and the proposed optimal evolutionary models for gene and codon partitions as estimated in PartitionFinder v1.0.1. Supplementary Table S4. Genetic divergence of the genus Ablepharus in 16S and cyt b gene sequences.