Frank Tillack

The Himalaya and Hindu Kush remain among the least scientifically explored mountain regions of Asia. Within their vertebrate fauna, pitvipers of the Gloydius himalayanus (Günther, 1864) complex have long presented taxonomic challenges due to limited sampling in remote areas and the absence of integrative analyses. Here, we present a comprehensive taxonomic revision of the G. himalayanus complex using an integrative framework that combines genetic and distributional data, external morphology, osteology, and ecological evidence. Our dataset also uniquely incorporates both newly collected material and DNA sequences obtained from 19 ^th - and early 20 ^th -century museum specimens, including one of the syntypes of G. himalayanus (BMNH 1946.1.19.64), underscoring the enduring value of historical collections in modern taxonomy. Analyses of four mitochondrial and three nuclear genes from populations across the western and central Himalaya and the Hindu Kush, many of which were genetically studied for the first time, recover five well-supported monophyletic lineages: G. himalayanus sensu stricto, G. chambensis , and three previously unrecognised lineages from the Hindu Kush of north-western Pakistan, the Hazara region of north-eastern Pakistan at the western margin of the Himalaya, and the Himalaya of western and central Nepal. Genetic distances in protein-coding mtDNA markers indicate species-level divergence among these lineages, ranging from 9.2–12.6% in cyt b and 8.1–14.1% in ND4, with additional support from partial allelic differentiation in phased nuclear markers, morphology, and osteology. Based on these results, we formally describe three new species, substantially expanding the known diversity of Gloydius along the southern Himalaya. To stabilise the nomenclature, we designate a lectotype for Halys himalayanus Günther. By integrating modern and historical data, this study demonstrates the lasting scientific relevance of natural history collections and emphasises the urgent need for conservation of these narrowly distributed and potentially regionally threatened mountain snakes.

The genus Ptyctolaemus Peters, 1864 is currently represented by three species; P. gularis , P. collicristatus and P. chindwinensis . Previous molecular phylogenetic studies have highlighted additional lineages within P. gularis. Phylogenetic analysis using the ND2 mitochondrial gene recovered three distinct lineages in India. Multivariate analyses using morphological data placed the type specimen of P. gularis (ZMB 5004) with the samples collected from Meghalaya. We assign this lineage as P. gularis and describe the other two lineages as new species. Ptyctolaemus siangensis sp. nov. found west of Siang River in Arunachal Pradesh and Ptyctolaemus namdaphaensis sp. nov. found in Namdapha, Changlang district of Arunachal Pradesh. Although phenotypically similar the two new species differ from its congeners in gular colouration and subtle morphological differences.

We conduct a review of the known species of Cyrtodactylus from Nepal based on type and topotypical material, resulting in the synonymization of one species and description of three additional species from Central Nepal. The five species of Cyrtodactylus from Nepal are members of the Indo-Burma clade and are at least 11.7 % divergent from known congeners based on the mitochondrial ND2 gene. The results show that C. nepalensis and one of the new species are allied to the C. fasciolatus group while the other two new species and C. martinstolli are members of the mountain subclade within the khasiensis group. The new species can be readily distinguished from congeners in the Himalaya by a combination of body size, meristic characters including number of dorsal tubercle rows and mid ventral scales across the belly, the number and arrangement of pores in males and females, and subcaudal condition.

We discovered a historical specimen in the collection of the ZMB that fits an overlooked description of Agama gutturosa by Merrem (1819) and possibly represents a specimen on which Seba (1734) based one of his illustrations. Morphologically, the specimen held in the ZMB collection and that described by Merrem (1819) agrees with Bronchocela jubata Duméril & Bibron, 1837 and would constitute a senior synonym of that taxon. An earlier lectotype designation for the taxa Agama cristatella Kuhl, 1820 and Agama gutturosa Merrem, 1820 by Figueroa (2021) is discussed and corrected accordingly. In order to stabilize nomenclatural usage, we consider Agama gutturosa Merrem, 1819 as a nomen oblitum, so that the long standing nomina Bronchocela cristatella (Kuhl, 1820) and Bronchocela jubata Duméril & Bibron, 1837 can prevail.

From the beginning of the 19th century to the present, the authorship of the description of the Balkan wall lizard, Podarcis tauricus (Pallas, 1814) was considered undoubted. Recently, Schmidtler (2022) has been concluded that the description of this species should be attributed to J.G. Georgi. Therefore we have chosen to discuss this issue. The description of Lacerta taurica by Georgi in 1801 was based on earlier works by C.L. Hablitz (1785, 1789). However, his description does not align with our current understanding of the morphological and ecological characteristics of the taxon currently recognized as P. tauricus. Some details contained in this description suggest that it was based not on the species recently known as Podarcis tauricus, but on another lizard inhabiting the same place and recently known as Darevskia lindholmi (Szczerbak, 1962). If recognize authorship of the name Lacerta taurica as belonging to Georgi (1801), this would require transferring that name to the species currently known as D. lindholmi and the need to introduce a new name for the species currently known as P. taurica, that will cause numerous confusions. To provide the stability of the names, we suggest to continue to regard Pallas (1814) as the author of the name Lacerta taurica and will apply to the Commissuion of Zoological Nomenlature for conservation of this status. Regrettably, a specimen collected by Pallas himself in Crimea has not been located until now. However, the search for preserved historic specimens continues.

In response to the recent in passing (“en passant”) taxonomic decision to split Naja naja (Linnaeus) and recognise the Sri Lankan populations as a separate species, N. polyocellata Deraniyagala, we analyse the evidence underlying the proposal and its nomenclatural implications. The proposed split is weakly supported by the available evidence, so that retaining N. naja as a single species seems appropriate until further analysis. Moreover, the proposal raises several issues concerning types, type locality and nomenclature. Linnaeus’ description of Coluber naja was based on a single preserved specimen seen by him (now lost) and several illustrations in Seba’s Thesaurus. The specimens that were the basis of these illustrations constitute part of the type series. Two of the latter specimens, ZMB 2795 and 2796, have been rediscovered in the collections of the Museum für Naturkunde, Berlin. Here, we describe them, and determine that both are of Sri Lankan origin. To settle the question of the type and type locality of this iconic taxon, we designate ZMB 2796 as lectotype for the species, thereby implicitly restricting the type locality to Sri Lanka. The name “polyocellata” thus becomes a subjective junior synonym of Coluber naja, and the name Naja brasiliensis Laurenti, 1768 an objective junior synonym thereof. Any taxonomic recognition of additional diversity within N. naja would thus require the renaming of Indian, not Sri Lankan spectacled cobras, but should await a significant body of convincing evidence. We caution against taxonomic decisions taken “in passing”, based on limited evidence and without in-depth assessment of their nomenclatural implications.

The analyses of molecular genetic data (mtDNA markers 16S, ND4, CYTB, and the nuclear marker c-mos) provided evidence that the Asian cat snake taxa Boiga multomaculata and B. ochracea actually represent a single species. They form mixed clades of low intraclade genetic differentiation. This evidence for conspecificy is supported by the lack of differentiation in all examined pholidotic and morphometric characters. Therefore, we formally place Dipsas ochracea Theobald, 1868 in the synonymy of Dipsas multomaculata Boie, 1827. We provide a summary of the tangled taxonomic history of the taxa involved in this study. Also, we resurrect Dipsadomorphus stoliczkae Wall, 1909 from synonymy of B. ochracea, for specimens exhibiting 21 midbody dorsal scale rows. Boiga stoliczkae is found in the Himalayas north and west of the Brahmaputra valley. Finally, based on the detection of historical genetic lineages within the newly defined species Boiga multomaculata we recognize three subspecies: Boiga multomaculata multomaculata (Boie, 1827), Boiga multomaculata ochracea (Theobald, 1868), and Boiga multomaculata septentrionalis n. ssp. which is distributed in northern Myanmar and Assam and Nagaland, India. We designate BMNH 1946.1.2.60 (1) as neotype of Dipsas ochracea Theobald, (2) as lectotype of D. ochraceus Günther, and (3) as lectotype of Boiga ochracea walli Smith, thereby making these names objective synonyms. Finally, we designate BMNH 94.12.31.55 as lectotype of Dipsadomorphus stoliczkae Wall.

Colour polymorphism has been previously reported in several colubrid snakes including Boiga spp. In this paper, we report colour variations within the poorly known southern Indian Boiga dightoni, provide the first molecular data for this species, from two localities (including the type locality) and compare them with data from other congeners. Additionally, we provide detailed dentition and hemipenis descriptions for B. dightoni. Molecular data for B. dightoni show very little difference (0.2–0.4% 16S; 0.9–1.2% cyt b) to the recently described Boiga whitakeri, also from southern India. We have re-examined and present new information on the pholidosis of the type specimens of B. whitakeri and reconsider its taxonomic status. On the basis of molecular data and overlapping morphological characteristics, we argue that Boiga whitakeri and Boiga dightoni are conspecific, and place B. whitakeri under the subjective synonymy of the latter. Furthermore, we show that colour polymorphism in B. dightoni is a gender-independent character and that both colour morphs are found in high as well as low elevations and partly in sympatry. A revised key to the Boiga ceylonensis complex is provided.