2016-12-23T18:52:10

In this first paper of a series of three, the taxonomy of the Asian pitvipers of the genus Tropidolaemus is re-evaluated on the basis of morphological analyses. Variation in morphological characters was investigated on the basis of specimens from the whole range of the pitviper currently known as Tropidolaemus wagleri (Boie, 1827). Our results, based on morphological univariate and multivariate analyses, define three clusters of populations that are morphologically diagnosable and which are here considered to represent distinct species following the Biological Species Concept and the Phylogenetic Species Concept. After a review of available names among the list of synonyms created during the confused taxonomical history of the genus Tropidolaemus, it appears that Tropidolaemus wagleri (Boie, 1827) is the valid name of the first cluster which includes populations inhabiting Southern Thailand, West Malaysia, Sumatra, Nias, Mentawei Archipelago and Bangka Island (but not Belitung). In order to stabilize the binomen, we select and describe a neotype for Tropidolaemus wagleri. A second cluster, for which the binomen Tropidolaemus subannulatus (Gray, 1842) isavailable, includes in this preliminary step populations from Borneo, Sulawesi, Sulu Archipelago and the Philippines. Its detailed taxonomy will be addressed in the second paper of the series. Lastly, the third cluster includes specimens from Mindanao Island, Philippines, recognized here as Tropidolaemus philippensis (Gray, 1842).KEY WORDS: Indonesia, Thailand, West Malaysia, Sumatra, Borneo, Sulawesi, Philippines, Serpentes, Viperidae, Tropidolaemus, Tropidolaemus wagleri, Tropidolaemus subannulatus, Tropidolaemus philippensis, Tropidolaemus laticinctus, Tropidolaemus huttoni, taxonomy, neotypeAmong pitvipers of tropical Asia, members of the genus Tropidolaemus Wagler, 1830 are among the most widespread and often commonly encountered venomous snakes in many islands of the Indo-Malayan Archipelago. Long regarded as a synonym or a subgenus of Trimeresurus (see, for example, Brattstrom, 1964), the genus Tropidolaemus was resurrected by Burger (1971) to then accommodate the sole species formerly called Trimeresurus wagleri. The validity of the genus is accepted by all recent authors. This genus is characterized by the absence of a nasal pore, upper surfaces of the snout and head covered with distinctly keeled small scales, strongly keeled gular scales, second supralabial not bordering the anterior margin of the loreal pit and topped by a prefoveal, and a green coloration in juveniles which may or may not change with growth. For long, Tropidolaemus wagleri was the sole species included in the monotypic genus, but David & Vogel (1998) showed that the Indian species Trimeresurus huttoni Smith, 1949 was clearly a member of this genus. In this first paper of a series of three, we address the rather confused nomenclatural history and taxonomy of Tropidolaemus wagleri (Boie, 1827) sensu auctorum (see, for example, David & Ineich, 1999; McDiarmid et al., 1999; Gumprecht et al., 2004). Members of this species complex are widespread throughout the IndoMalayan part of Asia, with an isolated population in Southern Vietnam. Besides this latter country, it is distributed from southern Thailand to the Philippines and Sulawesi Island, including West Malaysia, and the islands of Sumatra, Borneo, Bangka, Nias, the Mentawai Archipelago, and Belitung. Although a common and conspicuous, very variable species, few authors tried to investigate its taxonomy, most probably following Boulenger (1896) who synonymised the various names under the sole specific name Lachesis wagleri. Nevertheless, Taylor (1917, 1922) examined Philippine populations and recognized three subspecies, of which two were considered endemic to the Philippine islands, Tropidolaemus wagleri alboviridis (Taylor, 1917) and T. wagleri subannulatus (Gray, 1842). This position was not accepted by Leviton (1964), who investigated the taxonomy of the Philippine populations and considered again Tropidolaemus wagleri to be monotypic. However, Leviton added: “The exact status of the nominal species and subspecies I have placed into the synonymy of T. wagleri cannot be settled until the type specimens and additional material from scattered localities can be examined.” The monotypic status of Tropidolaemus wagleri was accepted by subsequent authors (Harding & Welch, 1980; Hoge & Romano-Hoge, 1981; Alcala, 1986; Welch, 1988; Golay et al., 1993; David & Vogel, 1996, Manthey & Grossmann, 1997; McDiarmid et al., 1999), although some noted that the taxonomy of the species was unsatisfactory (David & Ineich, 1999). David & Vogel (1998) discussed the taxon described as Trimesurus philippensis Gray, 1842, regarded as valid by Taylor (1922) and Maslin (1942) as Trimeresurus philippinensis, but placed in the synonymy of Tropidolaemus wagleri by Leviton (1964), who, however, seemingly did not examine its holotype. David & Vogel (1998) examined two specimens, namely the holotypes of Trimeresurus philippensis Gray, 1842 and Tropidolaemus hombronii Jacquinot & Guichenot, 1848, clearly a synonym of the former one. David & Vogel (1998) and David & Ineich (1999) noted that both specimens displayed notable morphological differences (scalation of head and body and coloration) with Tropidolaemus wagleri.

Slug-eating snakes of the subfamily Pareinae are an insufficiently studied group of snakes specialized in feeding on terrestrial mollusks. Currently Pareinae encompass three genera with 34 species distributed across the Oriental biogeographic region. Despite the recent significant progress in understanding of Pareinae diversity, the subfamily remains taxonomically challenging. Here we present an updated phylogeny of the subfamily with a comprehensive taxon sampling including 30 currently recognized Pareinae species and several previously unknown candidate species and lineages. Phylogenetic analyses of mtDNA and nuDNA data supported the monophyly of the three genera Asthenodipsas, Aplopeltura, and Pareas. Within both Asthenodipsas and Pareas our analyses recovered deep differentiation with each genus being represented by two morphologically diagnosable clades, which we treat as subgenera. We further apply an integrative taxonomic approach, including analyses of molecular and morphological data, along with examination of available type materials, to address the longstanding taxonomic questions of the subgenus Pareas, and reveal the high level of hidden diversity of these snakes in Indochina. We restrict the distribution of P. carinatus to southern Southeast Asia, and recognize two subspecies within it, including one new subspecies proposed for the populations from Thailand and Myanmar. We further revalidate P. berdmorei, synonymize P. menglaensis with P. berdmorei, and recognize three subspecies within this taxon, including the new subspecies erected for the populations from Laos and Vietnam. Furthermore, we describe two new species of Pareas from Vietnam: one belonging to the P. carinatus group from southern Vietnam, and a new member of the P. nuchalis group from the central Vietnam. We provide new data on P. temporalis, and report on a significant range extension for P. nuchalis. Our phylogeny, along with molecular clock and ancestral area analyses, reveal a complex diversification pattern of Pareinae involving a high degree of sympatry of widespread and endemic species. Our analyses support the “upstream” colonization hypothesis and, thus, the Pareinae appears to have originated in Sundaland during the middle Eocene and then colonized mainland Asia in early Oligocene. Sundaland and Eastern Indochina appear to have played the key roles as the centers of Pareinae diversification. Our results reveal that both vicariance and dispersal are responsible for current distribution patterns of Pareinae, with tectonic movements, orogeny and paleoclimatic shifts being the probable drivers of diversification. Our study brings the total number of Pareidae species to 41 and further highlights the importance of comprehensive taxonomic revisions not only for the better understanding of biodiversity and its evolution, but also for the elaboration of adequate conservation actions.

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The group of Asian colubrid species morphologically similar to Oligodon taeniatus (Günther, 1861), previously containing only O. taeniatus, Oligodon mouhoti (Günther, 1864) and Oligodon barroni (Smith, 1916), is revised on the basis of variation in external morphology and dentition of 175 specimens. The confused nomenclatural history of O. taeniatus and its name bearing type is discussed. A neotype is described for Simotes quadrilineatus Jan & Sordelli, 1865, a synonym of O. taeniatus. The holotype of Simotes taeniatus var. mouhoti Boulenger, 1914 is identified. Three new species within this group are described. Oligodon pseudotaeniatus spec. nov. is described on the basis of specimens from central Thailand. This species is morphologically similar to Oligodon taeniatus, but differs by the combination of 17 dorsal scale rows at midbody, 8 supralabials, the absence of dark dorsal and tail blotches and the presence of a vertebral stripe edged with black but no dorsolateral stripes. Oligodon deuvei spec. nov. is described on the basis of specimens from southern Vietnam and Laos; it differs from other known species of the group by the combination of 12–15 maxillary teeth, 17 dorsal scale rows at midbody, usually seven supralabials, the absence of dark dorsal and tail blotches and the presence of a broad vertebral stripe, often conspicuously orange or rusty red. This species is most similar to Oligodon barroni but differs from the latter by a higher number of maxillary teeth and the absence of dark dorsal and tail blotches. Lastly, Oligodon moricei spec. nov. is described on the basis of a single specimen from southern Vietnam. It differs from other species by the combination of a broad rusty brown vertebral stripe edged with two broad black stripes, 12 maxillary teeth, 17 dorsal scale rows, a high number of ventral scales, seven supralabials and a dark cloudy or smoky venter. These new species are compared with other species known from the Indo-Chinese Region. The diagnoses of O. taeniatus, O. mouhoti and O. barroni are revised. A key to members of the group is given.

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Kraits of the genus Bungarus Daudin 1803 are widely known venomous snakes distributed from Iran to China and Indonesia. Here, we use a combination of mitochondrial DNA sequence data and morphological data to describe a new species from Yingjiang County, Yunnan Province, China: Bungarus suzhenaesp. nov. Phylogenetically, this species forms a monophyletic lineage sister to the Bungarus candidus/multicinctus/wanghaotingi complex based on cyt b and ND4 genes but forms a sister species pair with the species B. magnimaculatus Wall & Evans, 1901 based on COI gene fragments. Morphologically, B. suzhenaesp. nov. is similar to the B. candidus/multicinctus/wanghaotingi complex but differs from these taxa by a combination of dental morphology, squamation, coloration pattern, as well as hemipenial morphology. A detailed description of the cranial osteology of the new species is given based on micro-CT tomography images. We revised the morphological characters of B. candidus/multicinctus/wanghaotingi complex and verified the validity of three species in this complex. The distribution of these species was revised; the records of B. candidus in China should be attributed to B. wanghaotingi. We also provide an updated key to species of Bungarus.

This paper deals with three nomenclatural and taxonomic problems affecting two species groups of the colubrid snake genus Oligodon Fitzinger, 1826: (i) A neotype is formally designated for Coronella cyclura Cantor, 1839, associating this specific nomen with populations from India, Bangladesh and Myanmar with 19 scale rows at midbody; (ii) Oligodon kheriensis Acharji & Ray, 1936 is shown to be a valid species of the Oligodon cyclurus group occurring in northern India and Nepal; (iii) The type-locality of Simotes multifasciatus Jan & Sordelli, 1865 is shown to be Sultanpur, India. This taxon is considered a synonym of Oligodon cinereus (Günther, 1864). The range of this species in India is extended. The status of specimens of Oligodon cinereus from India and Myanmar is briefly discussed. Specimens from Thailand identified as Oligodon cinereus multifasciatus and Oligodon cinereus swinhonis (Günther, 1864) are referred to Oligodon joynsoni (Smith, 1917). India is home to at least 21 species of the genus Oligodon, an updated list of which is provided.