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On the Occurrences of Japalura kumaonensis and Japalura tricarinata (Reptilia: Sauria: Draconinae) in China

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Although the recognized distribution of Japalura kumaonensis is restricted largely to western Himalaya, a single, isolated outlier population was reported in eastern Himalaya at the China-Nepal border in southeastern Tibet, China in Zhangmu, Nyalam County. Interestingly, subsequent studies have recognized another morphologically similar species, J. tricarinata, from the same locality in Tibet based on photographic evidence only. Despite these reports, no studies have examined the referred specimens for either record to confirm their taxonomic identifications with robust comparisons to congener species. Here, we examine the referred specimen of the record of J. kumaonensis from southeastern Tibet, China; recently collected specimens from the same locality in southeastern Tibet; type specimens; and topotypic specimens of both J. kumaonensis and J. tricarinata, to clarify the taxonomic identity of the focal population from southeastern Tibet, China. Our results indicate that individuals of the referred Tibetan population differ from J. kumaonensis in external morphology, but match descriptions and specimens of J. tricarinata. We conclude that the previous record of J. kumaonensis from Tibet was a misidentification of J. tricarinata, and J. kumaonensis should be recognized as occurring in western Himalaya only. To facilitate future taxonomic studies of the genus Japalura sensu lato in the Himalaya, we provide a detailed description of J. tricarinata and an updated diagnostic key to the genus from the Himalaya.
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... We examined a total of 217 specimens representing 27 species of Japalura s.l. and 31 specimens representing 11 species from seven genera of outgroup agamid diversity (Supporting Information, Appendix S3). Following previous studies (Moody, 1980;Mahony, 2010;Grismer et al., 2016a), and our prior knowledge on morphological characteristics of Draconinae (Mahony, 2010;Wang et al., 2018b), we chose a suite of pholidosis characteristics to examine for all specimens, as they are shown to be most useful in delimiting generic-level diversity (abbreviations given in parentheses): presence of modified scales on dorsal body (MDS), defined as presence of scales on dorsal body that are modified in shape, size or texture (e.g. enlarged, keeled, flat or raised in conical shape); degree of modification of dorsal modified scales (DMDS), defined as the level of modification compared to the ground or normal scales on dorsal body, defined as strongly modified or weakly modified; presence of dorsolateral ridges (PDR), defined as presence of regularly and closely arranged, lateral rows of enlarged, keeled dorsal scales on the body; presence of V-shaped ridges on dorsal body (PVR), defined as presence of enlarged, keeled scales that are arranged in V-shape ridges on dorsal body; and presence of head spines (PS), defined as presence of elongated spines on the post-orbital, occipital and supratympanic regions of head. ...
... Although species of both clades are from the Himalayan region, they are recovered in distantly related groups within Draconinae with strong support (Fig. 2). Statistical topology tests further support the phylogenetic results, rejecting the assumed monophyly of the Himalayan congeners (PP = 0.00; Table 4; Stuart-Fox & Owens, 2003;Mahony, 2010;Ananjeva et al., 2011;Wang et al., 2018b). Second, we reject the previously hypothesized sister relationship between island Japalura species and Malayodracon (Pyron et al., 2013). ...
... However, we recognize Japalura sensu stricto to represent this clade for the following reasons: (1) Japalura sensu stricto has temporal priority over the other two generic names; (2) recognizing the synonym Oriotiaris as a valid genetic name would render itself and Japalura sensu stricto both paraphyletic (Fig. 2); (3) for Oreocalotes, although we do not have access to the genetic materials of the type species, the morphology of the type species (O. major or J. major as currently recognized) falls within the morpho-cluster of species of Japalura sensu stricto, and the type species is morphologically most similar to J. kumaonensis (Wang et al., 2018b), which is a confirmed member of Japalura sensu stricto (Figs 2, 3). Therefore, we treat both Oriotiaris and Oreocalotes as junior synonyms of Japalura sensu stricto. ...
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Although the genus Japalura s.l. has long been recognized as paraphyletic based on limited genetic sampling, its problematic taxonomy has not been revised, and phylogenetic relationships among the majority of congeners remain unknown. Here we utilize a densely sampled dataset of both multilocus genetic and morphological data to provide the first phylogenetic inference of relationships among Japalura s.l.species. Our results show that Japalura s.l. is paraphyletic, consisting of four major clades that are scattered across the phylogeny of the subfamily Draconinae: the first clade from the western, central and middle-eastern Trans-Himalayas, the second clade from the far eastern Trans-Himalayas, the third clade from East Asia and the last clade from Indochina. To address this widespread paraphyly of the genus and to stabilize the taxonomy within the family Draconinae, we revise the current taxonomy and split Japalura s.l. into four genera. By doing so, we recognize two existing generic names, Japalura sensu stricto and Pseudocalotes, resurrect one name available in the literature, Diploderma, and describe one new genus, Cristidorsa gen. nov. We discuss phylogenetic relationships and taxonomy within Japalura s.l. and present a diagnostic key to all recognized genera of the subfamily Draconinae.
... We examined a total of 217 specimens representing 27 species of Japalura s.l. and 31 specimens representing 11 species from seven genera of outgroup agamid diversity (Supporting Information, Appendix S3). Following previous studies (Moody, 1980;Mahony, 2010;Grismer et al., 2016a), and our prior knowledge on morphological characteristics of Draconinae (Mahony, 2010;Wang et al., 2018b), we chose a suite of pholidosis characteristics to examine for all specimens, as they are shown to be most useful in delimiting generic-level diversity (abbreviations given in parentheses): presence of modified scales on dorsal body (MDS), defined as presence of scales on dorsal body that are modified in shape, size or texture (e.g. enlarged, keeled, flat or raised in conical shape); degree of modification of dorsal modified scales (DMDS), defined as the level of modification compared to the ground or normal scales on dorsal body, defined as strongly modified or weakly modified; presence of dorsolateral ridges (PDR), defined as presence of regularly and closely arranged, lateral rows of enlarged, keeled dorsal scales on the body; presence of V-shaped ridges on dorsal body (PVR), defined as presence of enlarged, keeled scales that are arranged in V-shape ridges on dorsal body; and presence of head spines (PS), defined as presence of elongated spines on the post-orbital, occipital and supratympanic regions of head. ...
... Although species of both clades are from the Himalayan region, they are recovered in distantly related groups within Draconinae with strong support (Fig. 2). Statistical topology tests further support the phylogenetic results, rejecting the assumed monophyly of the Himalayan congeners (PP = 0.00; Table 4; Stuart-Fox & Owens, 2003;Mahony, 2010;Ananjeva et al., 2011;Wang et al., 2018b). Second, we reject the previously hypothesized sister relationship between island Japalura species and Malayodracon (Pyron et al., 2013). ...
... However, we recognize Japalura sensu stricto to represent this clade for the following reasons: (1) Japalura sensu stricto has temporal priority over the other two generic names; (2) recognizing the synonym Oriotiaris as a valid genetic name would render itself and Japalura sensu stricto both paraphyletic (Fig. 2); (3) for Oreocalotes, although we do not have access to the genetic materials of the type species, the morphology of the type species (O. major or J. major as currently recognized) falls within the morpho-cluster of species of Japalura sensu stricto, and the type species is morphologically most similar to J. kumaonensis (Wang et al., 2018b), which is a confirmed member of Japalura sensu stricto (Figs 2, 3). Therefore, we treat both Oriotiaris and Oreocalotes as junior synonyms of Japalura sensu stricto. ...
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Although the genus Japalura s.l. has long been recognized as paraphyletic based on limited genetic sampling, its problematic taxonomy has not been revised, and phylogenetic relationships among the majority of congeners remain unknown. Here we utilize a densely sampled dataset of both multilocus genetic and morphological data to provide the first phylogenetic inference of relationships among Japalura s.l.species. Our results show that Japalura s.l. is paraphyletic, consisting of four major clades that are scattered across the phylogeny of the subfamily Draconinae: the first clade from the western, central and middle-eastern Trans-Himalayas, the second clade from the far eastern Trans-Himalayas, the third clade from East Asia and the last clade from Indochina. To address this widespread para-phyly of the genus and to stabilize the taxonomy within the family Draconinae, we revise the current taxonomy and split Japalura s.l. into four genera. By doing so, we recognize two existing generic names, Japalura sensu stricto and Pseudocalotes, resurrect one name available in the literature, Diploderma, and describe one new genus, Cristidorsa gen. nov. We discuss phylogenetic relationships and taxonomy within Japalura s.l. and present a diagnostic key to all recognized genera of the subfamily Draconinae.
... Because species of the genus Diploderma are known to be sexually dimorphic in morphometric measurements Wang et al., 2015Wang et al., , 2018, morphometric data of each sex are treated separately for analyses and comparisons. ...
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... All specimens were measured by using a digital caliper to the nearest 0.1 mm, except for tail length, which was measured to the nearest 1 mm using a ruler. The morphometric characters were measured following Wang et al. (2018), with some additional morphological characteristics added (Supplementary material (Methods)). Color names and codes follow Köhler (2012). ...
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