Monday, October 15, 2018

[Herpetology • 2018] Multilocus Phylogeny and Revised Classification for Mountain Dragons of the Genus Japalura s.l. (Agamidae: Draconinae) from Asia


Japalura s.l. 

in Wang, Che, Lin, Deepak, Aniruddha, et al., 2018. 

Abstract
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.

Keywords: China, India, integrative taxonomy, lizard, new genus, Tibetan Plateau

Figure 2. Phylogenetic relationships among Japalura sensu lato based on both Maximum Likelihood and Bayesian Analyses of two mitochondrial genes (COI and ND2), tRNAs and three nuclear genes (BDNF, CMOS, and R35). Maximum Likelihood bootstrap and Bayesian posterior probability values are included at all nodes, except (1) terminal nodes that unify multiple individuals of the same species [which all have 1.00/100 support (Bayesian/Maximum Likelihood)]; and (2) non-conflicting (either kind of analysis yielded significant supports) yet inconsistent nodes between two types of analyses (e.g. nodes unifying clades I and J), in which only the Bayesian posterior probability is given. Photos of selected species of Japalura s.l. are included (scaled to reflect relative sizes of each species) with their current taxonomic names (vs. revised taxonomic names, summarized in taxonomic accounts).

Taxonomic accounts 
Japalura Gray, 1853
 Etymology: The Latin name ‘Japalura’ may be derived from a locality name in India, and the term is feminine gender. We suggest the English common name as ‘Himalayan Dragon’, and the Chinese name as ‘攀蜥’ (pronounced as ‘Pan-Xi’). 

Type species: Japalura variegata, Gray, 1853.

Included species: Based on our phylogenetic results, we assign the following species to the genus Japalura sensu stricto: J. andersoniana, J. kumaonensis, J. tricarinata and J. variegata. Following our morphological results and proposed morphological diagnoses, we also assign J. dasi, J. major and J. sagittifera into this genus, pending future phylogenetic studies.


Cristidorsa Wang, Deepak, Datta-Roy, Lin, Jiang, Che & Siler gen. nov.

 Etymology: The Latin term ‘Cristidorsa’ means ‘ridged dorsum’, which describes the distinct, characteristic ridges on the dorsal surface of the body in the new genus. The generic name is feminine and it consists of two parts, namely ‘Cristi-’ (meaning ‘ridged’) and ‘-dorsa’ (meaning ‘dorsum’). We suggest the English common name as ‘Ridged Dragons’ and the Chinese name as ‘棱背蜥’ (pronounced as ‘Leng-Bei-Shi’). 

Type species: Cristidorsa otai (Mahony, 2009).

Included species: Based on our phylogenetic results, we assign C. otai and C. planidorsata to the genus Cristidorsa.


Diploderma Hallowell, 1861 

Etymology: The Latin generic name ‘Diploderma’ consists of two parts, ‘Diplo-’ means ‘double’ or ‘many’, and ‘-derma’ means ‘skin’, and the whole word is in a neuter gender. As the previous generic name ‘Japalura’ and most species names of the genus s.l. are feminine, most names of species that are now assigned to Diploderma need their gender changed to neutral (except for existing neutral-gender names like brevipes or flaviceps, Latin nouns like vela or names derived from peoples’ names, i.e. dymondi, luei, makii, swinhonis, varcoae and zhaoermii). We suggest the English common name of the genus as ‘Mountain Dragon’, and the Chinese common name as ‘龙蜥’ (pronounced as ‘Long-Xi’). 

Type species: Diploderma polygonatum Hallowell, 1861.

Included species: Based on our phylogenetic results, we assign the following species into the genus Diploderma: D. batangense, D. brevipes, D. chapaense, D. dymondi, D. flaviceps, D. laeviventre, D. luei, D. makii, D. micangshanense, D. polygonatum (and all of its subspecies), D. slowinskii, D. splendidum, D. swinhonis, D. varcoae, D. vela, D. yulongense, D. yunnanense and D. zhaoermii

According to our proposed morphological diagnoses, we also assign Diploderma brevicaudum, D. fasciatum, D. grahami, D. hamptoni and D. iadinum to this genus, pending on future phylogenetic confirmations. In total, 24 species of Japalura s.l. are reclassified into the genus Diploderma (for discussion on the taxonomic status of D. ngoclinense see below).


Pseudocalotes Fitzinger, 1843 

Type species: Pseudocalotes tympanistriga (Gray, 1831).

Included species: A single species of Japalura s.l. Pseudocalotes kingdonwardi bapoensis, is reclassified into the genus Pseudocalotes. Currently, the genus includes 22 recognized species in total (Grismer et al., 2016a; Harvey et al., 2017).


Kai Wang, Jing Che, Simin Lin, V. Deepak, Datta-Roy Aniruddha, Ke Jiang, Jieqiong Jin, Hongman Chen and Cameron D. Siler. 2018. Multilocus Phylogeny and Revised Classification for Mountain Dragons of the Genus Japalura s.l. (Reptilia: Agamidae: Draconinae) from Asia. Zoological Journal of the Linnean Society. zly034.  DOI:  10.1093/zoolinnean/zly034