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| Calotes vultuosus (Harlan, 1825), Calotes farooqi Auffenberg & Rehman, 1993 in Gowande, Pal, Jablonski, ... et Shanker, 2021. DOI: 10.3897/vz.71.e62787 |
Abstract
The genus Calotes Cuvier, 1817 (Agamidae: Draconinae) is highly diverse, with species occurring in South and Southeast Asia, and Oceania. Most species of the subfamily except C. versicolor have narrow geographic distributions. Calotes versicolor is distributed from western Iran in the west to south China and Indonesia in the east and has been introduced to parts of Africa and North America. The species has had a complicated taxonomic history; multiple species and subspecies related to C. versicolor were described from India and adjoining regions, which were synonymized in subsequent revisions. However, a study of Burmese C. versicolor yielded two new species, C. htunwini and C. irawadi, indicating that C. versicolor is a species complex. Such integrative taxonomic studies have not been carried out in India, the supposed type locality of C. versicolor. Hence, we studied C. versicolor sensu lato from the Indian subcontinent and generated sequences of mitochondrial 16S and COI fragments from tissues sampled from multiple localities in the region, including the type localities of its synonyms. Phylogenetic analyses revealed four well-supported, deeply-divergent lineages, supported by morphological data. These lineages represent (i) C. versicolor sensu stricto, from South India and parts of the east coast, (ii) C. irawadi sensu lato from northeast India and Southeast Asia, (iii) a synonym from the eastern Indo-Gangetic Plains which we resurrect here, and (iv) a subspecies from Pakistan which we elevate to species level. We provide re-descriptions for the resurrected or elevated species, and a diagnostic key to the species of the C. versicolor complex. The study shows that C. versicolor sensu stricto is endemic to parts of southern and eastern India, and not widely distributed, though it may have been introduced to other parts of the world.
Key words: 16S, COI, distribution, mtDNA, phylogenetics, species complex, species delimitation, systematics
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| A map of south and Southeast Asia representing the localities from which morphological and/or primary or secondary genetic data were collected. Red circles indicate the localities from which C. versicolor sensu stricto (Clade1) originate, yellow samples indicate those of C. irawadi sensu lato (Clade2), blue circles represent the localities of C. vultuosus comb. nov. (Clade3), green circles indicate Clade4, with C. farooqi stat. nov. (77, 83, 85, 86) and C. aff. farooqi (78, 79). A single sequence of Calotes irawadi from Huanan Province, China (locality 73) is not marked on the map, since it was very distant to all other localities, as also those whose precise localities were unknown. Triangles indicate type localities of the respective species. See supplementary Table S1 for details on molecular and/or morphological vouchers used, and their respective localities. |
Calotes versicolor (Daudin, 1802)
Etymology: The specific epithet is an adjective in Latin referring to variable or to turn (versi, derived from versare) and color (color) of the species in life.
Distribution: The species appears to be endemic to India, occurring largely on the Southern Granulite Terrain and the eastern coast of India. The species was found in the Southern Western Ghats, the south-west coast of India, the southern Eastern Ghats, the eastern coast of India till Mahanadi basin in the north, and in the low-elevation areas of Peninsular India between the Eastern and the Western Ghats during this study. The species also has been introduced to the Maldives (Figs 1–3).
Calotes htunwini Zug and Vindum, 2007
Taxonomic comments:
We did not include this species in our analyses; however, the species is distinct from members of the C. versicolor group due to the horizontal orientation of the scale rows of the neck and the supra-axillary region. The species occurs throughout the lower elevations of Myanmar’s Central Dry Zone (Zug et al. 2007).
Calotes irawadi Zug, Brown, Schulte and Vindum, 2007
Taxonomic comments:
The species can be diagnosed by its posteriorad or vertical orientation of the scale rows of the neck and the supra-axillary region, this separates the species from C. htunwini; detailed comparisons with C. versicolor, the species resurrected and elevated in this communication are provided in the diagnoses and comparisons sections of those species. The species was thought to have a narrower distribution in the Central Dry Zone of Myanmar, in comparison to C. htunwini (Zug et al. 2007). It was subsequently reported from the northeast Indian states of Assam and Tripura (Das et al. 2009, Anonymous 2020), and has recently been reported from Western Yunnan, China (Liu et al. 2021). Our samples from northeast Indian states of Arunachal Pradesh and Tripura belonged to Clade2 (Fig. 2), thus confirming the occurrence of the species in northeast India.
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| An uncollected male of Calotes vultuosus comb. nov., from Burdwan, West Bengal, India, in life. Photograph by Ayan Mondal. |
Calotes vultuosus (Harlan, 1825) comb. nov.
Etymology: The specific epithet ‘vultuosa’ could refer to the grim or frowning look of the species when viewed frontally, which it gets due to the flattened scales of the CanthR and the supercilium. The specific epithet vultuosa is feminine singular and is here changed to the masculine gender vultuosus in agreement with the generic epithet, Calotes, which is masculine in gender.
Distribution: The species appears to be widely distributed, occupying parts of the Dharwar Craton till Brahmagiri, Deccan Volcanic Province, the Central Highlands, the Gangetic Plains, and the Indian Deserts (Fig. 1).
Calotes farooqi Auffenberg and Rehman, 1993, stat. nov.
Etymology: The specific epithet is a patronym in genitive singular case, dedicated to Farooq Ahmed, former Director, Zoological Survey Department, Pakistan.
Conclusion:
An integrated molecular-morphological investigation demonstrates that C. versicolor on the Indian subcontinent is composed of multiple lineages. It is not as widely distributed as was earlier believed, but occurs in southern India and on the east coast, with the possibility of its occurrence on the island of Sri Lanka. The study led to the revalidation of the species C. vultuosus comb. nov., a species potentially endemic to the Deccan Volcanic Province, Central Highlands, arid and semi-arid regions of India and Gangetic Plains, which was under the synonymy of C. versicolor; and the promotion of the subspecies C. versicolor farooqi from Pakistan to species rank, as C. farooqi stat. nov. We further confirm the presence of C. cf. irawadi in northeast India. The taxonomic status of the populations introduced to Maldives was also resolved. Our results lend further support to the argument that many widely distributed species represent species complexes, demanding urgent taxonomic attention.
Gaurang Gowande, Saunak Pal, Daniel Jablonski, Rafaqat Masroor, Pushkar U. Phansalkar, Princia Dsouza, Aditi Jayarajan and Kartik Shanker. 2021. Molecular Phylogenetics and Taxonomic Reassessment of the Widespread Agamid Lizard Calotes versicolor (Daudin, 1802) (Squamata, Agamidae) Across South Asia. Vertebrate Zoology. 71: 669-696. DOI: 10.3897/vz.71.e62787
