Molecular phylogenetics and taxonomic reassessment of the widespread agamid lizard Calotes versicolor (Daudin, 1802) (Squamata, Agamidae) across South Asia

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.

A checklist of herpetofauna of Telangana state, India

“…For the present checklist, we researched and critically analysed all published (both peer-reviewed and nonpeer-reviewed) literature, online databases (including iNaturalist, India Biodiversity Portal, HerpMapper) and also relied on field surveys conducted since 1995 in various parts of Telangana State (Srinivasulu 2003;Srinivasulu et al 2006Srinivasulu et al , 2009Srinivasulu et al , 2011Srinivasulu et al , 2014Srinivasulu et al , 2016Seetharamaraju et al 2008Seetharamaraju et al , 2011Seetharamaraju et al , 2019Srinivasulu & Das 2008;Seetharamaraju & Srinivasulu 2013;Kumar et al , 2017aSeetharamaraju 2015;Kumar 2018). We confirmed the species identities by consulting standard references (Daniel 2002;Das 2002;Whitaker & Captain 2004), incorporating further updates by more recent literature (Deepak et al 2016(Deepak et al , 2018(Deepak et al , 2021Lajmi et al 2016;Ganesh et al 2017;Mirza & Patel 2018;Pal et al 2018;Mallik et al 2020Mallik et al , 2021Bisht et al 2021;Gowande et al 2021;Bandara et al 2022). We have also provided explanation for deletion of the taxa earlier reported in literature, and appeal to future workers to collect voucher specimens or photographs to report additions to the herpetofauna diversity of Telangana.…”

Invasive risk assessment and expansion of the realized niche of the Oriental Garden Lizard Calotes versicolor species complex (Daudin, 1802)

“…However, the majority of populations is still assigned to C. versicolor sensu lato (below abbreviated s.l.). Recently, Gowande et al (2021) published a phylogeny based on the mitochondrial 16S and CO1 genes and supported this by morphological data to disentangle the Indian taxa of the C. versicolor complex. According to the latter authors, the species complex contains at least four distinct lineages: (1) C. versicolor s.str.…”

“…Further, we georeferenced six occurrence records from the literature (see Table 1) using GEOLocate (https://www.geo-locate.org/) to increase the number of records for the invasive range. Furthermore, we used 93 genetically confirmed records from Gowande et al (2021) to increase the number of records. After checking the dataset for outliers in ArcMap (ESRI 2020)(see distribution in introduction), we corrected the dataset for potential sampling bias (see systematic sampling in Fourcade et al 2014) and spatial autocorrelation using the raster package (Hijmans 2016) and the ecospat.mantel.correlogram() function in the ecospat package (Di Cola et al 2017) for R (R Core Team 2021).…”

“…However, it must be mentioned that at least C. irawadi is still known to be a species complex and also C. farooqi includes another distinct phylogenetic lineage and further surveys are required to clarify these issues. Furthermore, Gowande et al (2021) found that C. calotes (Linnaeus, 1758) is a phylogenetic sister taxon to C. farooqi and is therefore also included in the C. versicolor complex.…”

Reptilian assemblages in the wetlands of Amboli hill complex, northern Western Ghats, Maharashtra, India during the monsoon season

“…Reptiles were photographed using a DSLR camera (Canon 600D); identification characters were based on consultation of standard literature (Daniel 2002;Das 2008;Whitaker & Captain 2008). For nomenclature, we followed Khandekar et al ( 2021), effecting other nomenclatural updates from recent publications (Ganesh et al 2020(Ganesh et al , 2021Mallik et al 2020Mallik et al , 2021Deepak et al 2021;Gowande et al 2021). Jaccard's similarity index of species richness between wetlands was calculated using PAST software (Hammer et al 2001).…”