Organisms Diversity & Evolution

, Volume 16, Issue 3, pp 659–677 | Cite as

Molecular data in conjunction with morphology help resolve the Hemidactylus brookii complex (Squamata: Gekkonidae)

  • Aparna LajmiEmail author
  • Varad B. Giri
  • K. Praveen Karanth
Original Article


Molecular data are increasingly being used to resolve cryptic species complexes; however, subsequent formal species description and taxonomic revisions often remain incomplete. Given that most species are described based on morphology-based alpha taxonomy, one cannot resolve nomenclatural issues of species complexes without the aid of morphology. In this study, we examined the taxonomic status of a long-known human commensal and species complex, Hemidactylus brookii. To this end, samples of H. cf. brookii and related species were collected across India. We analyzed molecular as well as morphological data to resolve the taxonomy of this species complex. Seven deeply divergent, well-supported clades were recovered using the mitochondrial phylogeny, five of which were also retrieved in the nuclear tree. One of these consists of five morphologically distinct species of ground-dwelling Hemidactylus. The genetic distances across each clade of putative species of H. brookii sensu lato were comparable to that between morphologically distinct species of ground-dwelling Hemidactylus. Meristic characters such as number of precloacal-femoral pores, number of non-pore bearing scales interrupting the series of pored scales, dorsal pholidosis, and presence/absence of divided lamellae can be used to distinguish these putative species from each other. However, morphological characters of H. brookii sensu stricto did not correspond to any of the putative species studied. The study also revealed that the “H. brookii complex” in India includes two commensal species, Hemidactylus parvimaculatus and Hemidactylus murrayi. Furthermore, these two lineages have independently acquired adaptations that could have assisted them in exploiting human habitat. An identification key to diagnose species within this complex and rest of the Hemidactylus in India is proposed.


Cytochrome b RAG1 Cryptic species Invasive species Phylogeny 



We would like to thank Ishan Agarwal, Aniruddha Datta-Roy, Aakarsh, Harshil Patel, Saunak Pal, Mrugank Prabhu, Pankaj Lad, Kshamata Gaikwad, Navendu Page, Ashok Kumar Mallik, Deepak Veerappan, Manjunath Reddy, Jahnavi Joshi, Diptarup Nandi, Rochishnu Dutta, Shreekant Deodhar, N P I Das, Sartaj Ghuman and Rohini Bansal for their help in sampling. Aaron Bauer and Indraneil Das for helping us find required literature. Kavita Isvaran, Diptarup Nandi, and Rittik Deb for discussions and comments regarding the statistical analyses, Navendu Page and Viraj Torsekar for their help in making figures. We are grateful to Aaron Bauer, and the anonymous reviewers for giving their valuable comments on the manuscript. Thanks to Asad R Rahmani, Bombay Natural History Society, for his support and encouragement. All the lab members of Karanth lab and staff at the Collection Department, Bombay Natural History Society. Department of Science and Technology, and Ministry of Environment and Forest for funding fieldwork and molecular work. Partial funding for fieldwork also came from National Science Foundation (U.S.A.) grants DEB 0844523 and DEB 1019443 to Aaron M. Bauer. VG would like to thank Uma Ramakrishnan (DAE Outstanding Scientist Grant to Uma Ramakrishnan) and Krushnamegh Kunte for their support.

Compliance with ethical standards


Molecular work was funded by the Department of Science and Technology, India (grant no. SR/SO/AS-57/2009). Fieldwork was mainly funded by the Ministry of Environment and Forest (India), and partly by National Science Foundation (U.S.A.) grants DEB 0844523 and DEB 1019443 to Aaron M. Bauer.

Conflict of interest

The authors declare that they have no competing interests.

Ethical approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed. This article does not contain any studies with human participants performed by any of the authors.

Supplementary material

13127_2016_271_MOESM1_ESM.pdf (60 kb)
Supplementary Table 1 Morphometric data in millimeter (PDF 60 kb)
13127_2016_271_MOESM2_ESM.docx (48 kb)
Supplementary Table 2 Tukey multiple comparisons of means across each clade with adjusted p value. The clades that were significantly different are in bold. (DOCX 47 kb)
13127_2016_271_MOESM3_ESM.pdf (125 kb)
Supplementary Table 3 Meristic data and descriptive characters for individuals within the H. brookii complex. FP (L, R) – Femoral pore (left, right); SFP – non-pore bearing scales between pored series; NDT – Number of enlarged dorsal tubercle rows; SL – Supralabials; IL – infralabials; (PDF 124 kb)
13127_2016_271_MOESM5_ESM.pdf (210 kb)
Supplementary Fig. 1 Maximum likelihood (ML) tree based on mitochondrial cyt b dataset. The values on each node represent ML bootstrap value / Bayesian posterior probability. Support values below 50 % have been denoted as ‘-’. In the case of identical sequences from multiple individuals, a single exemplar sequence was used. The sample number of the exemplar sequence is shown and the sample numbers of other identical sequences are given in parenthesis (PDF 210 kb)
13127_2016_271_MOESM6_ESM.pdf (216 kb)
Supplementary Fig. 2 Maximum likelihood (ML) tree based on nuclear RAG1 dataset. The values on each node represent ML bootstrap value / Bayesian posterior probability. Support values below 50 % have been denoted as ‘-’ (PDF 215 kb)
13127_2016_271_MOESM7_ESM.pdf (346 kb)
Supplementary Fig. 3 Haplotype network of RAG1 sequences showing clusters corresponding to each of the clades retrieved in the phylogeny (PDF 345 kb)
13127_2016_271_MOESM4_ESM.txt (2 kb)
Supplementary Material 1 Likelihood support values for each of the putative species identified by the PTP method (TXT 1 kb)


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Copyright information

© Gesellschaft für Biologische Systematik 2016

Authors and Affiliations

  • Aparna Lajmi
    • 1
    Email author
  • Varad B. Giri
    • 2
    • 3
  • K. Praveen Karanth
    • 1
  1. 1.Centre for Ecological SciencesIndian Institute of ScienceBangaloreIndia
  2. 2.National Centre for Biological SciencesTIFRBangaloreIndia
  3. 3.Bombay Natural History SocietyWestern Ghats Regional StationKolhapurIndia

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