Organisms Diversity & Evolution

, Volume 19, Issue 2, pp 341–361 | Cite as

The hills are alive with geckos! A radiation of a dozen species on sky islands across peninsular India (Squamata: Gekkonidae, Hemiphyllodactylus) with the description of three new species

  • Ishan AgarwalEmail author
  • Akshay Khandekar
  • Varad B. Giri
  • Uma Ramakrishnan
  • K. Praveen Karanth
Original Article


Sky Islands are high-elevation environments that are separated by warmer, low elevations, forming natural patches of unique montane habitat that often persist through changing climates. Peninsular India was ancestrally forested and has gradually become more arid since at least the Oligocene, and open landscapes have dominated since the middle-late Miocene. Mesic forests today are largely restricted to coastal mountains and some other montane habitats. A mitochondrial phylogeny and fossil-calibrated timetree of Indian Hemiphyllodactylus reveal an Indochinese origin and an endemic radiation with 12 species-level lineages, where a single species was known, that diversified in the Oligocene-Miocene across montane forest habitats in the Eastern Ghats and south India. The phylogeny also suggests the discontinuous Eastern Ghats mountain range encompasses two distinct biogeographic entities: north and south of the Pennar/Krishna-Godavari River basins. This study highlights the deep history of the region and the importance of montane habitats as islands of unique biodiversity that have persisted through millions of years of changing climates. We describe three new species: Hemiphyllodactylus arakuensis sp. nov., H. jnana sp. nov. and H. kolliensis sp. nov. from montane habitats above 1000 m. The montane habitats of these species are emerging hotspots of reptile endemism, and this study emphasizes the need for systematic biodiversity inventory across India to uncover basic patterns of diversity and distribution.


Biogeography Divergence dating Eastern Ghats Systematics Western Ghats 



We thank Pratyush P Mohapatra, Aparna Lajmi, R Chaitanya and the Evolutionary Ecology Lab (CES, IISc; Saunak Pal, SP Vijayakumar and Kartik Shanker) for contributing tissues used in this study. We also thank the Andhra Pradesh and Tamil Nadu Forest Departments for collection permits and hospitality and Tarun Khichi, Aniruddha Datta-Roy, V Deepak, R Chaitanya, MS Chaitra, R. Padmawathe and Nikhil Gaitonde for assistance in the field. Lee Grismer helped with r scripts and discussion on multivariate analyses, Maitreya Sil with Lagrange analyses, SR Ganesh with sampling locations in the Shevaroys, and Saunak Pal provided data on BNHS specimens. Joshua Muyiwa, Luis Ceriaco, and Shreya Yadav contributed to nomenclatural discussions. The two reviewers provided useful inputs.

Funding information

Partial funding came from the Ministry of Environment and Forests, the Department of Atomic Energy (2012/21/06/BRNS to Uma Ramakrishnan), the Department of Science and Technology (DST grant SR/SO/AS-57/2009 to Praveen Karanth), Government of India and National Science Foundation (USA) grant DEB 0844523 to Aaron M Bauer.

Compliance with ethical standards

Ethical approval

No live animals were used in experiments, and specimens were collected with permits from the Andhra Pradesh and Tamil Nadu forest Departments, besides from private land and other non-forest areas. Specimens collected for this study were euthanized with halothane, tissue vouchers stored in ethanol and whole specimens fixed with formalin. This study was approved by the National Centre for Biological Sciences, Bangalore (NCBS) animal ethics committee.

Supplementary material

13127_2019_392_MOESM1_ESM.xlsx (21 kb)
Table S1 Sequences used in this study. Museum abbreviations as follows: ABTC, Australian Biological Tissue Collection, South Australian Museum, Adelaide; ADS, Anslem de Silva field series; AMB, Aaron M. Bauer field series; AMS, Australian Museum; BPBM, Bernice P. Bishop Museum; BNHS, Bombay Natural History Society, Mumbai; CAS, California Academy of Sciences; CD, Charles Daugherty field series; CES G (Karanth lab field series) and CES L (Kartik Shanker lab field series), Centre for Ecological Sciences, Bangalore; CHUNB, Coleção Herpetológica da Universidade de Brasília; CJS, Christopher J. Schneider field series; CUMZ, Chulalongkorn University Museum of Zoology; DB, Don Buden; DJH, D. James Harris; EBG, Eli B. Greenbaum field series; FG/MV, Frank Glaw and Miguel Vences; FGZC, Frank Glaw; FLMNH, Florida Museum of Natural History; FT, frozen tissue collection of Victoria University of Wellington; GVH, Gerald V Haagner; IEBR, Institute of Ecology and Biological Resources (Hanoi); IRSNB, Institute des Sciences Naturelles du Belgique, Brussels; ITB, Institute of Tropical Biology Collection of Zoology; JB, Jon Boone; JEM, John E. Measey; KU, University of Kansas Natural History Museum, Lawrence; LJAMM, Luciano J. Avila and Mariana Morando; LLG, L. Lee Grismer field series; LSHUC, La Sierra University Herpetological Collection, L. Lee Grismer; MCZ, Museum of Comparative Zoology, Harvard University; MTSN, Trento Museum of Natural Sciences; MV, Museum of Victoria; MVZ, Museum of Vertebrate Zoology, Berkeley; MZUSP, Museu de Zoologia da Universidade de São Paulo; NCBS, National Centre for Biological Sciences, Bangalore; NJNUh, Nanjing Normal University, Jiangsu, China; NMZ, National Museum of Zimbabwe; PEM, Port Elizabeth Museum; PMNH, Pakistan Museum of Natural History; QM, Queensland Museum; RAH, Rod A. Hitchmough; RMB, Rafe M. Brown; ROM, Royal Ontario Museum; SAM, South Australian Museum; SC, Salvador Carranza; SYS, the Museum of Biology, Sun Yat-sen University (SYS), Guangzhou; TG, Tony Gamble; USNM, National Museum of Natural History, Smithsonian Institution; WBJ, W. Bryan Jennings; WDH, Wulf D. Haacke; YPM, Yale Peabody Museum, New Haven; ZCMV, Miguel Vences; ZMKU TM, Zoological Museum, Kasetsart University, Bangkok, Thailand; ZRC, Zoological Reference Collection, Raffles Museum; ZSM, Zoologische Staatssammlung München. (XLSX 21.2 kb)
13127_2019_392_MOESM2_ESM.xlsx (10 kb)
Table S2 Pairwise uncorrected ND2 sequence divergence across Indian Hemiphyllodactylus, numbers in bold along diagonal represent intraspecific diversity. (XLSX 9 kb)
13127_2019_392_MOESM3_ESM.pdf (258 kb)
Figure S1 Complete BEAST MCC tree from divergence dating analyses. (PDF 258 kb)
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Figure S2

Hemiphyllodactylus aurantiacus in life (adult male AK 245). (PNG 5854 kb)

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Figure S3

Hemiphyllodactylus jnana sp. nov. in life (adult male BNHS 1936). (PNG 5181 kb)

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High Resolution Image (TIF 6279 kb)
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Figure S4

Dorsal and ventral view of adult male holotype (AQ 191) of Hemiphyllodactylus jnana sp. nov. (PNG 8816 kb)

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High Resolution Image (TIF 14443 kb)
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Figure S5

Type series of Hemiphyllodactylus jnana sp. nov. (PNG 1417 kb)

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High Resolution Image (TIF 7195 kb)
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Figure S6

Hemiphyllodactylus kolliensis sp. nov. in life (from top to bottom: dorsal and ventral view of adult male holotype CES G138, dorsal view of adult female paratype AK 277). (PNG 7698 kb)

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High Resolution Image (TIF 9160 kb)
13127_2019_392_Fig9_ESM.png (11.5 mb)
Figure S7

Dorsal and ventral view of adult male holotype (CES G138) of Hemiphyllodactylus kolliensis sp. nov. (PNG 11794 kb)

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High Resolution Image (TIF 19231 kb)
13127_2019_392_Fig10_ESM.png (2.2 mb)
Figure S8

Type series of Hemiphyllodactylus kolliensis sp. nov. (PNG 2235 kb)

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High Resolution Image (TIF 6240 kb)
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Figure S9

Hemiphyllodactylus arakuensis sp. nov. in life (from top to bottom: dorsal view of adult male holotype CES G446, dorsal and ventral view of adult male paratype CES G068). (PNG 7907 kb)

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Figure S10

Dorsal and ventral view adult male holotype (CES G446) of Hemiphyllodactylus arakuensis sp. nov. (PNG 9519 kb)

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High Resolution Image (TIF 15777 kb)
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Figure S11

Type series of Hemiphyllodactylus arakuensis sp. nov. (PNG 1801 kb)

13127_2019_392_MOESM13_ESM.tif (9.5 mb)
High Resolution Image (TIF 9694 kb)


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Copyright information

© Gesellschaft für Biologische Systematik 2019

Authors and Affiliations

  1. 1.National Centre for Biological SciencesTata Institute of Fundamental ResearchBangaloreIndia
  2. 2.Centre for Ecological SciencesIndian Institute of ScienceBangaloreIndia
  3. 3.Foundation for Biodiversity ConservationNanded City, PuneIndia

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