High cryptic diversity of endemic Indirana frogs in the
Western Ghats biodiversity hotspot
A. Nair
1
, S. V. Gopalan
2
, S. George
2
, K. S. Kumar
2
, A. G. F. Teacher
1,3
& J. Merilä
1
1 Ecological Genetics Research Unit, Department of Biosciences, University of Helsinki, Helsinki, Finland
2 Chemical Biology, Rajiv Gandhi Centre for Biotechnology, Thiruvananthapuram, Kerala, India
3 Centre for Ecology and Conservation, University of Exeter, Cornwall Campus, Tremough, Penryn TR10 4BE, UK
Keywords
amphibia; Ranixalidae; Indirana; cryptic
diversity; biodiversity hotspot; Western
Ghats.
Correspondence
Abhilash Nair, Ecological Genetics Research
Unit, Department of Biosciences, University
of Helsinki, PO Box 65, Biocenter 3,
Viikinkaari, Helsinki FIN-00014, Finland.
Tel: +358919157807; Fax: +358 919157694
Email: abhilash.nair@helsinki.fi
Editor: Trent Garner
Associate Editor: James Austin
Received 6 November 2011; accepted 28
February 2012
doi:10.1111/j.1469-1795.2012.00539.x
Abstract
Amphibians are rapidly declining worldwide, but recent studies have shown that
their diversity may be heavily underestimated, and many new species have been
recently reported from biodiversity hotspots. For successful conservation and
management strategies to be implemented within such hotspots, a better under-
standing of the species diversity and their evolutionary relationships is required.
We used three mitochondrial (16S, 12S and CO1) and two nuclear (rag1 and
rhodopsin) gene fragments to investigate the genetic diversity within the endemic
Indirana genus from the Western Ghats biodiversity hotspot. The species diversity
within Indirana was found to be much higher than previously anticipated. Instead
of the expected six species within this region, our analyses identified 11 clades with
high (4.2–17.1%) sequence divergence. Each of these clades is likely to represent a
distinct species. Particularly pronounced polyphyly was found within Indirana
beddomii, which consisted of four highly supported monophyletic clades with high
genetic divergence. Similarly, Indirana diplosticta was divided into two highly
divergent monophyletic clades. We also report a new candidate species within the
genus from Vellarimala in Kerala, which we believe is yet to be described. Our
results suggest the existence of multiple unrecognized cryptic lineages within
Indirana, all of which are likely to have more narrow distribution ranges and lower
abundances than the taxonomic units into which they are currently assigned.
Hence, the International Union for Conservation of Nature (IUCN) Red List
statuses of Indirana frogs are likely to be in need of substantial revision, and
detailed genetic studies across the Western Ghats might uncover additional new
candidate species from this poorly studied endemic genus. Together with several
earlier amphibian studies describing cryptic species from tropics, our results high-
light the importance of proper species identification efforts before it is possible to
reliably determine the IUCN conservation status of tropical amphibians described
on the basis of morphological criteria.
Introduction
Species delimitation is a fundamental part of ecological and
evolutionary studies, and also plays an important role in
the development of conservation management programs.
The methodological problem of identifying the boundaries
between a set of species has recently attracted renewed
attention (de Quieroz, 2007; Wiens, 2007). Cryptic species,
which can be defined as two or more genetically distinct
species that are (mistakenly) classified as single species on
the basis of similar morphology, pose substantial taxonomic
challenges as they are hard to describe on the basis of mor-
phological or typological species concepts. The advent of
relatively inexpensive and rapid DNA sequencing, and
advances in molecular phylogenetic methods over the past
decades have given biologists new tools for identifying pre-
viously unrecognized cryptic species (Bickford et al., 2007).
Amphibian systematics and taxonomy provide several
examples of these and frequently highlight incongruence
between morphology-based taxonomies and genetic line-
ages (Hanken, 1999; Lemmon, Lemmon & Cannatella,
2007; Pauly, Piskurek & Shaffer, 2007). In recent years,
many new cryptic amphibian lineages have been described
from various biodiversity hotspots (e.g. Fouquet et al.,
2007; Vieites et al., 2009), indicating that the amphibian
diversity within these hotspots may be heavily under-
estimated. If so, it follows that allocation of accurate
International Union for Conservation of Nature (IUCN)
conservation statuses and development of sound conserva-
tion strategies in such regions is severely compromised.
Therefore, there is an urgent need for efforts to identify
cryptic species, especially given the increasing number of
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