1.
THEME PAPER ON
AMPHIBIANS OF THE WESTERN GHATS
N.A. Aravind* and K.V. Gururaja**
*SuriSehgal Centre for Biodiversity and Conservation
Ashoka Trust for Research in Ecology and the Environment (ATREE),
Royal Enclave, Sriramapura, Jakkur PO., Bangalore 560064
aravind@atree.org
**Centre for infrastructure, Sustainable Transportation and Urban Planning (CiSTUP)
Indian Institute of Science, Bangalore 560012
gururaj@cistup.iisc.ernet.in
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2.
1. Introduction
Amphibians are the first vertebrate animals to have two completely different life stages, one as tadpoles
that generally occur in water and other as adults dwelling more on land (Duellman and Trueb 1994).
Frogs, toads, caecilians and salamanders together constitute amphibians. Among the known 6639
species of amphibians in the world (Frost 2010), nearly one third are threatened with extinction (IUCN
2009; Hamer and McDonnell 2008). Amphibians are vital components of a healthy environment. Semipermeable skin, anamniotic eggs and biphasic life style make them particularly vulnerable to changes
and contamination of their habitats on land and in water. Presence of a good population of amphibians
in a region is indication of a healthy environment (Gururaja et al., 2008). Amphibians play major role in
ecosystem functioning, as prey and predator, especially as consumers of pest insects. They are also food
for snakes, birds and small mammals. They have great therapeutical value, be in development of
painkillers (Bufotonin, Epibatidine, etc.) or antimicrobials (refer to detailed review by Erspamer 1994) or
regenerating limbs (Tseng et al., 2010).
The amphibian fauna of India comprising of caecilians, frogs and toads and salamanders is extremely
diverse (Dinesh et al., 2009) and has high-level endemism in the Western Ghats and Eastern Himalayas
(Inger 1999; Biju and Bossuyt 2003; Roelants et al., 2004; Aravind et al., 2004). India harbours 311
species of amphibians belonging to all the extant orders, namely Caudata, Anura and Gymnophiona,
while the Western Ghats harbours 161 species of which 138 are frogs and toads (Anura) and 23 are
caecilians (Gymnophiona) belonging to 11 families and 30 genera. An overview of biodiversity and
ecology of hill streams of the Western Ghats has been reviewed by Subramanian (2010).
In this paper, we review the current status of amphibian research in the Western Ghats and assess the
gaps in our knowledge and provide solution for the conservation of amphibians in this global
biodiversity hotspot.
2. Species richness and Endemism
Richness and Endemism: Indian amphibian fauna belong to 3 orders, 14 families, 54 genera and 311
species (277- Anura, 33-Gymnophiona, 1-Caudata). The Western Ghats harbours 161 species of which
137 are Anurans (frogs and toads) and 24 are Gymnophiona (caecilians) belonging to 11 families and 30
genera (Table 1). The families Micrixalidae, Nasikabatrachidae and Ranixalidae are exclusive to the
Western Ghats represented by only one genus, whereas the Family Nyctibatrachidae is endemic to both
Western Ghats and Sri Lanka with genus Nyctibatrachus endemic to the Western Ghats and genus
Lankanectus to Sri Lanka.
The amphibians of the Western Ghats are very diverse and unique, with nearly 87 percent (138 species)
of the 158 species are endemic to this region, which is highest for any group of animal in the Western
Ghats. Most of the endemic species have restricted distribution, confined to the rainforests of the
Western Ghats. The anuran genera, which are endemic, include - Ghatophryne, Xanthophryne,
Micrixalus, Melanobatrachus, Ramanella, Nasikabatrachus, Nyctibatrachus, Indirana, Ghatixalus and
Raorchestes. Among gymnophiona, the endemic genera include Indotyphlus and Uraeotyphlus.
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Three families viz., Dicroglossidae, Rhacophoridae and
Nyctibatrachidae form 50 percent of the total species
richness in the Western Ghats (Table 2). The recent
discovery of new frog, Nasikabatrachus sahyadrensis
belongs to new family Nasikabatrachidae, is believed to be
a living fossil. This Gondwana relict frog is more closely
related to species found in the Seychelles Islands in the
Indian Ocean. The endemic torrent toads Ghatophryne
ornata and Ghatophryne rubigina are found only in hill
streams between Nilgiris and Kudremukh. The high level of
endemism in the Western Ghats might have caused
because of discontinuity in the chain of mountains which
limits the dispersal (Vasudevan et al., 2006), thus
promoting local endemism (Bossuyt et al., 2004), both in
terms of ‘out-of-India’ and ‘in-to-India’ hypothesis
(Bossuyt and Milinkovitch 2001; Bocxlaer et al., 2006).
Pseudophilautus amboli
Table 1: Endemic and Non-endemic amphibians of India and the Western Ghats.
Group
Family
Genus
Species
Anura (Frogs and Toads)
Caudata (Salamanders)
Gymnophiona (Caecilians)
India (total)
11
01
02
14
49
01
04
54
278
01
33
312
Anura (Frogs and Toads)
Caudata (Salamanders)
Gymnophiona (Caecilians)
Western Ghats (total)
09
0
02
11
25
0
04
29
135
0
23
158
Endemic
(Non Endemic)
Raorchestes akroparallagi
115 (20)
0
23 (0)
138 (20)
Rhacophorus malabaricus
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Table 2: List of families of amphibians reported from India and corresponding distribution of genera and
species in India and the Western Ghats.
Group/Family
Genera In India
Anura
Bufonidae
Dicroglossidae
Hylidae
Megophryidae
Micrixalidae
Microhylidae
Nasikabatrachidae
Nyctibatrachidae
Ranidae
Ranixalidae
Rhacophoridae
Caudata
Salamandridae
Gymnophiona
Caeciliidae
Ichthyophiidae
Total
Species In India
Species in the WG
7
12
1
4
1
7
1
1
6
1
8
27
58
1
16
11
23
1
16
30
10
84
14
24
0
0
11
13
1
16
4
10
42
1
1
0
2
2
54
12
21
311
11
12
158
Threatened species: According to 2006 IUCN red list assessment, 30 percent of the Western Ghats
amphibians are threatened, 4 percent is near threatened, 29 percent is data deficient and 22 percent is
least concern (Figure 1). The species Raorchestes travancoricus is assessed as extinct. Twenty-two
species of globally threatened species of amphibians found in the Western Ghats are highly restricted
and occurs only in one or two locations (Das et al., 2006). Since year 2000, over 50 new species have
been described from the Western Ghats, status of which has not been assessed yet. Highest number of
threatened species is found in the genera Raorchestes (11 species) and Nyctibatrachus (9 species). This
is mainly because of their specialised habitat requirements and restricted distribution.
Ghatophryne ornata
Pedostibes tuberculosus
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Figure 1: Number of species of amphibians in different IUCN categories.
Source: www.redlist.org
3. Discovery Patterns
Greater interest in amphibian taxonomy in last two decade has a tremendous impact on the discovery of
new species from the Western Ghats. Ever since 2000, 50 new species belonging to 40 anurans and 10
caecilians have been described. A recent analysis on temporal discovery patterns (Aravind et al., 2004,
2007; Gower et al., 2004; Dinesh et al., 2009) suggest that the discovery curve of amphibians of the
Western Ghats region is not yet saturated and there has always been a chance for new species to be
discovered. Further, most of those discovered are likely or invariably endemic. It reinforces that fact that
the discovery of an unusually high number of new species of amphibians from the Western Ghats
hotspot is rather not surprising (Aravind et al., 2004). Species discovery trend in Raorchestes,
Nyctibatrachus and Fejervarya, still shows there are many more species to be described from these
genera.
First species to be discovered from the Western Ghats dates back to 1799 by Schneider. On plotting
species discovery over time (Figure 2) for both India as well as for the Western Ghats, they exhibit three
spurs in the trend; first one around 1880, the second one in 1935 and the last one is after 2000. The
pattern exhibited in species discovery of India is more or less influenced by the species discovered from
the Western Ghats. The first one is because of G.A. Boulenger (1858-1937) followed by C.R.N .Rao
(1882-1960) (Figure 4) and the last one is by many authors, but significantly by S.D. Biju (1963-). Till
date, 73 authors have contributed to the discovery of species in the Western Ghats, either alone or with
others. Among these, 27 descriptions have single author.
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Figure 2: Temporal discovery patterns for amphibians of the Western Ghats.
Figure 4: Pioneers of Indian Batrachology: George A. Boulenger (1858-1937) and
C.R. Narayan Rao (1882-1960). Image courtesy: Wikipedia and Current Science
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4. Ecological studies
Ecological studies on amphibians of the Western Ghats are still in nascent stage, more so for the
tadpoles, looking at the number of publications against issues and sheer diversity the Western Ghats
have. Shifting from listing of species to looking at their habitat and distribution took very long time in
the Western Ghats. The earliest study emphasising the importance of community ecology of amphibians
date backs to 1987 by Inger et al. in Ponmudi area. A detailed review of ecological studies have been
categorised into varied heads below.
Tadpole Ecology: The early stage of an amphibian is a tadpole stage, be it in frogs, salamanders or in
caecilians. Tadpole survival needs maximum attention for the successful establishment of adult anuran
populations. From a purely aquatic to terrestrial mode during their development, studies on tadpoles
offers wide array of scientific opportunity to look into ecological, evolutionary and complex biological
processes that shapes their survival and growth and their transition from water to land. The success of
survival is dependent on habitat, morphological features, metamorphosis and ecological conditions and
varies within and between the species. The study of tadpole ecology is quite essential not only to know
the habitat requirement of tadpoles but also to formulate better conservation priority regions.
According to Saidapur (2001), studies on ecology of amphibian tadpoles in general, behavioural ecology
in particular, are in its infancy in India. Barring direct developing frogs (Genera Psuedophilautus and
Raorchestes with 32 species) as unique group with no free living tadpoles, all other anurans (129
species) and caecilians (23 species) of the Western Ghats, theoretically at least, must have tadpoles (106
species). Yet only 31 species of tadpoles
have been described, either partially or
fully so far from the Western Ghats, even
after 211 year of first species described
from the Indian peninsular (Annandale
1918; Hampson and Bennet 2002;
Saidapur 2001 and the literature there in;
Daniel 2002; Daniels 2005). Tadpole
research among caecilians is known from a
recent study on Gegeneophis seshachari,
which has viviparity (Gower et al., 2008)
and on developmental stages in
Ichthyophis cf. malabarensis (Jadav et al.,
2007). Among the direct developing
(endotrophic) frogs, which have evolved
independently in Philautus and Taylorana
group (Marmayou et al., 2000), description
of eggs, clutch size, habitat and stages of
development are described in 9 species
(Gururaja and Ramachandra 2006
references there in; Biju et al., 2010).
Summary of different studies on tadpole
ecology is given in the Table 3.
Tadpoles of Rhacophorus malabaricus
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Table 3: Ecological researches in the Western Ghats’ anuran tadpoles are about two decades old and
many of them have been carried out in laboratory conditions.
Theme
Species
Reference
Prey-predatory detection
Prey: Sphaerotheca breviceps,
Duttaphrynus melanostictus
Predator: Hoplobatrachus tigerinus
Indirana beddomii
D. melanostictus, D. scaber, S.
Breviceps
Saidapur et al., 2009
Jumping
Kin recognition
Food and foraging
S. breviceps, Hylarana temporalis
Chemical cues
Kinship and density on
growth
In: S. breviceps To: H. tigerinus
D. scaber, S. breviceps, H. temporalis,
D. melanostictus
Inter-population variation
Pattern of growth
Clinotarsus curtipes
H. tigerinus, Euphlyctis cyanophlyctis,
C. curtipes, Polypedates maculatus
and D. melanostictus
H. tigerinus, E. cyanophlyctis, C.
curtipes, P. maculatus and D.
melanostictus, D. scaber, S. breviceps,
Nasikabatrachus sahyadrensis
Limnonectus limnocharis*
Nyctibatrachus major
N. major
Microhabitat choice
Pesticide Impact
Population structure
Veeranagoudar et al., 2009
Saidapur and Girish 2000
Eluvathingal et al., 2009
Gramapurohit et al., 2006
Sugur et al., 2008
Veeranagoudar et al., 2004a,b
Sharma et al., 2008
Gramapurohit et al., 2008, 2004;
Girish and Saidapur 2003, 1999;
Saidapur and Girish 2001
Saidapur et al., 2003
Gramapurohit et al., 1998
Hiragond and Saidapur 2001
Raj et al., 2011
Gurushankara et al., 2006, 2007
Girish and Krishnamurthy 2009
Species richness: Studies have documented the species richness in different habitat type (Ganesh et al.,
2002), Myristica swamps (Chandran et al., 2010), protected areas (Gururaja 2002; Krishnamurthy 2003;
Vasudevan et al., 2001; Ganesh et al., 2002; Ali et al., 2007; Pillai 1986; Ravichandran and Pillai 1990;
Radhakrishnan 1996; Ravichandran 1996; Easa 1998; Abraham and Easa 1999; Cherian et al., 1999) or
for a region (Easa 1998, Ramachandra et al, 2008; Padhye and Ghate 2002; Daniels 2000; Table 4).
Majority of these studies concentrate on listing amphibians present in the region and hardly any
ecological information has been provided. It is interesting to note that studies in different hill ranges
have reported around 30–40 species with in a hill range in central and southern Western Ghats
(Krishnamoorthy, pers. commun. In Naniwadekar and Vasudevan 2007), Anamalai hills (Vasudevan et
al., unpublished data in: Naniwadekar and Vasudevan 2007) and Ashambu hills (Naniwadekar and
Vasudevan 2007). Comparison of Species richness in different habitat types or forest types suggests that
evergreen forest has highest diversity followed by moist deciduous and scrub forest and plantations
being least diverse (Ganesh et al., 2002).
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Table 4: Number of amphibian species reported in different study areas in the Western Ghats.
Scale
India
Western Ghats
State
Karnataka
Kerala
Maharashtra
Goa
Gujarat
Protected areas
Dandeli Anshi Tiger Reserve
Kalakkad-Mundanthurai
Tiger
Reserve
Sharavathi Wildlife Sanctuary
Parambikulam Wildlife Sanctuary
Periyar Tiger Reserve
Kudremukh National Park
Biligiri Rangaswamy Temple
Wildlife Sanctuary
Kalakad Wildlife Sanctuary
Nagarahole National Park
Others
Wayanad
Sringeri Taluk
Silent Valley
Pune
No of Species
205; 311
117; ~120;
~219; 158
72
117
References
Chanda 2002; Dinesh et al., 2010
Daniels 1992; Vasudevan et al., 2001;
Gunawardane et al., 2007;
Biju (Pers Comm.)
43
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22
Daniels 2000
Easa 2003;
www.kerenvis.nic.in/biodiversity/fauna.htm
Padhye and Ghate 2002
Abdulali and Sekar 1988
Vyas 2009
29
32
Gururaja and Ramachandra 2010
Vasudevan et al., 2001
24
23
14
29
33
19
Ali et al., 2007
Radhakrishnan 1996
Ravichandran and Pillai 1990
Dinesh and Radhakrishnan 2007
Krishnamurthy and Hussain 2000
Ravichandran and Krishnan 2006
29
14
Ravichandran 1996
Ganesh et al., 2002
30
35
19
11
31
Abraham et al., 2001
Krishnamurthy and Shakuntala 1993
Pillai 1986; Easa and Shaji 1999
Yazdani and Mahabal 1976
Padhye et al., 2000
Species turnover: A study by Vasudevan et al., (2006) show that there is species turnover and changes in
the abundance at local scale, from one to other drainage within a hill range. They showed that turnover
is highest between drainages separated by greater distances, such as those between two hill ranges
(between Ashambu hills and Anamalai hills; Vasudevan et al., (2006). Krishna et al., (2005) compared
species composition and abundance between streams of forests and adjoining cardamom and coffee
plantations. They found that there is significant variation in all three habitats. Another study by Ganesh
et al., (2002) shows that, similarity between dry deciduous and moist deciduous forest and between
moist deciduous and teak plantations is very low. But the similarity between dry deciduous and teak
plantation is high indicating that many species are common to both the habitat types and are generalist
species. Apart from these two studies there are hardly any work that looked at species turnover
between altitudinal and latitudinal gradients.
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Distributional patterns: Our knowledge on the latitudinal and altitudinal distributional patterns of
amphibians at the scale of the Western Ghats still relies majorly on the work by Daniels (1992) which
included 117 species. Since then, nearly 50 new species has been described, new localities and range
extension has been reported and also there has been splitting and lumping of many species. This calls
for a fresh analysis of distribution pattern in amphibians of Western Ghats, emphasising the importance
of ecological niches of species using Ecological niche modelling.
At level of the Western Ghats (regional scale), Southern half of the Western Ghats are richer than
northern half of the Ghats (Daniels 1992). The latitudinal division between 12-13°N has maximum
species richness and endemic species compared to other divisions. This is attributed to extended rainfall
regime and relative stable climatic condition. Along altitude, mid latitude range (800-1200m asl) is more
diverse than high altitudes (Daniels 1992; Naniwadekar and Vasudevan 2007). Among the habitats,
species those inhabiting wet evergreen forests shows patchy distribution (Daniels 1992).
At local scale, the community species richness (alpha level) increases linearly with increase in altitude,
while at the landscape level (gamma level) there are three peaks. Even at the local scale, the maximum
species richness was observed at the mid elevation (1200 m) for both the alpha and gamma levels
(Naniwadekar and Vasudevan 2007). This study also shown that with increase in altitude there is a
deceasing body size of frogs and number of endemic and unique species were greater at the elevations
above 700 m asl. The proportion of arboreal frogs increased from low to high elevation. Higher elevation
also have greater proportion of frogs those exhibits direct development (Genus: Pseudophilautus and
Raorchestes; Naniwadekar and Vasudevan 2007).
Habitat Association: Among different species of amphibians of the Western Ghats, D. melanostictus, E.
cyanophlyctis, H. tigerinus, F. caperata, Microhyla ornata, Polypedates maculatus, etc., are generalist
species, commonly found in and around human habitation and homestead. Species belonging to genera
Micrixalus are generally found in the slow flowing to fast flowing streams and also near waterfalls
(Gururaja 2010). All Nyctibatrachus spp. are confined to the streams most of the time, whereas,
Fejervarya spp. and Minervarya spp. are mostly found in lentic water bodies, Indirana is found along the
streams and in the forest floor. Raorchestes spp., Polypedates spp. and Rhacophorus spp. are generally
found on the bushes, shrubs and even at high canopy (arboreal and semi arboreal). Figure 5 shows
percent number of species preferring different habitat types in the Western Ghats. It is evident from the
graph that nearly 62 percent of the species are present in either aquatic or semi aquatic habitat.
Figure 5: Percent number of species present in different habits.
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Among host of environmental variables, high relative humidity, alkalinity, CO2 and high dissolved oxygen
(DO), under low air, water and soil temperature are the major influencing factors for the habitat
occupancy of Nyctibatrachus major and Micrixalus saxicola (Gururaja et al., 2003; Reddy et al., 2002;
Naniwadekar and Vasudevan 2007). Habitat parameters that influence the occupancy of amphibians are
rock cover, litter cover, litter depth and distance from water (Vasudevan et al., 2001; Gururaja 2002;
Krishnamurthy 2003). Any change in the habitat and environmental parameters will have negative
impact on the amphibian community. Krishnamurthy (2003) has shown that in Kudremukh National Park
the habitat with high disturbance has species poor community compared to undisturbed habitats.
Reduction in canopy cover increases light level and air and water temperature of the streams
discouraging the occurrence of adult frogs and tadpoles (Girish and Krishnamurthy 2009).
Nyctibatrachus major is very susceptible to changes in the land use around the habitat where they occur
(Krishnamurthy 1997).
Amphibians as indicator taxa: Amphibians are considered as ‘Canaries in a coal mine’. There are
debates on whether amphibians truly act as canaries or others species have more sensitivity than
amphibians (Kerby 2010). Despite setting aside whether they are canaries or not, they are still ‘in a
coalmine’. Recent studies in the Western Ghats also have emphasized that amphibians are the best
surrogates for conservation priority regions (Das et al., 2006) and Philautus is considered as an indicator
of fragmented forests for its patchy distribution (Gururaja et al., 2007). Impact of dam construction on
distribution of amphibians has been shown in Sharavathi river basin (Ramachandra et al., 2007).
5. Biological and Behavioural studies
Vocalization: Vocalization in anurans is mainly for the purpose of advertisement, but there are
circumstances when it is used for threat, territory and distress (Duellman and Trueb 1994). Although,
anuran communication system has more relevance in evolutionary significance of call production, sexual
selection, behavioural and reproductive ecology, visual cues and studying impact of urbanization (Roy
1997; Rodriguez-Prieto and Fernandez-Juricic 2005; Baker and Richardson, 2006; Bee and Swanson
2007; Lengagne 2008; Hamer and McDonnell 2008; Parris et al., 2009). Analysis of call pattern among
the anurans of the Western
Ghats are used predominantly
in the context of taxonomy and
systematics (Kanamadi 1996
and references there in; Roy
1996; Roy and Elepfandt 1993;
Kuramoto and Joshy 2001,
2006; Gururaja et al., 2007a,b;
Kuramoto et al., 2007; Joshy et
al., 2009). Krishna and Krishna
(2006) have analysed acoustic
pattern of Micrixalus saxicola,
a unique diurnal species
inhabiting torrential streams
with background noise and a
novel reproductive mode (Gururaja 2010).
Foot flagging in Micrixalus saxicola
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Visual communication: Apart from advertisement calls, amphibians are known to exhibit visual and
chemical cues. Micrixalus saxicola and M. fuscus of Micrixalidae, a family of 11 species of torrent frogs
are known to exhibit visual communication through foot flagging behaviour (Malhotra and Davis 1991;
Vasudevan 2001; Reddy et al., 2002; Krishna and Krishna 2006). Foot flagging behaviour is used as visual
communication, since these frogs inhabits torrential streams with high background noise. Similar visual
communication has evolved independently in three continents among different species (Walter Hödl
Pers. Comm.). Further study on the evolution and diversification of visual signalling will be a remarkable
research waiting to happen in the Western Ghats.
Breeding Behaviour and Natural History: Natural history studies on amphibians’ dates back to Vedic
period around 1000 BCE, where in 10 shlokas of Rig Veda (Mandala 7, Hymn 103) mention two species
of frogs, their behaviour, calling pattern and colour pattern. Scientific and systematic studies on
breeding behaviour of amphibians in the Western
Ghats are limited to very few species. Breeding activity
in Nasikabatrachus sahyadrensis is has been reported
recently (Raj et al., 2011), as an explosive breeder in
torrential streams. A new reproductive mode, 40 of its
kind is described from the Western Ghats recently. In
this reproductive more, M. saxicola, a torrential frog,
makes a cavity inside a stream and after keeping the
fertilized eggs, covers it with gravels and stones
(Gururaja 2010). Natural history in Nyctibatrachus
petraeus having pseudo or no amplexus was reported
by Kunte (2004). Biju (2009) reported novel nesting
behaviour in Rhacophorus lateralis, which folds leaf
over the egg mass to avoid desiccation. The nesting
behaviour of R. calcadensis, R. pseudomalabaricus and
R. malabaricus (Girish and Kanamadi 2000),
Xanthophryne tigerinus and Pedostibes tuberculosus
Live embryos of Nyctibatrachus petraeus
(Bocxlaer et al., 2009; Biju et al., 2009) was also
described recently. Mating and nesting behaviour was
observed in Polypedates maculatus (Girish and Saidapur 1999). Among the direct developing frogs,
reproductive behaviour and natural history is known in 9 species (Gururaja and Ramachandra 2006
references there in; Biju et al., 2010). Breeding and mating behaviour in Ramanella montana, which lays
eggs inside tree holes were observed by Krishna et al., (2004). Daniel (2002) and Daniels (2005) also
provided information on breeding and life history of amphibians of Western Ghats.
6. Origin of amphibians of the Western Ghats
Geological, ecological and evolutionary history of Western Ghats and associated adaptive radiations in
amphibians has shaped current amphibian distribution in the Western Ghats. First attempt in this
direction came from Bossuyt and Milinkovitch (2000). Remarkable similarities between Madagascar and
Asian ranids based on molecular phylogeny using nuclear and mitochondrial DNA and morphological
traits provided evidence on convergence of these two independent lineages. In addition, several adult
and larval characters allowed them invade similar adaptive zones despite being independent of one
other. This supported the ‘Out-of-India’ hypothesis, where the moving Indian sub-continent plate after
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splitting from Gondwana super continent carried Gondwanan life forms to Asia. Amphibians are one of
the best model organisms to test such hypothesis as they are poor dispersers and completely unable to
disperse over salt water. This is
further supported from fossil
records of ranids from peninsular
India (Roy and Karanth 2009 and
references there in). Bossuyt and
Milinkovitch (2001) demonstrated
that three lineages among the
Western
Ghats
amphibians,
namely members of frog families
Micrixalidae,
Nyctibatrachidae
(except of genus Lankanectus) and
Ranixalidae, are endemic to the
Western Ghats having not
dispersed to any other part of
Indian sub-continent and hence
considered Gondwan relicts. While
Nasikabatrachus sahyadrensis
members
of
families
(Photo credits: Subramanian K.A.)
Rhacophoridae,
Ranidae
and
Dicroglossidae,
which
have
dispersed out of India (after Indian subcontinent docked with Asian plate about 50 mya), are examples
of out- of-India hypothesis. This is again supported by the fossil evidence of ranidae in Eurasian land
mass dating back to 23-34mya. (Later on the discovery of a burrowing frog Nasikabatrachus
sahyadrensis (Biju and Bossuyt 2003; Dutta et al., 2004), which is not only a new species but a new
family, called Nasikabatrachidae, described from the Western Ghats, put strong evidence on IndoMadagascan linkage as the family was found closer to Sooglossidae from Seychelles Island than any
other family from the Western Ghats). Thus Nasikabatrachus sahyadrensis was resultant of vicariant
divergence from Sooglossids and a Gondwan relict of the Western Ghats (Roy and Karanth 2009).
Not only anurans, but members of apodans also supported ‘out-of-India’ hypothesis. Gegeneophis
ramaswami is the sister taxon of all Seychel caeciliids, which is a Gondwan relict in the Western Ghats
(Wilkinson et al., 2002). Gower et al., (2002) showed that among three families found in the Western
Ghats, Caeciliidae and Uraeotyphlidae are Gondwan relicts, while Ichtyophiidae dispersed up to southEast Asia. All the while, when biodiversity hotspot issue was raised, the Western Ghats and Sri Lanka
together constituted such hotspot. Bossuyt et al., (2004) showed that despite land mass connectivity
between peninsular India and Sri Lanka existed about 500,000 year ago, both Indian and Sri Lankan
fauna were quite distinct from each other. Such local endemism was overwhelmingly more in
amphibians (especially rhacophorids) of these two land masses. This eventually led to splitting of
Western Ghats-Sri Lankan hotspot into Western Ghats and Sri Lanka biogeographic regions.
At the time, when more evidences were garnered for out-of-India, Bocxlaer et al., (2009) provided
evidence on in-to-India hypothesis based on amphibians of the Western Ghats. Evolutionary history and
biogeography of true toad (Bufonidae) revealed that they colonized Indian subcontinent after the IndoAsia collision. Species belonging to genera Ghatophryne, Xanthophryne and Pedostibes have diversified
within the Western Ghats after their arrival from the south East Asia and thus enhancing species
diversity (Biju et al., 2009).
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14.
7. Threats
Habitat disturbance, loss and fragmentation: Habitat disturbance, loss and fragmentation are the most
important threat to amphibians in India. A vast majority of Indian amphibians occupy regions that are
increasingly being used for agricultural purposes. In addition to this, a vast majority of amphibian
species dwell in regions that are undergoing urban development, logging and industrialization that have
resulted in a drop in stable amphibian habitats. The building of dams and other barriers to stream and
rivers creates disturbance in the aquatic environments by altering the natural flow regime. Increased
human activities also alter light, temperature and moisture regimes (Krishnamurthy 1996). These
activities might gradually change the structure and composition of mid and understory vegetation,
resulting in more open canopy in the forest, possibly increasing fluctuations in habitat variables of
stream. Thus, these activities and modification hamper amphibian population structure and community.
Except for few species, most of the amphibians show patchiness in their distribution. This patchy and
restricted distribution makes them highly vulnerable to extinction, and has major implications in the
context of habitat fragmentation (Kumar et al., 2002). Several studies in the Western Ghats on
disturbance, loss and fragmentation of the habitat have shown to have negative impact on the
amphibian communities Krishnamurthy 2003; Gururaja et al., 2007; Gururaja 2002; Kumar et al., 2002;
Ghate and Padhye 1996). Other kinds of
disturbance such as roads passing through
the protected areas or next to aquatic habitat
or in the forest areas have a serious impact
on amphibian population and species
richness. A few studies have looked into
impact of these on amphibians. The study
indicated the high levels of amphibian
mortality (Seshadri et al., 2009; Vijaykumar et
al., 2001; Baskaran and Boominathan 2010).
Also, among all animals killed, the amphibians
contribute more than 50 percent of the total
animal mortality on road (Baskaran and
Boominathan 2010).
A moist deciduous Forest patch in the Western Ghats
Land-use and land cover change: There are very few studies that have actually looked at the impact of
land use and land cover changes on the amphibian communities. The study at Sharavathi, Aghanashini
and Bedti river basins in central Western Ghats shows that sub-catchments with lesser percentage of
forest, low canopy cover, higher amount of agricultural area, low rainfall have low species richness, less
endemic species and abundant non-endemic species. Whereas endemism, species richness and
abundance of endemic species are more in the sub-catchments with high tree density, endemic trees,
canopy cover, rainfall and lower amount of agriculture fields. This analysis aided in prioritising regions in
the Sharavathi river basin for further conservation measures using amphibians as the indicator taxa
(Gururaja et al., 2008). Ganesh et al., 2002 has shown that the teak plantations harbour significantly
lower number of species and individuals when compared to surrounding natural forests. A study done
on the effects of tea plantations on amphibians in Valparai has shown only 13 species of amphibians,
whereas 30 to 40 species were recorded in a relatively undisturbed forest (Daniels 2003).
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15.
Table 5: Summary of impact of different disturbance on the amphibian communities.
Disturbance type
Response
Species/Community
References
Fragmentation
Species and Population
Decrease,
Change in community
composition
Species and Population
Decrease
community
Gururaja et al., 2008
Gururaja 2002
Community
Vijaykumar et al., 2002
Seshadri et al., 2009
Baskaran and
Boominathan 2010
Ghate and Padhye 1996
Krishnamurthy 1996
Krishnamurthy 2003
Roads
Disturbance (Fuel wood Species and Population
extraction, timber,
Decrease,
organic much collection) Change in community
composition
Pesticides and Pollution Organ Malformation
Land use-land cover
change
Species and Population
Decrease,
Change in community
composition
N. major and N. aliciae
N. major,
L. limnocharis*
Community
Krishnamurthy et al.,
2008
Gururaja et al., 2008
Ganesh et al., 2002
Daniels 2003
Pollution, Pesticides and Malformation: Pollution
plays a major role in creating an unstable
environment for amphibians in India. Excessive
use of pesticides such as DDT, Dieldrin and
Malathion have been shown to affect the immune
system of certain amphibian species (Kleiner 2002)
while use of herbicides such as Atrazine has an
effect on their reproductive ability by inducing sex
reversal. Thirty different pesticides are used to
combat several species of arthropods, fungi and
plants that infest tea (Daniels 2003). Recent
studies have shown that insecticides such as DDT,
dieldrin and malathion affect the immune system
of frogs and herbicides. For example, atrazine in
Fejervarya caperata without an eye
quantities as minute as 0.1 parts per billion can
cause sex reversal. Ironically, the permitted residue of atrazine in India is 1000 times higher than what is
safe for an amphibian. Further studies need to be undertaken to substantiate the claim of a suspected
change in the amphibian population structure (Daniels 2003).
Of late, several reports have reported the presence of abnormalities or deformities in frogs from Indian
region (Kurulkar and Deshpande 1932; Nair and Kumar 2007; Mathew and Sen 2006). Field studies
indicate that the abnormalities/ deformities have become more prevalent in recent times (Mathew and
Sen 2006). Several factors which have been implicated as to be the causative agents for various
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16.
abnormalities/deformities in amphibians include UV radiation, cosmic rays, chemical contamination of
water, parasitic infestation and ground level ozone (Reaser and Johnson 1997; Nair and Kumar 2007).
In the laboratory condition, the exposure to malathion has decreased tadpole survival from 20 to 6
tadpoles and decrease in growth in Limnonectus limnocharis* (Indian cricket frog) (Gurushankara et al.,
2007, 2006). Mohanty-Hejmadi and Dutta (1981) found that exposure to malathion increases the time
required for completion of tadpole metamorphosis. The drastic decrease in growth, development, and
food consumption of tadpoles prevents them from reaching metamorphosis. This clearly depicts
physiologic changes in the tadpole as well as the possible threat to this species in the agroecosystems of
the Western Ghats (Gurushankara et al., 2007). Another study looked at nitrate poisoning in the
tadpoles of two ranid frogs (Nyctibatrachus major and Limnonectus limnocharis*) again studied under
laboratory conditions. The tadpoles of N. major showed protruded mouth, swollen body and head,
paralysis, lateral and vertical swimming, restlessness, and wriggling movement. On the other hand,
tadpoles of L. Limnocharis* showed swollen body and head, bent tail, paralysis, restlessness,
depigmentation and intestinal haemorrhage. In both species, abnormalities increased with the increase
in nitrate concentration (Krishnamurthy et al., 2008)
What is more worrying is the recent report of malformation of frogs in the undisturbed habitats. Recent
study in Kudremukh National Park (Vasudevan et al., 2006) has shown very high levels of deformities in
frogs. They have recorded deformities in 58 individuals belonging to 9 species with 12 different types.
Among them, brachydactyly and polydactyly are most common deformities. The parts which is most
affected are limbs and eyes (Vasudevan et al., 2006; Hippargi et al., 2010). The studies have also shown
that the deformities also occur in advanced tadpole stage. Interesting to note that, in the study by
Vasudevan et al., (2006), more than 50 percent of the species that have malformation are from the
Genus, Nyctibatrachus spp. The malformation in the population ranges from 12 to 55 percent. This
clearly shows that the endemic and less dispersed frog species are more prone to malformation than
non-endemic species (Vasudevan et al., 2006). The question as to what caused deformation in absence
of human interference remains unanswered and needs to be studied further.
Table 6: Different types of malformation in frogs reported in different studies in the Western Ghats
Malformed organ
Left eye
Brachydactyly and
polydactyly
Protruded mouth,
swollen body and head
Swollen body and head
Brachydactyly
Micro-ophthalmia
Deformed limb
Limb and eye deformity
Stage
Adult
Adult
Species
Sphaerotheca rufescens
Nyctibatrachus major
References
Nair and Kumar 2007
Vasudevan et al., 2006
Tadpoles
Nyctibatrachus major
Gurushankara et al., 2007
Tadpoles
Adult
Adult
Adult
Adult
Limnonectus limnocharis*
D. melanostictus
D. melanostictus
Ramanella Montana
Clinotarsus curtipes
D. scaber
Nyctibatrachus petraeus
Hylarana temporalis
F. caperata
Minervarya syhadris
Gurushankara et al., 2007
Hippargi et al., 2010
Hippargi et al., 2010
Seshadri Pers. Comm.
Seshadri Pers. Comm.
Gururaja Pers. Obs.
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17.
Invasive species: Invasive species are another greatest threat to native biodiversity. Introduction of
exotic fishes like Gambusia and Tilapia in water bodies (for human consumption), which serve as
amphibian breeding sites, has also wiped out amphibian populations (Ghate and Padhye 1996). Tilapia
has shown to be voracious feeders, hence it might have an impact on the amphibian population. No
studies have been carried out to quantify the possible spread and impact of these alien species on
amphibians.
Chytrid fungus infection: Globally, one of the major reasons for amphibian decline is Chytridiomycosis
fungus infection. However, there are no official confirmed reports of fungal infection from Indian region.
This does not rule out the possibility of total absence of this threat from this region. However, for the
first time in India, a fungal infected frog individual of Fejervarya caperata from Dandeli is recorded
(Gururaja, Patil and Cunningham, Under preparation). It has been identified as fungal infection through
microtome sections. We were not able to identify the fungi, either through culture or through molecular
analysis. Intensive and more exhaustive study needs to be done to ascertain the presence of infectious
fungal pathogens.
Figure 5: Fejervarya caperata with the lesion on the body (left) and section of the lesion showing fungal
spores (Photo credits: Gururaja K.V. and Rajashekhar K. Patil).
Frog leg trade: Our knowledge on the magnitude of trade on frog leg from India dates back to 1981
(Abdulali 1981). India's trade in frog leg was highest in 1981, during that year India exported 4,368 tons
of frog limbs, earning about 9.3 million USD. In spite of the ban on trade of frog legs, it has not been
particularly effective. Hoplobatrachus tigerinus and Euphlyctis hexadactylus are among the main frogs
traded in India and their biology, notably breeding biology, needs to be studied. The quantum of illegal
trade and local consumption of frog leg needs to be documented.
8. Gap Analysis
Region of high diversity and endemism: Amphibians within the Western Ghats hotspot shows very high
level of point endemism or local endemism especially in the Genera Raorchestes and Nyctibatrachus.
Compared to other bio-geographic regions of India, the Western Ghats harbour high diversity for
amphibians. There are several regions with in the Western Ghats hotspots which sustains high diversity
and endemism. Within the central Western Ghats, Gundia-Hassan-Kottigehara region from where C.R.N.
Rao had reported 17 new species in 1937 (Rao 1937) and from Waynad, where recent discoveries of
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18.
several species of Raorchestes sp. and others have been described. In Southern Western Ghats,
southern most regions of the Western Ghats comprising KMTR, Ponmudi and surrounding areas are
exceptionally rich in endemic and restricted range species such as D. beddomi, N. vasanthi, Rhacophorus
calcadensis, etc.
Ecologically sensitive regions in the Western Ghats: The study by Das et al., (2006) has shown that
existing protected areas fail to represent endemic and threatened amphibians of the Western Ghats.
However, there is no exhaustive study on the distribution of amphibians in the Western Ghats. Unless
we survey entire extent of the Western Ghats and distribution in different habitat types it is not possible
to delineate the ecologically sensitive regions keeping amphibians in mind. First and foremost thing we
need to do is to assess the geographical distribution patterns of amphibians of the Western Ghats
incorporating extensive field data. Earlier, Daniels (1992) have assessed this, but since then more than
50 species have been described, several species has been synonymised and many local and regional
level distribution data are available. Hence, we need to freshly look at it and come up with the
ecologically sensitive region with the Western Ghats hotspot. Use of ecological niche modelling
techniques in understanding the geographical distribution of amphibians in the Western Ghats can be
thought of.
Natural history, Ecology and Biology: What we know is much less than what we don’t know about
natural history, ecology and biology of amphibians of the Western Ghats. The life history, microhabitat
preference, and the factors affecting the distribution of most the Indian amphibian species are not
known. Many studies on amphibian population occur most commonly at single or few sites that too in
protected areas. Studies on population genetic structure of the fragmented population needs to be
carried out for better conservation of those which are cryptic and threatened species. Tadpole ecology
of most species is not known. We have very little or no data on the community ecology of the
amphibians. Studies also focus on the seasonal changes in the population, community structure, niche
overlap, etc. We strongly recommend research should occur often at the larger landscape level and
conservation efforts should focus on suitable habitat.
Taxonomic problems: Taxonomy in the present age is considered as an out dated subject (Daniels 1999),
yet it is fundamental to all ecological studies. The consequences of bad taxonomy in ecological research
have cascading effect which is neither quantified nor scrutinized (Bortolus 2008). The funding for
taxonomic research in India and elsewhere is very meagre. There is an urgent need to train researchers
on taxonomy, specifically on amphibians. At the same time, taxonomy and ecology must integrate to
have better scientific understanding of the organism. The importance for ecological and
biotechnological research has a great bearing on the taxonomic research among Indian students and in
the universities and institutions. Since many of the known amphibians had been described during the
pre-independence period, nearly 49 percent of type specimens are housed in different
museums/depositories outside India (Dinesh et al., 2009), which is not easily accessible for majority of
Indian researchers. The old Indian collections which are deposited in some of the national museums are
in very bad state or totally lost. This non-availability of type specimen and bad curation of the specimens
has further aggravated the taxonomic studies in India. So, what we need is a good state of art national
repository of type specimens in our country which can aid taxonomic research.
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19.
Raorchestes ochlandrae
The taxonomic status of many species, of the
genus
Philautus,
Micrixalus
and
Nyctibatrachus and of Ranids is even now
unresolved (Inger 1996). Members of highly
specious genus Raorchestes are also highly
variable in terms of colour (Inger et al., 1984;
Bossuyt and Dubois 2001) and patterns on the
body. This group is particularly difficult for
identification not only for taxonomists but
also for non-taxonomists and field
researchers. There are recent attempts to
resolve taxonomic issues in the country
mainly by the efforts of S.D.Biju from Delhi
University. But still lot more needs to be done.
Good field keys can solve the problem
partially if not completely.
The available field guides on amphibians of
Western Ghats (Chanda, 2002; Daniel,
2002; Daniels, 2005; Ganesh et al, 2009;
Bhatta 1998; Ravichandran and Pillai 1999)
require substantial revision and more
refined field identification keys. Dynamic
web based database on the available
species, literature, distribution and images
would go long way in conservation and
management efforts. Few of them started
the
initiative
(http://wgbis.ces.iisc.ernet.in/biodiversity/
database_new/?q=node/148). As of now
they are either static (both in terms of
query as well as update) or cease to exist.
Colour variations in Pseudophilautus wynaadensis
Studies outside Protected areas: Less than 5 percent of the total forest area in the Western Ghats is
under protected area network (PA). There are many studies on amphibian ecology and conservation
which are concentrated around PAs such as Kalakkad-Mundanthurai Tiger Reserve (KMTR), Sharavathi
WLS, Nagarahole NP, Silent Valley NP, Dandeli WLS, Kudremukh NP, etc. (Krishnamurthy 2003;
Vasudevan et al., 2001; Ganesh et al., 2002; Gururaja et al., 2008; Pillai 1986; Ravichandran and Pillai
1990; Radhakrishnan 1996; Ravichandran 1996; Easa 1998; Abraham and Easa 1999; Cherian et al.,
1999). Considerable amount of amphibian biodiversity do exist outside PAs. To reinforce this fact,
recently, two new species of Philautus - P. anili and P.dubois has been discovered from the urban areas
in the Western Ghats (Biju and Bossuyt 2006), P. ochlandrae in the reserve forest (Gururaja et al., 2007),
and Nyctibatrachus dattatreyaensis in high altitude stream (Dinesh et al., 2008). Hence, future research
in terms of inventorying and monitoring amphibian diversity should concentrate more outside the PA
network as these areas are more vulnerable to disturbance from the anthropogenic activities.
Long-term monitoring: In India, we do not have a national long-term monitoring program for any
groups. The need of the hour is to have national program on long-term monitoring program for studying
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20.
amphibian communities. The program should be based on statistically sensitive monitoring approach as
suggested by Storfe (2003). Studies have noted that populations of most of the amphibians generally
fluctuate and also the population tend to decline between years than they increase (Storfe 2003). Longterm monitoring program will be able to address cause and consequence of amphibian population
fluctuation in the wake of climate change and increased human disturbance on to its habitats. Here, in
this context we propose a national program on “Monitoring Amphibian Populations” (MAP). This
initiative should be a mandate of the central government. This monitoring program should not only
include scientific community but also amateurs, kids and lay public to make this program more inclusive
rather than being confined to scientific community. Long-term monitoring of frogs using automated
sound recording- non-invasive method has to be tried out (Seshadri Pers. Comm).
Innovative conservation measures: In some agricultural regions, natural wetlands are scarce, and
constructed agricultural ponds may represent important alternative breeding habitats for amphibians.
Properly managed, these agricultural ponds may effectively increase the total amount of breeding
habitat and help to sustain populations (Knutson et al., 2004). Batracharium, a unique concept for
conservation of amphibians, where in few localities with high species richness are identified, habitats
are marked and provisions are made for ‘frog watch’ with sign boards indicating the species to look for,
kind of habitat and regular monitoring programmes are scheduled.
Popularizing amphibian conservation: Awareness and education is the key for any successful
biodiversity conservation program. This is especially true in Indian scenario. In case of amphibians, of
late there has been some effort in creating awareness and education among various sections of the
society and stakeholders. But this effort has to be magnified for better conservation. The major hurdle
in popularising amphibian watching is the lack of good illustrated/ photographic guide of amphibians
unlike birds or butterflies. Hence, efforts are required to produce the coloured guide that can be used by
non-scientific community. Earlier attempts in making amphibian research popular was made by Daniel
(1963, 1975), Daniel and Sekar (1989), Daniels (1997a,b,c) and for frogs by Pillai and Ravichandran
(1999) for caecilians. Recently, Ranjit Daniels’ (2004) book on Amphibians of Peninsular India helped in
creating interest among many non-scientific communities. However, we need better guide than existing
ones with good photographs, distribution maps and natural history information.
With the advent of web technology the communication has become easier and can be effectively used
for
awareness
and
education
program.
The
web-based
discussion
group,
AmphibianIndia@yahoogroups.com is in this direction. This group also conducts AmphibianIndia Meet
to teach, educate and create awareness regarding amphibians of India. A new Initiative by Delhi
University along with host of National and international partners has set foot in search of those species
that are thought to be extinct. The initiative named “Lost! Amphibians of India” has listed 48 species
from India of which, 28 are from the Western Ghats. The initiative is soliciting volunteers and amphibian
researchers from India to join this program in search of so-called extinct species of amphibians. This
program has received great response from the scientists and public alike. The year 2008 was declared as
‘The Year of the Frog’, by the Amphibian Ark - a joint effort of the World Association of Zoos and
Aquariums (WAZA), the IUCN/SSC Conservation Breeding Specialist Group (CBSG) and the IUCN/SSC
Amphibian Specialist Group (ASG) to create awareness and to understand global amphibian extinction
crisis and conserve global amphibians. We need more such innovative approaches locally for educating
public.
*Note: Existence of Limnonectus limnocharis in the Western Ghats is questionable (Kuramoto et al., 2007), however, we
retained its name as mentioned in the publication till further confirmation is done.
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21.
Acknowledgments:
We thank Subramanian KA for providing photograph of Nasikabatrachus. We express our sincere
gratitude to Ms. Preeti for commenting on the manuscript.
21

22.
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