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AMPHIBIANS OF THE WESTERN GHATSAMPHIBIANS OF THE WESTERN GHATSAMPHIBIANS OF THE WESTERN GHATSAMPHIBIANS 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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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. Semi-permeable 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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Pseudophilautus amboli

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).

Table 1: Endemic and Non-endemic amphibians of India and the Western Ghats.

Group Family Genus Species Endemic

(Non Endemic)

Anura (Frogs and Toads) 11 49 278

Caudata (Salamanders) 01 01 01

Gymnophiona (Caecilians) 02 04 33

India (total) 14 54 312

Anura (Frogs and Toads) 09 25 135 115 (20)

Caudata (Salamanders) 0 0 0 0

Gymnophiona (Caecilians) 02 04 23 23 (0)

Western Ghats (total) 11 29 158 138 (20)

Raorchestes akroparallagi 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 Species In India Species in the WG

Anura

Bufonidae 7 27 14

Dicroglossidae 12 58 24

Hylidae 1 1 0

Megophryidae 4 16 0

Micrixalidae 1 11 11

Microhylidae 7 23 13

Nasikabatrachidae 1 1 1

Nyctibatrachidae 1 16 16

Ranidae 6 30 4

Ranixalidae 1 10 10

Rhacophoridae 8 84 42

Caudata

Salamandridae 1 1 0

Gymnophiona

Caeciliidae 2 12 11

Ichthyophiidae 2 21 12

Total 54 311 158

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

Saidapur et al., 2009

Jumping Indirana beddomii Veeranagoudar et al., 2009

Kin recognition D. melanostictus, D. scaber, S.

Breviceps

Saidapur and Girish 2000 Eluvathingal et al., 2009 Gramapurohit et al., 2006

Food and foraging S. breviceps, Hylarana temporalis Sugur et al., 2008 Veeranagoudar et al., 2004a,b

Chemical cues In: S. breviceps To: H. tigerinus Sharma et al., 2008

Kinship and density on

growth

D. scaber, S. breviceps, H. temporalis,

D. melanostictus

Gramapurohit et al., 2008, 2004; Girish and Saidapur 2003, 1999; Saidapur and Girish 2001

Inter-population variation Clinotarsus curtipes Saidapur et al., 2003

Pattern of growth H. tigerinus, Euphlyctis cyanophlyctis,

C. curtipes, Polypedates maculatus

and D. melanostictus

Gramapurohit et al., 1998

Microhabitat choice H. tigerinus, E. cyanophlyctis, C.

curtipes, P. maculatus and D.

melanostictus, D. scaber, S. breviceps,

Nasikabatrachus sahyadrensis

Hiragond and Saidapur 2001 Raj et al., 2011

Pesticide Impact Limnonectus limnocharis*

Nyctibatrachus major

Gurushankara et al., 2006, 2007

Population structure N. major 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.

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.

Scale No of Species References

India 205; 311 Chanda 2002; Dinesh et al., 2010

Western Ghats 117; ~120; ~219; 158

Daniels 1992; Vasudevan et al., 2001; Gunawardane et al., 2007; Biju (Pers Comm.)

State

Karnataka 72 Daniels 2000

Kerala 117 Easa 2003; www.kerenvis.nic.in/biodiversity/fauna.htm

Maharashtra 43 Padhye and Ghate 2002

Goa 11 Abdulali and Sekar 1988

Gujarat 22 Vyas 2009

Protected areas

Dandeli Anshi Tiger Reserve 29 Gururaja and Ramachandra 2010

Kalakkad-Mundanthurai Tiger Reserve

32 Vasudevan et al., 2001

Sharavathi Wildlife Sanctuary 24 Ali et al., 2007

Parambikulam Wildlife Sanctuary 23 Radhakrishnan 1996

Periyar Tiger Reserve 14 Ravichandran and Pillai 1990

Kudremukh National Park 29 33

Dinesh and Radhakrishnan 2007 Krishnamurthy and Hussain 2000

Biligiri Rangaswamy Temple Wildlife Sanctuary

19 Ravichandran and Krishnan 2006

Kalakad Wildlife Sanctuary 29 Ravichandran 1996

Nagarahole National Park 14 Ganesh et al., 2002

Others

Wayanad 30 Abraham et al., 2001

Sringeri Taluk 35 Krishnamurthy and Shakuntala 1993

Silent Valley 19 Pillai 1986; Easa and Shaji 1999

Pune 11 31

Yazdani and Mahabal 1976 Padhye et al., 2000

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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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Live embryos of Nyctibatrachus petraeus

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 (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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Nasikabatrachus sahyadrensis

(Photo credits: Subramanian K.A.)

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 members of families 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 Indo-Madagascan 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 south-East 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 Indo-Asia 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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A moist deciduous Forest patch in the Western Ghats

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). 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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Fejervarya caperata without an eye

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

community Gururaja et al., 2008 Gururaja 2002

Roads Species and Population Decrease

Community Vijaykumar et al., 2002 Seshadri et al., 2009 Baskaran and Boominathan 2010

Disturbance (Fuel wood

extraction, timber,

organic much collection)

Species and Population Decrease, Change in community composition

N. major and N. aliciae

Ghate and Padhye 1996 Krishnamurthy 1996 Krishnamurthy 2003

Pesticides and Pollution Organ Malformation N. major,

L. limnocharis* Krishnamurthy et al., 2008

Land use-land cover

change

Species and Population Decrease, Change in community composition

Community 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 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

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 Stage Species References

Left eye Adult Sphaerotheca rufescens Nair and Kumar 2007

Brachydactyly and

polydactyly

Adult Nyctibatrachus major Vasudevan et al., 2006

Protruded mouth,

swollen body and head

Tadpoles Nyctibatrachus major Gurushankara et al., 2007

Swollen body and head Tadpoles Limnonectus limnocharis* Gurushankara et al., 2007

Brachydactyly Adult D. melanostictus Hippargi et al., 2010

Micro-ophthalmia Adult D. melanostictus Hippargi et al., 2010

Deformed limb Adult Ramanella Montana Seshadri Pers. Comm.

Limb and eye deformity Adult Clinotarsus curtipes

D. scaber

Nyctibatrachus petraeus

Hylarana temporalis

F. caperata

Minervarya syhadris

Seshadri Pers. Comm.

Gururaja Pers. Obs.

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

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.

19

Raorchestes ochlandrae

Colour variations in Pseudophilautus wynaadensis

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.

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. 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

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). Long-term 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.

21

Acknowledgments:

We thank Subramanian KA for providing photograph of Nasikabatrachus. We express our sincere

gratitude to Ms. Preeti for commenting on the manuscript.

22

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