JoTT R E V I E W 4(11): 3029–3037

Land snaffs (Moffusca: ) of Indfa: status, threats and conservatfon strategfes

Sandeep Sen 1 , G. Ravfkanth 2 & N.A. Aravfnd 3

1,2,3 Surf Sehgaf Centre for Bfodfversfty and Conservatfon, Ashoka Trust for Research fn Ecofogy and the Envfronment (ATREE), Royaf Encfave, Srframpura, Jakkur PO, Bengafuru, 560064, Indfa Emaff: 1 [email protected], 2 [email protected], 3 [email protected] (correspondfng author)

Date of pubffcatfon (onffne): 26 September 2012 Abstract: Land snaffs form an fmportant component fn the forest ecosystem. In terms Date of pubffcatfon (prfnt): 26 September 2012 of number of specfes, the phyfum Moffusca, to whfch fand snaffs befong, fs the fargest ISSN 0974-7907 (onffne) | 0974-7893 (prfnt) phyfum after Arthropoda. Moffusca provfde unfque ecosystem servfces fncfudfng recycffng of nutrfents and they provfde a prey base for smaff mammafs, bfrds, snakes Edftor: Fred Naggs and other reptffes. However, fand snaffs have the fargest number of documented Manuscrfpt detaffs: extfnctfons, compared to any other taxa. Tfff date 1,129 specfes of fand snaffs are Ms # o2722 recorded from Indfan terrftory. But onfy basfc fnformatfon fs known about thefr taxonomy Recefved 03 March 2011 and ffttfe fs known of thefr popufatfon bfofogy, ecofogy and thefr conservatfon status. In Ffnaf recefved 18 Jufy 2012 thfs paper, we brfefy revfew status, threats and conservatfon strategfes of fand snaffs Ffnaffy accepted 24 August 2012 of Indfa. Cftatfon: Sen, S., G. Ravfkanth & N.A. Aravfnd (2012). Land snaffs (Moffusca: Gastropoda) of Keywords: Bfodfversfty, conservatfon, fand snaffs, taxonomfc bfas, refntroductfon, Indfa: status, threats and conservatfon strategfes . Journaf of Threatened Taxa 4(11): 3029–3037.

Copyrfght: © Sandeep Sen, G. Ravfkanth & N.A. Aravfnd 2012. Creatfve Commons Attrfbutfon INTRODUCT ION 3.0 Unported Lfcense. JoTT affows unrestrfcted use of thfs artfcfe fn any medfum for non-proft purposes, reproductfon and dfstrfbutfon by provfdfng adequate credft to the authors and the The tropfcs have faced massfve bfodfversfty foss due to fntensfve source of pubffcatfon. anthropogenfc actfvftfes such as changes fn fand use and degradatfon of Author Detaffs: See end of thfs artfcfe envfronment. Recent reports suggest that the tropfcs are fosfng bfodfversfty

Author Contrfbutfon: Aff authors have at an afarmfng rate (Sodhf 2008). Much of thfs bfodfversfty foss has contrfbuted equaffy to thfs manuscrfpt been reported for vertebrates and pfants. However, there fs very ffttfe Acknowfedgements: The authors greatfy knowfedge on the extent of foss fn fesser known groups, especfaffy the acknowfedge the Darwfn Inftfatfve vfa DEFRA, Government of UK for fundfng. We are afso fnvertebrates. In thfs paper, we hfghffght the fmportance of fand snaffs gratefuf to Drs. T. Ganesh, Soubadra Devy and the need for thefr conservatfon. and K.V. Gururaja for crftfcaf comments on the earffer versfon of the manuscrfpt. We afso Land snaffs fncfude severaf dfstfnct ffneages of terrestrfaf gastropods thank two anonymous revfewers for thefr crftfcaf comments that greatfy hefped fn fmprovfng the and befong to the second fargest phyfum after arthropods fn terms of manuscrfpt. number of specfes wfth more than one fakh descrfbed specfes (Lydeard et af. 2004). Land snaffs constftute about sfx per cent of the totaf specfes on Earth (Cfark & May 2002). A farge part of moffuscan fauna fn many tropfcaf regfons of the worfd fs stfff poorfy known. They form an fmportant component of the forest ecosystem by recycffng nutrfents (Gravefand et af. 1994; Dunk et af. 2004) and are the prey base for a number of smaff mammafs, bfrds, reptffes, amphfbfans and other fnvertebrates, fncfudfng carnfvorous snaffs (Deepak et af. 2010). In cafcfum poor habftats fand snaffs can form an fmportant source of cafcfum for other anfmafs. Land snaffs afso serve as an fndfcator of ecofogfcaf condftfons, and are very sensftfve to cffmatfc and ecofogfcaf change (Shfmek 1930; Sfmone 1999; Čejka & Hamerfík 2009 ). Thus, they are usefuf for reconstructfng past envfronments (Bar-Yosef Mayer 2002; Gümüş 2009). As earfy as 1839 Charfes Darwfn showed the vafue of fand snaffs fn studyfng

OPEN ACCESS | FREE DOWNLOAD envfronmentaf change (Naggs et af. 2006). Terrestrfaf snaffs prove to be

Journaf of Threatened Taxa | www.threatenedtaxa.org | September 2012 | 4(11): 3029–3037 3029 Land snails of S. Sen et al. valuable research subjects for studies in evolutionary These malacological pioneers laid the foundation of biology, biogeography, phylogeography, biodiversity, our knowledge on the taxonomy and distribution of ecology and conservation biology (Schilthuizen et Indian land snails. Following this period of intensive al. 2007; Davison et al. 2008; Richards & Davison study, there was a drastic decline in studies on Indian 2010). With their generally low dispersal powers, land snails. More recent studies in India, have mainly land snails tend to exhibit conservative distribution concentrated on inventorying regional snail faunas patterns, making them valuable subjects in studying (like state or protected areas) and less on species historical biogeography (Solem 1984; Naggs & description, ecology and conservation (Aravind et Raheem 2005; Wade et al. 2006). Highly diverse and al. 2010). While globally, there has been a renewed narrowly distributed, land snails are good indicators interest in land snail research, in India the research has of areas of conservation importance and endemicity truly been at a snail’s pace (Aravind et al. 2005, 2008, when compared to widely distributed groups such as 2010; Aravind & Naggs 2012). Little information is vertebrates (Moritz et al. 2001). available on species limits, distribution ranges and The distribution and activity of land snails depends patterns of diversity. Recent analysis of Indian land on several factors including precipitation, soil pH, soil and freshwater molluscan literature has confirmed Ca content, canopy density, etc. Calcium availability that that there are hardly any studies on the ecology in the soil is a major limiting factor for their survival and conservation of Indian land snails compared to as it is required for their shell formation. Several the wide range of historical literature available on studies have shown that Ca is positively correlated taxonomy (Aravind et al. 2010). There are no studies with species richness and density (Burch 1955; Hotopp on the population status, phylogeny and taxonomic 2002; Aravind 2005). However, in the regions such as revision of different families or genera of Indian land the Western Ghats, where the soil is usually acidic the snails. snail richness is usually high but abundance is low. The past two decades have seen a large number of Species diversity and rarity in land snails studies highlighting the need for mollusc conservation Globally, nearly 35,000 species of land snails have globally (Bouchet 1992; Ponder 1997; Herbert 1998; been described and there may be 30,000 to 60,000 Killeen & Seddon 2004; Budha 2005; Solymos additional species yet to be described (Lydeard et & Feher 2005; Régnier et al. 2009). Killeen & al. 2004). Within modern India’s boundaries 1129 Seddon (2004) notably edited a volume with global species belonging to 140 genera and 26 families of coverage on molluscan biodiversity and conservation, land snails have been recorded (Ramakrishna et al. highlighting the importance of molluscan ecology 2010). The Western Ghats hotspot has 270 species of and conservation. However, very little information land snails of which 76% are endemic to this region is available on the status and threats of land snails (Aravind 2005) and 40% are micro-gastropods (i.e. in India. Here, we review current status of ecology, <5mm on greatest dimension) (Aravind et al. 2008). conservation and threats to land snails with particular Unlike most other systematic groups many land snail reference to India and discuss the strategies required species have restricted range distributions with some for conserving this important group. endangered species having a range of less than 5km2 and many endemic species having ranges less than Early studies on land snails in India 10km2 (Cameron 1998; Dunk et al. 2004). According Indian malacology was pioneered by William to Solem (1984) nearly half of all terrestrial molluscs Henry Benson (1803–1870), who contributed have a species range of less than 100km2. Within the significantly to our knowledge on Indian land snails Western Ghats, species distributed in the southern in the mid 19th century (Naggs 1997). The Blanford region are absent in the northern region. Further, there bothers-William and Henry, H. Theobald, L. Pfeiffer, is very little overlap between the southern and central, G.K. Gude, H.H. Godwin-Austen and R. Beddome, and central and northern regions (Table 1; Aravind led Indian land snail research until the early 20th 2005). Nearly 75% of land snails from the Western century, but Gude and Pfeiffer’s research was based Ghats have been reported from less than three sites entirely on museum material as neither visited India. (Fig. 1). This data clearly indicates how vulnerable this

3030 Journal of Threatened Taxa | www.threatenedtaxa.org | September 2012 | 4(11): 3029–3037 Land snaffs of Indfa S. Sen et af.

Tabfe 1. Percent specfes shared between dffferent regfons 40 of the Western Ghats (Dfvfsfon of the Western Ghats fs 35 based on Aravfnd et af. 2005) 30 Regfon South Centraf North 25 South 100 20 E-17.19 Centraf 100 NE-11.11 15 Percent specfes Percent E-0.00 E-1.56 10 North 100 NE-2.78 NE-1.39 5

0 1 2 3 4 5 >5 Number of focatfons group fs to any smaff scafe change fn the ecosystem. Ffgure 1. Dfstrfbutfon pattern of fand snaffs fn the Western Ghats The sftuatfon fn other regfons of Indfa fs afso cause for concern. Northeastern Indfa harbours a rfch mfxture of Indfan and Burmese/Mafayan snaff groups resuftfng between forest patches (Aravfnd 2005; Raheem et af. fn the hfghest specfes dfversfty fn thfs regfon, but there 2008). Cffmate change fs consfdered to be a threat are hardfy any studfes on thefr dfstrfbutfon and threats. to many specfes (Thomas et af. 2004), and fand snaffs Our fnformatfon on the fand snaffs of northeastern are partfcufarfy vufnerabfe (Pounds & Crump 1994; Indfa, fs basfcaffy from the Fauna of Brftfsh Indfa and Pounds et af. 1999; Sternberg 2000; Bezemer & Knfght a few Zoofogfcaf Survey of Indfa reports. Conversefy, 2001; Gerfach 2007). Changes fn rafnfaff patterns though fess rfch, the fand snaff fauna of the Western and fuctuatfon fn soff temperature coufd fead to the Ghats fs better known when compared to other regfons death of juvenffe snaffs and fmpafr mobfffty across a of Indfa (Rao 1924; Sathyamaurthf 1960; Subbarao & fragmented mosafc of naturaf and transformed habftats Mftra 1979; Ramakrfshna & Mftra 2002 ; Madhyastha fn response to cffmate change. Extfnctfon of fand snaff et af. 2004; Mavfnkurve et af. 2004a,b; 2005; Aravfnd specfes due to change fn cffmatfc condftfons such as a 2005; Aravfnd et af. 2005, 2008; Rajashekhar & decrease fn rafnfaff and gfobaf warmfng has afso been Aravfnd 2012). reported (Baur & Baur 1993; Gerfach 2007). In Indfa, we do not have any fnformatfon on potentfaf fmpact of Threats cffmate change on fand snaffs. The major threats to the natfve fand snaff fauna A vast extent of the Western Ghats fs covered fn fncfude habftat foss and fragmentatfon as a resuft of pfantatfons such as tea, coffee, areca, rubber etc. anthropogenfc actfvftfes such as fntense fand use, A wfde varfety of pestfcfdes/herbfcfdes are used to constructfon of roads, dams, pfantatfons, poffutfon and controf severaf specfes of arthropods, fungf and pfants the spread of fnvasfve specfes (Aravfnd 2005; Aravfnd that fnfest these pfantatfon crops (Dfptf & Vefho 2007). et af. 2005) whfch reduce dfversfty and change the The fmpact of these chemfcafs on the focaf fora and communfty structure of fand snaffs (Aravfnd 2005; fauna fncfudfng endemfc fand snaffs as weff as other Rajashekhar & Aravfnd 2012). Between 1973 and fnvertebrates fs unknown. Forest fres (even surface 1995 the southern Western Ghats fost nearfy 25 percent fres) and reductfon fn vegetatfon cover are afso major of forest cover ( Jha et af. 2000). Thfs regfon of the threats to fand snaffs. Ghats harbours hfgh specfes dfversfty and endemfsm fn fand snaffs (Aravfnd 2005) . Poor dfspersaf and Negfected taxa fn conservatfon smaff dfstrfbutfon ranges of many fand snaffs have The scarcfty of mafacofogfsts fn Indfa has had a undoubtedfy contrfbuted to hfgh fevefs of extfnctfon. serfous fmpact on studfes refated to taxonomy, dfversfty, Because of strfct habftat preference, any fragmentatfon dfstrfbutfon, endemfsm, threats and conservatfon of of popufatfons coufd affect thefr gene fow. Even, fand snaffs. Lack of fnterest fn fand snaff research fs cuttfng of roads wfthfn a protected area coufd fragment afso due to poor fundfng opportunftfes for taxonomy popufatfons of snaffs. However, for a subset of forest and basfc bfofogy, unavaffabfffty of good taxonomfc snaff specfes, man-made habftats such as home gardens keys and fefd gufdes. The scfence of taxonomy fs and pfantatfons can act as efther refugfa or corrfdors practfsed on an fnternatfonaf fevef. Indfan Bfodfversfty

Journaf of Threatened Taxa | www.threatenedtaxa.org | September 2012 | 4(11): 3029–3037 3031 Land snaffs of Indfa S. Sen et af.

Act of 2002 does not permft exchange of specfmens 2006). Despfte havfng the fargest extfnctfon rates and wfth fnternatfonaf scfentffc communftfes, whfch hfghest number of threatened specfes (Ffg. 2), fand further hfnders the taxonomfc studfes on fand snaffs snaffs are stfff not consfdered worthy for conservatfon (Prathapan et af. 2006) . Further, most researchers and efforts despfte havfng deep fndependent phyfogenetfc conservatfon bfofogfsts show consfderabfe fnterest ffneages fn many groups. fn “charfsmatfc conservatfon” (Burner et af. 2001). Moffuscs have the fargest number of documented extfnctfons sfnce 1500AD (www.redffst.org). Non- CON SE RVAT ION OF LAND SNA ILS IN IND IA marfne specfes (terrestrfaf and freshwater) constftute 99 percent of aff moffuscan extfnctfons. Among the The rofe of communftfes and organfsatfons 566 extfnct moffuscs, the fargest proportfon fs of Land snaffs have a very poor fmage among the the fand snaffs (422 specfes) foffowed by freshwater pubffc, forest managers and poffcy makers (Seddon moffuscs (140 specfes). Tfff date there are onfy four 1998) and a fack of pubffc support. Recentfy, an recorded extfnctfons of marfne moffuscs (Lydeard et attempt was made to popufarfse fand snaffs to a wfder af. 2004; Regnfer et af. 2009 ). In the fast 300 years, audfence fn Indfa fn the form of an fffustrated gufde, the Indfan Ocean Isfands of Maurftfus, Rodrfgues and produced by the Naturaf Hfstory Museum, London Reunfon have fost 30 specfes of fand snaffs (Burner et fn assocfatfon wfth ATREE, Bengafuru on fand snaffs af. 2001) and St. Hefena and Madefra fn the Atfantfc of the Western Ghats (Appendfx 1). Thfs gufde not Ocean have fost 36 specfes of fand snaffs (WCMC onfy assfsts conservatfon bfofogfsts but afso amateur 1992). Afthough terrestrfaf vertebrate extfnctfons are naturaffsts, students and the fay pubffc affke to fdentffy weff documented, fnvertebrate extfnctfons often go fand snaffs of the Western Ghats (Raheem et af. unnotfced. Onfy a smaff fractfon, f.e., <2% of known 2009). moffuscan specfes have had thefr conservatfon status Effectfve conservatfon afso requfres awareness properfy assessed (Lydeard et af. 2004; Naggs et af. regardfng fmportance of thfs taxa fn the ecosystem

Ffgure 2. Status of Moffusks. Data from 2010 IUCN Red Lfst of Threatened Specfes (www.redffst.org)

3032 Journaf of Threatened Taxa | www.threatenedtaxa.org | September 2012 | 4(11): 3029–3037 Land snails of India S. Sen et al. and the services they provide. Communicating the Habitat conservation importance of snails and their role in ecosystem health, In India, many forested areas, which are devoid could also attract interest of wider ranging audience. of large and charismatic mammal species, have high An organization called Buglife in Europe was set up land snail diversity. However, these reserve forests to conserve rare invertebrates in Britain (www.buglife. generally receive little protection status as they fall org.uk). Such efforts are needed in this country for outside the protected area network. These reserve effective conservation of invertebrates in general and forests are open access and hence they are prone to a land snails in particular. variety of anthropogenic disturbances such as collection of minor forest products, grazing, fire, etc. Significant Species-specific conservation plans proportions of endemic species are distributed in A notable exception to the general apathy towards these non-protected areas and hence are vulnerable snail conservation is the International Partula to extinction. Current habitat conservation practice Conservation Programme, a unique ex-situ breeding is focussed on encompassing iconic but generally programme initiated by Prof. Bryan Clarke at the widely distributed/low risk species. Conversely University of Nottingham and coordinated by the habitats with the highest total of biological diversity Zoological Society of London. About 25 Partula snail are not targeted. Land snails are of potential value as species were exterminated in the wild by the predatory indicators of high diversity habitats for a wide range of snail, Euglandina rosea that was introduced from plant and groups. Thus, snails can be utilised Florida in a misguided attempt to control the introduced for identifying biodiversity rich habitats that should Giant African Snail Lissachatina fulica. This program be given high conservation priorities. In addition, the involved a managed breeding programme for 25 high geographical turnover of many land snail species species of Partula in 15 collaborating zoos globally exemplifies the value of local scale conservation in along with local conservationists, local communities capturing biological diversity in general (Ponder 1997; and the French Polynesian Government. To date more Raheem et al. 2009). For example, the low elevation than 30 species of Partula have been rescued and evergreen reserve forests of Agumbe, , etc., captive breeding populations have been established in the central Western Ghats have lower levels of in 18 zoos and laboratories throughout the world protection but harbour high species diversity of land (Pearce-Kelly et al. 1997). In another collaborative snails (Aravind 2005). The top down approach of effort of the Natural History Museum, Bermuda conserving large mammals will exclude the majority and London Zoo, 56 species of highly endangered of land snails, other invertebrates and plants from land snails from Bermuda were captive bred and protection (Aravind et al. 2005). Hence, for effective reintroduced as a part of their species conservation conservation of land snails, some of the species rich action plan (www.zsl.org/zsl-london-zoo//news/snail- areas surrounding the protected areas such as Agumbe, mail-from-bermuda,123,NS.html). In the USA plans Hulikal and similar areas rich in land snail diversity for reintroductions to conserve the Stock Island Tree need to be given additional protection. In India and Snail Orthalicus reses have been taken up (Boresma other developing countries, where information is et al. 2001; Froys et al. 2001). In India the lack of seriously lacking, an alternative approach to maximise information base, studies on the distribution, ecology, the conservation of rare and endemic species is by genetics and threats, has hindered the development of identifying sites with high diversity and endemism an informed conservation plan. However, a captive and protecting the habitat itself (Gaston 1996). breeding program for selected species would help in increasing public awareness. Hence, setting up of Need for future research “snailariums” for breeding of endemic snails could Research on land snails in India should focus on usefully be taken up by national parks and zoos. This their distribution patterns, taxonomy and ecology. might raise public interest in snail conservation across Taxonomic expertise is a basic foundation for the country. estimation of global biodiversity and formulation of policy on conservation of biological diversity (Golding & Timberlake 2002; Budha 2005). One of

Journal of Threatened Taxa | www.threatenedtaxa.org | September 2012 | 4(11): 3029–3037 3033 Land snails of India S. Sen et al. the greatest impediments for malacological research in Glessula through molecular phylogenetic applications India has been a severe lack of trained malacologists. or through DNA barcoding should be initiated. The initiative such as All India Coordinated Project on Taxonomy (AICOPTAX) by Ministry of Environment and Forests (MoEF) produced little work Conclusion on molluscan taxonomy. The recent collaborative project “Developing land snail expertise in South For informed conservation measures to be and Southeast Asia” funded under Darwin Initiative implemented, detailed studies on land snail Project by DEFRA, UK, helped build capacity on systematics, on threats to survival and on identifying land snail taxonomy not only in India but also in Sri “hot-spots” for narrow range endemics are urgently Lanka, Nepal, Laos, Thailand, Malaysia and Vietnam needed. Serious attention needs to be paid towards (Naggs et al. 2006). However, more funds need protecting remaining forested areas, maintaining to be allocated for capacity building in the areas of and possibly restoring connectivity, especially in the taxonomy, natural history, ecology and biology of the tropical rain forests which support rich snail diversity species in India. Developing databases on ecology, (Emberton 1996). More funds need to be allocated to breeding behaviour, distribution and other details of capacity building in malacology. Land snail expertise land snails should be encouraged and made accessible is urgently needed for economic reasons; awareness of in the public domain, which could change the esoteric native species will certainly help in recognizing newly status of malacology. In recent years, there have been introduced exotic species allowing effective control accidental introductions of alien and invasive species or management before they become invasive. Efforts into the Indian subcontinent. The impact of these should be made to establish snailariums in zoos in invasive snails and slugs such as Lissachatina fulica, order to create awareness about snails among people. Derocerus leave, Semiperula sp. on native land snail Priority should be given to conserve critical habitat populations needs to be monitored. The introduced for conservation of land snails. India should also start species generally occupy transformed habitats and an initiative such as the Frozen Arc Project (www. their agricultural/horticultural pest status is more of an frozenark.org), where the viable cells of a number of issue than being a threat to the native snails. However, near extinct species could be stored for possible use in impact of invasive and pest species on native land snails the future. and on the economic damage they incur to agriculture and horticultural crops needs to be assessed. For the exotic invasive malacofauna of the Indian sub- References continent to be controlled in a timely way its status Aravind, N.A. (2005). Ecology of Land Snails of Western urgently needs to be assessed. Land snails such as the Ghats. PhD Thesis, Department of Applied Zoology, African Giant Snail and others are hosts for the rat-lung University, Mangalore, 182pp. worm, which can transmit meningitis to humans. In Aravind, N.A. & F. Naggs (2012). Snailing up the Canopies India, we hardly have any information on what percent of Western Ghats, pp. 43–46. In: Devy, M.S., T. Ganesh of human population is affected by this parasite; what & A. Tripaty (eds.), Canopies of South Asia - A Glimpse. are the health implications and how many species are ATREE, . Aravind, N.A., K.P. Rajashekhar & N.A Madhyastha carriers for the same. Hence, research is needed in this (2005). Species diversity, endemism and distribution of direction as well. Indian land snails include lineages land snails of Western Ghats, India; Records of Western ( and Caenogastropda) that diverged as far Museum Supplement 68: 31–38. back as the Devonian (416 to 359.2 million years ago) Aravind, N.A., K.P. Rajashekhar & N.A. Madhyastha and there are a number of deep independent lineages (2008). Micromolluscs of Western Ghats, India: Diversity, of ancient groups some of which are unique to South distribution and threats. Zoosymposia 1: 281–294. Aravind, N.A., K.P. Rajashekhar & N.A Madhyastha (2010). Asia (Wade et al. 2006). Thus, research should also A review of ecological studies on patterns and processes focus on the biogeographical patterns, phylogeny and of distribution of land snails of the Western Ghats, India. evolution of Indian land snail fauna. Further, studies to Proceeding of World Congress of Malacology, 222pp. resolve the taxonomically problematic groups such as Bar-Yosef Mayer, D.E. (2002). Archaeomalacology: Molluscs

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Appendix 1. An illustrated guide to the land snails in India (Raheem et al. 2009)

Author Details: Mr. Sa n d e e p Se n is a research intern at the Conservation Genetics Department at ATREE. Basically a geneticist by training, He is studying phylogenetics and population genetics of land snails of the Western Ghats. He is also developing species pages for Indian mollusk fauna. Dr. Ra v i k a n t h leads the conservation Genetics Lab at ATREE. He has been actively involved in working out genetic structure of threatened species in the Western Ghats. He has undertaken a number of research projects related to the conservation and species recovery of economically important and/or endangered species. Dr. Ar a v i n d is a Fellow at ATREE. He has been working on ecology and taxonomy of non-marine mollusk of the Western Ghats for a decade. He is collaborating with Natural History Museum, London on non-marine mollusc taxonomy.

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