Distribution, threats and conservation status of the Wayanad Mahseer, Neolissochilus wynaadensis (Day, 1873) (Teleostei: Cyprinidae): an endemic large barb of the Western Ghats, India
Anvar Ali 1, Neelesh Dahanukar 2, Siby Philip 3, K. Krishnakumar 4 & Rajeev Raghavan 5
1,3,4,5 Conservation Research Group (CRG), St. Albert’s College, Kochi, Kerala 682018, India
2 Indian Institute of Science Education and Research (IISER), Dr. Homi Bhabha Road, Pashan, Pune, Maharashtra 411008, India
2,5 Systematics, Ecology & Conservation Laboratory, Zoo Outreach Organization (ZOO), 96 Kumudham Nagar, Vilankurichi Road, Coimbatore, Tamil Nadu 641035, India
3 Department of Zoology, Nirmalagiri College, Koothuparambu, Kannur, Kerala 670701, India
5 Mahseer Trust, c/o The Freshwater Biological Association, East Stoke River Laboratory, Wareham, Dorset, BH20 6BB, United Kingdom
1 firstname.lastname@example.org, 2 email@example.com, 3 firstname.lastname@example.org, 4 email@example.com, 5 firstname.lastname@example.org (corresponding author)
Abstract: The Wayanad Mahseer Neolissochilus wynaadensis (Day, 1873) is an endemic cyprinid fish that occurs in the upland streams and rivers of the southern region of the Western Ghats. This species has been listed as ‘Critically Endangered’ on the IUCN Red List of Threatened Species due to its restricted distribution and heavy declines in populations. Like many large cyprinids of the Western Ghats, N. wynaadensis is poorly known and documented, with very few verified records and voucher specimens. Based on specimens recently collected from Wayanad, Kerala, the type locality, as well as two additional locations in the Kodagu District of Karnataka; we provide information on the current distribution, phylogenetic position, threats and conservation. An updated conservation assessment of this species following the IUCN Red List categories and criteria is also provided.
Keywords: Barbodes, Coorg, freshwater fish, Vulnerable.
Abbreviations: BMNH - Natural History Museum, London; CRG-SAC - Conservation Research Group, St. Albert’s College, Kochi; FMNH - Field Museum, Chicago.
The cyprinid genus, Neolissochilus Rainboth, 1985, comprises of 25 species of medium to large sized barbs, of which 22 are known to occur in Southeast Asia (see Kottelat 2013) and seven in the Indian subcontinent (Table 1). They are important game fishes in parts of South and Southeast Asia (Jhingran 1977; Khaironizam 2010) and are also known to occur in the aquarium pet trade (Khaironizam 2010). Sixteen species of Neolissochilus have been assessed for their conservation status, of which, two species have been assessed as ‘Critically Endangered’, two as ‘Vulnerable’, three as ‘Near Threatened’, four as ‘Least Concern’ and five as ‘Data Deficient’ (see Table 1; and also species specific accounts on IUCN Red List of Threatened Species at www.iucnredlist.org).
The Wayanad Mahseer, Neolissochilus wynaadensis (Day, 1873) is one of two species within the genus that occurs in peninsular India; the other being N. bovanicus, found in the Bhavani River. Day (1873; p528) described Barbus (Barbodes) wynaadensis, from Vythiri, Wayanad and reported it to be a common species in the larger streams of the region, growing up to a length of 200 mm (also see Day 1878, p 568; 1889, p 313). The species was considered to be endemic to the streams in the Wayanad region of Kerala, until Manimekalan (1998) and subsequently, Yazdani et al. (2001) recorded it from Mudumalai, Tamil Nadu and Arunachalam et al. (2005) recorded it from Abbey Falls near Madikeri, Kodagu, Karnataka.
Neolissochilus wynaadensis has been considered to be a rare species with several surveys in and around the type locality, failing to collect this species, and very few records available in the recent ichthyological literature which are backed by voucher specimens (see Abraham 2011). Due to its restricted distribution and severe population declines (up to 80%) in the last decade, N. wynaadensis has been listed as ‘Critically Endangered’ in the IUCN Red List of Threatened Species (Abraham 2011).
As part of the on-going project on the ‘Lost fishes of Western Ghats’, which aims to document the status and distribution of the region’s rarest species (see Ali et al. 2013 a,b,c) we obtained specimens of N. wynaadensis (Image 1 a,b,c) from its type locality (Wayanad) as well as several locations in Kodagu District, Karnataka. This paper serves to document these records, and discuss the current distribution, threats and conservation status of this species. In the wake of this new information, we also propose a revised Red List status for N. wynaadensis.
Materials and Methods
Neolissochilus wynaadensis, CRG-SAC.2013.109-110, 07.x.2013, 2 ex., 126.45–139.83 mm SL, off the road from Mukkodlu to Surlabi, Kodagu, 12.5010N & 75.7570E, Cauvery River, Karnataka, India, coll. Raghavan et al.; Neolissochilus wynaadensis, 08.x.2013, CRG-SAC.2013.111-112, 2 ex., 114.85–130.03 mm SL, off the road from Bhagamandala to Napoklu, 12.3290N & 75.6590E, Cauvery River, Karnataka, India, coll. R. Raghavan et al.; Neolissochilus wynaadensis, CRG-SAC.2013.113-114, 11.x.2013, 2 ex., 83.97–84.48 mm SL, Thirunelli, 11.9020N & 75.9980E, Kalindi Stream of Kabini River, Wayanad, Kerala, India, coll. A. Ali & K. Krishnakumar.
Photographs: Barbus wynaadensis, F. Day, BMNH 18188.8.131.521, 1 ex, Vythiri, Wayanad, India; Barbus wynaadensis, F. Day, FMNH 2318, 1 ex, Vythiri, Wayanad, India (both paralectotypes) (Image 2 a,b).
Counts and measurements follow Pethiyagoda et al. (2008, 2012). Measurements were taken using a digital calliper to the nearest 0.1mm. Subunits of body are presented as percent of standard length (SL) and subunits of head are provided as percent of head length (HL) (see Table 2).
DNA isolation and molecular phylogeny
Fin clips were extracted from fresh specimens collected from three different locations (one of which was from the type locality) in the Cauvery drainage (see Table 3) and was preserved in absolute ethanol. Laboratory protocols for DNA isolation, PCR amplification of co1 and cytb genes, and molecular phylogeny are similar to those detailed in Ali et al. (2013a,c).
Results and Discussion
Taxonomy and generic status
Neolissochilus wynaadensis has been placed in different genera, viz.: Barbus (Barbodes) (Day 1873, 1878, 1889), Barbus (Puntius) (Hora & Law 1941), Puntius (Tonapi & Mulherkar 1963; Jayaram 1981; Manimekalan 1998), Neolissochilus (Rainboth 1985; Talwar & Jhingran 1991; Devi et al. 1996; Pethiyagoda et al. 2012), and Barbodes (Wu 1977; Zhu 1995; Chen et al. 1999).
Our study shows that the correct generic allocation of the species is Neolissochilus (see Fig. 1) and not Barbodes s.s. (after Kottelat 2013) as mentioned in public databases like Catalog of Fishes (Eschemeyer 2014) and FishBase (Froese & Pauly 2014). The N. wynaadensis sequences (co1 and cytb) formed a distinct clade closely related to other Neolissochilus species such as N. hexagonolepis, N. hexastichus and N. stracheyi. The closest genus related to Neolissochilus is Tor (Fig. 1a,b). Endemic large barbs of the Western Ghats (Hypselobarbus and Lepidopygopsis) also formed a monophyletic group along with Tor and Neolissochilus.
Rainboth (1985) suggested the possibility of the presence of more than one species of Neolissochilus per drainage based on the information available from the eastern part (i.e., Southeast Asia) of its distribution. However, our results on the biometrics as well as mitochondrial co1 and cytb gene sequences of N. wynaadensis collected from Wayanad (type locality) and two locations in Kodagu (=Coorg) are similar (see Table 2, 4 and Fig 1a,b) and suggest that there is only one species within the upper Cauvery drainage (Wayanad and Kodagu). The morphometric characters of the fish analysed in the present paper, are also consistent with those recorded by Arunachalam et al. (2005) from Abbey Falls. It is to be noted here that, similar to other large barbs of Western Ghats (see Knight et al. 2013 a,b; Ali et al. 2013 a,b), N. wynaadensis also has a wide range of lateral line scale counts (26–29).
Currently, only two species of Neolissochilus are known from peninsular India and the Western Ghats, N. bovanicus and N. wynaadensis. The following names, viz., Neolissochilus tamiraparaniensis, N. acutirostris, N. microphthalmus, N. capudelphinus, N. minimus and N. anamalaiensis, which are mentioned in some scientific and non-scientific literature, are unavailable and represent ‘nomina nuda’ (for details see Raghavan et al. 2013).
Neolissochilus wynaadensis is endemic to the Western Ghats of India (Dahanukar & Raghavan 2013). The species has a restricted distribution (an area of occupancy of 1000km2) occurring in the east flowing streams and rivers part of the larger Cauvery drainage, i.e., Kalindi Stream of Kabini River in Thirunelli; Noolpuzha Stream of Kabini River inside the Wayanad Wildlife Sanctuary, Kakkanhalla and Moyar tributaries inside the Mudumalai Wildlife Sanctuary, as well as streams near Vythiri in Wayanad, and Muvathoklu, Mukkodlu, Surlabi, Bhagamandala and Abbey falls in Kodagu (see Image 3; Table 5). In Kodagu, N. wynaadensis has also been recorded in Hamyala (Images 4, 5) and Kakkehole (Steven Lockett, pers. comm. 2013). However, no voucher specimens are available. The species is currently known between the altitudinal range 400–1100 m.
The fact that the species has a very restricted distribution, and is confined to only a few tributaries/streams in its range is additionally evident from the studies of Johnson & Arunachalam (2009) (studies conducted in 2001–2002) where they did not record N. wynaadensis from two tributaries (Thalipuzha and Bavalipuzha) of the east flowing Cauvery in Wayanad District.
Abraham (2011), mentions that the fish also occurs in the Chooralmala and Kanthampara areas in Wayanad (part of the west flowing Chaliyar River system), without providing any reference. The author also mentions (a personal communication; see http://www.iucnredlist.org/details/172429/0) that the species occurs in the Periya Forest Range near Mananthavady, Wyanad. It is to be noted that, in a comprehensive study of the fishes of Nilgiri Biosphere Reserve including parts of Wayanad and upper reaches of Chaliyar (Easa & Basha 1995; Easa & Shaji 1997), not a single specimen of N. wynaadensis was collected from Chooralmala and Kanthampara (see Easa & Basha 1995; Table 10; p 29), suggesting that the species could be restricted to only the east flowing drainages (Easa & Shaji 1997). Therefore, in the absence of any references to the records, and actual voucher specimens, it is premature to conclude that N. wynaadensis occurs in the west flowing drainages as suggested by Abraham (2011).
Arunachalam et al. (2005) mentioned that N. wynaadensis occurs in Vattapoil in Wayanad District by citing Gopi (2000). This record may however be based on the personal examination of voucher specimens (ZSI/WRS Calicut 6868) collected from this location, rather than as mentioned by Gopi (2000). This is because, Gopi (2000), in his review on the fishes of Kerala, does not mention such a location and only records this species from Thirunelli and Vythiri (see p 62 and 71).
Abraham (2011) mentions about a record of N. wynaadensis from Bhadra River, citing Shahnawaz et al. (2009 sic) (note that the correct year should have been 2010). However a perusal of the original paper by Shahnawaz et al. (2010) reveals no such record. Similarly, although Pillai (1929) and Hora & Law (1941) recorded Barbus wynaadensis from Travancore in southern Kerala, it is highly unlikely that the species occurs in the region. The record by Pillai (1929) from Tenmalai was also doubted by Menon (2004). In all probability, the specimen misidentified as Barbus wynaadensis from Travancore could be the fish currently known as ‘Barbodes’ carnaticus.
Neolissochilus wynaadensis was reported from Mula-Mutha River of Pune by Tonapi & Mulherkar (1963). However, this record was doubted by Kharat et al. (2001, p 47). Similarly, the records of N. wynaadensis from China (Wu 1964, cited in Talwar & Jhingran 1991), especially from the Yunnan region (Wu 1977, cited in Chen et al. 1999; Yang & Chen 1994; Jing et al. 2013; also see cytb sequence KC696524 in GenBank) are ‘at best’ misidentifications, and may in all probability be N. hexagonolepis, N. stracheyi (see Fig 1) or an undescribed species similar to it. The examination of the specimen, KIZ8110101 (see Yang & Chen 1994) at the Kunming Institute of Zoology, China will help clear the exact identity of the ‘Barbodes wynaadensis’ recorded from the rivers in Yunnan.
Further studies are, however, required to understand the diversity of this genus especially from the southern regions of the Western Ghats from where many ‘nomina nuda’ under the genus Neolissochilus (see Raghavan et al. 2013) as well as previous records (Pillai 1929; Hora & Law 1941) of N. wynaadensis are available. There is specifically a need to examine specimens identified and catalogued in various museums, as Tor species/Mahseer from various regions south of the Palakkad Gap.
Neolissochilus wynaadensis has been considered to be a very rare species (Gopi 2000; Easa & Shaji 1997), as it was not recorded in many collections from Wayanad during 1999–2004 and in 2008 (Abraham 2011). Although not backed by any scientific data, global populations of this species were considered to have reduced by 80% during 2000–2010, and are still declining (See Abraham 2011).
Our field surveys in Thirunelli in April 2004 (Unpublished), and recently in October 2013 (vouchers mentioned in the present paper) led to the collection of only three specimens, while those in Vythiri in April 2004 did not yield any. Nevertheless, there are fairly good populations in Kodagu District of Karnataka State as evident from large shoals inhabiting the streams and rivers where they were occur. A large group (probably in hundreds) of N. wynaadensis were seen in a single splash pool near Muvathoklu where they are protected in a sacred grove (Devarkaadu).
Habitat and Ecology
Neolissochilus wynaadensis prefers fast flowing upland streams and rivers where they occur in both rocky pools (Abraham 2011) as well as riffles (Kurup et al. 2004). The pool-riffle habitats have a moderate flow velocity and provide good hiding places for the species. These habitats are also frequented by riffle beetles and chironomids that form the major food for N. wynaadensis (Manojkumar & Kurup 2002). Kurup et al. (2004) provides information on several microhabitat variables in the locations frequented by N. wynaadensis. Habitats in the sites from where N. wynaadensis was recorded in the present study are shown in Images 6, 7 and 8. It has also been suggested that N. wynaadensis are long range migrants that travel to the upper reaches of rivers for spawning (Menon 2004).
The Neolissochilus from peninsular India (N. wynaadensis) forms a monophyletic grouping with other Neolissochilus species from northeastern India and Southeast Asia (Fig 1a,b). Also, the specimens of Neolissochilus found in various locations in Kodagu (=Coorg, in Bhagamandala and Mukkodlu), were genetically similar to those that occur in the type locality of the species (i.e., Wayanad) (also see Table 4). The discontinuous distribution of the genus Neolissochilus (Sundaland, Indo Burma, Eastern Himalaya and Western Ghats) could also form the basis for interesting biogeographic hypothesis testing; for instance, to check the scenario of ‘true disjuncts’ (see Dahanukar et al. 2013 and references therein). On the other hand, detailed anatomical and morphological analysis is needed to ascertain if the Western Ghats representatives of the genus are ‘false disjuncts’ and need a separate generic allocation. Such biogeographic and evolutionary questions should however be validated with larger multi-locus datasets.
Threats and conservation
Anthropogenic threats including habitat destruction as well as pollution caused by the discharge of fertilizers, pesticides and other chemical effluents from surrounding plantations and sand mining have been reported from the habitats frequented by the species (Abraham 2011). There is no data to show that this species is being exploited as a food fish (Abraham 2011), but given its size, it is likely to be harvested by local communities. Overall, there is a need for greater understanding of the specific threats, their intensity and the impacts they have on the survival of the species.
Much of the known range of N. wynaadensis is outside protected areas. Currently, Wayanad Wildlife Sanctuary in Kerala and Mudumalai Wildlife Sanctuary in Tamil Nadu are the only protected areas where the species occur. However, an alternate protected area system in the form of sacred groves, (Devarkaadu) exists in Kodagu, Karnataka. Streams flowing through these sacred groves harbour good populations of N. wynaadensis which are protected (for e.g., in Mukkodlu, Muvathoklu and Surlabi regions) by the local communities.
Currently, N. wynaadensis has been listed as ‘Critically Endangered’ (Abraham 2011) in the IUCN Red List of Threatened Species. Additional information on distribution and threats as documented in the present paper has led to a scenario where the conservation status of the species needs to be revised. The proposed Red List Status of the species has been provided in Appendix 1.
Large cyprinids of the genus Tor, Neolissochilus, Hypselobarbus and ‘Barbodes’ comprise one of the most poorly known freshwater fishes of the Western Ghats, with very little information available on their diversity (both generic and specific) and distribution. Recent studies have helped to improve our knowledge on some genera (for e.g., Hypselobarbus) (see Ali et al. 2013a,b; Knight et al. 2013a,b,c), while others (Tor, Neolissochilus and Barbodes) remain understudied. Future research should be directed on these large barbs to understand in detail their taxonomy, diversity, distribution, population dynamics and stock structure and conservation. There is particularly a need to understand the identity of the barb described by Day (1877) as Barbus bovanicus, from the Bhavani River in the southern Western Ghats, which is currently considered to be a species of Neolissochilus (see Pethiyagoda et al. 2012).
Abraham, R. (2011). Barbodes wynaadensis. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.2. <www.iucnredlist.org>. Downloaded on 24 December 2013.
Ali, A., N. Dahanukar, A. Kanagavel, S. Philip & R. Raghavan (2013a). Records of the endemic and threatened catfish, Hemibagrus punctatus from the southern Western Ghats with notes on its distribution, ecology and conservation status. Journal of Threatened Taxa 5(11): 4569–4578; http://dx.doi.org/10.11609/JoTT.o3427.4569-78
Ali, A., S. Philip & R. Raghavan (2013b). Back from obscurity: notes on the current distribution, threats and conservation status of a poorly known cyprinid, Hypselobarbus lithopidos (Day, 1874) from the Western Ghats of India. Journal of Threatened Taxa 5(13): 4743–4751; http://dx.doi.org/10.11609/JoTT.o3655.4743-51
Ali, A., S. Philip, N. Dahanukar, C.R. RenjithKumar, A. Bijukumar & R. Raghavan (2013c). Distribution, threats and conservation status of Hypselobarbus thomassi (Day, 1874), a poorly known cyprinid fish of the Western Ghats freshwater ecoregion. Journal of Threatened Taxa 5(17): 5202–5213; http://dx.doi.org/10.11609/JoTT.o3838.5202-5213
Arunachalam, M., M. Muralidharan, P. Sivakumar, R. Soranam & M. Murugan (2005). New record of a rare barbin Neolissochilus wynaadensis (Day) from Abby Falls, Karnataka. Zoos’ Print Journal 20(11): 2073–2074; http://dx.doi.org/10.11609/JoTT.ZPJ.1342.2073-4
Chen, X.Y., J.X. Yang & Y.R. Chen (1999). A review of the cyprinoid fish genus Barbodes Bleeker, 1859 from Yunnan, China, with descriptions of two new species. Zoological Studies 38(1): 82–88.
Dahanukar, N. & R. Raghavan (2013). Freshwater fishes of Western Ghats: Checklist v 1.0 August 2013. MIN - Newsletter of IUCN SSC/WI Freshwater Fish Specialist Group - South Asia and the Freshwater Fish Conservation Network of South Asia (FFCNSA) 1: 6–16.
Dahanukar, N., S. Philip, K. Krishnakumar, A. Ali & R. Raghavan (2013). The phylogenetic position of Lepidopygopsis typus (Teleostei: Cyprinidae), a monotypic freshwater fish endemic to the Western Ghats of India. Zootaxa 3700(1): 113–139. http://dx.doi.org/10.11646/zootaxa.3700.1.4
Day, F. (1873). On some new fishes of India. The Journal of the Linnean Society of London 11: 524–530. http://dx.doi.org/10.1111/j.1096-3642.1873.tb01674.x
Day, F. (1878). The Fishes of India; Being A Natural History of The Fishes Known to Inhabit the Seas and Freshwaters of India, Burma, and Ceylon - Part 4. Bernard Quatrich, London, i-xx + 553-579, pls 139–195.
Day, F. (1889). Fishes. In: Blanford, W.T. (ed). The Fauna of British India including Ceylon and Burma. Taylor & Francis, London. v. 1: 1-xviii + 1-548.
Devi, K.R., T.J. Indra & K.G. Emiliyamma (1996). On the fish collections from Kerala deposited in the Southern Regional Station, Zoological Survey of India, by NRM, Stockholm. Records of the Zoological Survey of India 95(3–4): 129–146
Easa, P.S. & S.C. Basha (1995). A Survey of The Habitat and Distribution of Stream Fishes in the Kerala Part of Nilgiri Biosphere Reserve. Kerala Fisheries Research Institute Research Report No. 104, 87pp.
Easa, P.S. & C.P. Shaji (1997). Freshwater fish diversity in the Kerala part of the Nilgiri Biosphere Reserve. Current Science 73(2): 180–182.
Euphrasia, C.J., K.V. Radhakrishnan & B.M. Kurup (2006). The threatened freshwater fishes of Kerala, India, pp. 570–588. In: Kurup, B.M. & K. Ravindran (eds.). Sustain Fish. School of Industrial Fisheries, Cochin University of Science and Technology, Cochin, India.
Gopi, K.C. (2000). Freshwater fishes of Kerala State, pp. 56–76. In: Ponniah, A.G. & A. Gopalakrishnan (eds.). Endemic Fish Diversity of Western Ghats. NBFGR-NATP, India.
Hora, S.L. & N.C. Law (1941). The freshwater fishes of Travancore. Records of Indian Museum 43: 233–256.
Jayaram, K.C. (1981). The Freshwater Fishes of India, Pakistan, Bangladesh, Burma and Sri Lanka - A Handbook. Zoological Survey of India, i-xxii+475pp, pls. 1–13.
Jhingran, V.G. (1977). Fish and Fisheries of India. Hindustan Publishing Corporation, 953pp.
Jing, W., X.Y. Wu, Z.M. Chen, Z.P. Yue, W. Ma, S.Y. Chen, H. Xiao, R.W. Murphy, Y.P. Zhang, R.G. Zan & J. Luo (2013). Molecular phylogeny of European and African Barbus and their West Asian relatives in the Cyprininae (Teleostei: Cypriniformes) and orogenesis of the Qinghai-Tibetan Plateau. Chinese Science Bulletin 58 (31): 3738–3746. http://dx.doi.org/10.1007/s11434-013-5878-z
Johnson, J.A. & M. Arunachalam (2009). Diversity, distribution and assemblage structure of fishes in streams of southern Western Ghats, India. Journal of Threatened Taxa 1(10): 507-513; http://dx.doi.org/10.11609/JoTT.o2146.507-13
Khaironizam, M.Z. (2010). Some biological aspects of Neolissochilus spp. (Teleostei: Cyprinidae) from Peninsular Malaysia. PhD Thesis. University of Malaya, Kuala Lumpur, 215pp.
Kharat, S.S., N. Dahanukar & R. Raut (2001). Decline of fresh-water fish of Pune urban area. Journal of Ecological Society 13/14: 46–51.
Knight, J.D.M., A. Rai & R.K.P. D’souza (2013a). Re-description of Hypselobarbus lithopidos (Teleostei: Cyprinidae), based on its rediscovery from the Western Ghats, India, with notes on H. thomassi. Journal of Threatened Taxa 5(13): 4734-4742; http://dx.doi.org/10.11609/JoTT.o3602.4734-42
Knight, J.D.M., A. Rai & R.K.P. D’souza (2013b). On the identities of Barbus mussullah Sykes and Cyprinus curmuca Hamilton with notes on the status of Gobio canarensis Jerdon (Teleostei: Cyprinidae). Zootaxa 3750(3): 201–215; http://dx.doi.org/10.11646/zootaxa.3750.3.1
Knight, J.D.M., A. Rai & R.K.P. D’souza (2013c). Rediscovery of Hypselobarbus pulchellus, an endemic and threatened barb (Teleostei: Cyprinidae) of the Western Ghats, with notes on H. dobsoni and H. jerdoni. Journal of Threatened Taxa 5(17): 5194–5201; http://dx.doi.org/10.11609/JoTT.o3686.5194-201
Kottelat, M. (2013). The fishes of the inland waters of South East Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries. Raffles Bulletin of Zoology (Supplement) 27: 1–663.
Kurup, B.M., K.V. Radhakrishnan & T.G. Manojkumar (2004). Biodiversity Status of Fishes Inhabiting Rivers of Kerala (South India) With Special Reference to Endemism, Threats and Conservation Measures. In: Welcomme, R.L & T. Petr. (eds.). Proceedings of the Second International Symposium on the Management of Large Rivers for Fisheries 2 (LARS2), Cambodia, 310pp.
Manimekalan, A. (1998). The fishes of Mudumalai Wildlife Sanctuary, Tamil Nadu, South India. Journal of the Bombay Natural History Society 95(3): 431–443.
Manojkumar, T.G. (2006). Fish habitats and species assemblages in the selected rivers of Kerala and investigation on life history traits of Puntius carnaticus (Jerdon, 1849). PhD Thesis. Cochin University of Science and Technology, Kochi, India.
Manojkumar, T.G. & B.M. Kurup (2002). Fish habitat diversity and species assemblage structure with reference to five major river systems of Kerala, pp. 141–150. In: Riverine and Reservoir Fisheries of India. Boopendranath, M.R., B. Meenakumari., J. Joseph., T.V. Sankar., P. Pravin & L. Edwin (eds.). Society for Fisheries Technology, Cochin and Central Institute of Fisheries Technology, Cochin.
Menon, A.G.K. (1999). Check list-fresh water fishes of India. Records of the Zoological Survey of India. Miscellaneous Publication, Occasional Paper No. 175: i-xxviii+1–366.
Menon, A.G.K. (2004). Threatened fishes of India and their conservation. Zoological Survey of India, 170pp.
Pethiyagoda, R., M. Meegaskumbura & K. Maduwage (2012). A synopsis of the South Asian fishes referred to Puntius (Pisces: Cyprinidae). Ichthyological Exploration of Freshwaters 23(1): 69–95.
Pethiyagoda, R., A. Silva, K. Maduwage & M. Meegaskumbura (2008). Puntius kelumi, a new species of cyprinid fish from Sri Lanka (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters 19(3): 201–214.
Pillai, R.S.N. (1929). A list of fishes taken in Travancore from 1901– 1915. Journal of the Bombay Natural History Society 33(2): 347–379.
Raghavan, R., N. Dahanukar, S. Philip, K. Krishnakumar & A. Ali (2013). Phantom references, nomina nuda and the dilemma of freshwater fish taxonomy in India. Current Science 104(10): 1277–1279.
Rainboth, W.J. (1995). Neolissochilus, a new genus of South Asian cyprinid fishes. Beaufortia 35 (3): 25–35.
Shahnawaz, A., M. Venkateshwarlu, D.S. Somashekar & K. Santosh (2010). Fish diversity with relation to water quality of Bhadra River of Western Ghats (India). Environmental Monitoring and Assessment 161(1–4): 83–91; http://dx.doi.org/10.1007/s10661-008-0729-0
Shaji, C.P. & P.S. Easa (1995). Freshwater fish diversity in Wayanad, Kerala, south India. Journal of Zoological Society of Kerala 5(1&2): 30–36.
Talwar, P.K. & A.G. Jhingran (1991). Inland fishes of India and adjacent countries - 2 volumes. Oxford & IBH Publishing Co. New Delhi, 1158pp.
Tonapi, G.T. & L. Mulherkar (1963). Notes on the freshwater fauna of Poona. Proceedings of the Indian Academy of Sciences - Section B 58(4): 187–197.
Wu, X.W. (1964). The cyprinid fishes of China - Volume 1. Science Press, Shanghai, 228pp. [not referred to in original]
Wu, X.W. (1977). The Cyprinid Fishes of China - Volume 2. Science Technical Press, Shanghai, pp. 236–252 [not referred to in original]
Yazdani, G.M., K.R. Devi, M.B. Raghunathan & D.F. Singh (2001). Fauna of Nilgiri Biosphere Reserve - Pisces. Fauna of Conservation Area Series 11. Zoological Survey of India, 207–224.
Yang, J. & Y. Chen (1994). Phylogeny and zoogeography of the cyprind genus Spinibarbus (Pisces: Cyprinidae). Zoological Research 15: 13–25.
Zhu, S.Q. (1995). Synopsis of freshwater fishes of China, 549p. [not referred to in original]