Abstract
The taxonomy of the three native taxa assigned to the genus Labeo (L. dussumieri, L. fisheri and L. porcellus lankae) in Sri Lanka is reviewed. The population hitherto identified as L. dussumieri in Sri Lanka is shown to be a distinct species, here named L. heladiva. Labeo heladiva, new species, has a wide distribution in the low and mid-elevations of the island and is distinguished from its Indian congeners by the combination of having two pairs of barbels; 12–13 branched dorsal-fin rays; lateral line with 44–51 scales; ½8–½9+1+6–7 scales in transverse series; and 19–22 circumpeduncular scales. It differs from its closest relative, L. dussumieri, principally by having 44–51 vs. 50–60 lateral-line scales, 19–22 vs. 22–27 circumpeduncular scales, and by uncorrected pairwise genetic distances of 1.27–2.22% and 1.88–2.91% for the two mitochondrial genes COI and cytb, respectively. Labeo fisheri, which is endemic to the upper reaches of the Mahaweli River basin in the Knuckles mountain range and the central hills in the vicinity of Kandy, is distinguished from Indian congeners by having (in combination) only a single pair of barbels; dorsal fin with 10–12 branched rays; lateral line with 37–39 scales; 7+1+4½–6 scales in transverse series; and 17–20 circumpeduncular scales. Labeo lankae is recognized as a valid species endemic to Sri Lanka. Long suspected to have become extinct, or known only from spurious records, an extant population is reported from the northern dry zone of the island. Labeo lankae is the sister species of L. porcellus of peninsular India; it can be distinguished from its congeners by having, in combination, 10–12 branched dorsal-fin rays; 36–39 lateral-line scales; ½8+1+5–6½ scales in transverse series; and 21–24 circumpeduncular scales. It differs from L. porcellus principally by having ½8 (vs. ½6–½7) scales between the origin of the dorsal fin and the lateral line, 21–24 (vs. 20–21) circumpeduncular scales and uncorrected pairwise genetic distances of 1.27% and 1.41% for the mitochondrial genes COI and cytb, respectively. The three species of Labeo in Sri Lanka do not form a monophyletic group.
References
Anonymous (2011) “Labeo lankae” is back again in dumbara valley. Newsletter of the National Science Foundation of Sri Lanka, 2 (3), 1–3.
Anusha, J., Arunachalam, M. & Sivakumar, P. (2017) New record of a Sri Lankan endemic species Labeo fisheri (Jordan and Starks, 1917; Cypriniformes: Cyprindae) from Kalakkad Mundanthurai Tiger Reserve, Tamil Nadu, India. Trends in Fisheries Research, 6 (2), 19–23.
Bambaradeniya, C.N.B., Ekanayake, S.P., Kekulandala, L.D.C.B., Samarawickrama, V.A.P., Ratnayake, N.D. & Feranando, R.H.S.S. (2002) An assessment of the status of Biodiversity in the Muthurajawela Wetland Sanctuary. IUCN Sri Lanka, Country Office, Occasional Paper, 3, 1–48.
Batuwita, S., Maduwage, K. & Sudasinghe, H. (2015) Redescription of Pethia melanomaculata (Teleostei: Cyprinidae) from Sri Lanka. Zootaxa, 3936 (4), 575–583.
https://doi.org/10.11646/zootaxa.3936.4.7Bleeker, P. (1860) [De visschen van den Indischen Archipel beschreven en toegelicht. Deel 2]. Ordo Cyprini, karpers. Acta Societatis Regiae Scientiarum IndoNeêrlandicae, 7 (2), i–xiv + 1–492.
Bossuyt, F., Meegaskumbura, M., Beenaerts, N., Gower, D.J., Pethiyagoda, R., Roelants, K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K., Schneider, C.J., Oommen, O.V. & Milinkovitch, M.C. (2004) Local endemism within the western Ghats-Sri Lanka biodiversity hotspot. Science, 306, 479–481.
https://doi.org/10.1126/science.1100167Clement, M., Snell, Q., Walke, P., Posada, D. & Crandall, K. (2002) TCS: estimating gene genealogies. Proceedings of the 16th International Parallel and Distributed Processing Symposium, 2, 184.
https://doi.org/10.1109/IPDPS.2002.1016585Cuvier, G. and A. Valenciennes (1842) Histoire naturelle des poissons. Tome seizième. Livre dix-huitième. Les Cyprinoïdes, 16, i–xx + 1–472, pls. 456–487.
Day, F. (1878) Fishes of India. Part 4. Quaritch, London, xx + 226 pp. (pp. i–xx + 553–778), pls. 134–195.
Day, F. (1889) The Fauna of British India, including Ceylon and Burma. Fishes Vol. 1. Taylor & Francis, London, 548 pp.
De Silva, M.A, Hapuarachchi, N. & Jayaratne, T. (2015) Sri Lankan Freshwater Fishes. Wildlife Conservation Society, Galle, 391 pp.
Deraniyagala, P.E.P. (1929) Two new freshwater fishes. Spolia Zeylanica, 15 (2), 73–77.
Deraniyagala, P.E.P. (1930) The Eventognathi of Ceylon. Spolia Zeylanica, 16 (1), 1–4 1.
Deraniyagala, P.E.P. (1952) A coloured atlas of some vertebrates from Ceylon, Vol. 1. Fishes. National Museums of Ceylon, Colombo, 149 pp., 34 pls.
Duncker, G. (1912) Die Susswasserfische Ceylons. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, Beiheft 2, Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 29 (2), 241–272.
Eschmeyer, W. N., Fricke, R. & van der Laan, R. (Eds.) (2018) CATALOG OF FISHES: GENERA, SPECIES, REFERENCES. Electronic Version. Available from: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp (accessed 1 August 2017)
Goonatilake, S. de A., (2007) Freshwater Fishes of Sri Lanka. Ministry of Environment, Colombo, Sri Lanka, 135 pp.
Günther, A. (1868) Catalogue of the fishes in the British Museum. Vol. 7. Catalogue of the Physostomi containing the families Heteropygii, Cyprinidae, Gonorhynchidae, Hyodontidae, Osteoglossidae, Clupeidae, Chirocentridae, Alepocephalidae, Notopteridae, Halosauridae, in the collection of the British Museum. British Museum, London, 512 pp.
Heckel, J.J. (1844) Fische Kaschmir's, nebst einem Anhang von drei neuen Arten aus Indien, gesammelt von Freiherrn Carl v. Hügel. In: Hügel, C.A. von (Ed.), Kaschmir und das Reich der Siek. Hallberger'sche Verlagshandlung, Stuttgart, pp. 351–392.
Huelsenbeck, J.P., Ronquist, F., Nielsen, R. & Bollback, J.P. (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science, 294, 2310–2314.
https://doi.org/10.1126/science.1065889IUCN Sri Lanka and the Ministry of Environment and Natural Resources (2007) The 2007 Red List of Threatened Fauna and Flora of Sri Lanka. The World Conservation Union (IUCN) and Ministry of Environment and Natural Resources, Colombo, Sri Lanka, xiii + 148 pp.
IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. Second edition. Gland, Switzerland and Cambridge, UK: IUCN. iv + 32 pp.
Jayaram, K.C. & Dhas, J.J. (2000) Revision of the genus Labeo Cuvier from the Indian region with a discussion on its phylogeny and Zoogeography (Pisces: Cypriniformes Cyprinidae, Cyprininae). Records of the Zoological Survey of India. Occasional Paper, 183, 1–143.
Jordan, D.S & Starks, E.C. (1917) Notes on a collection of fishes from Ceylon with descriptions of new species. Annals of Carnegie Museum, 11 (344), 430–460.
Kottelat, M. (2013) The fishes of the inland waters of Southeast Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries. Raffles Bulletin of Zoology, 27 (Supplement), 1–663.
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33 (7), 1870–1874.
https://doi.org/10.1093/molbev/msw054Lal, K.K., Gupta, B.K., Punia, P., Mohindra, V., Saini, V.P., Dwivedi, A.K., Singh, R.K., Dhawan, S., Luhariya, R.K., Basheer, V.S. & Jena, J.K. (2015) Revision of gonius subgroup of the genus Labeo Cuvier, 1816 and confirmation of species status of Labeo rajasthanicus (Cypriniformes: Cyprinidae) with designation of a neotype. Indian Journal of Fisheries, 62 (4), 10–22.
Leigh, J.W. & Bryant, D. (2015) PopART: Full-feature software for haplotype network construction. Methods in Ecology and Evolution, 6 (9), 1110–1116.
https://doi.org/10.1111/2041-210X.12410McLoughlin, S. (2001) The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany, 49 (3), 271–300.
https://doi.org/10.1071/BT00023
Mendis, A.S. & Fernando, C.H. (1962) A guide to the freshwater fauna of Ceylon. Bulletin of the Fisheries Research Station, Ceylon, 12, 1–160.
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), 2010, 1–8.
https://doi.org/10.1109/GCE.2010.5676129MOE (2012) The National Red List 2012 of Sri Lanka; Conservation Status of the Fauna and Flora. Ministry of Environment, Colombo, Sri Lanka, 476 pp.
Munro, I.S.R. (1955) The Marine and Freshwater Fishes of Ceylon. Department of External Affairs, Canberra, 349 pp.
Ng, H.H. & Pethiyagoda, R. (2013) Mystus zeylanicus, a new species of bagrid catfish from Sri Lanka (Teleostei: Bagridae). Ichthyological Exploration of Freshwaters, 24 (2), 161–170.
Palumbi, S.R. (1996) Nucleic acids II: the polymerase chain reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular systematics. Sinauer Associates, Sunderland, pp. 205–247.
Patrick, A.E.S., Kuganathan, S. & Edirisinghe, U. (2017) Species composition and abundance of fishes with seasonal fluctuations of rainfall and water level in Vavuniya reservoir, Sri Lanka. International Research Journal of Environmental Sciences, 6 (6), 12–21.
Pethiyagoda, R. (1991) Freshwater fishes of Sri Lanka. Wildlife Heritage Trust, Colombo, Sri Lanka, 362 pp.
Pethiyagoda, R. (1994) Threats to the indigenous freshwater fishes of Sri Lanka and remarks on their conservation. Hydrobiologia, 285, 189–201.
https://doi.org/10.1007/BF00005666Pethiyagoda, R. (1996) Labeo lankae. The IUCN Red List of Threatened Species 1996.
https://doi.org/10.2305/IUCN.UK.1996.RLTS.T11072A3249745.enPethiyagoda, R. (2006) Conservation of Sri Lankan freshwater fishes. In: Bambaradeniya, C.N.B. (Ed.), Fauna of Sri Lanka: status of taxonomy, research and conservation. The World Conservation Union, Colombo, pp. 103–112.
Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer. Version 1.6, Available from: http://tree.bio.ed.ac.uk/software/tracer (accessed 30 July 2018)
Reid, G.M. (1985) A revision of African species of Labeo (Pisces: Cyprinidae) and a re-definition of the genus. Verlag von J. Cramer, Braunschweig, 322 pp.
Rohling, E.J., Fenton, M., Jorissen, F.J., Bertrand, P., Ganssen, G. & Caulet, J.P. (1998) Magnitudes of sea-level lowstands of the past 500,000 years. Nature, 394 (6689), 162–165.
https://doi.org/10.1038/28134Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic biology, 61 (3), 539–542.
https://doi.org/10.1093/sysbio/sys029Rüber, L., Britz, R., Hui Tan, H., Ng, P.K. & Zardoya, R. (2004) Evolution of mouthbrooding and life-history correlates in the fighting fish genus Betta. Evolution, 58 (4), 799–813.
https://doi.org/10.1554/03-364Samarawickrama, V.A.M.P.K., Samarawickrama, D.R.N.S. & Gunawardane, J. (2012) A new site record of Gadeya (Labeo fisheri) in Mahaweli Ganga at Hakkinda, Warathenna, Central Province, Sri Lanka. Proceedings of the Eighteenth Sessions of the Sri Lanka Association for Fisheries and Aquatic Resources, 2012, 60.
Senanayake, F.R. (1980) The Biogeography and ecology of the inland fishes of Sri Lanka. Unpublished Ph.D. dissertation, Department of Wildlife and Fisheries Biology, University of California, Davis, 421 pp.
Siddall, M., Rohling, E.J., Almogi-Labin, A., Hemleben, C., Meischner, D., Schmelzer, I. & Smeed, D.A. (2003) Sea-level fluctuations during the last glacial cycle. Nature, 423 (6942), 853–858.
https://doi.org/10.1038/nature01690Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30 (9), 1312–1313.
https://doi.org/10.1093/bioinformatics/btu033Sudasinghe, H. & Meegaskumbura, M. (2016) Ompok argestes, a new species of silurid catfish endemic to Sri Lanka (Teleostei: Siluridae). Zootaxa, 4158 (2), 261–271.
https://doi.org/10.11646/zootaxa.4158.2.7Sudasinghe, H., Pethiyagoda, R., Maduwage, K. & Meegaskumbura, M. (2016) Mystus nanus, a new striped catfish from Sri Lanka (Teleostei: Bagridae). Ichthyological Exploration of Freshwaters, 27 (2), 163–172.
Swofford, D.L. (2002) PAUP, Phylogenetic Analysis using Parsimony (and other methods), v. 4b10. Sunderland: Sinauer Associates.
Sykes, W. H. (1839) On the fishes of the Deccan. Proceedings of the Zoological Society of London, 1838 (6), 157–165.
Talwar, P.K. & Jhingran, A.G. (1991) Inland Fishes of India and Adjacent Countries. Vol. I & II. Oxford and IBH Publishing Co. Pvt. Ltd, New Delhi, 1158 pp.
Templeton, A.R., Crandall, K.A. & Sing, C.F. (1992) A cladistic analysis of phenotypic associations with haplotypes inferred from restriction endonuclease mapping and DNA sequence data. III. Cladogram estimation. Genetics, 132, 619–633.
Yang, L. & Mayden, R.L. (2010) Phylogenetic relationships, subdivision, and biogeography of the cyprinid tribe Labeonini (sensu) (Teleostei: Cypriniformes), with comments on the implications of lips and associated structures in the labeonin classification. Molecular Phylogenetics and Evolution, 54 (1), 254–265.
https://doi.org/10.1016/j.ympev.2009.09.027Yang, L., Arunachalam, M., Sado, T., Levin, B.A., Golubtsov, A.S., Freyhof, J., Friel, J.P., Chen, W.J., Hirt, M.V., Mancikam, R., Agnew, M.K., Simons, A.M., Saitoh, K., Miya, M., Mayden, R.L. & He, S. (2012) Molecular phylogeny of the cyprinid tribe Labeonini (Teleostei: Cypriniformes). Molecular Phylogenetics and Evolution, 65 (2), 362–379.
https://doi.org/10.1016/j.ympev.2012.06.007Ward, R.D., Zemlak, T.S., Innes, B.H., Last, P.R. & Hebert, P.D.N. (2005) DNA barcoding Australia's fish species. Philosophical Transactions of the Royal Society B: Biological Sciences, 360, 1847–1857.
https://doi.org/10.1098/rstb.2005.1716