Abstract
The taxonomic status of the large snakeheads of the Channa marulius group that occur in Sri Lanka is reviewed and clarified. Two species are recognized from the island, based on both morphological and molecular (cytochrome c oxidase subunit 1: cox1) differentiation: C. marulius Hamilton from the northern dry zone and C. ara Deraniyagala from the middle and lower regions of the Mahaweli basin. Channa ara is endemic to Sri Lanka and can be distinguished from its Marulius group congeners, C. marulius, C. aurolineata and C. auroflammea, by having fewer dorsal- and anal-fin rays, fewer lateral-line scales and fewer vertebrae; from C. marulioides by a different adult colour pattern; and from C. pseudomarulius by having more vertebrae. At the cox1 barcoding locus, Channa ara is at least 3.6% genetically different from C. marulius, and at least 8% different from the other described species in the group. Specimens collected from the southwestern wet zone in Sri Lanka are a puzzling third component of the Marulius group’s diversity, uncovered in this study, and identified here as C. cf. ara. Whilst genetically more similar to C. marulius, C. cf. ara possesses fewer dorsal- and anal-fin rays, fewer lateral-line scales and fewer vertebrae and is therefore morphologically more similar to C. ara. Channa ara can be distinguished from C. cf. ara, however, by differences in circumpeduncular scale count, adult colour pattern, and by an uncorrected pairwise genetic distance of 3.7% in cox1 sequences. A neotype is designated for Ophicephalus marulius ara Deraniyagala.
References
Adamson, E.A.S. & Britz, R. (2018) The snakehead fish Channa aurolineata is a valid species (Teleostei: Channidae) distinct from Channa marulius. Zootaxa, 4514 (4), 542–552.
https://doi.org/10.11646/zootaxa.4514.4.7
Adamson, E.A.S., Hurwood, D.A. & Mather, P.B. (2010) A reappraisal of the evolution of Asian snakehead fishes (Pisces, Channidae) using molecular data from multiple genes and fossil calibration. Molecular Phylogenetics and Evolution, 56 (2), 707–717.
https://doi.org/10.1016/j.ympev.2010.03.027
Adamson, E.A.S., Britz, R. & Lieng, S. (2019) Channa auroflammea, a new species of snakehead fish of the Marulius group from the Mekong River in Laos and Cambodia (Teleostei: Channidae). Zootaxa, 4571 (3), 398–408.
https://doi.org/10.11646/zootaxa.4571.3.7
Anonymous (1977) Admiralty chart, India and Sri Lanka: Cochin to Vishakapatnam. UK Hydrographic Office, Taunton.
Bandelt, H.J., Forster, P. & Röhl, A. (1999) Median-joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution, 16 (1), 37–48.
https://doi.org/10.1093/oxfordjournals.molbev.a026036
Benziger, A., Philip, S., Raghavan, R., Anvar Ali, P.H., Sukumaran, M., Tharian, J.C., Dahanukar, N., Baby, F., Peter, R., Devi, K.R., Radhakrishnan, K.V., Haniffa, M.A., Britz, R. & Antunes, A. (2011) Unraveling a 146 Years Old Taxonomic Puzzle: Validation of Malabar Snakehead, Species-Status and Its Relevance for Channid Systematics and Evolution. PLoS ONE, 6 (6), e21272.
https://doi.org/10.1371/journal.pone.0021272
Britz, R. (2008) Channa ornatipinnis and C. pulchra, two new species of dwarf snakeheads from Myanmar (Teleostei: Channidae). Ichthyological Exploration of Freshwaters, 18 (4), 335–344.
Britz, R. (2013) Channa andrao, a new species of dwarf snakehead from West Bengal, India (Teleostei: Channidae). Zootaxa, 3731 (2), 287–294.
https://doi.org/10.11646/zootaxa.3731.2.9
Britz, R. (2019) Hamilton’s Gangetic Fishes in Colour: A New Edition of the 1822 Monograph, with Reproductions of Unpublished Coloured Illustrations. The Ray Society, London, 641 pp.
Britz, R., Adamson, E., Raghavan, R., Ali, A. & Dahanukar, N. (2017) Channa pseudomarulius, a valid species of snakehead from the Western Ghats region of peninsular India (Teleostei: Channidae), with comments on Ophicephalus grandinosus, O. theophrasti and O. leucopunctatus. Zootaxa, 4299 (4), 529–545.
https://doi.org/10.11646/zootaxa.4299.4.4
Chaudhry, S. (2010) Channa marulius. The IUCN Red List of Threatened Species 2010: e.T172328A6868796.
https://doi.org/10.2305/IUCN.UK.2010-4.RLTS.T172328A6868796.en
Conte-Grand, C., Britz, R., Dahanukar, N., Raghavan, R., Pethiyagoda, R., Tan, H.H., Hadiaty, R.K., Yaakob, N.S. & Rüber, L. (2017) Barcoding snakeheads (Teleostei, Channidae) revisited: Discovering greater species diversity and resolving perpetuated taxonomic confusions. PLOS ONE, 12 (9), e0184017.
https://doi.org/10.1371/journal.pone.0184017
Courtenay, W.R. & Williams, J.D. (2004) Snakeheads (Pisces, Channidae): a biological synopsis and risk assessment. US Geological Survey Circular 1251, Denver, CO, 143 pp.
https://doi.org/10.3133/cir1251
Day, F. (1878) The fishes of India; being a natural history of the fishes known to inhabit the seas and fresh waters of India, Burma, and Ceylon. William Dawson & Sons, London, 778 pp.
https://doi.org/10.5962/bhl.title.55567
Day, F. (1889) The fauna of British India including Ceylon and Burma. Fishes. Vol. 1. Taylor & Francis, London, 548 pp.
https://doi.org/10.5962/bhl.title.100755
De Silva, M.A., Hapuarachchi, N. & Jayaratne, T. (2015) Sri Lankan Freshwater Fishes. Wildlife Conservation Society, Galle, 391 pp.
Deraniyagala, P.E.P. (1929) The Labyrinthici of Ceylon. Spolia Zeylanica, 15, 79–111.
Deraniyagala, P.E.P. (1945) New subspecies among Ophiocephali of Ceylon and India. Spolia Zeylanica, 24 (2), 93.
Deraniyagala, P.E.P. (1952) A coloured atlas of some vertebrates from Ceylon. Vol. 1. Fishes. Ceylon National Museum, Colombo, 147 pp.
Duncker, G. (1912) Die Süsswasserfische Ceylons. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, Beiheft 2, Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 29, 241–272.
Eken, G., Bennun, L., Brooks, T.M., Darwall, W., Fishpool, L.D.C., Foster, M., Knox, D., Langhammer, P., Matiku, P., Radford, E., Salaman, P., Sechrest, W., Smith, M.L., Spector, S. & Tordoff, A. (2004) Key Biodiversity Areas as Site Conservation Targets. BioScience, 54 (12), 1110–1118.
https://doi.org/10.1641/0006-3568(2004)054[1110:KBAASC]2.0.CO;2
Günther, A. (1861) Catalogue of the fishes in the British Museum. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Vol. 3. Gobiidae, Discoboli, Pediculati, Blenniidae, Labyrinthici, Mugilidae, Notacanthi. Taylor & Francis, London, 586 pp.
Hamilton, F. (1822) An account of the fishes found in the River Ganges and its branches. Constable, Edinburgh, 405 pp.
https://doi.org/10.5962/bhl.title.59540
IUCN (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, 148 pp.
IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. Second. IUCN, Gland, Switzerland and Cambridge, 32 pp.
IUCN (2019) The IUCN Red List of Threatened Species. Version 2019-1. Available from: http://www.iucnredlist.org (accessed 2 May 2019)
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, 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/msw054
Lalhlimpuia, D.V., Lalronunga, S. & Lalramliana, L. (2016) Channa aurantipectoralis, a new species of snakehead from Mizoram, north-eastern India (Teleostei: Channidae). Zootaxa, 4147 (3), 343–350.
https://doi.org/10.11646/zootaxa.4147.3.7
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.12410
Mendis, A.S. & Fernando, C.H. (1962) A guide to the freshwater fauna of Ceylon. Bulletin of the Fisheries Research Station, Ceylon, 12 (1), 1–160.
Miller, K.G., Kominz, M.A., Browning, J.V., Wright, J.D., Mountain, G.S., Katz, M.E., Sugarman, P.J., Cramer, B.S., Christie-Blick, N. & Pekar, S.F. (2005) The Phanerozoic Record of Global Sea-Level Change. Science, 310 (5752), 1293–1298.
https://doi.org/10.1126/science.1116412
MOE (2012) The National Red List 2012 of Sri Lanka; Conservation Status of the Fauna and Flora. Ministry of Environment, Colombo, 476 pp.
Munro, I.S.R. (1955) The Marine and Freshwater Fishes of Ceylon. Department of External Affairs, Canberra, 349 pp.
Musikasinthorn, P. (1998) Channa panaw, a new channid fish from the Irrawaddy and Sittang River basins, Myanmar. Ichthyological Research, 45 (4), 355–362.
https://doi.org/10.1007/BF02725188
Musikasinthorn, P. (2000) Channa aurantimaculata, a new channid fish from Assam (Brahmaputra River basin), India, with designation of a neotype for C. amphibeus (McClelland, 1845). Ichthyological Research, 47 (1), 27–37.
https://doi.org/10.1007/BF02674310
Myers, G. & Shapovalov, L. (1932) On the identity of Ophicephalus and Channa, two genera of labyrinth fishes. Peking Natural History Bulletin, 6, 33–37.
Ng, P.K.L. & Lim, K.K.P. (1990) Snakeheads (Pisces: Channidae): natural history, biology and economic importance. In: Chou, L.M. & Ng, P.K.L. (Eds.), Essays in Zoology. Department of Zoology, National University of Singapore, Singapore, pp. 127–152.
Pedraza-Marrón, C.del.R., Silva, R., Deeds, J., Van Belleghem, S.M., Mastretta-Yanes, A., Domínguez-Domínguez, O., Rivero-Vega, R.A., Lutackas, L., Murie, D., Parkyn, D., Bullock, L.H., Foss, K., Ortiz-Zuazaga, H., Narváez-Barandica, J., Acero, A., Gomes, G. & Betancur-R, R. (2019) Genomics overrules mitochondrial DNA, siding with morphology on a controversial case of species delimitation. Proceedings of the Royal Society B: Biological Sciences, 286 (1900), 20182924.
https://doi.org/10.1098/rspb.2018.2924
Pethiyagoda, R. (1991) Freshwater fishes of Sri Lanka. Wildlife Heritage Trust, Colombo, 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/BF00005666
Pethiyagoda, 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.
Rainboth, W. (1996) Fishes of the Cambodian Mekong. FAO, Rome, 265 pp.
Ramasamy, S.M. & Saravanavel, J. (2019) Drowned valleys of Vaigai and Tamiraparani rivers in the Gulf of Mannar region, India. Current Science, 16 (12), 1958–1960.
Rüber, L., Tan, H.H. & Britz, R. (2019) Snakehead (Teleostei: Channidae) diversity and the Eastern Himalaya biodiversity hotspot. Journal of Zoological Systematics and Evolutionary Research, 00, 1–31.
https://doi.org/10.1111/jzs.12324
Somasekaram, T., Perera, L.A.G., Perera, M.P., de Silva, M.B.G., Karunanayake, M.M. & Epitawatte, D.S. (1988) The National Atlas of Sri Lanka. Survey Department, Colombo. [1 atlas (x, 142 p.): col. ill., col. maps; 52 cm.]
Sudasinghe, 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.4158.2.7
Sudasinghe, H. (2017) Schistura madhavai, a new species of hill-stream loach from Sri Lanka, with redescription of S. notostigma (Teleostei: Nemacheilidae). Zootaxa, 4311 (1), 96–110.
https://doi.org/10.11646/zootaxa.4311.1.6
Sudasinghe, H. (2018) A new species of Schistura (Teleostei: Nemacheilidae) from the south-western lowlands of Sri Lanka. Zootaxa, 4422 (4), 478–492.
https://doi.org/10.11646/zootaxa.4422.4.2
Sudasinghe, H., Ranasinghe, R.H.T., Goonatilake, S.de.A. & Meegaskumbura, M. (2018a) A review of the genus Labeo (Teleostei: Cyprinidae) in Sri Lanka. Zootaxa, 4486 (3), 201–235.
https://doi.org/10.11646/zootaxa.4486.3.1
Sudasinghe, H., Herath, J., Pethiyagoda, R. & Meegaskumbura, M. (2018b) Undocumented translocations spawn taxonomic inflation in Sri Lankan fire rasboras (Actinopterygii, Cyprinidae). PeerJ, 6, e6084.
https://doi.org/10.7717/peerj.6084
Sudasinghe, H., Pethiyagoda, R., Maduwage, K. & Meegaskumbura, M. (2019a) The identity of the Sri Lankan Amblypharyngodon (Teleostei, Cyprinidae). ZooKeys, 820, 25–49.
https://doi.org/10.3897/zookeys.820.29632
Sudasinghe, H., Pethiyagoda, R. & Meegaskumbura, M. (2019b) A review of the genus Esomus in Sri Lanka (Teleostei: Cyprinidae). Ichthyological Exploration of Freshwaters, 00, 1–18.
https://doi.org/10.23788/IEF-1106
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.
Ward, 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 (1462), 1847–1857.
https://doi.org/10.1098/rstb.2005.1716
Willey, A. (1910) Notes on the fresh-water fisheries of Ceylon. Spolia Zeylanica, 7 (26), 88–105.