Abstract
The 22 Sri Lankan representatives of the family Platystictidae, all endemic to the island and belonging to the distinct endemic subfamily Platystictinae, are revised, and a new reconstruction of the phylogeny based on molecular characters is provided. Five new species are described: Ceylonosticta venusta sp. nov. (holotype ♂: Rambodde Falls, at the tunnel; Nuwara Eliya District, Central Province; N7.0489, E80.6961; 12-vii-2012; to be deposited at National Museum of Natural History, Colombo, Sri Lanka), C. inferioreducta sp. nov. (holotype ♂: Norton Bridge, stream on the B43 road 1.5km WNW of Norton Bridge; Nuwara Eliya District, Central Province; N6.9171, E80.5075; 28-vii-2009; to be deposited at National Museum of Natural History, Colombo, Sri Lanka), C. mirifica sp. nov. (holotype ♂: Uwella, primary forest on the road Uwella-Ratnapura, 11.5km NW of Balangoda; Ratnapura District, Sabaragamuwa Province; N6.6968, E80.6059; 16-vii-2012; to be deposited at National Museum of Natural History, Colombo, Sri Lanka), Platysticta secreta sp. nov. (holotype ♂: Hasalaka; Kandy District, Central Province; N7.3535, E80.9509; 31-v-1975; deposited at National Museum of Natural History, Smithsonian Institution, Washington, USA) and P. serendibica sp. nov. (holotype ♂: Kanneliya; Galle District, Southern Province; N6.2291, E80.3834; 8 & 9-vi-1975; deposited at National Museum of Natural History, Smithsonian Institution, Washington, USA). Additionally, a determination key, figures showing morphological details and coloration in life, as well as distribution maps for all species are presented. Based on molecular analysis of 21 taxa, the phylogeny of Platystictinae is presented and discussed from the zoogeographical and paleogeographical point of view. Sri Lankan species, traditionally placed in the genera Platysticta Selys and Drepanosticta Laidlaw / Ceylonosticta Fraser, separated into distinct clades within the subfamily as presently defined, but the monophyletic nature of the Platystictinae and its Sri Lankan endemicity is confirmed. For the South Indian species, formerly known as Platysticta deccanensis, morphological and molecular analyses demonstrated that it does not belong to the Sri Lankan clade and a new genus Indosticta gen. nov. is erected to accommodate it.
References
Aberer, A.J., Kobert, K. & Stamatakis, A. (2014) ExaBayes: Massively parallel Bayesian tree inference for the whole-genome era. Molecular Biology and Evolution, 31, 2553–2556.
http://dx.doi.org/10.1093/molbev/msu236Ali, J.R. & Aitchison, J.C. (2008) Gondwana to Asia: Plate tectonics, paleogeography and the biological connectivity of the Indian sub-continent from the Middle Jurassic through latest Eocene (166–35 Ma). Earth-Science Reviews 88 (3–4), 145–166.
http://dx.doi.org/10.1016/j.earscirev.2008.01.007Babu, R., Subramanian, K.A. & Supriya, N. (2013) Endemic Odonates of India. Records of the Zoological Survey of India, Occasional Paper No. 347, 1–60.
Bedjanič, M. (2002) Dragonflies collected in Sri Lanka during January and February 1995. Opuscula zoologica fluminensia, 205, 1–22.
Bedjanič, M. (2009) Drepanosticta starmuehlneri St. Quentin, 1972 from Sri Lanka, a synonym of D. lankanensis (Fraser, 1931) (Zygoptera: Platystictidae). Notulae odonatologicae, 7 (4), 38–39.
Bedjanič, M. (2010) Three new Drepanosticta species from Sri Lanka (Zygoptera: Platystictidae). Odonatologica, 39 (3), 195–215.
Bedjanič, M. (2012) On the synonymy of three endemic dragonfly species from Sri Lanka (Zygoptera: Platystictidae, Protoneuridae). Notulae odonatologicae, 7 (9), 77–80.
Bedjanič, M., Conniff, K. & De Silva Wijeyeratne, G. (2007) Gehan’s Photo Guide: A Photographic guide to the dragonflies of Sri Lanka. Jetwing Eco Holidays, Colombo, 248 pp.
Bedjanič, M., Conniff, K., van der Poorten, N. & Šalamun, A. (2014) Dragonfly fauna of Sri Lanka: distribution and biology, with threat status of its endemics. Pensoft Publishers, Sofia, 321 pp.
Biswas, S. (2008) Did biotic impoverishment facilitate phenomenal diversification in Sri Lanka? Current Science, 95 (8), 1021–1025.
Bossuyt, F., Meegaskumbura, M., Beenaerts, N., Gower, D.J., Pethiyagoda, R., Roelants, K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K., Ng, P.K.L., Schneider, C.J., Oommen, O.V. & Milinkovitch, M.C. (2004) Local endemism within the Western Ghats - Sri Lanka Biodiversity Hotspot. Science, 306, 479–481.
http://dx.doi.org/10.1126/science.1100167Chatterjee, S., Goswami, A. & Scotese, C.R. (2013) The longest voyage: Tectonic, magmatic, and paleoclimatic evolution of the Indian plate during its northward flight from Gondwana to Asia. Gondwana Research, 23 (1), 238–267.
http://dx.doi.org/10.1016/j.gr.2012.07.001de Fonseka, T. (2000) The dragonflies of Sri Lanka. Wildlife Heritage Trust, Colombo, 303 pp., color pls. 1–20 excl.
de Silva Wijeyeratne, G. (2015) A naturalist's guide to the butterflies and dragonflies of Sri Lanka. John Beaufoy Publishing, Oxford, 176 pp.
Dijkstra, K.-D.B., Bechly, G.S., Bybee, M., Dow, R.A., Dumont, H.J., Fleck, G., Garrison, R.W., Hämäläinen, M., Kalkman, V.J., Karube, H., May, M.L., Orr, A.G., Paulson, D., Rehn, A.C., Theischinger, G., Trueman, J.W.H., van Tol, J., von Ellenrieder, N. & Ware, J. (2013) The classification and diversity of dragonflies and damselflies (Odonata). In: Zhang, Z.-Q. (Ed.), Animal Biodiversity: An Outline of Higher level Classification and Survey of Taxonomic Richness (Addenda 2013). Zootaxa, 3703 (1), 36–45.
http://dx.doi.org/10.11646/zootaxa.3703.1.9Dijkstra, K.-D.B., Kalkman, V.J., Dow, R.A., Stokvis, F.R. & van Tol, J. (2014) Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata). Systematic Entomology, 39 (1), 68–96.
http://dx.doi.org/10.1111/syen.12035Doorenweerd, C. (2016) PhylOStack: a phylogenetic analyses pipeline in Linux Ubuntu 14.04 using OpenStack cloud computing. Available from: http://cdoorenweerd.github.io/PhylOStack/ (accessed 7 September 2016)
Dow, R.A. (2013) Drepanosticta burbachi spec. nov. from Sarawak, Borneo, a new species allied to D. dulitensis Kimmins, with notes on related species (Zygoptera: Platystictidae). Odonatologica, 42 (3), 203–210.
Dow, R.A. (2014) Telosticta iban sp. nov. from Sarawak (Odonata: Zygoptera: Platystictidae). Zootaxa, 3784 (1), 74–78.
http://dx.doi.org/10.11646/zootaxa.3784.1.4Dow, R.A., Afendy, A. & Rahman, H. (2016) Telosticta fugispinosa sp. nov. from Sabah (Odonata: Zygoptera: Platystictidae). Zootaxa, 4103 (4), 390–395.
http://dx.doi.org/10.11646/zootaxa.4103.4.7Dow, R.A. & Orr, A.G. (2012) Telosticta, a new damselfly genus from Borneo and Palawan (Odonata: Zygoptera: Platystictidae). The Raffles Bulletin of Zoology, 60 (2), 361–397.
Emiliyamma, K.G. (2014) Systematic studies on Odonata (Insecta) of southern Western Ghats. Records of the Zoological Survey of India, 114 (Part-1), 57–87.
Emiliyamma, K.G., Radhakrishnan, C. & Palot, M.J. (2007) Odonata (Insecta) of Kerala. Zoological Survey of India, Kolkata, 243 pp.
Erdelen, W.R. (2012) Conservation of biodiversity in a hotspot: Sri Lanka’s amphibians and reptiles. Amphibian and Reptile Conservation, 5 (2), 33–51.
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome coxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5), 294–299.
Foster, P.G. (2004) Modeling compositional heterogeneity. Systematic Biology, 53, 485–495.
http://dx.doi.org/10.1080/10635150490445779Fraser, F.C. (1924) A survey of the Odonata fauna of western India with special remarks on the genera Macromia and Idionyx and description of thirty new species. Records of the Indian Museum, 26 (5), 423–522, pls. 25–27 excl.
Fraser, F.C. (1931a) Indian dragonflies, XXXVII. The Journal of the Bombay Natural History Society, 35, 66–76, 1 pl. excl.
Fraser, F.C. (1931b) Indian dragonflies, XXXVIII. The Journal of the Bombay Natural History Society, 35, 325–341, pls. 1 & 2 excl.
Fraser, F.C. (1933a) The Fauna of British India including Ceylon and Burma: Odonata. Vol. I. Taylor & Francis, London, xiii + 423 pp.
Fraser, F.C. (1933b) The Platystictas of Ceylon (Order Odonata). Ceylon Journal of Science (B), 17 (3), 201–224.
Hagen, H.A. (1858) Synopsis der Neuroptera Ceylons. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 8, 471–488.
Hagen, H.A. (1859) Synopsis der Neuroptera Ceylons. (Pars II.). Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 9, 199–212.
Hebert, P.D.N., Penton, E.H., Burns, J.M., Janzen, D.H. & Hallwachs, W. (2004) Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America, 101 (41), 14812–14817. http://dx.doi.org/10.1073/pnas.0406166101
Huang, D.Y., Azar, D., Cai, C.Y. & Nel, A. (2015) New damselfly genera in the Cretaceous Burmese amber attributable to the Platystictidae and Platycnemididae Disparoneurinae (Odonata: Zygoptera). Cretaceous Research, 56, 237–243.
http://dx.doi.org/10.1016/j.cretres.2015.05.004Kalkman, V.J., Clausnitzer, V., Dijkstra, K.-D.B., Orr, A.G., Paulson, D.R. & van Tol, J. (2008) Global diversity of dragonflies (Odonata) in freshwater. Hydrobiologia, 595, 351–363.
http://dx.doi.org/10.1007/s10750-007-9029-xKatoh, K., Asimenos, G. & Toh, H. (2009) Multiple alignment of DNA sequences with MAFFT. Methods in Molecular Biology, 537, 39–64.
http://dx.doi.org/10.1007/978-1-59745-251-9_3Kimmins, D.E. (1966) A list of odonata types described by F. C. Fraser, now in the British Museum (Natural History). Bulletin of the British Museum (Natural History), Entomology, 18 (6), 173–227.
Kiran, C.G., Kalesh, S. & Kunte, K. (2015) A new species of damselfly, Protosticta ponmudiensis (Odonata: Zygoptera: Platystictidae) from Ponmudi Hills in the Western Ghats of India. Journal of Threatened Taxa, 7(5), 7146–7151.
http://dx.doi.org/10.11609/JoTT.o4145.7146-51Kirby, W.F. (1891) On some Neuroptera (Odonata) collected by Mr. E. E. Green in Ceylon. Proceedings of the Zoological Society of London, 1891, 203–206, pl. 20.
Kirby, W.F. (1894) Catalogue of the described Neuroptera Odonata (Dragonflies) of Ceylon, with Description of New Species. The Journal of the Linnean Society of London, Zoology, 24 (157), 545–566, pls. 41, 42 excl.
Laidlaw, F.F. (1915) Notes on oriental dragonflies in the Indian Museum. No. 3. Indian species of the "legion" Protoneura. Records of the Indian Museum, 11 (5), 387–391.
Laidlaw, F.F. (1917) A list of the dragonflies recorded from the Indian empire, with special reference to the collection of the Indian Museum. Part II. The family Agrionidae. The sections Podolestes, Platycnemis, Platysticta and Protoneura. Records of the Indian Museum, 13 (6), 321–348, pls. 13–14 excl.
Laidlaw, F.F. (1924) A catalogue of the dragonflies (Odonata) recorded from Ceylon, based on material collected by Mr. E. E. Green, with description of a new species. Spolia zeylanica, 12 (47/48), 335–374.
Laidlaw, F.F. (1951) A note on the Derivation of the Odonate Fauna of the Island of Ceylon. Entomological News, 62 (2), 77–83.
Lanfear R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution, 29 (6), 1695–1701.
http://dx.doi.org/10.1093/molbev/mss020Larsson, A. (2014) AliView: a fast and lightweight alignment viewer and editor for large data sets. Bioinformatics, 30 (22), 3276–3278.
http://dx.doi.org/10.1093/bioinformatics/btu531Li, J.-T., Li, Y., Klaus, S., Rao, D.-Q., Hillise, D.M. & Zhang, Y.-P. (2013) Diversification of rhacophorid frogs provides evidence for accelerated faunal exchange between India and Eurasia during the Oligocene. Proceedings of the National Academy of Sciences of the United States of America, 110 (9), 3441–3446.
http://dx.doi.org/10.1073/pnas.1300881110Lieftinck, M.A. (1940) On some Odonata collected in Ceylon, with description of new species and larvae. Ceylon Journal of Science (B), 22 (1), 79–117.
Lieftinck, M.A. (1955) Synopsis of the dragonflies (Odonata) of Ceylon. Zoologische mededeelingen, 34 (5), 67–87.
Lieftinck, M.A. (1971) Odonata from Ceylon. Entomologica Scandinavica Supplement, 1, 188–207.
Machado, A.B.M. (2009) Palaemnema brasiliensis spec. nov., first Platystictidae record from Brazil (Zygoptera). Odonatologica, 38 (3), 255–260.
Myers, N., Mitermeier, R.A., Mitermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858.
http://dx.doi.org/10.1038/35002501Naggs, F. & Raheem, D. (2005) Sri Lankan snail diversity: faunal origins and future prospects. Records of the Western Australia Museum, Supplement, 68, 11–29.
Orr, A.G. & Dow, R.A. (2015) Description of two final stadium platystictid larvae from Borneo, including that of Drepanosticta attala Lieftinck, identified using DNA barcoding (Odonata: Zygoptera: Platystictidae). Zootaxa, 3985 (4), 565–574.
Peña, C. & Malm, T. (2012) VoSeq: a Voucher and DNA Sequence Web Application. PLoS ONE, 7 (6), e39071.
http://dx.doi.org/10.1371/journal.pone.0039071Pethiyagoda, R. (2005) Introduction: Exploring Sri Lanka’s Biodiversity. The Raffles Bulletin of Zoology, 12 (Supplement), 1–4.
Phan, Q.T. & Kompier, T. (2016) A study of the genus Protosticta Selys, 1855, with descriptions of four new species from Vietnam (Odonata: Platystictidae). Zootaxa, 4098 (3), 529–544.
http://dx.doi.org/10.11646/zootaxa.4098.3.6Pyron, R.A., Dushantha Kandambi, H.K., Hendry, C.R., Pushpamal, V., Burbrink, F.T. & Somaweera, R. (2013) Genus-level phylogeny of snakes reveals the origins of species richness in Sri Lanka. Molecular Phylogenetics and Evolution, 66 (2013), 969–978.
http://dx.doi.org/10.1016/j.ympev.2012.12.004Rambaut, A. (2014) FigTree. Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed 7 September 2016)
Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer v1.6. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 7 September 2016)
Ratnasingham, S. & Hebert, P.D.N. (2007) BOLD: the barcode of life data system (www.barcodinglife.org). Molecular Ecology Notes, 7, 55–364.
http://dx.doi.org/10.1111/j.1471-8286.2007.01678.xRoelants, K., Jiang, J. & Bossuyt, F. (2004) Endemic ranid (Amphibia: Anura) genera in southern mountain ranges of the Indian subcontinent represent ancient frog lineages: Evidence from molecular data. Molecular Phylogenetics and Evolution, 2004 (31), 730–740.
http://dx.doi.org/10.1016/j.ympev.2003.09.011Schorr, M. & Paulson, D. (2015) World Odonata list. Available from: http://www.pugetsound.edu/academics/academic-resources/slater-museum/biodiversity-resources/dragonflies/world-odonata-list2/ (accessed 30 April 2016)
Selys-Longchamps, E.M. de (1860) Synopsis des Agrionines, derniere légion: Protoneura. Bulletin de l’ Academie royale de Belgique, des Lettres et des Beaux-Arts de Belgique, Series 2, 10, 431–462.
Selys-Longchamps, E.M. de, 1886. Revision du synopsis des Agrionines, 1e partie comprenant les légions Pseudostigma-Podagrion-Platycnemis et Protoneura. Mémoires Couronnés et autres Mémoires publiés par l'Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique, 38 (4), i–iv, 1–233.
Stamatakis, A. (2014) RAxML Version 8: A tool for Phylogenetic Analysis and Post-Analysis of Large Phylogenies. Bioinformatics, 30 (9), 1312–1313.
http://dx.doi.org/10.1093/bioinformatics/btu033St. Quentin, D. (1972) Results of the Austrian Ceylonese Hydrobiological Mission 1970 of the 1st Zoological Institute of the University of Vienna (Austria) and the Department of Zoology of the Vidyalankara University of Ceylon, Kelaniya. Part VI: A new Drepanosticta from Ceylon (Order: Odonata; Insecta). Bulletin of the Fisheries Research Station, Sri Lanka (Ceylon), 23 (1&2), 137–139.
Subramanian, K.A. (2009) Dragonflies of India—A Field Guide. Vigyan Prasar, New Delhi, 168 pp.
Subramanian, K.A. (2013) Platysticta deccanensis. The IUCN Red List of Threatened Species 2013: e.T175200A7121240.
http://dx.doi.org/10.2305/IUCN.UK.2011-1.RLTS.T175200A7121240.enSubramanian, K.A., Kakkassery, F. & Nair, M.V. (2011) Chapter 5. The status and distribution of dragonflies and damselflies (Odonata) of the Western Ghats. In: Molur, S., Smith, K.G., Daniel, B.A. & Darwall, W.R.T. (Comp.), The Status and Distribution of Freshwater Biodiversity in the Western Ghats, India. IUCN and Coimbatore, Cambridge and Gland & Zoo Outreach Organisation, Coimbatore, Tamil Nad, pp. 63–72,
Suhling, F., Sahlen, G., Gorb, S., Kalkman, V.J., Dijkstra, K.-D.B. & van Tol, J. (2015) Chapter 35. Order Odonata. In: Thorp, J.H., Rogers, D.C. (Eds.), Ecology and General Biology: Thorp and Covich's Freshwater Invertebrates. Ecology and General Biology, 2015, pp. 893–932.
http://dx.doi.org/10.1016/b978-0-12-385026-3.00035-8Van Bocxlaer, I., Biju, S.D., Willaert, B., Giri, V.B., Shouche, Y.S. & Bossuyt, F. (2011) Mountain-associated clade endemism in an ancient frog family (Nyctibatrachidae) on the Indian subcontinent. Molecular Phylogenetics and Evolution, 62 (3), 839–847.
http://dx.doi.org/10.1016/j.ympev.2011.11.027van Hinsbergen, D.J.J., Lippert, P.C., Dupont-Nivet, G., McQuarrie, N., Doubrovine, P.V., Spakman, W. & Torsvik, T.H. (2012) Greater India Basin hypothesis and a two-stage Cenozoic collision between India and Asia. Proceedings of the National Academy of Sciences of the United States of America, 109 (20), 7659–7664.
http://dx.doi.org/10.1073/pnas.1117262109van Tol, J. (2005) Revision of the Platystictidae of the Philippines (Odonata), excluding the Drepanosticta halterata-group, with description of twenty-one new species. Zoologische Mededelingen, 79 (2), 195–282.
van Tol, J. (2009) Phylogeny and biogeography of the Platystictidae (Odonata). Ph.D. thesis, University of Leiden, Leiden, x + 294 pp.
Varghese, A.P., Nikesh, P.R. & Mathew, J. (2014) Odonata (Insecta) diversity of Salim Ali Bird Sanctuary and its adjacent areas in Thettekkad, Kerala, India. Journal of Threatened Taxa, 6 (6), 5887–5893.
http://dx.doi.org/10.11609/JoTT.o3395.5887-93Villanueva, R.J.T., van der Ploeg, J. & van Weerd, M. (2011) Drepanosticta hamalaineni spec. nov and Sulcosticta sierramadrensis spec. nov from the Northern Sierra Madre Natural Park, Luzon, the Philippines (Zygoptera: Platystictidae). Odonatologica, 40 (4), 333–338.
Villanueva, R.J.T. & Schorr, M. (2011) Two new damselfly species from Polillo Island, Philippines (Odonata: Platystictidae). Zootaxa, 3017, 46–50.
Watson, J.A.L. & O’Farrell, A.F. (1991) Odonata (dragonflies and damselflies). In: Naumann, I.D., Carne, P.B., Lawrence, J.F., Nielsen, E.S., Spradbery, J.P., Taylor, R.W., Whitten, M.J. & Littlejohn, M.J. (Eds.), The insects of Australia. 2nd Edition. Melbourne University Press, Melbourne, pp. 294–310.
Zheng, D., Zhang, Q., Chang, S.-C. & Wang, B. (2016) A new damselfly (Odonata: Zygoptera: Platystictidae) from mid-Cretaceous Burmese amber. Cretaceous Research, 63, 142–147.
http://dx.doi.org/10.1016/j.cretres.2016.03.006Yu, X. & Bu, W. (2009) Description of two new damselflies, Protosticta zhengi and Sinosticta sylvatica, from China (Odonata: Zygoptera: Platystictidae). Zootaxa, 2245, 54–58.