Abstract
The taxonomy and biogeography of the western representatives of the largely Papuan-Australian genus Hemicordulia are discussed and compared with other alate fauna including butterflies, birds, bats and other dragonflies. Specimens from Malawi, Mozambique, Réunion, South Africa, Tanzania and Uganda were compared with Indian specimens of H. asiatica, with which they were previously regarded conspecific. They are found to be distinct and are described as the continental H. africana n. sp. and those from Réunion as H. atrovirens n. sp. The three species were compared with H. similis of Madagascar and H. virens of Mauritius. Insufficient material of the Seychelles taxon H. similis delicata was available; it may represent another insular endemic species. The distribution of Hemicordulia is discussed in the light of the dispersal capacity of Odonata and the biogeography of taxa with similar distributions in the region, with an emphasis on the survival of ‘oceanic’ species on the continent. Recent (i.e. in the last few million years) trans-oceanic airborne dispersal aided by westward storms, is the most likely explanation for the distribution of the genus in Africa and the Indian Ocean islands, as well as for other winged animals of Asian affinities in the region. The world range of Hemicordulia is largely insular, broadly excluding continents, and H. africana n. sp. demonstrates ‘inverted insularity’: all continental sites are in proximity to large water bodies, such as the great African lakes. This pattern may be related to the climatological instability of these sites, which offer suitable cool habitat where competition is (temporarily) reduced. Hemicordulia prefer cool conditions, but may be vulnerable to overheating and competition with more warm-adapted species.
References
Ackery, P.R. & Vane-Wright, R.I. (1984) Milkweed butterflies, their cladistics and biology, British Museum (Natural History), London, 425 pp.
Armstrong, J.S. (1978) Colonisation of New Zealand by Hemicordulia australiae, with notes on its displacement of the indigenous Procordulia grayi (Odonata: Corduliidae). New Zealand Entomologist, 6, 381–384.
Asahina, S. (1940) Odonata-Anisoptera of Micronesia. Tenthredo, 3, 1–23.
Asahina, S. (1970) Burmese Odonata collected by Dr. Arthur Svihla with supplementary notes on Asiatic Ceriagrion species. Japanese Journal of Zoology, 16, 99–126.
Asahina, S. (1987) A list of Odonata recorded from Thailand, Part XVIII. Corduliidae 2. Kontyû, 55, 699–720.
Austin, J.J., Arnold, E.N. & Jones, C.G. (2004) Reconstructing an island radiation using ancient and recent DNA: the extinct and living day geckos (Phelsuma) of the Mascarene islands. Molecular Phylogenetics and Evolution, 31, 109–122.
Belle, J. (1988) A record of the Old World species Tramea basilaris burmeisteri Kirby from Suriname (Odonata: Libellulidae). Zoologische Mededelingen, 62, 1–3.
Bergmans, W. (1990) Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 3. The genera Scotonycteris Matschie, 1894, Casinycteris Thomas, 1910, Pteropus Brisson, 1762, and Eidolon Rafinesque, 1815. Beaufortia, 40, 111–177.
Bergmans, W. (1997) Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899, and Megaloglossus Pagensteicher, 1885; general remarks and conclusions; annex: key to all species. Beaufortia, 47, 11–90.
Blackman, R.A.A. & Pinhey, E. (1967) Odonata of the Seychelles and other Indian Ocean island groups, based primarily on the Bristol University Expedition of 1964–1965. Arnoldia Rhodesia, 12(3), 1–38.
Buden, D.W. & Paulson, D.R. (2003) The Odonata of Kosrae, Eastern Caroline Islands, Micronesia. Pacific Science, 57, 399–407.
Chippindale, P.T., Davé, V.K., Whitmore, D.H. & Robinson, J.V. (1999) Phylogenetic relationships of North American damselflies of the genus Ischnura (Odonata: Zygoptera: Coenagrionidae) based on sequences of three mitochondrial genes. Molecular Phylogenetics and Evolution, 11, 110–121.
Clausnitzer, V. (2003) Teinobasis alluaudi Martin, 1896 from mainland Africa: notes on ecology and biogeography (Zygoptera: Coenagrionidae). Odonatologica, 32, 321–334.
Colgan, D.J. & da Costa, P. (2002) Megachiropteran evolution studied with 12S rDNA and c-mos DNA sequences. Journal of Mammalian Evolution, 9, 3–22.
Corbet, P.S. (1979) Pantala flavescens (Fabricius) in New Zealand (Anisoptera: Libellulidae). Odonatologica, 8, 115– 121.
Corbet, P.S. (1999) Dragonflies: behaviour and ecology of Odonata, Harley Books, Colchester, 829 pp.
Cordero Rivera, A., Lorenzo Carballa, M.O., Utzeri, C. & Vieira, V. (2005) Parthenogenetic Ischnura hastata (Say), widespread in the Azores (Zygoptera: Coenagrionidae). Odonatologica, 34, 1–9.
Couteyen, S. (2006) Etude de l’explotation des sites de reproduction par les Anisoptères à l’île de la Réunion (Odonata). Bulletin de la Société entomologique de France, 111, 65–71.
Couteyen, S. & Papazian, M. (2000) Contribution à la connaissance des Odonates de la Réunion 3. Hemicordulia asiatica Sélys, 1878, une espèce nouvelle pour l’île (Odonata: Corduliidae). Martinia, 16, 107–110.
Diamond, A.W. & Hamilton, A.C. (1980) The distribution of forest passerine birds an Quaternary climatic change in tropical Africa. Journal of Zoology, London, 191, 379–402.
Dijkstra, K.-D.B. (2005) Taxonomy and identification of the continental African Gynacantha and Heliaeschna (Odonata: Aeshnidae). International Journal of Odonatology, 8, 1–32.
Dijkstra, K.-D.B., Clausnitzer, V. & Martens, A. (2007) Tropical African Platycnemis damselflies and the biogeographical significance of a new species from Pemba Island, Tanzania (Odonata: Platycnemididae). Systematics and Biodiversity (in press).
Donnelly, N. (1994) Thailand collecting, July 1994. Malangpo, 11, 83–86.
Donnelly, T.W. & Parr, M.J. (2003) Odonata, Dragonflies and Damselflies. In: Goodman, S.M. & Benstead, J.P. (Ed.), The natural history of Madagascar. University of Chicago Press, Chicago and London, 645–654.
Dumont, H.J. (1980) The dragonfly fauna of Egypt and the role of the Nile in its origin and composition. Water Supply and Management, 4, 29–34.
Dumont, H.J. & Verschuren, D. (1991) Atypical ecology of Pantala flavescens (Fabricius) on Easter Island (Anisoptera: Libellulidae). Odonatologica, 20, 45–51.
Farkas, T. (1985) Notes on the biology and taxonomy of the Madagascar robin-chat (Copsychus albospecularis Eydoux & Gervais). In: Schuchmann, K.-L. (Ed.), Proceedings of the International Symposium on African Vertebrates: systematics, phylogeny and evolutionary ecology. Museum Alexander Koenig, Bonn, 375–394.
Feng, H.-Q., Wu, K.-M., Ni, Y.-X., Cheng, D.-F. & Guo, Y.Y. (2006) Nocturnal migration of dragonflies over the Bohai Sea in northern China. Ecological Entomology, 31, 511–520.
Fjeldså, J. & Lovett, J.C. (1997) Geographical patterns of old and young species in African forest biota: the significance of specific montane areas as evolutionary centres. Biodiversity and Conservation, 6, 325–346.
Fonseka, T. de (2000) The dragonflies of Sri Lanka, WHT Publications, Colombo, 303pp.
Fraser, F.C. (1936) The Fauna of British India, Odonata. Vol. III, Taylor and Francis, London, 461pp.
Fraser, F.C. (1949) Notes on the Odonata of Mauritius. 1. The genus Hemicordulia Selys. Proceedings of the Royal entomological Society of London (B), 18, 130–134.
Fraser, F.C. (1950) The Odonata of Mauritius, suborder Anisoptera, and description of a new species of Zygoptera. Proceedings of the Royal entomological Society of London (B), 19, 70–77.
Gerlach, J. (2003) The biodiversity of the granitic islands of Seychelles. Phelsuma Supplement, 11(B), 1–47.
Giannini, N.P. & Simmons, N.B. (2003) A phylogeny of megachiropteran bats (Mammalia: Chiroptera: Pteropodidae) based on direct optimization analysis of one nuclear and four mitochondrial genes. Cladistics, 19, 496–511.
Gibon, F.-M. (2003) Trichoptera, Caddisflies. In: Goodman, S.M. & Benstead, J.P. (Ed.), The natural history of Madagascar. University of Chicago Press, Chicago and London, 740–748.
Gillespie, R.G. & Roderick, G.K. (2002) Arthropods on islands: colonization, speciation, and conservation. Annual Review of Entomology, 47, 595–632.
Grand, D. (2004) Compte rendu odonatologique d’un voyage à l’île de la Réunion. Martinia, 20, 67–75.
Graves, T. (1999) East African Odonata. WDA’s Agrion, 3, 15–16.
Griffiths, C.J. (1993) The geological evolution of East Africa. In: Lovett, J.C. & Wasser, S.K. (Eds), Biogeography and ecology of the rain forests of eastern Africa. Cambridge University Press, Cambridge, 9–21.
Groombridge, J.J., Jones, C.G., Nichols, R.A., Carlton, M. & Bruford, M.W. (2004) Molecular phylogeny and morphological change in the Psittacula parakeets. Molecular Phylogenetics and Evolution, 31, 96–108.
Hamilton, A., Taylor, D. & Howard, P. (2001) Hot spots in African forests as Quaternary refugia. In: Weber, W., White, L.J.T., Vedder, A. & Naughton-Treves, L. (Eds), African rain forest ecology and conservation. Yale University Press, New Haven and London, 57–67.
Hamilton, A.C. (1981) The Quaternary history of African forests: its relevance to conservation. African Journal of Ecology, 19, 1–6.
Haritonov, A.Y. & Malikova, E.I. (1998) Odonata of the Russian Far East: a summary. Odonatologica, 27, 375–381.
Jansa, S.A., Goodman, S.M. & Tucker, P.K. (1999) Molecular phylogeny and biogeography of the native rodents of Madagascar (Muridae: Nesomyinae): a test of the single-origin hypothesis. Cladistics, 15, 253–270.
Jønsson, K.A. & Fjeldså, J. (2006) A phylogenetic supertree of oscine passerine birds (Aves: Passeri). Zoologica Scripta, 35, 149–186.
Juste B., J., Álvarez, Y., Tabarés, E., Garrido-Pertierra, A., Ibáñez, C. & Bautista, J.M. (1999) Phylogeography of African fruitbats (Megachiroptera). Molecular Phylogenetics and Evolution, 13, 596–604.
Keith, S. (1980) Origins of the avifauna of the Malagasy Region. In: Johnson, D.N. (Ed.), Proceedings of the fourth pan- African ornithological congress. Southern African Ornithological Society, Johannesburg, 99–108.
Kirby, W.F. (1890) A synonymic catalogue of Neuroptera Odonata, or dragonflies. With an appendix of fossil species, Gurney & Jackson, London, 202pp.
Lieftinck, M.A. (1942) The dragonflies (Odonata) of New Guinea and neighbouring islands. Part VI. Results of the Third Archbold Expedition 1938–’39 and the Le Roux Expedition 1939 to Netherlands New Guinea (I. Anisoptera). Treubia, 18, 441–608.
Lieftinck, M.A. (1962) Insects of Micronesia Odonata. Insects of Micronesia, 5, 1–95.
Lindley, P. (1974) The dragonflies of Korhogo, Ivory Coast. Bulletin de l’Institut fondamental d’Afrique noire (A), 36, 682–698.
Louette, M. (1987) Size increase as a result of successive bird colonisations: a hypothesis. Revue de Zoologie Africaine, 101, 257–264.
Lovett, J.C. (1993) Climatic history and forest distribution in eastern Africa. In: Lovett, J.C. & Wasser, S.K. (Eds), Biogeography and ecology of the rain forests of eastern Africa. Cambridge University Press, Cambridge, 23–29.
Machet, P. & Duquef, M. (2004) Un visiteur inattendu, et de taille!... Hemianax ephippiger (Burmeister, 1839) capture a la Guyane francaise. Martinia, 20, 121–124.
Martin, R. (1896) Odonates des iles Séchelles. Memoires de la Société Zoologique de France, 9, 101–112.
Martin, R. (1907) Cordulines. Collections Zoologiques du Baron Edm. de Selys Longchamps. Catalogue Systématique et Descriptif, 17, 1–94.
Martin, R. (1914) Odonata, fam. Libellulidae, subfam. Corduliinae. Genera Insectorum, 155, 1–32.
McGlone, M.S. (2005) Goodbye Gondwana. Journal of Biogeography, 32, 739–740.
Meurgey, F. (2004) Premiere observation d’Anax junius (Drury, 1773) en France (Odonata, Anisoptera, Aeshnidae) Naturelle de Nantes. Martinia, 20, 13–15.
Moore, N.W. (1993) Behaviour of imaginal Pantala flavescens (Fabr.) on Easter Island (Anisoptera: Libellulidae). Odo natologica, 22, 71–76.
Moyle, R.G. & Marks, B.D. (2006) Phylogenetic relationships of the bulbuls (Aves: Pycnonotidae) based on mitochondrial DNA sequence data. Molecular Phylogenetics and Evolution, 40, 687–695.
Muñoz, J., Felicísimo, A.M., Cabezas, F., Burgaz, A.R. & Martínez, I. (2004) Wind as a long-distance dispersal vehicle in the Southern Hemisphere. Science, 304, 1144–1147.
O’Brien, J. & Hayden, T.J. (2004) The application of genetic research to the conservation of fruit bats in the western Indian Ocean. Phelsuma, 12, 141–146.
Paulian, R. & Viette, P. (2003) An introduction to terrestrial and freshwater invertebrates. In: Goodman, S.M. & Benstead, J.P. (Ed.), The natural history of Madagascar. University of Chicago Press, Chicago and London, pp. 503– 511.
Paulson, D.R. & Buden, D.W. (2003) The Odonata of Pohnpei, Eastern Caroline Islands, Micronesia. International Journal of Odonatology, 6, 39–64.
Pellow, K. (1999) Common green darner Anax junius (Drury) in Cornwall and Isles of Scilly – the first British and European records. Journal of the British Dragonfly Society, 15, 21–22.
Pinhey, E. (1961) A survey of the dragonflies (order Odonata) of Eastern Africa. British Museum (Natural History), London, 214pp.
Pinhey, E. (1962) A descriptive catalogue of the Odonata of the African continent. Publicacoes Culturais Companhia de Diamantes de Angola, 59, 11–322.
Pinhey, E. (1966) Check-list of dragonflies (Odonata) from Malawi, with description of a new Teinobasis Kirby. Arnoldia Rhodesia, 33(2), 1–24.
Pinhey, E. (1976) Further notes on the Odonata of Mauritius. Arnoldia Rhodesia, 8(6), 1–6.
Pinhey, E. (1981) Checklist of the Odonata of Mocambique. Occasional Papers. National Museum of Southern Rhodesia (B), 6, 557–631.
Pinhey, E. (1984) A checklist of the Odonata of Zimbabwe and Zambia. Smithersia, 3, 1–64.
Pinhey, E. (1985) A survey of the dragonflies (Odonata) of South Africa. Part 2. Anisoptera. Journal of the Entomological Society of Southern Africa, 48, 1–48.
Prasad, M. & Varshney, R.K. (1995) A check-list of the Odonata of India including data on larval studies. Oriental Insects, 29, 385–428.
Queiroz, A. de (2005) The resurrection of oceanic dispersal in historical biogeography. Trends in Ecology and Evolution, 20, 68–73.
Rambur, J.P. (1842) Histoire naturelle des insectes névroptères. Roret’s Suite à Buffon, Paris, 534 pp.
Raxworthy, C.J., Forstner, M.R.J. & Nussbaum, R.A. (2002) Chameleon radiation by oceanic dispersal. Nature, 415, 784–787.
Samways, M.J. & Osborn, R. (1998) Divergence in a transoceanic circumtropical dragonfly on a remote island. Journal of Biogeography, 25, 935–946.
Sanmartín, I. & Ronquist, F. (2004) Southern Hemisphere biogeography inferred by event-based models: plant versus animal patterns. Systematic Biology, 53, 216–243.
Scholz, C.A. & Rosendahl, B.R. (1988) Low lake stands in Lake Malawi and Tanganyika, East Africa, delineated with multifold seismic data. Science, 240, 1645–1648.
Selys-Longchamps, E. de (1870) Sous-famille des Cordulines, Sélys (1). In: Comptes-rendus des séances de la société entomologique de Belgique: Assemblée mensuelle du 5 novembre 1870. Annales de la Société entomologique de Belgique, 14, iv–vii.
Selys-Longchamps, E. de (1871) Synopsis des Cordulines. Bulletins de l’Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique (2), 31, 238–565 (bulletin); 5–128 (reprint).
Selys-Longchamps, E. de (1878) Secondes additions au Synopsis des Cordulines. Bulletins de l’Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique (2), 45, 183–222 (bulletin); 5–44 (reprint).
Shapiro, B., Sibthorpe, D., Rambaut, A., Austin, J., Wragg, G.M., Bininda, E., Olaf, R.P., Lee, P.L.M. & Cooper, A. (2002) Flight of the dodo. Science, 295, 1683.
Stager, J.C. (2001) Climatic change and African rain forests in the twenty-first century. In: Weber, W., White, L.J.T., Vedder, A. & Naughton-Treves, L. (Eds), African rain forest ecology and conservation. Yale University Press, New Haven and London, 140–147.
Thomassen, H.A. (2005) Evolutionary development of echolocation in swiftlets (Aves: Apodidae) in association with their biogeographical distribution. In: Thomassen, H.A., Swift as sound: design and evolution of the echolocation system in swiftlets (Apodidae: Collocaliini). PhD thesis, Leiden University, Leiden, 157–174.
Thomassen, H.A., den Tex, R.-J., de Bakker, M.A.G. & Povel, G.D.E. (2005) Phylogenetic relationships amongst swifts and swiftlets: a multi locus approach. Molecular Phylogenetics and Evolution, 37, 264–277.
Times Books (1997) The Times atlas of the world – ninth comprehensive edition, Times Books, London, 218pp.
Tol, J. van (1997) The genus Procordulia Martin in western Malesia (Odonata, Corduliidae). Tijdschrift voor Entomologie, 140, 133–146.
Torres, E., Lees, D.C., Vane-Wright, R.I., Kremen, C., Leonard, J.A. & Wayne, R.K. (2001) Examining monophyly in a large radiation of Madagascan butterflies (Lepidoptera: Satyrinae: Mycalesina) based on mitochondrial DNA data. Molecular Phylogenetics and Evolution, 20, 460–473.
Tsuda, S. (2000) A Distributional List of World Odonata 2000, Private publication, Osaka, 430pp.
Tuxen, S.L. (1976) Odonata. The Zoology of Iceland 3(39a): 1–7.
Ugai, S. (1996) The first migrate record of Hemianax ephippiger from Japan. Tombo, 39, 45–46.
Warren, B.H., Bermingham, E., Bowie, R.C.K., Prys-Jones, R.P. & Thébaud, C. (2003) Molecular phylogeography reveals island colonization history and diversification of western Indian Ocean sunbirds (Nectarinia: Nectariniidae). Molecular Phylogenetics and Evolution, 29, 67–85.
Wells, N.A. (2003) Some hypotheses on the Mesozoic and Cenozoic paleoenvironmental history of Madagascar. In: Goodman, S.M. & Benstead, J.P. (Ed.), The natural history of Madagascar. University of Chicago Press, Chicago and London, 16–34.
White, F. (1981) The history of the Afromontane archipelago and the scientific need for its conservation. African Journal of Ecology, 19, 33–54.
White, F. (1993) Refuge theory, ice-age aridity and the history of tropical biotas: an essay in plant geography. Fragmenta Floristica et Geobotanica Supplement, 2, 385–409.
Wildermuth, H. (2006) Verhaltensgesteuerte Thermoregulation bei Somatochlora flavomaculata (Odonata: Corduliidae). Libellula, 25, 31–46
Zakharov, E.V., Smith, C.R., Lees, D.C., Cameron, A., Vane-Wright, R.I. & Sperling, F.A.H. (2004) Independent gene phylogenies and morphology demonstrate a Malagasy origin for a wide-ranging group of swallowtail butterflies. Evolution, 58, 2763–2782.