Abstract
The globally distributed ant genus Tetramorium Mayr is especially diverse in the Afrotropical, Oriental, and Indo-Australian regions, while the Malagasy fauna, by contrast, seemed to be comparatively species-poor. However, recent ant inventories in Madagascar and its surrounding island systems have generated an immense amount of new material. As a result, there is now a great need of an updated taxonomic revision for the genus Tetramorium in this region. The present study represents an introduction to the genus, and treats the species groups encountered in the Malagasy region. All former species groups were redefined, and several new groups were established, in order to accommodate the substantial amount of new material. We propose 14 species groups of Tetramorium for this region. Morphological boundaries between these groups and their biogeographic affinities are discussed, and an illustrated identification key to species groups is provided. Diagnoses of all species groups are presented, as well as images of typical group members. Additionally, the species level taxonomy of the T. bicarinatum, T. obesum, T. sericeiventre and T. tosii species groups is revised with descriptions and images of all species, and an identification key to the species of the T. bicarinatum group is provided. No nomenclatorial changes are documented from the T. bicarinatum, T. obesum, and T. tosii groups. However, within the T. sericeiventre species group, one new species, T. mahafaly sp. n., is described and T. quadrispinosum Emery, 1886 and all its former synonyms are proposed as junior synonyms of T. sericeiventre Emery, 1877. At present, there are still 39 valid species of Tetramorium for the Malagasy region, but this number is expected to increase significantly with upcoming taxonomic revisions of the species groups not revised in this study.References
Arnold, G. (1926) A monograph of the Formicidae of South Africa. Appendix. Annals of the South African Museum, 23, 191–295.
Astruc, C., Malosse, C. & Errard, C. (2001) Lack of intraspecific aggression in the ant Tetramorium bicarinatum: a chemical hypothesis. Journal of Chemical Ecology, 27, 1229–1248.
Bernard, F. (1948) Les insectes sociaux du Fezzân. Comportement et biogéographie. In Mission scientifique du Fezzân (1944–1945). Zoologie, 5, 87–200.
Blard, F., Dorow, W.H.O. & Delabie, J.H.C. (2003) Les fourmis de l'ile de la Reunion (Hymenoptera: Formicidae). Bulletin de la Société Entomologique de France, 108, 127–137.
Bolton, B. (1976) The ant tribe Tetramoriini (Hymenoptera: Formicidae). Constituent genera, review of smaller genera and revision of Triglyphothrix Forel. Bulletin of the British Museum (Natural History) Entomology, 34, 281–379.
Bolton, B. (1977) The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Oriental and Indo-Australian regions, and in Australia. Bulletin of the British Museum (Natural History) Entomology, 36, 67–151.
Bolton, B. (1979) The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Malagasy region and in the New World. Bulletin of the British Museum (Natural History) Entomology, 38, 129–181.
Bolton, B. (1980) The ant tribe Tetramoriini (Hymenoptera: Formicidae). The genus Tetramorium Mayr in the Ethiopian zoogeographical region. Bulletin of the British Museum (Natural History) Entomology, 40, 193–384.
Bolton, B. (1985) The ant genus Triglyphothrix Forel a synonym of Tetramorium Mayr. (Hymenoptera: Formicidae). Journal of Natural History, 19, 243–248.
Bolton, B. (1995) A new general catalogue of the ants of the world. Cambridge: Harvard University Press, 504 pp.
Bolton, B., Alpert, G. & Ward, P.S. (2007) Bolton's Catalogue of Ants of the World: 1758–2005. Cambridge: Harvard University Press.
Bolton, B. (2010) Bolton's catalogue and synopsis. Global Ant Project. Available from http://gap.entclub.org, version 1 July 2010 [accessed 1 January 2011].
Brown, W.L. (1957) Is the ant genus Tetramorium native in North America? Breviora, 72, 1–8.
Creighton, W.S. (1950) The ants of North America. Bulletin of the Museum of Comparative Zoology, 104, 1–585.
Brown, W.L. (1974) A remarkable new island isolate in the genus Proceratium (Hymenoptera: Formicidae). Psyche, 81, 70–83.
Csösz, S., Radchenko, A. & Schulz, A. (2007) Taxonomic revision of the Palaearctic Tetramorium chefketi species complex (Hymenoptera: Formicidae). Zootaxa, 1405, 1–38.
Csösz, S. & Schulz, A. (2010) A taxonomic review of the Palaearctic Tetramorium ferox species-complex (Hymenoptera, Formicidae). Zootaxa, 2401, 1–29.
Deyrup, M., Davis, L. & Cover, S. (2000) Exotic ants in Florida. Transactions of the American Entomological Society, 126, 293–326.
Donisthorpe, H.S.J.K. (1946) New species of ants (Hym., Formicidae) from the island of Mauritius. Annals and Magazine of Natural History, 11, 776–782.
Donisthorpe, H.S.J.K. (1948) A fourth installment of the Ross Collection of ants from New Guinea. Annals and Magazine of Natural History, 12, 131–143.
Donisthorpe, H.S.J.K. (1949) A new Camponotus from Madagascar and a small collection of ants from Mauritius. Annals and Magazine of Natural History, 12, 271–275.
Dorow, W.H.O. (1996) Review and bibliography of the ants of the Seychelles (Hymenoptera: Formicidae). Journal of African Zoology, 110, 73–94.
Emery, C. (1877) Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte prima. Formiche provenienti dall Viaggio dei signori Antinori, Beccari e Issel nel Mar Rosso e nel paese dei Bogos. Annali del Museo Civico di Storia Naturale Giacomo Doria (Genova), 9, 363–381.
Emery, C. (1886) Alcune formiche africane. Bollettino della Societa Entomologica Italiana, 18, 355–366.
Emery, C. (1889) Viaggio di Leonardo Fea in Birmania e regioni vicine. XX. Formiche di Birmania e del Tenasserim raccolte da Leonardo Fea (1885–87). Annali del Museo Civico di Storia Naturale Giacomo Doria (Genova), 7, 485–520.
Emery, C. (1893) Voyage de M. E. Simon à l'île de Ceylan (janvier - février 1892). 3e Mémoire. Formicides. Annales de la Société Entomologique de France, 62, 239–258.
Emery, C. (1895a) Voyage de M. E. Simon dans l'Afrique australe (janvier-avril 1893). Formicides. Annales de la Société Entomologique de France, 64, 15–56.
Emery, C. (1895b) Mission scientifique de M. Ch. Alluaud dans le territoire de Diego-Suarez (Madagascar-Nord). (Avril-août 1893). Annales de la Société entomologique de Belgique, 39, 336–345.
Emery, C. (1899) Formiche di Madagascar raccolte dal Sig. A. Mocquerys nei pressi della Baia di Antongil (1897–1898). Bollettino della Societa Entomologica Italiana, 31, 263–290.
Emery, C. (1914) Les fourmis de la Nouvelle-Calédonie et des îles Loyalty. In: Sarasin, F. & Roux, J. (Eds), Nova Caledonia. Zoologie. 1, Wiesbaden, pp. 393–437.
Evenhuis, N.L. (2009) The insect and spider collections of the world website. Available from http://hbs.bishopmuseum.org/codens [accessed 1 January 2010].
Fisher, B.L. (1996) Origins and affinities of the ant fauna of Madagascar. In: Lourenço, W.R. (Ed), Biogéographie de Madagascar. Editions ORSTOM, Paris, pp. 457–465.
Fisher, B.L. (1997) Biogeography and ecology of the ant fauna of Madagascar (Hymenoptera: Formicidae). Journal of Natural History, 31, 269–302.
Fisher, B.L. (1998) Ant diversity patterns along an elevational gradient in the Reserve Speciale d'Anjanaharibe-Sud and on the western Masoala Peninsula , Madagascar. Fieldiana: Zoology, 90, 39–67.
Fisher, B.L. (1999a) Improving inventory efficiency: a case study of leaf litter ant diversity in Madagascar. Ecological Applications, 9, 714–731.
Fisher, B.L. (1999b) Ant diversity patterns along an elevational gradient in the Reserve Naturelle Integrale d'Andohahela, Madagascar. Fieldiana: Zoology, 94, 129–147.
Fisher, B.L. (2000a) Ant inventories along elevational gradients in tropical wet forests in Eastern Madagascar. In: Agosti, D., Majer, J., Alonso, A. & Schultz, T. (Eds), Sampling ground-dwelling ants: case studies from the worlds' rain forests. Curtin University School of Environmental Biology Bulletin, Perth, Australia, pp. 41–49.
Fisher, B.L. & Girman, D.J. (2000b) Biogeography of ants in eastern Madagascar. In: Lourenco, W.R. & Goodman, S.M. (Eds), Diversity and Endemism in Madagascar. Memoires de la Societe de Biogeographie, Paris, pp. 331–344.
Fisher, B.L. (2002) Ant diversity patterns along an elevational gradient in the Reserve Speciale de Manongarivo , Madagascar. Boissiera, 59, 311–328.
Fisher, B.L. (2003) Ants (Formicidae: Hymenoptera). In: Goodman, S.M. & Benstead, J.P. (Eds), The natural history of Madagascar. University of Chicago Press, Chicago, pp. 811–819.
Fisher, B.L. (2005a) A model for a global inventory of ants: A case study in Madagascar. Proceedings of the California Academy of Sciences, 56, 86–97.
Forel, A. (1887) Fourmis récoltées à Madagascar, par le Dr. Conrad Keller. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 7, 381–289.
Forel, A. (1891) Histoire naturelle des Hyménoptères. Deuxième partie: Les Formicides. In: A. Grandidier (Ed), Histoire Physique, Naturelle et Politique de Madagascar. L'Imprimerie Nationale, Paris, pp. 1–280.
Forel, A. (1892) Nouvelles espèces de formicides de Madagascar. (Récoltées par M. Sikora). Annales de la Société entomologique de Belgique, 36, 516–535.
Forel, A. (1894) Abessinische und andere afrikanische Ameisen, gesammelt von Herrn Ingenieur Alfred Ilg, von Herrn Dr. Liengme, von Herrn Pfarrer Missionar P. Berthoud, Herrn Dr. Arth. Müller etc. Mitteilungen der Schweizerischen Entomologischen Gessellschaft, 9, 64–100.
Forel, A. (1895) Quelques fourmis du centre de Madagascar. Annales de la Société entomologique de Belgique, 39, 485–488.
Forel, A. (1900) Fourmis du Japon. Nids en toile. Strongylognathus Huberi et voisins. Fourmilière triple. Cyphomyrmex Wheeleri. Fourmis importées. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 10, 267–287.
Forel, A. (1902a) Fourmis nouvelles d'Australie. Rev. Suisse Zool., 10, 405–548.
Forel, A. (1902b) Myrmicinae nouveaux de l'Inde et de Ceylan. Revue Suisse Zoologie, 10, 165–249.
Forel, A. (1907) Fourmis d'Ethiopie. Recoltees par M. le baron Maurice de Rothschild en 1905. Revue d'Entomologie. Caen, 26, 129–144.
Forel, A. (1912) The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership of Mr. J. Stanley Gardiner, M.A. Volume 4, No. 11. Fourmis des Seychelles et des Aldabras, reçues de M. Hugh Scott. Transactions of the Linnean Society of London, Zoology, 2, 159–167.
Forel, A. (1914) Formicides d'Afrique et d'Amérique nouveaux ou peu connus. I. Afrique (Envois de M. G. Arnold à Bulawayo, Rhodesia). Bulletin de la Société Vaudoise des Sciences Naturelles, 50, 211–288.
Guérin-Méneville, F.E. (1852) Notice sur une nouvelle espéce de fourmi découverte á Saint-Domingue par M. Auguste Sallé, et qui fait son nid dans des plaines marécageuses, sur les buissons. Revue et Magasin de Zoologie Pure et Appliquee, 2, 73–79.
Güsten, R., Schulz, A. & Sanetra, M. (2006) Redescription of Tetramorium forte Forel, 1904 (Insecta: Hymenoptera: Formicidae), a western Mediterranean ant species. Zootaxa, 1310, 1–35.
Hita Garcia, F., Fischer, G. & Peters, M.K. (2010a) Tetramorium snellingi sp. n. – a new leaf-litter ant species (Hymenoptera: Formicidae) from a Western Kenyan rainforest. Myrmecological News, 13, 141–146.
Hita Garcia, F., Fischer, G. & Peters, M.K. (2010b) Taxonomy of the Tetramorium weitzeckeri species group (Hymenoptera: Formicidae) in the Afrotropical zoogeographical region. Zootaxa, 2704, 1–90.
Hita Garcia, F., Fischer, G., Kück, P., Thormann, B. & Peters, M.K. (2010c) Tetramorium boehmei sp. n. - a new ant (Hymenoptera: Formicidae) species from the Kakamega Forest, Western Kenya. Bonn zoological Bulletin, 57, 359–366.
Mann, W.M. (1921) The ants of the Fiji Islands. Bulletin of the Museum of Comparative Zoology of Harvard College, 64, 401–499.
Mayr, G. (1853) Beschreibungen einiger neuer Ameisen. Verhandlungen des Zoologisch-Botanischen Vereins in Wien, 3, 277–286.
Mayr, G. (1855) Formicina austriaca. Beschreibung der bisher im oesterreichischen Kaiserstaate aufgefundenen Ameisen nebst Hinzufuegung jener in Deutschland, in der Schweiz und in Italien vorkommenden Ameisen. Verhandlungen des Zoologisch-Botanischen Vereins in Wien, 5, 273–478.
Mayr, G. (1870) Neue Formiciden. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 20, 939–996.
Mayr, G. (1890) Aenictus-Typhlatta découverte de M. Wroughton. Nouveaux genres de formicides. Annales de la Société entomologique de Belgique, 34, 102–114.
Mcglynn, T.P. (1999) The worldwide transfer of ants: geographical distribution and ecological invasions. Journal of Biogeography, 26, 535–548.
Nylander, W. (1846) Additamentum adnotationum in monographiam formicarum borealium Europae. Acta Societatis Scientiarum Fennicae, 2, 1041–1062.
Roberts, D.L. & McGlynn, T.P. (2004) Tetramorium insolens Smith (Hymenoptera: Formicidae): a new record for Mauritius, Indian Ocean. African Entomology, 12, 265–267.
Roger, J. (1857) Einiges über Ameisen. Berliner Entomologische Zeitschrift, 1, 10–20.
Santschi, F. (1910) Formicides nouveaux ou peu connus du Congo Français. Annales de la Société Entomologique de France, 78, 349–400.
Santschi, F. (1911) Nouvelles fourmis de Madagascar. Revue Suisse Zoologie, 19, 117–134.
Santschi, F. (1918) Nouveaux Tetramorium africains. Bulletin de la Société d'Histoire Naturelle de l'Afrique du Nord, 9, 121–132.
Santschi, F. (1926) Description de nouveaux formicides éthiopiens (IIIme partie). Revue de Zoologie Africaine, 13, 207–267.
Santschi, F. (1928) Descriptions de nouvelles fourmis éthiopiennes (suite). Revue de Zoologie et de Botanique Africaines, 16, 191–213.
Santschi, F. (1930) Formicides de l'Angola. Résultats de la Mission scientifique suisse en Angola, 1928–1929. Revue Suisse Zoologie, 37, 53–82.
Santschi, F. (1933) Contribution à l'étude des fourmis de l'Afrique tropicale. Bulletin et Annales de la Société Entomologique de Belge, 73, 95–108.
Santschi, F. (1937) Fourmis angolaises. Résultats de la Mission scientifique suisse en Angola (2me voyage) 1932–1933. Revue Suisse Zoologie, 44, 209–248.
Schlick-Steiner, B.C., Steiner, F.M., Moder, K., Seifert, B., Sanetra, M., Dyreson, E., Stauffer, C. & Christian, E. (2006a) A multidisciplinary approach reveals cryptic diversity in Western Palearctic Tetramorium ants (Hymenoptera: Formicidae). Molecular Phylogenetics and Evolution, 40, 259–273.
Schlick-Steiner, B.C., Steiner, F.M. & Zettel, H. (2006b) Tetramorium pacificum Mayr, 1870, T. scabrum Mayr, 1879 sp. rev., T. manobo (Calilung, 2000) (Hymenoptera: Formicidae) - three good species. Myrmecological News, 8, 181–191.
Smith, F. (1860) Catalogue of hymenopterous insects collected by Mr. A. R. Wallace in the Islands of Bachian, Kaisaa, Amboyna, Gilolo, and at Dory in New Guinea. Journal of the Proceedings of the Linnean Society of London, Zoology, 5, 93–143.
Smith, F. (1861) Catalogue of hymenopterous insects collected by Mr. A. R. Wallace in the Islands of Ceram, Celebes, Ternate, and Gilolo. Journal of the Proceedings of the Linnean Society of London, Zoology, 6, 36–48.
Smith, F. (1862) Descriptions of new species of aculeate Hymenoptera, collected at Panama by R. W. Stretch, Esq, with a list of described species, and the various localities where they have previously occurred. Transactions of the Entomological Society of London, 3, 29–44.
Smith, M.A., Fisher, B.L. & Hebert, P.D.N. (2005b) DNA Barcoding for effective biodiversity assessment of a hyperdiverse arthropod group: the ants of Madagascar. Philosophical Transactions of the Royal Society of London B Biological Sciences, 360, 1825–1834.
Steiner, F.M., Schlick-Steiner, B.C., Sanetra, M., Ljubomirov, T., Antonova, V., Christian, E. & Stauffer, C. (2005) Towards DNA-aided biogeography: An example from Tetramorium ants (Hymenoptera: Formicidae). Annales Zoologici Fennici, 42, 23–35.
Steiner, F.M., Seifert, B., Moder, K. & Schlick-Steiner, B.C. (2010) A multisource solution for a complex problem in biodiversity research: Description of the cryptic ant species Tetramorium alpestre sp. n. (Hymenoptera: Formicidae). Zoologischer Anzeiger, 249, 223–254.
Stitz, H. (1910) Westafrikanische Ameisen. Mitteilungen aus dem Zoologische Museum in Berlin, 5, 127–151.
Stitz, H. (1912) Ameisen aus Ceram und Neu-Guinea. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1912, 498–514.
Stitz, H. (1923) Beitraege zur Kenntnis der Land- und Suesswasserfauna Deutsch-Suedwestafrikas. Ergebnisse der Hamburger deutsch-suedwestafrikanischen Studienreise 1911. Hymenoptera 7. Formicidae, 143–167.
Taylor, R.W. & Wilson, E.O. (1961) Ants from three remote oceanic islands. Psyche, 68, 137–144.
Ward, P.S. (1990) The endangered ants of Mauritius: doomed like the dodo? Notes from Underground, 4, 3–5.
Wetterer, J.K. (2010) Worldwide spread of the wooly ant, Tetramorium lanuginosum (Hymenoptera: Formicidae). Myrmecological News, 13, 81–88.
Wheeler, W.M. (1916) An Indian ant introduced into the United States. Journal of Economic Entomology, 9, 566–569.
Wheeler, W.M. (1929) Ants collected by Professor F. Silvestri in Formosa, the Malay Peninsula and the Philippines. Bollettino del Laboratorio di Zoologia Generale e Agraria della R. Scuola Superiore d'Agricultura, 24, 27–64.
Wheeler, W.M. (1934) Formicidae of the Templeton Crocker Expedition, 1933. Proceedings of the California Academy of Sciences, 4, 173–181.
Wilson, E.O. & Taylor, R.W. (1967) Ants of Polynesia. Pacific Insects Monographs, 14, 1–109.