Abstract
Only one species of the genus Allodia has been previously recorded from the Afrotropical region, Allodia (Brachycampta) flavorufa Matile, 1978. Six new species are described here, all representing the nominotypical subgenus, Allodia s.s. The new species are described from material collected in different mountainous areas in south and east Africa; A. jaschhofi sp. nov., A. karkloofensis sp. nov., A. drakensbergensis sp. nov., A. nyeriensis sp. nov., A. mazumbaiensis sp. nov. and A. keurbosensis sp. nov. The species are morphologically very similar, and can only be separated based on minor differences in wing venation and characters of the male terminalia. The genetic differences between the species in the DNA barcode region (CO1), however, support delimitation. The origin and distribution of these Afrotropical taxa, in relation to each other and to their Holarctic relatives, is discussed.
References
Bechev, D. (1999) The zoogeographic classification of the Palaearctic genera of fungus gnats (Diptera: Sciaroidea, excluding Sciaridae). Studia dipterologica, 6 (2), 321–326.
Brower, A.V. (1994) Rapid morphological radiation and convergence among races of the butterfly Heliconius erato inferred from patterns of mitochondrial DNA evolution. Proceedings of the National Academy of Sciences of the United States of America, 91 (14), 6491–6495.
Brunetti, E. (1912) XI. Diptera. In: Annandale, N. (Ed.), Zoological results of the Abor expedition 1911–12. Vol. 8. Records of the Indian Museum, Calcutta, pp.149–190.
https://doi.org/10.5962/bhl.part.1186Colless, D.H. (1966) Diptera: Mycetophilidae. In: Insects of Micronesia. 12 (6). Bernice P. Bishop Museum, Honolulu, pp. 637–667.
Collins, R.A., Boykin, L.M., Cruickshank, R.H. & Armstrong, K.F. (2012) Barcoding's next top model: an evaluation of nucleotide substitution models for specimen identification. Ecology and Evolution, 3 (3), 457–465.
https://doi.org/10.1111/j.2041-210x.2011.00176.xEdgar, R.C. (2004) MUSCLE: a multiple sequence alignment method with reduced time and space complexity. BMC Bioinformatics, 5 (1), 113.
https://doi.org/10.1186/1471-2105-5-113Edwards, F.W. (1928) Diptera Nematocera from the Federated Malay States Museum. Journal of the Federated Malay States Museum, 14, 1–139.
Fattorini, S. (2007) Historical relationships of African mountains based on cladistic analysis of distributions and endemism of flightless insects. African Entomology, 15 (2), 340–355.
https://doi.org/10.4001/1021-3589-15.2.340Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39 (4), 783–791.
https://doi.org/10.2307/2408678Fjeldså, 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 (3), 325–346.
https://doi.org/10.1023/a:1018356506390Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5), 294–299.
Hebert, P.D.N., Cywinska, A., Ball, S.L. & deWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society Biological Sciences, Series B, 270 (1512), 313–321.
https://doi.org/10.1098/rspb.2002.2218Jürgenstein, S., Kurina, O. & Põldmaa, K. (2015) The Mycetophila ruficollis Meigen (Diptera, Mycetophilidae) group in Europe: elucidating species delimitation with COI and ITS2 sequence data. ZooKeys, 508, 15–51.
https://doi.org/10.3897/zookeys.508.9814Kingdon, J. (1990) Island Africa:The Evolution of Africa's Rare Animals and Plants. Collins, London, 278 pp.
Kirk-Spriggs, A.H. & Stuckenberg, B.R. (2009) Afrotropical Diptera – Rich savannas, poor rainforests. In: Bickel, D.J., Pape, T. & Meier, R. (Eds.), Diptera Diversity: Status, Challenges and Tools. Brill Academic Publishers, Leiden, pp. 155–196.
https://doi.org/10.1163/ej.9789004148970.i-459.31Kjærandsen, J. (2006) Review of fungus gnats of the genus Tarnania Tuomikoski, with a phylogeny of the Rymosia s.l. genus group (Diptera: Mycetophilidae). Insect Systematics and Evolution, 37 (2), 121–148.
https://doi.org/10.1163/187631206788831146Kjærandsen, J. (2007) Two new species of Allodia subgenus Brachycampta Winnertz from Norway and Sweden (Diptera: Mycetophilidae). Entomologica Fennica, 18 (1), 1723.
Kjærandsen, J. (2009) The genus Pseudexechia Tuomikoski re-characterized, with a review of European species (Diptera: Mycetophilidae). Zootaxa, 2056, 1–45.
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/msw054Kurina, O. (1997) A new species of fungus gnats of the genus Allodia Winnertz, 1863 (Diptera, Mycetophilidae) from Estonia. Studia dipterologica, 4 (2), 275–279.
Kurina, O., Õunap, E. & Põldmaa, K. (2015) Two new Neuratelia Rondani (Diptera, Mycetophilidae) species from Western Palaearctic: a case of limited congruence between morphology and DNA sequence data. ZooKeys, 496, 105–129.
https://doi.org/10.3897/zookeys.496.9315Lachaise, D. & Chassagnard, M. (2001) Seven new montane species of Drosophila in the Eastern Arc mountains and Mt Kilimanjaro in Tanzania attesting to past connections between eastern and western African mountains (Diptera: Drosophilidae). European Journal of Entomology, 98 (3), 351–366.
https://doi.org/10.14411/eje.2001.056Lim, G., Balke, M., & Meier, R. (2012) Determining species boundaries in a world full of rarity: Singletons, species delimitation methods. Systematic Biology, 61 (1), 165–169.
https://doi.org/10.1093/sysbio/syr030Matile, L. (1978) Dipteres Mycetophilidae de l'Archipel des Comores. Memoires du Museum National d'Histoire Naturelle Serie A Zoologie, 109, 247–306.
Matile, L. (1980) 15. Family Mycetophilidae. In: Crosskey, R.W. (Ed.), Catalogue of the Diptera of the Afrotropical region. British Museum, London, pp. 217–230.
McAlpine, J.F. (1981) Morhology and terminology – adults. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockheroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Vol. 1. Canada department of Agriculture Research Branch Monograph, Ottawa, pp. 89–124.
Nei, M. & Kumar, S. (2000) Molecular Evolution and Phylogenetics. Oxford University Press, New York, 328 pp.
Rindal, E., Søli, G.E.E., Kjærandsen, J. & Bachmann, L. (2007) Molecular phylogeny of the fungus gnat tribe Exechiini (Mycetophilidae, Diptera). Zoologica Scripta, 36 (4), 327–335.
https://doi.org/10.1111/j.1463-6409.2007.00285.xSaitou, N. & Nei, M. (1987) The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4 (4), 406–425.
Sasakawa, M. & Ishizaki, H. (2003) Fungus gnats of the genera Anatella, Allodia and Cordyla in Japan (Diptera: Mycetophilidae). Entomological Science, 6 (2), 97–109.
https://doi.org/10.1046/j.1343-8786.2003.00013.xSenior-White, R.A. (1922) New Ceylon Diptera (Part II). Spolia Zeylanica 12 (45), 195–206.
https://doi.org/10.5962/bhl.part.12654Ševčík, J., Kaspřák, D., & Rulik, B. (2016) A new species of Docosia Winnertz from Central Europe, with DNA barcoding based on four gene markers (Diptera, Mycetophilidae). ZooKeys, 549, 127–143.
https://doi.org/10.3897/zookeys.549.6925Sivathsan, A. & Meier, R. (2011) On the inappropriate use of Kimura-2-parameter (K2P) divergences in the DNA-barcoding literature. Cladistics, 28 (2), 190–194.
https://doi.org/10.1111/j.1096-0031.2011.00370.xSøli, G.E.E. (1997) The adult morphology of Mycetophilidae (s. str.), with a tentative phylogeny of the family (Diptera, Sciaroidea). Entomologica Scandinavica Supplement, 50, 5–55.
Søli, G. (2017) Mycetophilidae (Fungus gnats). In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.), Manual of Afrotropical Diptera. Vol. 2. Nematocerous Diptera and lower Brachycera. Suricata 5. South African National Biodiversity Instuitute, Pretoria, pp. 533–555.
Tuomikoski, R. (1966) Generic taxonomy of the Exechiini (Diptera, Mycetophilidae). Suomen Hyonteistieteellinen Aikakauskirja, 32, 159–194.
Winnertz, J. (1863) Beitrag zu einer monographie der pilzmücken. Braumüller, Wien, 332 pp.
https://doi.org/10.5962/bhl.title.9961Wu, H., Zheng, L.Y. & Xu, H.C. (2003) First record of Allodia Winnertz (Diptera, Mycetophilidae) in China with descriptions of seven new species. Acta Zootaxonomica Sinica, 28 (2), 349–355.
Zaitzev, A. (1983) A review of Holarctic species of the subgenus Allodia s. str. (Diptera, Mycetophilidae). Zoologicheskii Zhurnal, 62, 1915–1920.
Zaitzev, A. (1984) A review of species of the subgenus Brachycampta (Diptera, Mycetophilidae) of the Holarctic fauna. Zoologicheskii Zhurnal, 63 (10), 1504–1515.
Zaitzev, A. (2003) Fungus gnats (Diptera, Sciaroidea) of the fauna of Russia and adjacent regions. Part II. Dipterological Research, 14, 77–386.