Re-description and range extension of the Afrotropical mayfly Cloeon perkinsi (Ephemeroptera, Baetidae)

  • Zohar Yanai - Musée Cantonal de Zoologie, Palais de Rumine, Lausanne. - Department of Ecology and Evolution, Biophore, University of Lausanne, 1015 Lausanne. - School of Zoology, Tel Aviv University, Tel Aviv 6997801, Israel.
  • Wolfram Graf University of Natural Resources and Applied Life Sciences, Institute of Hydrobiology and Aquatic Ecology Management, Vienna.
  • Yonas Terefe - University of Natural Resources and Applied Life Sciences, Institute of Hydrobiology and Aquatic Ecology Management, Vienna. - Department of Biology, College of Natural and Computational Sciences, Ambo University, Ambo, Ethiopia.
  • Michel Sartori - Musée Cantonal de Zoologie, Palais de Rumine, Lausanne. - Department of Ecology and Evolution, Biophore, University of Lausanne, 1015 Lausanne.
  • Jean-Luc Gattolliat - Musée Cantonal de Zoologie, Palais de Rumine, Lausanne. - Department of Ecology and Evolution, Biophore, University of Lausanne, 1015 Lausanne.
Keywords: Afrotropics, Arabian Peninsula, biogeography, Cloeon, Levant


Cloeon perkinsi was described from South Africa in 1932 by Barnard. Despite being relatively common in Africa, it was mentioned in the literature quite rarely, and its known distribution to date includes most of sub-Saharan Africa. Material collected recently in Ethiopia, Israel, Saudi Arabia, and Yemen extends its distribution in East Africa, Arabian Peninsula and the Levant. We examined this material, and provide a re-description of adults (females and males) and nymphs of the species. It represents a much-needed urge mainly due to inconsistencies in literature reports regarding colouration, and sometimes incomplete morphological description of all stages. We demonstrate the intraspecific morphological variability that we have witnessed, and provide information regarding the range of habitats colonised by C. perkinsi. Based on geological and climatic history of the studied region, taken together with among countries genetic distances of the mitochondrial barcoding gene COI, we propose colonisation mechanisms for the north-easternmost limit of distribution. The fragmented distribution pattern of the species highlights the conservation importance of isolated aquatic habitats in the region, as well as current knowledge gaps.


Ali R. 1970. Certain mayflies (order: Ephemeroptera) of west Pakistan. Pakistan Journal of Science 22: 119–124.

Ball S.L., Hebert P.D.N., Burian S.K. & Webb J.M. 2005. Biological identifications of mayflies (Ephemeroptera) using DNA barcodes. Journal of the North American Benthological Society 24: 508–524.

Barber-James H.M., Gattolliat J.-L., Sartori M. & Hubbard M.D. 2008. Global diversity of mayflies (Ephemeroptera, Insecta) in freshwater. Hydrobiologia 595: 339–350.

Barnard K.H. 1932. South African may-flies (Ephemeroptera). Transactions of the Royal society of South Africa 20: 201–259.

Barnard K.H. 1940. Additional records, and descriptions of new species, of South African alder-flies (Megaloptera), May-flies (Ephemeroptera), caddis-flies (Trichoptera), stone-flies (Perlaria), and dragon-flies (Odonata). Annals of the South African Museum 32: 609–661.

Bauernfeind E. & Soldán T. 2012. The Mayflies of Europe (Ephemeroptera). Apollo Books, Ollerup.

Botosaneanu L. 1992. Fauna Palaestina, Insecta VI: Trichoptera of the Levant. The Israel Academy of Sciences and Humanities, Jerusalem.

Botosaneanu L. 1999. Caddisflies (Trichoptera) – what can they tell about biogeography of the Levant? Israel Journal of Zoology 45: 75–78.

Botosaneanu L. 2002. A classical case of insular radiation: the Hydroptila species of La Réunion. Proceedings of the 10th International Symposium on Trichoptera, Nova Supplementa Entomologica 15: 323–330.

Chopra B. 1924. The fauna of an island in the Chilka Lake. The Ephemeroptera of Barkuda Island. Records of the Indian Museum 26: 415–422.

Demoulin G. 1965. LXXXVIII. – Ephemeroptera. in: Basilewsky P. & Lelup N.(eds) Mission zoologique de l’I.R.S.A.C. en Afrique orientale: 91–114. Royal Museum for Central Africa, Tervuren.

Dimentman C., Bromley H.J. & Por F.D. 1992. Lake Hula – reconstruction of the fauna and hydrobiology of a lost lake. The Israel Academy of Sciences and Humanities, Jerusalem.

Dumont H.J. 1991. Fauna Palaestina, Insecta V: Odonata of the Levant. The Israel Academy of Sciences and Humanities, Jerusalem.

Flemming N.C., Bailey G.N., Courtillot V., King G., Lambeck K., Ryerson F. & Vita-Finzi C. 2003. Coastal and marine palaeo-environments and human dispersal points across the Africa-Eurasia boundary. In: Brebbia CA, Gambin T (eds.) The maritime and underwater heritage: 61–74. Wessex Institute of Technology, Southampton.

Folmer 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: 294–299.

Frumkin A. & Comay O. 2019. The last glacial cycle of the southern Levant: Paleoenvironment and chronology of modern humans. Journal of Human Evolution.

Gattolliat J.-L. & Rabeantoandro S.Z. 2002. The genus Cloeon (Ephemeroptera, Baetidae) in Madagascar. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 74: 195–209.

Gattolliat J.-L. & Staniczek A.H. 2011. New larvae of Baetidae (Insecta: Ephemeroptera) from Espiritu Santo, Vanuatu. Stuttgarter Beiträge zur Naturkunde A, Neue Serie 4: 75–82.

Gattolliat J.-L., Vuataz L. & Sartori M. 2012. First contribution to the mayflies of Jordan. Zoology in the Middle East 56: 91–110.

Gattolliat J.-L., Cavallo E., Vuataz L. & Sartori M. 2015. DNA barcoding of Corsican mayflies (Ephemeroptera) with implications on biogeography, systematics and biodiversity. Arthropod Systematics & Phylogeny 73: 3–18.

Gattolliat J.-L., Kondratieff B.C., Kaltenbach T. & Al Dhafer H.M. 2018. Labiobaetis from the Kingdom of Saudi Arabia (Insecta, Ephemeroptera, Baetidae). ZooKeys 774: 77–104.

Gillies M.T. 1980. An introduction to the study of Cloeon Leach (Baetidae, Ephemeroptera) in West Africa. Bulletin de l’Institut Francaise d’Afrique Noire 42: 135–156.

Gillies M.T. 1985. A preliminary account to the East African species of Cloeon Leach and Rhithrocloeon gen.n. (Ephemeroptera). Aquatic Insects 7: 1–17.

Goren M. & Ortal R. 1999. Biogeography, diversity and conservation of the inland water fish communities in Israel. Biological Conservation 89: 1–9.

Hassan F.A. 1997. Holocene palaeoclimates of Africa. African Archaeological Review 14: 213–230.

Hassan S.A. & Abdel Fattah M.A. 2007. A revision of the order Ephemeroptera from Egypt. Journal of the Egyptian-German Society of Zoology 53: 189–214.

Heller J. 2007. A historic biogeography of the aquatic fauna of the Levant. Biological Journal of the Linnean Society 92: 625–639.

Kimmins D.E. 1947. New species of Indian Ephemeroptera. Proceedings of the Royal Entomological Society of London (B) 16: 92–100.

Kimmins D.E. 1955. Ephemeroptera from Nyasaland, with description of three new and some interesting nymphal forms. Annals and Magazine of Natural History 12: 859–880.

Kimmins D.E. 1960. Notes on East African Ephemeroptera, with descriptions of new species. Bulletin of the British Museum (Natural History), Entomology 9: 337–355.

Kluge N.J. 2016. A new subgenus Oculogaster subgen.n. for viviparous representatives of Procloeon, with discussion about status of the generic name Austrocloeon Barnard, 1932 and the species name africanum Esben-Petersen, 1913 [Cloeon] (Ephemeroptera, Baetidae). Zootaxa 4107: 491–516.

Kluge N.J. 2019. Ephemeroptera of the World. Available from [accessed 20 June 2019]

Kumar S., Stecher G., Li M., Knyaz C. & Tamura K. 2018. MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms (Version 10.0.2). Molecular Biology and Evolution 35: 1547–1549.

Landa V. & Soldán T. 1986. Invasive behavioural patterns of mayflies (Insecta, Ephemeroptera) in some man-influenced aquatic biotopes in Czechoslovakia. Ekologia 3: 239–246.

Lugo-Ortiz C.R. & McCafferty W.P. 1998. New species of Cloeon and Demoulinia (Ephemeroptera: Baetidae) from Madagascar. Entomological News 109: 357–362.

Malicky H. 1986. The caddisflies of Saudi Arabia and adjacent regions (Insecta, Trichoptera). Fauna of Saudi Arabia 8: 233–245.

Monaghan M.T., Gattolliat J.-L., Sartori M., Elouard J.-M., James H., Derleth P., Glaizot O., de Moor F. & Vogler A.P. 2005. Trans-oceanic and endemic origins of the small minnow mayflies (Ephemeroptera, Baetidae) of Madagascar. Proceedings of the Royal Society B 272: 1829–1836.

Navás L. 1926. Algunos insectos del Museo de Paris. Serie 3. Brotéria (Série Zoológica) 23: 95–115.

Pachur H.J. & Kröpelin S. 1987. Wadi Howar: Paleoclimatic evidence from an extinct river system in the southeastern Sahara. Science 237: 298–300.

Papadopoulou A., Anastasiou I. & Vogler A.P. 2010. Revisiting the insect mitochondrial molecular clock: the Mid-Aegean trench calibration. Molecular Biology and Evolution 27: 1659–1672.

Por F.D. 1975. An outline of the zoogeography of the Levant. Zoologica Scripta 4: 5–20.

Ritchie J.C. & Haynes C.V. 1987. Holocene vegetation zonation in the eastern Sahara. Nature 330: 645–647.

Rohling E.J., Fenton M., Jorissen F.J., Bertrand P., Ganssen G. & Caulet J.P. 1998. Magnitudes of sea-level lowstands of the past 500,000 years. Nature 394: 162–165.

Rutschmann S., Gattolliat J.-L., Hughes S.J., Báez M., Sartori M. & Monaghan M.T. 2014. Evolution and island endemism of morphologically cryptic Baetis and Cloeon species (Ephemeroptera, Baetidae) on the Canary Islands and Madeira. Freshwater Biology 59: 2516–2527.

Rutschmann S., Detering H., Simon S., Funk D.H., Gattolliat J.-L., Hughes S.J., Raposerio P.M., Desalle R., Sartori M. & Monaghan M.T. 2017. Colonization and diversification of aquatic insects on three Macaronesian archipelagos using 59 nuclear loci derived from a draft genome. Molecular Phylogenetics and Evolution 107: 27–38.

Salles F.F., Gattolliat J.-L., Angeli K.B., De-Souza M.R., Gonçalves I.C., Nessimian J.L. & Sartori M. 2014. Discovery of an alien species of mayfly in South America (Ephemeroptera). ZooKeys 399: 1–16.

Samocha M. 1972. Ephemeroptera of Israel. Master’s thesis, Tel Aviv University, Israel.

Sanlaville P. 1992. Changements climatiques dans la Péninsule Arabique durant le Pléistocène supérieur et l’Holocène. Paléorient 18: 5–26.

Sartori M. & Gillies M.T. 1990. Further records of mayflies (Insecta: Ephemeroptera) from the Arabian Peninsula. Leptophlebiidae and Baetidae. Fauna of Saudi Arabia 11: 10–17.

Ståhls G. & Savolainen E. 2008. mtDNA COI barcode reveal cryptic diversity in the Baetis vernus group (Ephemeroptera, Baetidae). Molecular Phylogenetics and Evolution 46: 82–87.

Soldán T. & Bojková J. 2015. New species of mayflies (Ephemeroptera) from Cape Verde. Zootaxa 3926: 561–575.

Sowa R. 1975. What is Cloeon dipterum (Linnaeus, 1761)? The nomenclatural and morphological analysis of a group of the European species of Cloeon Leach (Ephemerida: Baetidae). Entomologica Scandinavica 6: 215–223.

Thomas J.D. 1966. Some preliminary observations on the fauna and flora of a small man-made lake in the West African savanna. Bulletin de l’IFAN 28: 542–562.

Thunell R.C., Locke S.M. & Williams D.F. 1988. Glacio-eustatic sea-level control on Red Sea salinity. Nature 334: 601–604.

Tobias D. & Tobias W. 2019. Trichoptera Africana. Available from [accessed 8 July 2019]

Vuataz L., Sartori M., Wagner A. & Monaghan M.T. 2011. Toward a DNA taxonomy of alpine Rhithrogena (Ephemeroptera: Heptageniidae) using a mixed yule-coalescent analysis of mitochondrial and nuclear DNA. PLoS ONE 6: e19728.

Webb J.M., Jacobus L.M., Funk D.H., Zhou X., Kondratieff B., Geraci C.J., DeWalt R.E., Baird D.J., Richard B., Phillips I. & Hebert P.D.N. 2012. A DNA barcode library for North American Ephemeroptera: progress and prospects. PLoS ONE 7: e38063.

How to Cite
Yanai, Z., Graf, W., Terefe, Y., Sartori, M., & Gattolliat, J.-L. (2020). Re-description and range extension of the Afrotropical mayfly Cloeon perkinsi (Ephemeroptera, Baetidae). European Journal of Taxonomy, (617).