Abstract
The 12 species previously placed in Tortopus together with 3 species newly described here, are revised and included in a phylogenetic analysis. Based on synapomorphic characters on the nymphs and adults of both sexes, Tortopus is restricted to T. igaranus Needham & Murphy, T. circumfluus Ulmer, T. harrisi Traver, T. zottai (Navás), T. bellus Lugo-Ortiz & McCafferty, and T. arenales sp. nov., and the genus is defined by: female parastyli receptors with long furrows anterior to sockets; penes entirely flattened; male ninth abdominal sternum almost separated in two portions by a median notch; mesosternum with furcasternal protuberances contiguous only on basal corner; and nymphs with two subapical tubercles on mandibular tusks. Tortopsis is newly described for T. bruchianus (Navás), T. limoncocha sp. nov., T. obscuripennis (Domínguez), T. parishi (Banks), T. primus (McDunnough), T. puella (Pictet), T. sarae (Domínguez), T. spatula sp. nov., and T. unguiculatus (Ulmer). Tortopsis is characterized by: R sector of female fore wing without additional veins between R 2 and IR; female parastyli receptors C or V-shaped, with sockets opening towards median line; male gonopore associated with a claw-like structure; penes separated from the base; parastyli more than 5 times length of pedestals; parastyli curved in lateral view; nymphs with a single subapical tubercle on mandibular tusks. The study of available type material permitted inclusion of comparative diagnoses, with figures and redescriptions as needed. The male imago of the type species of Tortopus (T. igaranus Needham & Murphy) is described for the first time, as are the female adults of Tortopus bellus Lugo-Ortiz & McCafferty and T. harrisi Traver. Three new Neotropical species based on male and female adults are described: Tortopus arenales and Tortopsis limoncocha from Ecuador, and Tortopsis spatula from Colombia. Keys to separate the adults and nymphs of the genera of Polymitarcyidae, and for male and female adults of all the species of Tortopus and Tortopsis are presented, as well as line drawings, pictures and SEM photographs of important structures.References
Banks, N. (1918) New neuropteroid insects. Bulletin of the Museum of Comparative Zoology, 62 (1), 1–22.
Brittain, J.E. (1982) Biology of mayflies. Annual Review of Entomology, 27, 119–147.
Domínguez, E. (1985) El género Tortopus Needham y Murphy (Ephemeroptera: Polymitarcyidae) en la Argentina. Physis, B 43, 69–72.
Domínguez, E. (1987) Dos nuevas combinaciones de Polymitarcyidae (Ephemeroptera), descriptas por L. Navás. Physis, B 45, 9–12.
Domínguez, E., Molineri, C., Pescador, M., Hubbard, M.D. & Nieto, C. (2006) Ephemeroptera of South America in Aquatic Biodiversity in Latin America (ABLA, Vol. 2). Pensoft. Sofia-Moscow, 646 pp.
Eaton, A.E. (1871) A monograph on the Ephemeridae. Transactions of the Entomological Society of London, 1871, 1–164.
Eaton, A.E. (1883–1888) A revisional monograph of recent Ephemeridae or mayflies. Transactions of the Linnean Society of London, Zoology 3, 1–352.
Goloboff, P.A., Farris, J.S., Källersjö, M.,Oxelman, B., Ramírez, M.J.& Szumik, C.A. (2003) Improvements to resampling measures of group support. Cladistics, 19, 324–332.
Goloboff, P.A., Farris, J.S. & Nixon, K. (2008) T.N.T., a free program for phylogenetic analysis. Cladistics, 24, 774–786.
Ide, F.P. (1941) Mayflies of two tropical genera, Lachlania and Campsurus, from Canada with descriptions. Canadian Entomologist, 73, 153–156.
Kluge, N.Yu. (2004) The phylogenetic system of Ephemeroptera. Kluwer, Dordrecht, 442 pp.
Knight, S.S. & Cooper, C.M. (1989) New records for Tortopus incertus (Ephemeroptera) in Mississippi and notes on microhabitat. Entomological News, 100, 21–26.
Lugo-Ortiz, C. R. & McCafferty, W. P. (1994) The mayflies (Ephemeroptera) of Texas and their biogeographic affinities. In: L. Corkum and Ciborowski J. (eds.), Proceedings of the seventh international conference on Ephemeroptera. Sandhill Crane Press, Gainesville, pp. 151–169.
Lugo-Ortiz, C.R. & McCafferty W.P. (1996) Central American Tortopus (Ephemeroptera: Polymitarcyidae): a unique new species and new country records. Entomological News, 107, 23–27.
Maés, J.M. (1988) Catálogo de los Ephemeroptera y Plecoptera de Nicaragua. Revista Nicaraguense de Entomologia, 2, 49–50.
McCafferty, W.P. (1975) The burrowing mayflies (Ephemeroptera: Ephemeroidea) of the United States. Transactions of the American Entomological Society, 101, 447–504.
McCafferty, W.P. (1994) Distributional and classificatory supplement to the burrowing mayflies (Ephemeroptera: Ephemeroidea) of the United States. Entomological News, 105, 1–13.
McCafferty, W.P. & Bloodgood, D.W. (1989) The female and male coupling apparatus in Tortopus mayflies. Aquatic Insects, 11, 141–146.
McDunnough, J. (1924) New Ephemeridae from Illinois. The Canadian Entomologist, 56, 7–9.
Molineri, C. (2008) The larvae of the burrowing mayfly genus Tortopus (Ephemeroptera: Polymitarcyidae). Aquatic Insects, 30, 7–19.
Molineri, C. & Emmerich, D. (in press) New species and new stage descriptions of Campsurus major species-group (Polymitarcyidae, Campsurinae), with first report of silk-case construction in mayfly nymphs. Aquatic Insects.
Molineri, C., Siegloch A. & Righi-Cavallaro, K.O. (2010) The nymph of Tortopus harrisi Traver (Ephemeroptera: Polymitarcyidae). Zootaxa, 2436, 65–68.
Navás, L. (1920) Insecta nova. VI Series. Memorie della Pontificia Accademia Romana dei Nuovi Lincei, 5(2), 11–29.
Navás, L. (1922) Insectos nuevos o poco conocidos. Memorias de la Real Academia de Ciencias y Artes de Barcelona, 17 (15), 383–400.
Navás, L. (1926) Insectos de la Argentina y Chile. Seguna Serie. Estudios, 31, 103–111.
Needham, J.G. & Murphy, H.E. (1924) Neotropical mayflies. Bulletin of the Lloyd Library, 24, Entomological Series, 4, 1–79.
Needham, J. G., Traver, J. R. & Hsu, Y. C. (1935) The Biology of Mayflies. Comstock Pub. Inc. New York, 759 pp.
Nixon, K.C. (2002) WinClada. Program and documentation available at www.zmuc.dk/public/phylogeny.
Pictet, F.J. (1843–1845) Histoire naturelle générale et particulière des insectes névroptères. Famille des Éphémérines. Baillière édit., Paris (Kessmann et Cherbuliez édit., aussi à Genève), 300 pp.
Randolph, R. P. & McCafferty, W. P. (2002) First report of the genus Tortopus from Mexico, with additional new records of Mexican Ephemeroptera. Entomological news, 113, 34–36.
Sattler, W. (1967) Über die Lebensweise, insbesondere das bauverhalten, neotropischer Eintagsfliegen-Larven (Ephemeroptera, Polymitarcidae). Beiträge zur Neotropischen Fauna, 5(2), 89–110.
Scott, D.C., Berner, L., Hirsch, A. (1959) The nymph of the mayfly genus Tortopus (Ephemeroptera: Polymitarcidae). Annals of the Entomological Society of America, 52, 205–213.
Traver, J.R. (1950) Notes on Neotropical mayflies. Part. IV. Family Ephemeridae (continued). Revista de Entomologia, 21, 593–614.
Ulmer, G. (1920) Neue Ephemeropteren. Archiv für Naturgeschichte, 85(A), 1–80.
Ulmer, G. (1932) Bemerkungen über die seit 1920 neu aufgestellten Gattungen der Ephemeropteren. Stettiner Entomologische Zeitung, 93, 204–219.
Ulmer, G. (1933) Aquatic insects of China. Article VI. Revised key to the genera of Ephemeroptera. Peking Natural History Bulletin, 7, 195–218.
Ulmer, G. (1942) Alte und neue Eintagsfliegen (Ephemeropteren) aus Süd- und Mittelamerika. Stettiner Entomologische Zeitung, 103, 98–128.