DOI: 10.15468/m94fqt
title: Taxonomic revision of the Afrotropical species of the tribe Eudorylini (Diptera, Pipunculidae)

description: This dataset contains the digitized treatments in Plazi based on the original journal article FÖLDVÁRI, MIHÁLY (2013): Taxonomic revision of the Afrotropical species of the tribe Eudorylini (Diptera, Pipunculidae). Zootaxa 3656 (1): 1-121, DOI: 10.11646/zootaxa.3656.1.1, URL: http://dx.doi.org/10.11646/zootaxa.3656.1.1ABSTRACTThis paper revises the Afrotropical species of the tribe Eudorylini. The four genera (Claraeola, Clistoabdominalis, Dasydorylas, Eudorylas) present in the Afrotropical Region comprise 77 species (4, 4, 8 and 61 species respectively), 15 of them are new to science: Clistoabdominalis namibiensis, Dasydorylas bodocsi, D. okongoensis, Eudorylas angolae, E. barracloughi, E. brandbergensis, E. femoralis, E. gabela, E. hirsutus, E. lobus, E. pectinatus, E. pilulus, E. protumidus, E. rooibergensis and E. skorpionensis. Six new species name combinations are published: Claraeola hadrosoma, Clistoabdominalis crassus, Dasydorylas africanus, D. evanidus, D. minymerus and D. turneri.Type material for all available species was studied and species are (re)described in detail. Drawings of male and female terminalia are presented and an identification key to the males (only males were available in the majority of the species) is provided as well as individual species diagnoses for an easier species recognition.Eight new synonymies are proposed: Dorilas (Eudorylas) dorsalis and Dorilas (Eudorylas) apiculatus to Dasydorylas evanidus, Pipunculus (Eudorylas) fractus to Eudorylas amitinus, Dorilas (Eudorylas) pusillus to Eudorylas diversus, Dorilas (Eudorylas) modicus to Eudorylas encerus, Dorilas (Eudorylas) megacanthus to Eudorylas garambensis and Dorilas (Eudorylas) quadratus and Pipunculus (Eudorylas) eremnoptera to Pipunculus mutillatus.A discussion on earlier methodology of species description within this group, in particular that of D. E. Hardy’s works, is given.INTRODUCTIONPipunculidae or big-headed flies are usually small or medium sized, inconspicuous flies (except for Nephrocerus spp.) and are characterised by their large compound eyes occupying most of the hemispherical head. They are closely related to hover flies (Syrphidae), but can easily be differentiated by their wing venation, the cell r4+5 being open and the vena spuria being absent (Papp and Schumann 2000: 177). A recent paper by Wiegmann et al. (2011) based on molecular data (incl. nuclear gene and total mitochondrium sequences) suggests that Pipunculidae are the sister group to Schizophora.About 1,400 species have been described world-wide so far, but the fauna is still considered to be poorly known and an estimate of well over 2,000 is given by Skevington and Yeates (2001). During the larval stage pipunculids are almost exclusively endoparasitoids of Auchenorrhyncha; hosts have been recorded from Cicadellidae, Cixiidae, Cercopidae, Flatidae, Fulgoridae, Delphacidae and Membracidae (ferrar 1987). Nephrocerus is the only genus to develop in adult tipulidae according to Koenig and Young (2007) and Kehlmaier and Floren (2010). These authors reported larvae of North American and European Nephrocerus species and described their life cycle and developmental stages. All species of Eudorylas are parasitoids of the homopteran family Cicadellidae with the only exception being australian E. helluo (attacking Flatidae) according to Coe (1966). Together with Dryinidae (Hymenoptera) and Strepsiptera, Pipunculidae are considered as the most important parasites of Auchenorrhyncha (Freytag 1985, Waloff and Jervis 1987), and the family may have potential as biological control agents of pest species of leafhoppers. Considerable research has been done on those species that are parasites of rice leafhoppers (Hardy 1971, Jervis 1980, Koizumi 1959, Waloff and Jervis 1987, Yano 1979, Yano et al. 1984, May 1979, Morakote and Yano 1988).Within Pipunculidae the most problematic group is Eudorylini and particularly Eudorylas, since the tribe contains 515 species world-wide in nine genera (De Meyer 1996), and serious misinterpretations of what constitutes Eudorylas have led to an acute lack of stability in the genus. This was solved in two stages: Skevington and Yeates (2001) made a phylogenetic revision of the tribe with redefinitions of the genera, and the ICZN (2002) issued an opinion to stabilise the genus name Eudorylas.In their work Skevington and Yeates (2001) propose Congomyia and Moriparia as junior synonyms of Claraeola, as well as Metadorylas as junior synonym of Eudorylas; genus groups are defined based on the cladogram produced for sixty species and 137 characters, and they describe two new genera (Clistoabdominalis Skevington and Dasydorylas Skevington). The following genus groups were recognised within the tribe:- Allomethus genus group (Allomethus, Basileunculus, Claraeola)- Dasydorylas genus group (Dasydorylas)- Amazunculus genus group (Amazunculus, ElmoHardyia)- Eudorylas genus group (Eudorylas, Clistoabdominalis)Although revisionary work is proceeding in most biogeographical regions (Ale-Rocha 1996, De Meyer 1989 a, 1989 b, 1992, 1993, 1996, Földvári and De Meyer 2000, Kehlmaier 2005 a, 2008, Kehlmaier and Assmann 2008, Rafael 1993, 1995, 1996, Rafael and Ale-Rocha 1997, Rafael and Menezes 1999, Skevington and Marshall 1998, Skevington 1999, 2001, 2002, 2003, 2005, 2006, Skevington and Földvári 2007, Skevington and Yeates 2001), the Afrotropical Eudorylini has not been revised since Hardy's works (Hardy 1949 a, 1949 b, 1950, 1952, 1959 a, 1959 b, 1961, 1962, 1980), therefore it is necessary to deal with this group.Review of literature on Afrotropical EudoryliniThe Afrotropical pipunculid fauna was elaborated by numerous authors, however D. Elmo Hardy contributed most towards a better knowledge of the species and their recognition. In total, Hardy published 51 works dealing with Pipunculidae between 1939 and 1989. According to Skevington and De Meyer (2004) it is clear that his impact was substantial from 1938 onwards. By 1972 Hardy had doubled the number of described (and currently valid species): from 292 known in 1938, he described an additional 320. He tripled the known Neotropical fauna and doubled the known Nearctic and Oriental species. His contribution to the Australasian/Oceanian fauna is considerable (30.2% is described by Hardy, mostly from the Hawaiian Islands) and the only exceptions are the Palaearctic fauna (0.8%) and the Australian fauna (0%). The latter region is poorly understood, but apparently very species rich according to studies by Skevington (1999, 2001, 2002). Hardy’s largest impact was on the Afrotropical fauna, boosting the know species from 17 to 122. When compared with the currently known and valid species, 70.5% of the Afrotropical fauna is described by Hardy (38.3, 31.8 and 29.3% of the Nearctic, Oriental and Neotropical fauna, respectively) based on Skevington and De Meyer (2004).Hardy (1949 a, 1949 b, 1950, 1952, 1959 a, 1959 b, 1961, 1962, 1980) published numerous papers dealing with the Afrotropical species of the tribe Eudorylini, and he tried to summarise his knowledge by describing the species, naming what he believed to be the closest relative and providing an illustrated identification key. He mainly worked on material from the Democratic Republic of Congo (also known as Belgian Congo or Zaire), the republic of South Africa and Madagascar. These are the regions where most of the collections have specimens from. Hardy's descriptions are very detailed, since he described all external characters thoroughly, and noted most of the minor differences; however, he did not usually provide drawings of male genitalia, and when he did it is only the surstyli in dorsal view, which can be variable. Therefore in some cases he described a species under different names in the same publication (e.g. E. amitinus and E. fractus or E. diversus and E. pusillus).Other authors contributing to the African fauna include Becker (1914, 1919), Bezzi and Lamb (1926), Curran (1929), Lamb (1922), Lindner (1956), Loew (1858, 1860) and Rapp (1946), who described only few species, in many cases only one, and they usually did not give drawings or notes on possible relationships.The present taxonomic and systematic status (species names, types and type depositories) of Afrotropical Eudorylini is summarized in Table 1.TABLE 1. Present taxonomic and systematic status of African Eudorylini and their type depository (#: sex known; -: sex unknown; H: holotype; L: lectotype; N: neotype; S: syntypes).Name male female type depository (H, L, S) Claraeola Aczél, 1940 nigripennis group nigripennis (Hardy, 1949) # - H (ISNB) stuckenbergi Földvári sp. n. # - H (NMSA) unplaced species francoisi (Hardy, 1952) # - H (ISNB) hadrosoma (Hardy, 1962) comb. nov. # - H (MNHN) Clistoabdominalis Skevington, 2001 nitidifrons species group nitidifrons (Becker, 1900) # # L (BMNH) Pipunculus confusoides Lamb, 1922 [syn. Kehlmaier, 2005a: 23.] L (BMNH) Dorylomorpha lini Hardy, 1972 [syn. (with P. confusoides) De Meyer, 1995: 288.] H (BPBM) crassus (Becker, 1900) comb. nov. # # L (BMNH) lomholdti species group lomholdti Földvári, 2003 # # H (ZMUC) namibiensis Földvári sp. n. # # H (NMNW) Dasydorylas Skevington, 2001 evanidus species group bodocsi Földvári sp. n. # - H (NMSA) evanidus (Hardy, 1949) # # H (BMNH) Dorilas (Eudorylas) dorsalis Hardy, 1950, syn. nov. H (MRAC) Dorilas (Eudorylas) apiculatus Hardy, 1961, syn. nov. H (MRAC) turneri (Hardy, 1949) # - H (BMNH) sordidatus (Hardy, 1950) # # H (MRAC) africanus species group africanus (Lindner, 1956) comb. nov. # - H (SMNS) minymerus (Hardy, 1962) comb. nov. # - H (MNHN) okongoensis Földvári sp. n. # # H (NMNW) quasidorsalis (Hardy, 1961) # # H (MRAC)……continued on the next pageTABLE 1. (Continued)Name male female type depository (H, L, S) Eudorylas Aczél, 1940 umbrinus species group aemulus (Hardy, 1949) # # H (BMNH) aethiopicus (Hardy, 1949) # # H (BMNH) amani Földvári, 2003 # - H (ZMUC) pilulus Földvári sp. n. # - H (NMSA) umbrinus (Loew, 1858) # - H (NHRS) excisus species group barracloughi Földvári sp. n. # - H (NMSA) excisus (Hardy, 1949) # # H (BMNH) Pipunculus (Eudorylas) definitus Hardy, 1961 [syn. De Meyer and Földvári, 2008: 81] H (MRAC) unanimus species group brandbergensis Földvári sp. n. # # H (NMNW) hirsutus Földvári sp. n. # - H (BMNH) unanimus (Hardy, 1949) # - H (ISNB) aculeatus species group aculeatus (Loew, 1858) # # H (NHRS) gabela Földvári sp. n. # # H (BMNH) libratus (Hardy, 1949) # - H (BMNH) pondolandi Földvári, 2003 # - H (BMNH) setiformis species group angolae Földvári sp. n. # - H (BMNH) discretus (Hardy, 1952) # - H (ISNB) protumidus Földvári sp. n. # - H (NMSA) rooibergensis Földvári sp. n. # - H (NMNW) setiformis (Hardy, 1949) # # H (BMNH) skorpionensis Földvári sp. n. # - H (NMNW) swanengi Földvári, 2003 # - H (BMNH) semiopacus species group acroapex (Hardy, 1962) # - H (BMNH) amitinus (Hardy, 1962) # - H (MNHN) Pipunculus (Eudorylas) fractus Hardy, 1962, syn. nov. H (NHBM) diversus (Hardy, 1949) # - H (BMNH) Pipunculus (Eudorylas) pusillus Hardy, 1949, syn. nov. H (BMNH)……continued on the next page ……continued on the next pageTABLE 1. (Continued)Name male female type depository (H, L, S) facetus (Hardy, 1962) # # H (NHMB) garambensis (Hardy, 1961) # # H (MRAC) Pipunculus (Eudorylas) megacanthus Hardy, 1949, syn. nov. H (MRAC) inornatus (Hardy, 1949) # - H (BMNH) pectinatus Földvári sp. n. # - H (NMSA) semiopacus (Lamb, 1922) # # L (BMNH) encerus species group decorus (Hardy, 1950) # # H (MRAC) encerus (Hardy, 1949) # - H (BMNH) Pipunculus (Eudorylas) modicus Hardy, 1949, syn. nov. H (ISNB) luteopilus (Hardy, 1962) # # H (NHMB) tanzaniensis Földvári, 2003 # # H (ZMUC) mutillatus species group conformis (Hardy, 1959) # - H (MRAC) galeatus (Hardy, 1949) # # H (BMNH) mutillatus (Loew, 1858) # # N (BMNH) Pipunculus kumamotensis Matsumura, 1915 [syn. Yano et al., 1984: 56.] L (EIHU) Pipunculus hepaticolor Becker, 1900 [syn. Skevington, 2003: 660.] unknown Pipunculus cruciator Perkins, 1905 [syn. Skevington, 2003: 660.] L (BPBM) Pipunculus aequalis Becker, 1924 [syn. Hardy, 1968: 456.] H (DEI) Pipunculus matema Curran, 1936 [syn. Hardy, 1968: 457.] H (CAS) Dorilas (Eudorylas) quadratus Hardy, 1949, syn. nov. H (BMNH) Dorilas (Eudorylas) hiatus Hardy, 1956 [syn. Hardy, 1968: 457.] H (BPBM) Pipunculus (Eudorylas) eremnoptera Hardy, 1962, syn. nov. H (BMNH) Pipunculus (Eudorylas) distocruciator Hardy, 1966 [syn. Hardy, 1968: 457.] H (BMNH) Pipunculus (Eudorylas) ranikhetiensis Kapoor et al., 1977 [syn. Kapoor et al., 1987: 103.] unknown Pipunculus (Eudorylas) kumaonensis Kapoor et al., 1977 [syn. Kapoor et al., 1987: 103.] unknown natalensis (Hardy, 1949) # - H (AMNH) porrectus (Hardy, 1949) # - H (BMNH) rubrus (Hardy, 1950) # - H (MRAC) scharffi Földvári, 2003 # - H (ZMUC) vicarius (Hardy, 1949) # - H (BMNH) wittei (Hardy, 1950) # # H (MRAC)TABLE 1. (Continued)Name male female type depository (H, L, S) unplaced species abdominalis (Loew, 1858) - # H (NHRS) amuscarium (Hardy, 1959) # # H (ZML) angustus (Hardy, 1952) - # H (ISNB) bisetosus (Hardy, 1962) # - H (BMNH) bredoi (Hardy, 1949) - # H (ISNB) cupreiventris (Becker, 1914) # - H (ZMHB), lost denotatus (Hardy, 1959) # # H (ZML) falcatus (Hardy, 1949) - # H (BMNH) femoralis Földvári sp. n. # # H (NMSA) flexus (Hardy, 1949) # # H (BMNH) ghesquierei (Hardy, 1950) # # H (ISNB) katonae (Kertész, 1907) # - H (HNHM), lost liberia (Curran, 1929) - # H (AMNH) lobus Földvári sp. n. # - H (NMSA) meruensis (Hardy, 1949) # - H (USNM) mikenensis (Hardy, 1950) # # H (MRAC) parvifrons (Loew, 1858) # - S (MNHU), lost remiformis (Hardy, 1962) # # H (NHMB) sinuosus (Hardy, 1949) # # H (BMNH)MATERIALS AND METHODSThe following institutions and museums kindly put material at the author’s disposal (curators in brackets):AMNH — American Museum of Natural History, New York, USA (D. Grimaldi)BMNH — The Natural History Museum, London, England (N. Wyatt)BPBM — Bernice P. Bishop Museum, Honolulu, Hawaii, USA (N. Evenhuis)CAS — California Academy of Sciences, San Francisco, USA (N. Penny)EIHU — Hokkaido University, Sapporo, Hokkaido, Japan (M. Suwa)HNHM — Hungarian Natural History Museum, Budapest, Hungary (L. Papp)ISNB — Royal Belgian Institute of Natural Sciences, Brussels, Belgium (P. Grootaert)MNHN — Muséum National d'Histoire Naturelles, Paris, France (M. Baylac)MRAC — Royal Museum for Central Africa, Tervuren, Belgium (M. De Meyer)NHMB — Naturhistorisches Museum, Basel, Switzerland (D. Burckhardt)NHRS — Naturhistoriska riksmuseet, Stockholm, Sweden (Th. Pape)NMNW –National Museum of Namibia, Windhoek, Namibia (A. Kirk-Spriggs)NMSA — KwaZulu-Natal Museum, Pietermaritzburg, South Africa (D. A. Barraclough)SMNS — Staatliches Museum für Naturkunde, Stuttgart, Germany (H.-P. Tschorsnig)USNM — National Museum of Natural History, Washington D.C., USA (F. C. Thompson)ZMHB — Museum für Naturkunde der Humboldt–Universität, Berlin, Germany (M. Kotrba)ZML — Zoological Museum, Lund, Sweden (R. Danielsson)ZMUC — Zoological Museum, University of Copenhagen, Copenhagen, Denmark (R. Meier)Dry and alcohol preserved specimens were studied and dissected with an Olympus (SZ 60) stereoscopic microscope at magnifications of 10–112.5 times. Dissected genitalia were placed on microscope slides and studied with a Zeiss (Amplival) light microscope. Drawings were made with a drawing tube (“camera lucida”) attached to a Zeiss (Technival) microscope for stereoscopic drawings and a Zeiss (Amplival) for light microscopic drawings respectively. For some drawings (both stereo and light) a Leica (SM–LUX) was used, provided with a drawing tube as well. The original pencil drawings were copied in ink on tracing paper; scanned and then reduced digitally.Male genitalia were macerated in 10% NaOH or KOH for 24 hours. In some cases the vials containing the genitalia were heated to reduce reaction time. After clearing, the separated postabdomen was put into a drop of gelatine-glycerine on a microscopic slide. This mixture is solid at room temperature and becomes fluid after careful heating, thus, making it possible to change the orientation of the specimen and fix it until the drawing is finished. Eventually, genital parts were cleaned with NaOH or KOH, placed in a small plastic vial filled with glycerine and attached to the same pin, which bears the specimen. Following each treatment with KOH or NaOH all specimens were treated with lactic acid to neutralize the effect, and to stop the clearing process of the OH chemicals.The general terminology of morphological parts of Pipunculidae used in the taxonomic descriptions of the present work follows Albrecht (1990) and Földvári and De Meyer (2000). Definitions and expressions of the postabdomen of Diptera have been discussed by Griffith (1972) and by Sinclair (2000) and Merz and Haenni (2000). The pipunculid male postabdomen terminology follows Skevington (2001, 2002, 2003) and Kehlmaier (2005 a, 2005 b).In Pipunculidae, postabdominal segments 6 and 7 are largely reduced, whereas sternite 8 and tergite 8 are fused, forming the syntergosternite 8. Male genitalia (Fig. 1.) show a circumversion (rotation of 360 degrees along longitudinal axis) and a ventroflexion (rotation directed ventrally); they are also asymmetrical (due to deflexion) in ventral view. Thus the dorsal surface of the genitalia can be examined in the ventral view of the abdomen. An important part of the male genitalia is the epandrium, which is ontogenetically the 9th tergite. This envelops the bilobed hypandrium and accessory structures, i.e. cerci, surstyli, the outer and inner gonopods and the subepandrial sclerite. The trifid phallus and the phallic guide can be examined in ventral view. The form of the phallic guide in ventral or lateral view can be crucial in identifying species of Eudorylas. The phallus is a tubiform structure, which branches apically into ductuli, party sclerotised, partly membranous. Basally the ejaculatory duct ends in a sperm pump and ejaculatory apodeme (i.e. Fig. 3I). The latter serves as an attachment of the muscles that forces the seminal fluid into the duct. The sperm pump is connected to two testes, which are situated in the fifth abdominal segment.The female postabdomen comprises the ovipositor (Fig. 2.), formed by the base (fused tergite and sternite 7) and the piercer (tergite and sternite 9) (sg9 as in Fig. 2B).Descriptions are as detailed as possible in terms of listing all possible characters that can be helpful for species recognition and differentiation between the species. descriptions are uniform making it easier to compare single features. the same applies to the drawings, the six views provided per species being: genitalia in dorsal view; surstyli in dorsal view, enlarged; lateral view of the outer surstylus; lateral view of the inner surstylus; ventral view of genitalia without ST8 and sperm pump with ejaculatory apodeme.Label data are given as present on labels, remarks are between square brackets, and information in “” is the exact transcript of the original.