Diptera) Based on Morphological Data

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Zoological Journal of the Linnean Society, 2013, 167, 43–81. With 3 ﬁgures

Phylogeny and taxonomic revision of all genera of Conopidae (Diptera) based on morphological data

JOEL F. GIBSON1* and JEFFREY H. SKEVINGTON2,3

1Department of Integrative Biology, Biodiversity Institute of Ontario, University of Guelph, Guelph, ON, Canada N1G 2W1 2Agriculture and Agri-Food Canada, Canadian National Collection of Insects, Arachnids, and Nematodes, 960 Carling Ave., Ottawa, ON, Canada K1A 0C6 3Department of Biology, Carleton University, 209 Nesbitt Building, 1125 Colonel By Drive, Ottawa, ON, Canada K1S 5B6

Received 7 May 2012; revised 19 July 2012; accepted for publication 14 September 2012

All global genera of the fly family Conopidae are revised here. A cladistic analysis of 117 morphological characters recorded from 154 species, including representatives of 59 genera and subgenera, recovers a phylogenetic hypothesis for the family. This hypothesis is used as the basis of a new classification for the family. Both Sicini and Zodionini are removed from Myopinae and elevated to subfamilial status. A new tribe, Thecophorini, is proposed within Myopinae to accommodate Thecophora, Scatoccemyia, and Pseudoconops. Two genera, Pseudomyopa and Parazodion, are removed from Dalmanniinae and placed in Myopinae and Zodioninae, respectively. Conopinae is divided into 11 tribes, seven of which are newly described (Asiconopini, Caenoconopini, Gyroconopini, Micro- conopini, Neoconopini, and Siniconopini). Some examined species are transferred to different or new genera and subgenera. A new genus, Schedophysoconops gen. nov., and subgenus Asiconops (Aegloconops) subgen. nov. within Conopinae are described. A review of character evolution and phylogeography is included in light of the new classification. A catalogue of all genus-group names is included with new emendations noted.

ADDITIONAL KEYWORDS: classiﬁcation – genitalia – genus-group catalogue – morphology – synonym – thick-headed ﬂies.

INTRODUCTION hymenopteran pollinators (Freeman, 1966; Mei, 1999). Conopidae have been the subject of biological research Members of the family Conopidae are found in every with regard to host–parasite relationships (e.g. part of the world excluding Antarctica and the Pacific Schmid-Hempel, 2001; Otterstatter, 2004), mimicry islands. The complete life history of most species is (e.g. Waldbauer, 1988), and hilltopping mating behav- unknown, but in instances where it is known, the iour (e.g. Mei, Gibson & Skevington, 2010). Little larvae are obligatory endoparasitoids of other insects. research has been conducted on the phylogenetic rela- Aculeate Hymenoptera, crickets, and cockroaches have tionships between members of Conopidae. been confirmed as hosts. The most important economic Species of Conopidae have been the subject of and ecological impact of species of Conopidae is sporadic, but considerable taxonomic effort over the likely to be their deleterious effect on populations of past 250 years. Species of Conopidae were among the original species descriptions of Linnaeus (1758, 1761). The major figures in 18th and 19th century *Corresponding author. E-mail: [email protected] insect taxonomy all contributed in some way to the

© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167, 43–81 43 44 J. F. GIBSON AND J. H. SKEVINGTON systematics of Conopidae, including Fabricius (1775, of Conopidae are endemic to Australia (Schneider, 1781, 1787, 1794, 1805), Latreille (1797, 1802), Say 2010). To properly revise the present classification, (1823, 1829), Robineau-Desvoidy (1830, 1853), Wiede- as many genera, both global in range and locally mann (1830), Macquart (1834, 1835, 1840, 1843, endemic, as possible must be examined and included. 1846, 1851), Rondani (1843, 1845, 1856, 1857, 1865), Only then can biogeographical hypotheses be formu- Bigot (1859, 1887), and Loew (1847, 1853, 1863, lated for Conopidae. 1866). It was Williston (1882, 1883, 1885, 1888, Identification tools for Conopidae are limited to 1892a, b, 1893), however, who produced the first particular regions. Keys to genera exist for most major works focused on members of Conopidae. geographical regions (Chvála, 1961; Smith, 1969; Meunier (1899, 1912, 1916) described the first con- Zimina, 1970, 1975; Smith & Peterson, 1987; Schnei- firmed fossil specimens of Conopidae. Kröber took a der, 2010; Skevington, Thompson & Camras, 2010), global approach, describing over 20 genera and 200 excluding the Oriental and Afrotropical regions. species in a series of 39 taxonomic papers (e.g. Kröber, There presently exists no identification key to world 1915a, 1936, 1939a, 1940a) published in the first half genera of Conopidae. of the 20th century. The Afrotropical and Oriental Previous phylogenetic analyses based on molecular regions were given early taxonomic attention by and morphological data (Gibson, Skevington & Kelso, Brunetti (1912, 1923, 1925a, b, c, d, 1929) and Chen 2010, 2012) hypothesize at least five monophyletic (1939), respectively. Early regional taxonomic subfamilies of Conopidae: Conopinae, Dalmanniinae, revisions were completed for Stylogaster Macquart, Myopinae, Stylogastrinae, and Zodioninae. These 1835 (Aldrich, 1930; Lopes, 1937) and Dalmannia analyses offer characteristic apomorphies for clades Robineau-Desvoidy, 1830 (Bohart, 1938). Conopidol- including each subfamily as well as clades showing ogy in the second half of the 20th century was relationships between subfamilies. While suites of dominated by the work of Camras. Working on all characters are proposed that vary within subfamilies, subfamilies, from all parts of the world, Camras the taxon sampling of these previous studies does not described over a dozen genera and subgenera and 150 allow conclusions to be drawn about character evolu- species in over 40 publications (e.g. Camras, 1953a, tion within subfamilies. Existing characters must 1955b, 2000, 2008). Smith also contributed to our be supplemented with new characters selected to present knowledge of Conopidae, mainly through his clarify phylogenetic relationships within subfamilies contributions to world catalogues (1969, 1975, 1980, of Conopidae. 1989). Stuke has also been prolific, contributing a Gibson et al. (2012) conclude that members of handful of new descriptions and revisions, as well Zodion Latreille, 1797 and Parazodion Kröber, 1927 as numerous regional faunal lists (e.g. Stuke, 2003, form a monophyletic clade that is sister to Conopinae. 2004, 2005, 2008). The most important taxonomic This hypothesis differs from the traditional placement work on Conopidae in the past 20 years is the revision of Zodion within Myopinae and Parazodion within of Australian species by Schneider (2010), in which Dalmanniinae. The placement of Parazodion within five new genera and one new subfamily are described. Dalmanniinae is based almost exclusively on the Current catalogues (Camras, 1965; Papavero, 1971; shape of the female terminalia (e.g. Camras, 1953a). Smith, 1975, 1980, 1989; Chvála & Smith, 1988) offer The analyses of Gibson et al. (2012) suggest that a combined list of nearly 800 valid, extant species of the terminalia of Dalmannia and those of Parazo- Conopidae, organized into five subfamilies and 56 dion represent independent modifications of female genera and subgenera. Zimina (1960, 1974) divides abdominal segment 7. Baruerizodion Papavero, 1970 the subfamilies Conopinae and Myopinae each into and Pseudomyopa Pearson, 1974 were not included four tribes based on species found in the former in the analysis of Gibson et al. (2012) but have been USSR. Some of the subsequent regional catalogues included in Dalmaniinae based mainly on the female (Camras, 1965, 2000, 2001; Papavero, 1971; Smith, terminalia. 1980; Smith & Peterson, 1987) have adopted this Two genera of Conopidae are currently subdivided tribal classification, each with varying degrees of into subgenera. Physoconops Szilady, 1926 is divided modification. A summary of these tribal classifications into seven subgenera (Camras, 2004), with subgeneric is given in Table 1. Due to their geographical focus, status uncertain for four Afrotropical species (Smith, none of these previous classifications includes all 1980). Gibson et al. (2012) indicate that Physoconops world genera of Conopidae. (sensu lato, s.l.) represents a monophyletic clade, but Biogeographical theories for Conopidae are practi- only three species and two subgenera are tested in cally non-existent. This is probably due to the lack of that analysis. Conops Linnaeus, 1758 is divided into a robust classification. While some genera (e.g. Physo- six subgenera (Camras, 2000). Gibson et al. (2012) cephala Schiner, 1861) are found globally, others are include five species and three subgenera of Conops. more limited in their range. For example, 13 genera Each subgenus is recovered as monophyletic, but

Table 1. Past tribal classiﬁcations for genera of Conopidae

Zimina, 1960 Camras, 1965 Papavero, 1971

Conopidae of USSR Nearctic Conopidae Neotropical Conopidae Conopinae Conopinae Conopinae Brachyceraeini Conopini Conopini Brachyceraea Conops (1 subgenus) Conops (5 subgenera) Neobrachyceraea Physoconops (3 subgenera) Mallachoconops Conopini Physocephalini Physoconops (3 subgenera) Abrachyglossum Physocephala Physocephalini Conops Myopinae Physocephala Brachyglossum (now Leopoldius) Myopa Tropidomyiini Physocephala Robertsonomyia Tropidomyia Tropidomyiini Thecophora Myopinae Tropidomyia Zodion Myopa Myopinae Dalmanniinae Neozodion Myopini Dalmannia Robertsonomyia Melanosoma Stylogasterinae Scatoccemyia Myopa Stylogaster Thecophora Occemyia (now Thecophora) Zodion Sicini Dalmanniinae Sicus Baruerizodion Carbonosicus Parazodion Zodionini Stylogasterinae Zodion Stylogaster Dalmanniini Dalmannia

Smith, 1980 Chvála & Smith, 1988 Camras, 2000, 2001

Afrotropical Conopidae Palaearctic Conopidae Afrotropical Conopinae Conopinae Conopinae Conopinae Conopini Brachyceraeini Conopini Anticonops Brachyceraea Anticonops Archiconops Neobrachyceraea Caenoconops Caenoconops Conopini Conops (3 subgenera) Conops (2 subgenera) Abrachyglossum Euconops Dacops Archiconops Physoconops Euconops Conops (2 subgenera) Physocephalini Physocephalini Leopoldius Dacops Physocephala Macroconops Physocephala Physoconops Neobrachyglossum Pleurocerinella Pleurocerinella Physocephala Pseudophysocephala Pseudophysocephala Siniconops Tropidomyia Tropidomyia Pleurocerinellini Myopinae Pleurocerinella Paramyopa Tropidomyiini Pseudoconops Tropidomyia Thecophora Myopinae Zodion Myopini Stylogasterinae Melanosoma Stylogaster Myopa Myopotta Thecophora Sicini Carbonosicus Sicus Zodionini Zodion Dalmanniinae Dalmannia

Conops (s.l.) is not. Further analysis with members Müncheberg, Germany (SDEI); Tel Aviv University, of all subgenera is necessary to determine the status Tel Aviv, Israel (TAU); R.M. Bohart Museum of Ento- of these two genera. mology, University of California-Davis, Davis, CA, The present research revises all described genera of USA (UCDC); National Museum of Natural History, Conopidae. A phylogenetic hypothesis for the family, Washington, DC, USA (USNM); and personal collec- including representatives of as many genera as pos- tion of Dr Milan Chvála, Charles University, Prague, sible, is created using an analysis of morphologi- Czech Republic. cal characters. In addition to characters previously Colour of body structures is often employed in iden- shown to be phylogenetically informative (Gibson tification keys and phylogenetic analyses of insects. et al., 2012), new characters and character states are Even highly distinctive coloration, however, can be included. The resultant phylogenetic hypothesis is highly variable within some species of Diptera (Hol- used to produce a revised subfamilial and tribal clas- loway, 1993; Costa, Rohde & Valente, 2003). Instances sification. Characteristic apomorphies for newly pro- have been recorded where coloration is dependent on posed and existing groups are presented. season and temperature (Costa et al., 2003; Martín- Vega & Baz, 2011). This is particularly emphasized in species with wide geographical ranges (Holloway, MATERIALS AND METHODS 1993). In Conopidae, in particular, this has led to the TAXON SAMPLING description of a number of species complexes that Representatives of 154 taxa (Table 2), representing 59 include two or three species that vary only in colour genera and subgenera, have been explicitly coded for with intermediate colour morphs often noted (e.g. 117 characters (Table 3). An additional seven extant Camras, 1953b). For this reason, in this analysis, only and two extinct genera were not available as pinned instances of non-variable colours (e.g. wing pattern) specimens. These nine genera (Table 2) are not part are used as the basis of phylogenetic comparison. of the presented phylogenetic analysis, but are dis- Size measurements of various anatomical struc- cussed in light of available information from the lit- tures are also commonly employed in identification erature. Also included are seven species from seven keys and phylogenetic analyses of Diptera. While other dipteran families to serve as outgroup taxa. overall body length and absolute size of some struc- Based on previous analyses of Conopidae relation- tures are somewhat indicative of some genera of ships (Gibson et al., 2010, 2012), Lonchoptera tristis Conopidae, they are highly variable even within (Lonchopteridae) is used to root all trees. species. This may be due to the parasitoid nature of Conopidae. Past research has indicated that adult parasitoid body size is dependent on the size of the MORPHOLOGICAL ANALYSIS host (Kitthawee, 2000). To counteract the problem of Both male and female adult specimens of available variable absolute body measurements, only relative taxa were examined, where possible. Specimens were size comparisons between structures are used to examined under a dissecting stereomicroscope. Speci- determine morphological character states in this mens were obtained from and have been deposited analysis. at the following institutions: American Museum of Natural History, New York, USA (AMNH); Australian Museum, Sydney, New South Wales, Australia REGARDING CAENOCONOPS, CHRYSIDIOMYIA, (AMS); Australian National Insect Collection, CSIRO, AND CALLOSICONOPS Canberra, Australia (ANIC); The Natural History Evenhuis, Pape & Pont (2008) include Caenoconops in Museum, London, UK (BMNH); California Academy their list of post-1930 genera for which type species of Sciences, San Francisco, CA, USA (CAS); Cana- had not been designated. They determine that the dian National Collection of Insects, Arachnids, and proper authorship for the genus is Caenoconops Nematodes, Ottawa, ON, Canada (CNC); University Anonymous in Imperial Institute of Entomology, of Guelph Insect Collection, Guelph, ON, Canada 1940, and that the type species is Ca. subapicalis (DEBU); Utah State University, Logan, UT, USA Kröber, 1939a. They state that Smith’s (1980) desig- (EMUS); Field Museum of Natural History, Chicago, nation of Ca. rhodesiensis as the type species is incor- IL, USA (FMNH); Florida State Collection of rect. Evenhuis et al.’s explanation, while clear, is Arthropods, Gainesville, FL, USA (FSCA); Instituto complicated by a pair of prior taxonomic publications. Nacional de Biodiversidad, Santo Domingo de Kröber (1939a) designates Conops bicolor as a junior Heredia, Costa Rica (INBio); Muséum National synonym of Caenoconops rhodesiensis, itself originally d’Histoire Naturelle, Paris, France (MNHN); Natal described in Conops by Brunetti (1925a). Camras Museum, Pietermaritzburg, South Africa (NMSA); (2000) proposes a series of changes to classifica- Senckenberg Deutsches Entomologisches Institut, tion: he notes that the specimens of Ca. rhodesiensis

Table 2. List of taxa included in our analysis

Species Author Geographical origin Specimens examined Notes

Conopidae: Conopinae Abrachyglossum capitatum Loew, 1847 Switzerland 2M (CNCD79843, JSS25311); 1F (JSS25312) TSP Anticonops abdominalis Kröber, 1936 Zaire lit. only TSP; F only Archiconops insularis Kröber, 1936 Madagascar 2M (JSS25310, JSS25309) TSP; M only Archiconops pseudoerythrocephalus Stuke, 2004 Tanzania 1M (CNCD79844); 1F (CNCD79845) Atrichoparia curticornis (Kröber, 1940a) Australia (SA, WA) 3M (JSS25354, JSS25352, JSS25353); 1F (JSS25355) TSP Atrichoparia sp.A Australia (QLD) 2M (CNCD79828, CNCD82585); 2F (CNCD79826, CNCD82586) Atrichoparia sp.B Australia (NSW) 1M (JSS19811); 1F (JSS19815) Australoconops perbellum (Kröber, 1939b) Australia (WA) 1M (CNCD28476); 2F (JSS25308, JSS25307) Australoconops phaeomeros Schneider, 2010 Australia (QLD) 1M (JSS19800); 1F PT (JSS25357) Australoconops unicinctus (Kröber, 1939b) Australia (ACT, NSW, QLD, WA) 2M (JSS25304, JSS25479); 2F (JSS25305, JSS25306) 2013, Society, Linnean the of Journal Zoological Brachyceraea brevicornis Loew, 1847 Iran, Lebanon 1M (JSS25476); 1F (JSS25477) TSP Caenoconops bicolor (Kröber, 1931) South Africa 1M (JSS25474); 1F (JSS25475) TSP Caenoconops claripennis (Camras, 1962b) Nigeria 1F HT (JSS25472) F only Caenoconops friedbergi Camras, 2000 Kenya 1M HT (JSS25473) M only Camrasiconops ater (Camras, 1961) Australia (NSW, QLD) 1M (JSS19801); 1F (JSS25356) TSP Chrysidiomyia hirsuta (Kröber, 1940a) Australia (WA) 1M (JSS25482) Chrysidiomyia rufa Kröber, 1940a Australia (WA) 1M (JSS25481) TSP Chysidiomyia rugifrons Schneider, 2010 Australia (WA) 1M (CNCD28447) Conops (Asiconops) ater Macquart, 1843 Kenya 1M (JSS25471); 1F (JSS25470) Conops (Asiconops) aureomaculatus Kröber, 1933 China 1M (JSS25468) TSP Conops (Asiconops) australianus Camras, 1961 Australia (QLD) 1M (JSS25302); 2F (JSS19802; JSS25301) Conops (Asiconops) chinensis Camras, 1960 Japan 1M (CNCD79685) M only Conops (Asiconops) nubeculosus Bigot, 1887 Indonesia (W. Java) 1M (CNCD79847); 1F (CNCD79846) Conops (Ceratoconops) ornatus Williston, 1892a Brazil 1M HT (JSS25467) TSP; M only Conops (Conops) ﬂavipes Linnaeus, 1758 Germany, Italy 2M (JSS19803, CNCD79677); 2F (CNCD79678, CNCD79681) TSP

Conops (Conops) nigripes Kröber, 1913 Taiwan 2M (JSS25629, JSS25101); 1F (JSS25103) GENERA CONOPID OF REVISION Conops (Conops) verus Camras, 1955b Brazil 1M HT (JSS25466) M only Conops (Conops) vesicularis Linnaeus, 1761 Germany, Italy, South Korea 2M (CNCD79679, CNCD79682); 2F (CNCD79680, CNCD79684) Conops (Diconops) geminatus Camras, 1957a Costa Rica, Peru 2F (JSS25463, JSS25464) F only Conops (Diconops) trichus Camras, 1957a Brazil 1F PT (JSS25465) TSP; F only 43–81 167, Conops (Smithiconops) guineensis Kröber, 1915d Kenya 1M (JSS25461) M only Conops (Smithiconops) rondanii Bezzi, 1901 South Africa, Tanzania 2M (JSS20500, CNCD79686); 2F (CNCD79688, CNCD79689) TSP Conops (Sphenoconops) brunneosericeus Kröber, 1937 Brazil 1F (JSS25460) Conops (Sphenoconops) nobilis Williston, 1892a Argentina, Brazil 1M (JSS25462); 1F (CNCD79848) TSP Dacops abdominalis Kröber, 1915e Kenya, Mozambique, South Africa 1M (CNCD79849); 3F (JSS25452, JSS25450, JSS25456) TSP Dacops kaplanae Camras, 2001 Ethiopia 1M AT (JSS25451); 1F PT (JSS25454) Delkeskampomyia fasciata Kröber, 1940a Australia (WA) lit. only TSP; M only Euconops bellus Kröber, 1915e South Africa 2M (JSS19804, JSS25303); 2F (JSS25329, JSS25327) TSP Heteroconops antennatus Kröber, 1940a Australia (WA) 2M (JSS25449, JSS25448); 2F (JSS25457, JSS25453) Heteroconops gracilis Kröber, 1915e Australia (NT) 2M (JSS25445, CNCD79850); 1F (JSS25446) TSP Heteroconops sp. Australia (QLD) 2M (CNCD79829, CNCD79851); 1F (CNCD79830) Jelte neotropica (Camras, 2008) Bolivia 1F PT (JSS25444) TSP; F only Leopoldius coronatus (Rondani, 1857) Germany, Italy 2M (CNCD28439, CNCD79630); 2F (CNCD79691, CNCD79693) Leopoldius signatus (Wiedemann, 1824) England, Italy 2M (CNCD79866, CNCD79867); 2F (CNCD79868, CNCD79869) TSP 47 48 Table 2. Continued .F ISNADJ .SKEVINGTON H. J. AND GIBSON F. J. Species Author Geographical origin Specimens examined Notes

Macroconops helleri Kröber, 1927 China lit. only TSP; M only Mallachoconops atratulus (Malloch, 1933) Chile 1M HT (CNCD79852) TSP; M only Microbrachyceraea pendleburyi (Brunetti, 1927) Malaysia lit. only TSP; M only Microconops nigrithorax Kröber, 1940a Australia (WA) 3M (CNCD82591, JSS25484, JSS25485); 1F (CNCD82592) Microconops ornatus Kröber, 1915e Australia (ACT, QLD, WA) 2M (JSS25487, CNCD79825); 2F (JSS10483, JSS18162) TSP 03TeLnenSceyo London, of Society Linnean The 2013 © Microconops similis Kröber, 1940a Australia (SA) 2M (JSS25489, JSS25490); 2F (JSS25492, JSS25493) Microconops tasmaniensis Kröber, 1940a Australia (NSW, TAS, VIC) 2M (JSS25497, JSS25494); 2F (JSS25496, JSS25495) Neobrachyceraea elongata Stuke & Clements, 2005 Thailand 1F (JSS25442) Neobrachyceraea obscuripennis (Kröber, 1913) China 1M (JSS25443) Neobrachyglossum punctatum Kröber, 1915e Turkey? lit. only TSP; M only Neoconops brevistylus Schneider, 2010 Australia (QLD) 1M (CNCD6871) Neoconops longicornis Kröber, 1915e Australia (NSW, WA) 1M (JSS25483); 1F (CNCD82588) TSP Physocephala bimarginipennis Karsch, 1887 Angola, Congo, Tanzania 1M (CNCD79853); 2F (CNCD79854, JSS25455) Physocephala maculipes (Bigot, 1887) Tanzania, Tunisia, Uganda 3M (JSS19865, CNCD79696, CNCD79697); 1F (CNCD79695) Physocephala madagascariensis Kröber, 1915c Madagascar 2M (JSS19808, JSS25324); 2F (JSS25322, CNCD79823) Physocephala marginata (Say, 1823) Canada (ON, QC) 3M (CNCD349, JSS17567, JSS17621); 1F (CNCD79698) Physocephala ruﬁpes (Fabricius, 1781) Germany, Italy, Serbia 2M (CNCD79705, CNCD79824); 2F (CNCD79703, CNCD79704) TSP Physocephala tibialis (Say, 1829) Canada (ON), USA (VA) 2M (JSS17725, JSS18866); 2F (JSS18864, CNCD79706) Physoconops (Aconops) antennatus (Kröber, 1915f) Argentina, Brazil 1M (CNCD79856); 2F (CNCD79855, CNCD79859) TSP Physoconops (Aconops) costatus (Fabricius, 1805) Argentina, Venezuela 2M (CNCD79857, CNCD79858); 2F (JSS25441, JSS25440) Physoconops (Aureoconops) aureolus Camras, 2004 Peru 1M HT (JSS25439) TSP; M only Physoconops (Gyroconops) abbreviatus Camras, 1955b Mexico 1M (JSS25438); 1F (JSS25435) 2013, Society , Linnean the of Journal Zoological Physoconops (Gyroconops) parvus (Williston, 1892a) Brazil, Costa Rica, Mexico 1M (JSS25437); 1F (JSS18156) Physoconops (Gyroconops) sylvosus (Williston, 1882) Mexico, USA (AL, UT) 2M (CNCD79862, CNCD79861); 2F (CNCD79860, JSS25436) TSP Physoconops (Kroeberoconops) argentinus Camras, 2004 Argentina 1M HT (JSS19749) Physoconops (Kroeberoconops) hermanni (Kröber, 1915f) Brazil 1F (JSS25433) TSP Physoconops (Kroeberoconops) ruﬁpennis (Macquart, 1843) Argentina 1M (CNCD79864); 1F (CNCD79865) Physoconops (Pachyconops) (Macquart, 1843) Canada (ON), USA (MD, MI) 2M (JSS19812, CNCD79750); 2F (CNCD79748, CNCD79749) brachyrhynchus Physoconops (Pachyconops) bulbirostris (Loew, 1853) Guatemala, USA (AL, MS, SC) 2M (CNCD79870, JSS25434); 2F (CNCD79871, JSS25432) TSP Physoconops (Pachyconops) guianicus (Curran, 1934) Costa Rica, Venezuela 2M (JSS19954, JSS25317); 1F (JSS19813) Physoconops (Physoconops) discalis (Williston, 1892a) Brazil, Mexico, Venezuela, USA 2M (JSS19814, CNCD79752); 1F (CNCD79755) (AZ) Physoconops (Physoconops) fronto (Williston, 1885) Canada (MB), USA (IA, MI, NM) 2M (CNCD79872, CNCD79873); 2F (CNCD79874, CNCD79875) Physoconops (Physoconops) microvalvus (Kröber, 1930) Indonesia (W. Java) 2M (JSS25431, JSS25429); 2F (JSS25430, JSS25428) Physoconops (Physoconops) obscuripennis (Williston, 1882) Canada (BC), USA (AL, DE) 2M (JSS25420, CNCD79877); 2F (JSS25427, CNCD79878) TSP Physoconops (Physoconops) sepulchralis (Brunetti, 1912) India 1M (JSS25424) Physoconops (Shannonoconops) apicalis Camras, 1955b Brazil 1M (CNCD79879) TSP Physoconops notatifrons Camras, 1962b Kenya, Zambia 3F (JSS25384, JSS25423, JSS25425) F only Physoconops quadripunctatus (Kröber, 1915d) South Africa 1M (JSS25426) Physoconops rhodesiensis (Brunetti, 1925a) South Africa 1M (JSS25418); 1F (JSS25419) Pleurocerina brevis Schneider, 2010 Australia (QLD) 1M (JSS25350) 43–81 167, Pleurocerina fasciata Macquart, 1851 Australia (NSW) 1M (JSS25344); 1F (JSS25346) TSP Pleurocerina longicornis (Kröber, 1915e) Australia (NT) 1M (JSS25349); 1F (JSS25351) Pleurocerina luteiceps Schneider, 2010 Australia (WA) 3M (JSS25416, JSS25408, JSS25409); 1F (JSS25417) Pleurocerina turneri (Camras, 1961) Australia (WA) 2M (JSS25413, JSS25412); 2F (JSS25414, JSS25415) 03TeLnenSceyo London, of Society Linnean The 2013 © Pleurocerina vespiformis Schneider, 2010 Australia (NSW, QLD) 1M (JSS19810); 1F (JSS25345) Pleurocerinella albohalterata Smith, 1960 Botswana, South Africa 1M (JSS17437); 1F (JSS25109) Pleurocerinella copelandi Stuke, 2009 Botswana 1M HT (JSS25410) M only Pleurocerinella srilankai Stuke & Camras, 2009* Sri Lanka 1M HT (JSS25411) M only Pleurocerinella tibialis Chen, 1939 China 1F (JSS25108) Pleurocerinella sp. Nepal 3M (CNCD82589, CNCD82584, CNCD82590) Pseudophysocephala constricta (Kröber, 1915c) Lesotho, South Africa 1M (JSS25405); 2F (JSS25407, CNCD79880) Pseudophysocephala platycephala (Loew, 1853) South Africa 1M (JSS25406) TSP Setosiconops epixanthus Schneider, 2010 Australia (WA) 1M (JSS25320) M only Setosiconops robustus (Kröber, 1940a) Australia (WA) 1F (JSS25319) TSP Siniconops elegans Chen, 1939 China 1M (JSS25105); 1F (JSS25104) TSP Smartiomyia arena Schneider, 2010 Australia (QLD) 1M (CNCD79827); 1F (JSS19816) Smartiomyia obscura Kröber, 1940a Australia (WA) 1M (JSS25403); 1F (JSS25404) TSP Stenoconops niger Kröber, 1939b Australia (WA) lit. only TSP; F only Tammo rufa (Camras, 1955a) Mozambique 1M (JSS25402) TSP Tanyconops longicaudus Schneider, 2010 Australia (SA) 1F (JSS25401) TSP 2013, Society, Linnean the of Journal Zoological Tropidomyia alexanderi Camras, 1955a Argentina, Brazil 2M (CNCD79881, JSS25399); 1F (JSS25400) Tropidomyia aureifacies Kröber, 1915e Lebanon 1M (JSS25397) Tropidomyia bimaculata Williston, 1888 Costa Rica 2M (JSS25396, JSS25480); 2F (JSS25393, JSS25040) CT TSP Tropidomyia ornata Kröber, 1915e Côte d’Ivoire, Malawi, Namibia, 2M (JSS25395, CNCD79882); 2F (JSS25390, JSS25391) South Africa Tropidomyia sp. Brazil 1M (CNCD79883), 1F (CNCD79884) Conopidae: Dalmanniinae Baruerizodion steyskali Papavero, 1970 Brazil 1F (JSS18157) TSP; F only Dalmannia aculeata (Linnaeus, 1761) Italy, Spain 3M (JSS19818, CNCD79717, CNCD79716); 1F (CNCD79715) TSP Dalmannia nigriceps Loew, 1866 Canada (BC, ON, QC) 2M (JSS17637, CNCD79712) 2F (CNCD79708, CNCD79709) Dalmannia vitiosa Coquillett, 1892 Canada (AB, ON, QC) 2M (JSS17638, CNCD12810); 2F (CNCD79710, CNCD79713) Parazodion schmidti Kröber, 1927 Brazil, Guyana, French Guiana 1M (JSS19838); 3F (JSS25321, CNCD79699, CNCD79701) TSP Parazodion sp. Brazil 1F (JSS25338) Pseudomyopa camrasi Pearson, 1974 Brazil 1M PT (JSS25337) TSP; M only Conopidae: Myopinae

Carbonosicus carbonarius (Kröber, 1915e) China 1M CT (JSS25500); 1F CT (JSS25501) TSP GENERA CONOPID OF REVISION Melanosoma bicolor (Meigen, 1824) Italy, Syria 2M (CNCD79885, CNCD79887); 1F (CNCD79886) Melanosoma hyalipenne Kröber, 1915b Israel 1M (JSS25334); 1F (JSS25333) Myopa buccata (Linnaeus, 1758) England, Israel, Germany 2M (CNCD79718, CNCD79721); 2F (CNCD79722, CNCD79719) TSP Myopa clausa Loew, 1866 Canada (BC, QC, NS, SK) 2M (JSS17625, CNCD84918); 2F (CNCD79723, CNCD79725) 43–81 167, Myopa occulta Wiedemann, 1824 Czech Rep., Germany, Italy, Spain 2M (JSS19820, CNCD79727); 2F (CNCD79728, CNCD79730) Myopa vesiculosa Say, 1823 Canada (BC, ON, QC) 2M (JSS17525, CNCD12801); 2F (CNCD79731, CNCD79732) Myopa sp. Australia (NSW) 1M (JSS19821) Myopotta pallipes (Wiedemann, 1824) England? 1M (JSS25331); 2F (JSS25330, JSS25332) TSP Myopotta rubripes (Villeneuve, 1909) Germany 2M (CNCD79888, CNCD79889); 1F (JSS25389) Neozodion pruinosum Szilady, 1926 Peru lit. only TSP; F only Paramyopa oestracea (Loew, 1863) Lesotho, South Africa 3M (JSS25339, JSS25341, JSS25340); 1F (JSS25342) Pseudoconops antennatus Camras, 1962a Madagascar 4F (JSS25318, JSS25316, JSS25315, JSS19822) F only Robertsonomyia mexicana Camras, 1979 Mexico 2M (JSS25388, JSS25386) Robertsonomyia palpalis (Robertson, 1901) Canada (AB), USA (GA, MS, NC) 1M (CNCD79892); 3F (CNCD79891, CNCD79890, CNCD82596) TSP Robertsonomyia parva (Adams, 1903) Mexico, USA (CA, UT) 2M (JSS25506, CNCD79893); 2F (JSS25507, JSS25509) Robertsonomyia pearsoni (Camras, 1976) Brazil 2F PT (JSS25385, JSS25387) Scatoccemyia plaumanni Camras, 1957a Brazil 1M HT (JSS25502) TSP; M only Sicus abdominalis Kröber, 1915e Italy, Japan, Russia 2M (JSS25505, CNCD82594); 2F (CNCD82593, JSS25504) 49 Sicus ferrugineus (Linnaeus, 1761) France, Sweden 2M (CNCD79757, CNCD79758); 2F (CNCD79756, CNCD79759) TSP Table 2. Continued 50

Species Author Geographical origin Specimens examined Notes SKEVINGTON H. J. AND GIBSON F. J.

Thecophora africana (Brunetti, 1925b) Madagascar 1F (JSS19826) Thecophora atra (Fabricius, 1775) England, Czech Rep., Germany, 2M (CNCD79739, CNCD79738); 2F (CNCD79735, CNCD79737) TSP Spain Thecophora australiana (Camras, 1955a) Australia (NSW, QLD) 2M (JSS19823, CNCD79736) Thecophora metallica Camras, 1962a Madagascar 1F (JSS19824) F only Thecophora modesta (Williston, 1883) Canada (AB, BC, YT), USA (OR) 2M (CNCD1368, CNCD79740); 2F (CNCD79741, CNCD79742) 03TeLnenSceyo London, of Society Linnean The 2013 © Thecophora occidensis (Walker, 1849) Canada (ON, QC, SK) 2M (JSS17732, CNCD79745); 2F (CNCD79747, CNCD79743) Zodiomyia sumbaensis Camras, 1957b India, Sri Lanka 2F (JSS25336, JSS25335) TSP; F only Zodion cinereum (Fabricius, 1794) Italy, Turkey 2M (CNCD79793, CNCD79792); 2F (JSS25313, CNCD79791) TSP Zodion erythrurum Rondani, 1865 Israel, Italy 2M (JSS19837, CNCD79797); 1F (CNCD79798) Zodion fulvifrons Say, 1823 Canada (ON, QC), USA (TX) 2M (CNCD12799, CNCD79806); 2F (CNCD79800, CNCD79801) Zodion griseum Brunetti, 1923 India 1M (JSS25392); 1F (JSS25394) Zodion intermedium Banks, 1916 Canada (ON, QC), USA (MS) 2M (JSS17582, CNCD79803); 2F (CNCD79807, CNCD79804) Zodion pictum Schiner, 1868 Argentina, Ecuador, Mexico 1M (CNCD79796); 3F (JSS19839, CNCD79794, CNCD79795) Conopidae: Notoconopinae Notoconops alexanderi Schneider, 2010 Australia (NSW) 1M TSP Conopidae: Palaeomyopinae Palaeomyopa tertiaria Meunier, 1912 Baltic Amber fossil lit. only TSP; F only Conopidae: Stylogastrinae Stylogaster biannulata (Say, 1823) Canada (ON), USA (MS, TX) 2M (CNCD79763, CNCD85163); 2F (JSS19827, CNCD79761) Stylogaster breviventris Aldrich, 1930 Ecuador, Guyana, Peru 2M (JSS19828, CNCD79765); 2F (CNCD79766, CNCD79767) Stylogaster decorata Aldrich, 1930 Ecuador, Guyana, Peru 2M (CNCD79769, CNCD79770); 2F (JSS19829, CNCD79771)

2013, Society , Linnean the of Journal Zoological Stylogaster frauci Smith, 1979 Australia (QLD) 2M (JSS19830, JSS19831); 2F (CNCD79773, CNCD79774) Stylogaster inca Camras & Parrillo, 1985 French Guiana 1F (CNCD79764) F only Stylogaster neglecta Williston, 1883 Canada (ON), USA (MD, NM) 2M (CNCD79778, CNCD79785); 2F (CNCD79777, CNCD79780) Stylogaster pauliani Camras, 1962a Madagascar 2M (JSS198932, CNCD79772) Stylogaster rectinervis Aldrich, 1930 Brazil, Ecuador, French Guiana 2M (CNCD79782, CNCD79784); 2F (JSS19833, CNCD79781) Stylogaster stylata Fabricius, 1805 Brazil, Ecuador, French Guiana 3M (CNCD28441, CNCD79788, CNCD79789); 1F (CNCD28448) TSP Stylogaster westwoodi Smith, 1967 Congo, Kenya, Nigeria, Tanzania, 2M (CNCD79787, CNCD79790); 2F (CNCD79786, JSS19836) Zambia Stylogaster sp. Madagascar 2M (CNCD28498, JSS19834) Conopidae: subfam. unknown Hoffeinsia baltica Stuke, 2005 Baltic Amber fossil lit. only TSP; M only Outgroups Lonchopteridae: Lonchoptera tristis Meigen, 1824 Czech Rep., Denmark, England 2M (CNCD79832, CNCD79833); 2F (CNCD79834, CNCD79876) Phoridae: Conicera (Conicera) dauci Meigen, 1830 Canada (QC), USA (AZ) 2M (CNCD28440, CNCD79839) 2F (CNCD79840, CNCD79841) Syrphidae: Toxomerus marginatus (Say, 1823) Canada (ON, QC) 2M (CNCD28446, CNCD79820); 2F (CNCD79842, CNCD79821) Lauxaniidae: Minettia lupulina (Fabricius, 1787) Canada (QC), USA (OR) 2M (CNCD79816, CNCD79817); 2F (CNCD79831, CNCD79818) Psilidae: Chyliza scrobiculata Melander, 1920 Canada (BC) 2M (CNCD79813, CNCD79812); 2F (CNCD79815, CNCD79814) Pygotidae: Pyrgota undata Wiedemann, 1830 Canada (ON, QC) 2M (CNCD79836, CNCD79837); 2F (CNCD79835, CNCD79838) Strongylophthalmyiidae: Melander, 1920 Canada (BC, ON, QC) 2M (CNCD79809, CNCD79811); 2F (CNCD28445, CNCD79808) Strongylophthalmyia angustipennis 43–81 167, Proposed tribal classiﬁcation is not included. Genus, subgenus, and subfamily classiﬁcations are listed according to current catalogues prior to this analysis. Specimen voucher numbers provided: CNCD, CNC Diptera collection; JSS, Jeff Skevington Specimen; TSP, type species of genus or subgenus; M only, only males have been described for this species; F only, only females have been described for this species; HT, holotype; PT, paratype; AT, allotype; CTs, co-types; lit. only, only the literature description(s) were examined. *In Stuke (2009). 03TeLnenSceyo London, of Society Linnean The 2013 ©

Table 3. List of morphological characters included in phylogenetic analysis

Character Character states

1. Ptilinum (0) absent; (1) present 2. Central ocellus (0) present; (1) absent 3. Lateral ocelli (0) present; (1) absent 4. Ocellar tubercle (0) present, (1) absent 5. Divided vertex (0) present; (1) absent 6. Expanded vertex (0) absent; (1) present 7. Large, square, setose vertex (0) absent; (1) present 8. Broad longitudinal grooves on vertex (0) absent; (1) present 9. Fine longitudinal grooves on vertex (0) absent; (1) present 2013, Society, Linnean the of Journal Zoological 10. Texture of frons (0) almost entirely smooth; (1) with extensive lateral grooves 11. Posterior margin of compound eye (0) round; (1) with triangular, shiny notch 12. Fronto-orbital bristles (0) present; (1) absent 13. Convergent rows of frontal setae (0) absent; (1) present 14. Ocellar bristles (0) present; (1) absent 15. Postocellar bristles (0) present; (1) absent 16. Gena (0) small; (1) expanded to one-third of the total height of the head. 17. Long, dense, white occipital setae (0) absent; (1) present 18. Compound eye (0) ommatidia uniform across width; (1) ommatidia larger anteriorly 19. Facial fovea (0) absent; (1) present 20. Reduced facial fovea (0) absent; (1) present 21. Lunule (0) not developed; (1) expanded, visible between antennal bases and ptilinum. 22. Facial carina (0) absent; (1) present 23. Prominent facial carina (0) absent; (1) present GENERA CONOPID OF REVISION 24. Facial ridge (0) absent; (1) present 25. Pointed facial ridge (0) absent; (1) present 26. Row of dark setae on parafacial (0) absent; (1) present 43–81 167, 27. Scape (0) quadrate; (1) elongate 28. Pedicel (0) quadrate; (1) elongate 29. Transverse dorsal ridge on pedicel (0) absent; (1) present 30. Pedicel (0) narrow; (1) broad 31. Shape of first flagellomere (0) parallel sided or tapered; (1) distinctly rounded ventrally 32. Length of first flagellomere (0) short; (1) elongate 33. Location of arista (0) dorsal; (1) apical 34. Arista (0) filiform; (1) thickened 35. Retracted arista (0) absent; (1) present 36. Second aristomere (0) as wide as other segments; (1) expanded ventrally 37. Anterior margin of subcranial cavity (0) broad, straight or rounded; (1) pointed anteriorly

38. Bulb on anterior margin of subcranial cavity (0) absent; (1) present 51 Table 3. Continued 52

Character Character states SKEVINGTON H. J. AND GIBSON F. J.

39. Setae on maxillary palpi (0) present; (1) absent 40. Elongate maxillary palpus (0) present; (1) absent 41. Clypeus (0) reduced, bare; (1) broad, microsetose 42. Fused prementum (0) absent; (1) present 43. Greatly elongate prementum (0) absent; (1) present 03TeLnenSceyo London, of Society Linnean The 2013 © 44. Folded labellum (0) present; (1) absent 45. Shape of labella (0) broad, separate; (1) ﬁliform, fused 46. Expanded labella (0) absent; (1) present 47. Basisternum shape (0) broad; (1) short, narrow single sclerite; (2) narrow sclerite divided posteriorly 48. Posterolateral extensions of the basisternum (0) absent; (1) present 49. Setae on posterolateral extensions of the basisternum (0) absent; (1) present 50. Foretibial apical spurs (0) absent; (1) present 51. Shape of metafemur (0) parallel or spindle-shaped; (1) distinctly broadened basally 52. Double row of spines on ventral surface of femora (0) absent; (1) present 53. Apical shiny patch on metatibia (0) absent; (1) present 54. Prominent row of setae on mesofemur (0) absent; (1) present 55. Male pro- and mesocoxa (0) bare or with sparse setae; (1) with dense patch of long, pale setae 56. Diagonal, pale marking on metacoxa (0) absent; (1) present 57. Pleural sclerites (0) bare and shiny with setae and/or bristles; (1) entirely micropilose 2013, Society , Linnean the of Journal Zoological 58. Ventral half of proepisternum (0) with setae and/or bristles; (1) bare 59. Dorsal half of proepisternum (0) bare; (1) with patch of setae 60. Postpronotal bristles (0) present; (1) absent 61. Number of supra-alar bristles (0) zero or one; (1) two 62. Anepimeral bristles (0) absent; (1) present 63. Anepisternal bristles (0) absent; (1) present 64. Katepisternum (0) bare; (1) with bristles and/or setae 65. Scutellum (0) developed; (1) reduced, lacking bristles 66. Scutellar bristles (0) zero, one, or two pairs; (1) three or more pairs 67. Wing pattern (0) partly or completely opaque or darkened; (1) completely hyaline 68. Dark anterior wing pattern (0) absent; (1) present

69. M vein (0) parallel to vein R4+5; (1) deﬂected towards vein R4+5

70. Petiole of vein R4+5+M (0) absent or short; (1) elongate 71. Extended Sc vein (0) absent; (1) present

72. Extended R1 vein (0) absent; (1) present

73. Veins Sc and R1 (0) separate for their entire length; (1) fused before reaching costa

74. Costa (0) even in width along length; (1) thickened at insertion of Sc + R1 43–81 167, 75. Vein R2+3 (0) ends beyond endpoint of vein R1; (1) ends at or near endpoint of vein R1 76. Crossvein sc-r (0) absent; (1) present

77. Obtuse angle between vein CuA2 and cross-vein dm-cu (0) absent; (1) present

78. Elongate vein CuA2 + A1 (0) absent; (1) present 03TeLnenSceyo London, of Society Linnean The 2013 ©

79. Vein CuA2 (0) straight; (1) curved 80. Cell cup (0) narrow; (1) broad 81. Vena spuria (0) absent; (1) present 82. Broad alula (0) absent; (1) present 83. Expanded wing margin (0) absent; (1) present 84. Shiny, pollinose band on abdominal tergite 2 (0) absent; (1) present 85. Female abdominal sternites 2 and 3 (0) present; (1) absent 86. Patch of bristles on female abdominal sternite 2 (0) absent; (1) present 87. Female abdominal tergite 5 and sternite 5 (0) separate; (1) fused 88. Female abdominal tergite 6 and sternite 6 (0) separate; (1) fused 89. Female abdominal tergite 7 and sternite 7 (0) separate; (1) fused 90. Rows of spicules on female abdominal sternite 6 (0) absent; (1) present 91. Rows of spicules on female abdominal sternite 5 (0) absent; (1) present

2013, Society, Linnean the of Journal Zoological 92. Spicules conﬁned to tip of ventral genital plate (0) absent; (1) present 93. Medial lobe on ventral genital plate (0) absent; (1) present 94. Female abdominal syntergite 8+9 fused with cerci and elongate (0) absent; (1) present 95. Narrow female abdominal segments 5–7 (0) absent; (1) present 96. Female abdominal segment 7 deﬂected ventrally (0) absent; (1) present 97. Female abdominal segment 7 (0) absent or broad; (1) compressed laterally along entire length; (2) compressed laterally only in posterior half; (3) narrowed in posterior half 98. Female epiproct (0) present; (1) absent 99. Elongate, sclerotized hypoproct in female (0) absent; (1) present 100. Two square lobes on female abdominal sternite 8 (0) absent; (1) present 101. Lateral lobes on female abdominal sternite 8 (0) absent; (1) present 102. Sclerotized posteroventral hooks on female abdominal syntergite 8+9 (0) absent; (1) present 103. Male abdominal sternites 2–4 (0) partly or entirely membranous; (1) broad, sclerotized plates

104. Male abdominal tergites 3 and 4 (0) narrow and/or tapering; (1) broad and parallel-sided GENERA CONOPID OF REVISION 105. Male abdominal tergites 2 and 3 (0) broad, rectangular; (1) petiolate 106. Male abdominal sternite 5 (0) unmodiﬁed; (1) broad, with setae or spicules 107. Male abdominal sternite and tergite 5 (0) separate; (1) fused

43–81 167, 108. Male abdominal syntergosternite 7+8 (0) absent; (1) present 109. Broad, hemispherical male abdominal syntergosternite 7+8 (0) absent; (1) present 110. Apex of male abdomen (0) rounded or pointed; (1) square 111. Male abdominal sternite 8 (0) absent; (1) present 112. Dense, long, black, setae on male abdominal sternite 8 (0) absent; (1) present 113. Elongate male cerci (0) absent; (1) present 114. Male cercal attachment (0) broad; (1) narrow, sclerotized stalk 115. Epandrium (0) two separate lobes beyond cerci; (1) fused beyond cerci 116. Medial spine on epandrium (0) absent; (1) present 117. Posterior hump on epandrium (0) absent; (1) present

All character states are given. All multi-state characters are treated as unordered and non-additive. 53 54 J. F. GIBSON AND J. H. SKEVINGTON examined by Kröber were actually misidentified RESULTS Conops bicolor, thereby removing Co. bicolor from MORPHOLOGICAL CHARACTERS ANALYZED synonymy with Ca. rhodesiensis; he designates A list of all morphological characters included in Caenoconops subapicalis as a junior synonym of the phylogenetic analysis is included in Table 3. A Caenoconops bicolor; he transfers Ca. rhodesiensis to full character matrix for all included taxa is avail- Physoconops; he transfers Physoconops claripennis to able online as Table S1. Morphological terminology Caenoconops; and he describes Caenoconops fried- follows that of Cumming & Wood (2009) and the bergi. Accounting for this labyrinthine trio of papers, conopid-specific terminology of Gibson et al. (2012). In the following interpretation is made of the present instances where male or female specimens are not state of Caenoconops, in accordance with Article available (as noted in Table 2), characters are deter- 67.1.2. of the I.C.Z.N. (1999). mined from literature descriptions or else left as The type species of Caenoconops is subapicalis unknown. Notes on the initial definition and informa- Kröber, 1939a, which is now a junior synonym of Ca. tive qualities of each character, as well as the original bicolor (Kröber, 1931), by subsequent designation reference where applicable, are discussed in Gibson of Anonymous in Imperial Institute of Entomology et al. (2012), as well as in the Discussion section. (1940). The other valid species are Ca. claripennis A total of 117 characters are included in the analy- and Ca. friedbergi, with rhodesiensis currently placed sis. Characters included from each body region are as in Physoconops. follows: head – 46 characters (39.3%); thorax, includ- Also as part of their effort to clarify type species ing legs – 20 characters (17.1%); wing – 17 characters for genus group names, Evenhuis et al. (2008) note (14.5%); female abdomen, including terminalia – 19 that the original description of Chrysidiomyia Kröber, characters (16.2%); male abdomen, including termi- 1940a was a nomen nudum. The Australian catalogue nalia – 15 characters (12.8%). No characters are con- of Smith (1989) is the first to include a type species stant and all are parsimony informative. and therefore the correct authorship is Chrysidiomyia Smith, 1989. Following examination of many specimens, Schnei- der (2010) proposes that the differences between PARSIMONY ANALYSIS Chrysidiomyia and Callosiconops included by Kröber Parsimony analysis of all available pinned specimens (1940a) in his original descriptions are not sufficient (154 species of Conopidae plus seven outgroup species) to justify separate genera and synonymizes them. recovers 72 unique, equally parsimonious trees [length Unfortunately, she chooses Callosiconops as the 218; consistency index (CI) 0.550; retention index (RI) junior synonym, despite the correct author-date of 0.957]. The strict consensus of these trees is shown in Chrysidiomyia being Smith, 1989. This error is cor- Figure 1 with bootstrap and Bremer support values for rected here with Chrysidiomyia Smith, 1989 consid- each node. The full data matrix and all trees recovered ered a junior synonym of Callosiconops Kröber, are available from treebase.org (http://purl.org/phylo/ 1940a. treebase/phylows/study/TB2:S13087). The illustrated tree (Fig. 1) is well supported with 51 of 83 nodes (61%) having a bootstrap support value PARSIMONY ANALYSIS greater than 50%. New subfamilial and tribal clades All characters were equally weighted and treated are labelled and discussed below; full details of as unordered and non-additive. Unknown character generic membership are discussed in a later section. states were included as ‘?’ in the matrix. Parsimony A clade including all members of Schizophora is analysis was conducted using TNT (v1.1; March, recovered as monophyletic [boot – 70; TBS – 2; mor- 2011) (Goloboff, Farris & Nixon, 2008). A new tech- phological autapomorphies (MAA) – 1:1, 19:1, 22:1, nology search involved generating 1000 random- 24:1, 64:1, 89:1, 108:1; other morphological apomor- addition sequence Wagner trees, ratcheting, drift, and phies and reversals (MA) – 60:1, 82:1). Conopidae sectorial searches with default parameters. Bootstrap (except Notoconops) is recovered (boot – 82; TBS – 4; resampling support values (boot) were determined in MAA – 34:1, 42:1, 43:1, 45:1, 48:1, 54:1, 69:1, 88:1; TNT by a traditional search with 1000 replicates. MA – 28:1, 78:1, 98:1). Total Bremer support (TBS) values (Bremer, 1988, Conopinae is recovered (boot – 100; TBS – 11; 1994) were determined in TNT by performing a MAA – 33:1, 36:1, 115:1; MA – 5:1, 14:1, 15:1, 27:1, search for trees suboptimal by up to 1000 steps and a 37:1, 40:1, 49:0, 78:0, 81:1). Zodioninae is recovered relative fit difference of 1.0. Character optimizations (boot – 65; TBS – 1; MAA – 47:2; MA – 66:1). Sicinae were performed in MacClade 4.06 (Maddison & Mad- is recovered (boot – 62; TBS – 1; MAA – 112:1; MA – dison, 1992) using an accelerated transformation 49:0). Myopinae is recovered (boot – 79; TBS – 3; approach. MAA – 47:1, 73:1; MA – 52:1). Dalmanniinae is

REVISION OF CONOPID GENERA 55

Stylogastrina

Dalmanniinae

Myopinae

Sicinae

Zodioninae

) dae thalmyii

Thecophorini

Myopini

dae)

oph gyl on

dae)

(Str

(Phori

g. 1 g.

(Syrphi

(Psilidae)

lica

eyskali

(Pyrgotidae)

aumanni

eata

see F see

eviventris

griceps

tiosa

alexanderi alexanderi

ylata

arginata arginata

on st on

. . sp

yia pl yia

nae

ectanegl er

er inca er

er recti er nervis

er br er

er westwoodi er

frauci er

izodi

tipenngus an lmyia tha yloph is

occem

ong

decorata ylogaster

ylogast

ylogaster

iani paul ylogaster

st ylogaster

annulata bi ylogaster

onop

otoconops otoconops

almannia vi almannia

almannia ni almannia

almannia acul almannia

Zodiomyia sumbaensis sumbaensis Zodiomyia

Zodion pictum Zodion

Zodion intermedium Zodion

Zodion griseum Zodion

Zodion fulvifrons Zodion

Zodion erythrurum Zodion

Zodion cinereum Zodion

(Lonchopteri tristis Lonchoptera

cera (Conicera) dauci (Conicera) cera Coni

m Toxomerus

culata culata scrobi iza Chyl

Minettia(Lauxaniidae) lupulina Str

Pyrgota undata

Stylogast

Scat

Thecophora occidensis Thecophora

Thecophora modesta Thecophora

Thecophora metal Thecophora

Thecophora australiana Thecophora

Thecophora africana Thecophora

Baruer

Thecophora atra Thecophora

Pseudoconops antennatus Pseudoconops

Melanosoma hyalipenne Melanosoma

Melanosoma bicolor Melanosoma

Pseudomyopa camrasi Pseudomyopa

Myopotta rubripes Myopotta

Myopotta pallipes Myopotta

. . sp Myopa

Myopa vesiculosa Myopa

Myopa occulta Myopa

Myopa clausa Myopa

Myopa buccata Myopa

Paramyopa oestracea Paramyopa

Sicus ferrugineus Sicus

Sicus abdominalis Sicus

Carbonosicus carbonarius Carbonosicus

Robertsonomyiapearsoni

Robertsonomyiaparva

Robertsonomyiapalpalis

Robertsonomyiamexicana

sp. Parazodion

Parazodion schmidti Parazodion

112

101

114

113

111

109

106

102

100

69 1

1 are depicted as –). Taxon names are depicted in their previously accepted combinations. Character state transformations

1 1

4 82

50% and TBS

1 08

1 6

A, strict consensus of the 72 most-parsimonious cladograms (length – 218, CI – 0.550, RI – 0.957). Support values for each node are given (boot above;

2 69 TBS below; boot illustrated. Boxes depicted haveother character morphological number above apomorphiesclassiﬁcations and are and state given below. reversals. at Black the Character boxes top. represent states Continued morphological in for autapomorphies, Figure while 1B individual white on outgroup boxes next represent page. taxa B, are continued from not Figure 1A. depicted. Newly proposed subfamily and tribal A Figure 1.

Abrachyglossum capitatum B 93 4346 93 104 Leopoldius coronatus - 3 0 1 1 Leopoldius sigantus Conopini 103 1 1 53 Conops (Conops) flavipes Conops (Conops) vesicularis 1 1 86 83 103107 Archiconops insularis Archiconops pseudoerytrhocephalus 3 1 1 1 Conops (Asiconops) ater Conops (Asiconops) aureomaculatus Conops (Asiconops) australianus 10 Conops (Asiconops) chinensis 3 52 Conops (Asiconops) nubeculosus - 39 77 1 Asiconopini 1 - 1 Conops (Ceratoconops) ornatus 1 Conops (Conops) verus 1 1 1 65 - 11 - Conops (Diconops) geminatus Conops (Diconops) trichus 1 1 0 1 36 Conops (Sphenoconops) brunneosericeus 4 64 - 1 Conops (Sphenoconops) nobilis 0 Physoconops quadripunctatus 1 1 9 63 Conops (Smithiconops) guineensis Conops (Smithiconops) rondanii 21 1 1 65 - Euconops bellus incertae sedis 65 - 1 0 0 Mallachoconops atratulus 1 27 Brachyceraea brevicornis 1 60 38 Brachyceraeini 1 64 Neobrachyceraea elongata 0 1 Neobrachyceraea obscuripennis 1 Caenoconops bicolor 8 65 Caenoconops claripennis Caenoconopini 105 1 1 Caenoconops friedbergi - Physoconops rhodesiensis 94 1 1 62 Dacops abdominalis Dacops kaplanae 1 1 88 4 54 58 Physocephala bimarginipennis 3 Physocephala maculipes 11 68 1 0 1 51 Physocephala madagascariensis - 65 Physocephala marginata Physocephalini 1 1 1 1 1 Physocephala tibialis 2 Physocephala rufipes - 65 95 Pseudophysocephala constricta 1 1 1 Pseudophysocephala platycephala 1 Jelte neotropica Physoconops (Aconops) antennatus - 39 Physoconops (Aconops) costatus 1 Physoconops (Aureoconops) aureolus 1 Physoconops (Kroeberoconops) argentinus Physoconops (Kroeberoconops) hermanni - 2 Physoconops (Kroeberoconops) rufipennis 1 Physoconops (Pachyconops) brachyrhynchus 0 Physoconops (Pachyconops) bulbirostris Physoconops (Pachyconops) guianicus Tropidomyiini Physoconops (Physoconops) discalis - 3 Physoconops (Physoconops) fronto 1 Physoconops (Physoconops) obscuripennis 0 Physoconops (Shannonoconops) apicalis Physoconops notatifrons Tropidomyia alexanderi 96 20 23 24 Tropidomyia aureifacies 21 117 Tropidomyia bimaculata 52 3 1 1 0 Tropidomyia ornata 2 1 1 Tropidomyia sp. 84 Australoconops perbellum 61 Australoconops phaeomeros 1 Australoconops unicinctus 116 1 54 Microconops nigrithorax 25 67 Microconopini Microconops ornatus 1 1 79 Microconops similis 2 1 1 Microconops tasmaniensis 105 86 110 Physoconops microvalvus 55 1 Physoconops sepulchralis Siniconopini 1 1 1 Siniconops elegans Conops (Conops) nigripes Conopinae 7 Physoconops (Gyroconops) abbreviatus 63 Physoconops (Gyroconops) parvus Gyroconopini 1 1 Physoconops (Gyroconops) sylvosus Pleurocerinella albohalterata 5 14 15 27 3336 37 40 49 7881115 100 26 Pleurocerinella copelandi From Fig. 1A 62 Pleurocerinella srilankai Pleurocerinellini 11 1 1 1 1 1 1 1 1 0 0 1 1 43 65 53 1 1 Pleurocerinella tibialis Pleurocerinella sp. 2 0 1 Tammo rufa 70 31 - Atrichoparia curticornis 54 1 Atrichoparia sp. A 1 1 1 4370 Atrichoparia sp. B - Setosiconops epixanthus 2578 1 Setosiconops robustus 67 - 3 4 1 0 2 43 Tanyconops longicaudus 1 1 1 - 1 Heteroconops gracilis 70 92 1 1 Heteroconops antennatus - 1 1 0 2 Heteroconops sp. 7132 35 1 1 Smartiomyia arena Neoconopini 2 41 Smartiomyia obscura 1 1 65 Neoconops brevistylus 10 39 67 1 1 39 Neoconops longicornis - Camrasiconops ater 1 1 1 1 0 Pleurocerina brevis 57 6768 Pleurocerina longicornis 1 Pleurocerina luteiceps 0 1 Pleurocerina turneri Pleurocerina vespiformis 57 Callosiconops hirsutus 62 Callosiconops rufus 1 1 Callosiconops rugifrons

Figure 1. Continued

© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167, 43–81 REVISION OF CONOPID GENERA 57 recovered (boot – < 50; TBS – 1; MAA – 97:1; MA – No other species have been assigned to it since. Smith’s 14:1, 15:1, 88:0). Stylogastrinae is recovered (boot – (1989) Australasian catalogue includes the genus in 100; TBS – 13; MAA – 6:1, 18:1, 50:1, 85:1, 99:1; MA Conopinae, but does not include tribal classifications. – 19:0, 23:1, 37:1, 39:1, 40:1, 62:1, 64:0, 76:1, 82:0, Schneider (2010) could not locate the single identified 101:1, 113:1). specimen of this genus. Based on Schneider’s (2010) Relationships between newly defined subfamilies translation of Kröber’s original (1940a) description, of Conopidae are also determined. Conopinae + the following important character states can be Zodioninae is recovered (boot – 93; TBS – 7; MAA – determined: lateral ocelli and ocellar tubercle present; 44:1, 53:1, 87:1, 114:1; MA – 45:0, 76:1, 79:1). (Cono- central ocellus absent; frons with lateral grooves; pinae + Zodioninae) + Sicinae is recovered (boot – 79; first flagellomere elongate; arista stylate, retracted TBS – 2; MAA – 72:1; MA – 75:1). ((Conopinae + into first flagellomere; prementum shorter than head

Zodioninae) + Sicinae) + Myopinae is recovered (boot – width; wing completely hyaline; petiole of R4+5 + M 80; TBS – 4; MAA – 90:1, 91:1, 106:1, 111:1; MA present, elongate; male abdomen not petiolate. -109:1). Conopidae (excluding Notoconops and Sty- The genus Macroconops Kröber, 1927 was erected logastrinae) is recovered (boot – 65; TBS – 1; MAA – to accommodate the type species, M. helleri. Only 49:1, 71:1, 100:1, 102:1; MA – 12:1, 96:1). M. sinensis Ôuchi, 1942 has been assigned to it since. Some relationships within subfamilies are deter- Smith’s (1975) Oriental catalogue includes the genus mined. Conopini is recovered (boot – < 50; TBS – 1; in Conopinae, but does not include tribal classifica- MAA – 104:1). Asiconopini is recovered (boot – < 50; tions. Kröber’s (1927) original description states TBS – 1; MAA – 77:1; MA – 39:1). Brachyceraeini is that Macroconops is very similar to Siniconops macu- recovered (boot – 60; TBS – 1; MA – 27:0). Caeno- lifrons (Kröber, 1916b), but with a greatly enlarged conopini is recovered (boot – 65; TBS – 1; MAA – 8:1). arista. Kröber describes a small ventral genital plate, Physocephalini is recovered (boot – 88; TBS – 3; MA but from the illustration included, it is clear that the – 4:1, 54:0, 58:1). Tropidomyiini is recovered (boot – specimen is a male with a large abdominal sternite 5 < 50; TBS – 1; MA – 3:0). Microconopini is recovered and syntergosternite 7 + 8 produced into a point. (boot – 54; TBS – 1; MAA – 116:1). Siniconopini is The genus Microbrachyceraea Kröber, 1940b was recovered (boot – 55; TBS – 1; MAA – 86:1, 110:1). erected as a new genus for the type species, Micro- Gyroconopini is recovered (boot – 63; TBS – 1; MAA – conops pendleburyi. No other species have been 7:1). Pleurocerinellini is recovered (boot – 53; TBS – 2; assigned to it since. Smith’s (1975) Oriental catalogue MA – 43:0, 65:1). Neoconopini is recovered (boot – includes the genus in Conopinae, but does not include < 50; TBS – 1; MA – 10:1, 39:1, 67:1). Myopini is tribal classifications. Based on Brunetti (1927) and recovered (boot – 59; TBS – 1; MAA – 16:1). Theco- Kröber’s (1940b) original descriptions, the following phorini is recovered (boot – 59; TBS – 1; MAA – 74:1). important character states can be determined: facial fovea present; facial carina present; scape quadrate; pedicel narrow, elongate, and without transverse SYSTEMATICS dorsal ridge; first flagellomere slightly longer than pedicel; arista stylate, apical, not retracted into first CHARACTER STATES OF GENERA FOR WHICH PINNED flagellomere; vein CuA2 curved; vein CuA2 + A1 short; SPECIMENS ARE NOT AVAILABLE male abdomen petiolate. The genus Anticonops Kröber, 1936 was erected to The genus Neobrachyglossum Kröber, 1915e was accommodate the type species, A. abdominalis.No erected to accommodate the type species, N. punc- other species have been assigned to it since. It is tatum. No other species have been assigned to it included in the tribe Conopini in Smith’s (1980) Afro- since. Chvála & Smith’s (1988) Palaearctic catalogue tropical catalogue. Based on Kröber’s (1936) original includes it in Conopini. The sole specimen was depos- description and Camras’ (2000) redescription, the ited at the Hungarian Natural History Museum following important character states can be deter- (HNHM) and was probably destroyed during the mined: scape and pedicel elongate; first flagellomere Soviet attack and subsequent fire that destroyed the equal in length to scape and pedicel combined; collection in 1956 (Skevington & Marshall, 1998). arista apical, not retracted into first flagellomere; Based on Kröber’s (1915e) original description, the prementum twice as long as width of head; wing following important character states can be deter- not completely hyaline; female abdominal sternite 5 mined: prementum short, fleshy; first flagellomere with spicules, but ventral genital plate not present; as long as scape and pedicel combined; arista apical, female abdominal sternite 6 with spicules; female stylate, not retracted into first flagellomere; wing abdominal tergite 7 elongate. similar to Conops; abdomen slightly narrowed. All of The genus Delkeskampomyia Kröber, 1940a was these character states are present in all species of erected to accommodate the type species, D. fasciata. Leopoldius.

The genus Neozodion Szilady, 1926 was erected Asiconops, Physoconops quadripunctatus comb. nov., to accommodate the type species, N. pruinosum.No originally described by Kröber (1915a) in Conops and other species have been assigned to it since. Papave- transferred to Physoconops by Camras (2000), is now ro’s (1971) Neotropical catalogue includes it in Myopi- transferred to Asiconops as the type species of a new nae. The sole specimen was deposited at the HNHM subgenus, Aegloconops subgen. nov. Conops (Conops) and was probably destroyed during the Soviet attack verus is transferred to Asiconops (Asiconops) comb. and subsequent fire that destroyed the collection in nov. Conops (Conops) nigripes is transferred to Sini- 1956 (Skevington & Marshall, 1998). Szilady (1926) conops comb. nov. describes the single, female specimen as very similar Physoconops sepulchralis and Physoconops micro- to Zodion, but with abdominal segments 2–4 closely valvus are both transferred to Siniconops comb. nov. appressed and a ventral extension on abdominal ster- Jelte is transferred to a subgenus of Physoconops nite 5. These character states are present in many comb. nov. Gyroconops is removed from Physoconops species of Zodion, and no other characters are offered and elevated to genus status, stat. nov. Physoconops to distinguish Neozodion from Zodion. rhodesiensis is transferred to Caenoconops comb. rev. The genus Stenoconops Kröber, 1939b was erected A new genus, Schedophysoconops gen. nov., is erected to accommodate the type species, S. niger. No other with Physoconops notatifrons as its type. species have been assigned to it since. Smith’s Neobrachyglossum is designated a junior synonym (1989) Australasian catalogue includes the genus in of Leopoldius syn. nov. Neozodion is designated a Conopinae, but does not include tribal classifications. junior synonym of Zodion syn. nov. Schneider (2010) studied the only known specimen, the female holotype, and concluded that Stenoco- ASICONOPS (AEGLOCONOPS) GIBSON nops is a valid genus. Based on Schneider’s (2010) re-description, the following important character SUBGEN. NOV. (FIG.2) states can be determined: lateral ocelli and ocellar Type species: Conops quadripunctatus Kröber, 1915d: tubercle present; central ocellus absent [although 38, present designation. Kröber (1939b) describes three ocelli as present]; frons with lateral grooves; cephalic bristles absent; Diagnosis: Aegloconops can be distinguished from facial ridge not meeting subcranial cavity at 90° other members of Asiconops (s.l.) by the combination angle; first flagellomere not elongate; arista stylate, of narrow frons; fronto-facial spots; reduced facial not retracted into first flagellomere; maxillary palpus carina; short, wide, arista; short prementum; and absent; prementum elongate; apical shiny patches narrow second abdominal segment in the male. on metafemur present; scutellum developed, with two pairs of bristles; wing completely hyaline; petiole Description: Head. Occipital setae sparse, short, dark. of R4+5 + M present, length not stated; vein R1 ending Vertex round, not distinct. Frons slightly wider than near end of R2+3; angle between vein CuA1 and cross- long; no setae or bristles; smooth, concave, elevated at vein dm-cu acute; petiole of CuA2 + A1 short; vena anterior margin. Fronto-facial spot present. Ocellar spuria present; ventral genital plate present. tubercle and ocelli absent. Ocellar and postocellar bristles absent. Compound eye oval with triangular notch in posterior margin. Facial foveae deep, blends TAXONOMIC REVISION OF SOME SPECIES into gena; medial carina very small. Gena present, OF CONOPIDAE small, less than one-tenth of head height; anterior The following taxonomic revisions are proposed for margin of subcranial cavity pointed anteriorly at junc- some individual species and genera of Conopidae tion with medial carina. Lunule well developed, united based on phylogenetic analysis and the hypothesized with facial ridge. Scape elongate, length four times cladogram (Fig. 1). The proposed new status of these width; pedicel twice as long as scape; pedicel elongate, species and genera reflects morphological character length five times width; first flagellomere short, half states as recorded as part of this research. Complete length of scape and pedicel combined; first flagellomere descriptions for proposed tribal groups, including bare; arista short, wide; second aristomere expanded these taxa, are included in the complete classification ventrally. Maxillary palpi absent; prementum elon- that follows. All taxonomic revisions, as well as revi- gate, 1.25 times as long as width of head; labella short, sionary history of all genera, are summarized in the length one-tenth of length of prementum; labella catalogue of genus-group names (Appendix). ovate, pointed, without setae. Asiconops and Smithiconops are removed as subgenera of Conops and elevated to genus status, Cera- Thorax. Basisternum square, posterolateral exten- toconops stat. nov., Diconops, and Sphenoconops are sions pointed; proepimeron, anepimeron, and ane- transferred from subgenera of Conops to subgenera of pisternum bare; two proepisternal setae; small patch

A B

compound eye notch

2nd aristomere

apical shiny patch

2mm

Figure 2. Conopinae – Asiconopini – Asiconops (Aegloconops) quadripunctatus ǩ (Kröber, 1915d). A, lateral habitus; B, head. of bristles in posterodorsal portion of katepisternum; Measurements. Total length (excluding antennae) = dorsum of thorax without prominent bristles; scutel- 10 mm; wing length = 6 mm. lum reduced, without bristles. Shiny patches at apex of all tibia; prominent row of setae on posterior Geographical distribution: Afrotropical: Democratic surface of mesofemur. Republic of the Congo, South Africa, Zaire, Zimbabwe.

Wing. Dark pattern at apex of wing, anterior to vein Etymology: The name is constructed from the Greek R4+5, otherwise hyaline; petiole of R4+5 + M half as long term aeglo-, meaning brilliant or radiant, and Conops. as crossvein dm-cu; vein Sc ends at midpoint of costa; The name is masculine. vein R1 ends at a point seven-tenths length of the costa; vein R2+3 ends only slightly beyond the end of R1; SCHEDOPHYSOCONOPS GIBSON GEN. NOV. (FIG.3) crossvein sc-r present; petiole of CuA2++A1 half as long as crossvein dm-cu; vein CuA2 curved along its length; Type species: Physoconops notatifrons Camras, 1962b: vena spuria present; alula half as wide as wing. 219, present designation.

Female abdomen. Sternites 1–4 present, but narrow; Diagnosis: Similar to Physoconops (s.l.), but with tergite 5 and sternite 5 partially fused; ventral genital a fronto-facial spot, two ocelli, and a shorter plate half as long as height of tergite 6, rounded, prementum. broader than tall, covered with irregular rows of spicules; tergite 6 and sternite 6 fused anteriorly; Description: Head. Occipital setae sparse, short, sternite 6 with rows of spicules; tergite 7 and sternite dark. Vertex round, not distinct. Frons slightly wider 7 fused. than long; no setae or bristles; smooth with small rugose area at anterior margin. Fronto-facial spot Male abdomen. Tergite 2 equal in length to, but present. Ocellar tubercle round, non-distinct; two distinctly narrower than, tergites 1 and 3; sternite 2 ocelli present. Ocellar and postocellar bristles absent. and 3 present, narrow; spicules present on posterior Compound eye oval with triangular notch in posterior margin of sternite 5. Syntergosternite 7 + 8 round, margin. Facial foveae deep, blends into gena; medial hemispherical; sternite 8 broad, bare, shiny. Cerci carina very small. Gena present, small, less than large, rounded, and connected to the epandrium by a one-quarter of head height; anterior margin of sub- sclerotized stalk, densely setose. Epandrium strongly cranial cavity pointed anteriorly at junction with fused beyond cerci, quadrate, bare except for setae on medial carina. Lunule well developed, united with posterior margin, very small posterior hump where facial ridge. Scape elongate, length four times width; cerci attach. pedicel 1.5 times as long as scape; pedicel elongate,

A B

2mm

Figure 3. Conopinae – Tropidomyiini – Schedophysoconops notatifrons Ǩ (Camras, 1962b). A, lateral habitus; B, head; C, wing.

length four times width; ﬁrst ﬂagellomere short, less ends only slightly beyond end of R1; crossvein sc-r than half length of the scape and pedicel combined; present; petiole of CuA2++A1 half as long as crossvein

ﬁrst ﬂagellomere bare; arista short, wide; second aris- dm-cu; vein CuA2 curved along its length; vena spuria tomere greatly expanded ventrally, nearly reaching present; alula half as wide as wing. apex of third aristomere. Maxillary palpi absent; prementum elongate, 1.33 times as long as width of Female abdomen. Sternites 1–4 present, but narrow; head; labella short, length one-tenth length of pre- tergite 5 and sternite 5 partially fused; ventral mentum; labella ovate, pointed, without setae. genital plate two-thirds as long as height of tergite 6; rounded, broader than tall, covered with irregular Thorax. Basisternum square, posterolateral exten- rows of spicules; tergite 6 and sternite 6 fused ante- sions pointed; proepimeron, anepimeron, and ane- riorly; sternite 6 with rows of spicules; margin pisternum bare; one proepisternal seta; small patch of between tergite 6 and tergite 7 straight; tergite 7 and bristles in posterodorsal portion of katepisternum; sternite 7 fused; row of long setae on sternite 7; cerci dorsum of thorax without prominent bristles; scutel- broad, cupped, square, with long setae, attached nar- lum reduced, without bristles. Shiny patches at rowly; sternite 8 bilobed, square posteriorly, with long apex of all tibia; prominent row of setae on posterior setae; syntergite 8 + 9 with long, broad, posterior scle- surface of mesofemur. rotized hooks.

Wing. Dark pattern anterior to vein CuA2, hyaline Male abdomen. Males are not known for this genus. posterior; petiole of R4+5 + M half as long as crossvein dm-cu; vein Sc ends at point halfway along costa; vein Measurements. Total length (excluding antennae) =

R1 ends at point 0.7 of the way along costa; vein R2+3 9–11 mm; wing length = 5–7 mm.

Geographical distribution: Afrotropical: Cameroon, all members of Pleurocerinellini. The labella are fili- Kenya, Nigeria, Uganda, Zambia. form and fused (45) in Conopidae, but have been reversed to a broad and separate state in Conopinae Etymology: The name is constructed from the Greek and Zodioninae. There is a prominent row of setae on term schedo-, meaning near or almost, and Physo- the posterior surface of the mesofemur in Conopidae conops. The name is masculine. (54); this has been lost in Dacops + Physocephala + Pseudophysocephala. All members of Conopidae [except some Stylogaster – (S. biannulta + S. stylata) + DISCUSSION (S. frauci + S. pauliani + S. westwoodi + S. sp.)] are PROPOSED CLASSIFICATION OF CONOPIDAE lacking bristles on the postpronotum (60). The petiole Based on the present analysis, the following classifi- of vein CuA2 + A1 is longer than crossvein dm-cu (78) cation is proposed. Family, subfamily, and tribal in all members of Conopidae; this character has been ranks are designated based on an apomorphy-based reversed independently in three lineages: Callosi- phylogenetic concept. A list of apomorphic character conops, Pleurocerina + Camrasiconops, and Conopi- states defining each group is included. Previous nae excluding Neoconopini. Female abdominal tergite authors’ interpretations of character states are pro- and sternite 6 are at least partially fused in all vided using present terminology (sensu Cumming & members of Conopidae (88); this fusion has been Wood, 2009; Gibson et al., 2012). Most previous cata- lost in Dalmanniinae. Gibson et al. (2012) note the logues of Conopidae utilized tribal classifications complete fusion of tergite 6 and sternite 6 into a without diagnosis of tribal-group characters. These ring-like structure in all members of Stylogaster.In previous classifications are summarized in Table 1. the present analysis, the partial fusion of tergite 6 Gibson et al. (2012) examined phylogenetic rela- and sternite 6 is recorded instead. The male epiproct tionships within Conopidae using DNA sequence data is absent in all members of Conopidae (98); this state and many of the same morphological characters is also observed in Pyrgotidae. presented here, albeit with a smaller taxon set. That Gibson et al. (2012) recover Conopidae as mono- research recovered the same monophyletic clades phyletic with the same set of apomorphic character with the same morphological apomophies listed here. states as recovered in the present analysis. Latreille Also in Gibson et al. (2012), the past character inter- (1802) initially described Conopsariae, including pretations and proposed apomorphic character states Conops, Zodion, Myopa, and Stomoxys (now in Mus- for Conopidae and its component subfamilies are cidae), with the following characters: prementum reviewed. To avoid repetition of that work, only dis- elongate, cylindrical; labellum sometimes elongate, cussion of novel character states and tribal and sub- filiform, folded ventrally; maxillary palpi, when tribal relationships are included here. A list of species present, elongate; antennae sometimes longer than examined in the present analysis, either with speci- head; arista either apical or dorsal. According to the mens or through literature review, is included for present analysis, the cylindrical prementum is apo- each genus-group. Proposed taxonomic changes are morphic to Conopidae and the remaining characters also included. describe various states across conopid subfamilies. Schneider (2010) included Notoconops within Conopidae, as the type genus of the subfamily Noto- CONOPIDAE LATREILLE, 1802 conopinae, based on: the absence of a dorsal notch

Type genus Conops Linnaeus, 1758: 604. Included on the pedicel; a bare anepisternum; vein R1 bare; the extant subfamilies: Conopinae, Dalmanniinae, Myopi- absence of vibrissae; absence of mid coxal prongs; nae, Sicinae, Stylogastrinae, Zodioninae. Uncontro- costa lacking breaks; veins Sc and R1 separate; cell verted morphological autapomorphies: prementum cup acute; vein A1 complete. In the present analysis, fused into a tube (character 42); posterolateral none of these character states is found to be apomor- extensions of the basisternum present (48); and M phic to Conopidae. All mouthparts are completely vein deﬂected to meet or end near vein R4+5 (69). The absent in Notoconops. Although Schneider (2010) pedicel is elongate in all members of Conopidae (28), noted the presence of posterolateral extensions on the but this character state is also observed in Pyrgoti- basisternum in her description of Notoconops, they dae. The arista is thickened in all members of Conopi- were not present on the specimen available for the dae (34), but has been secondarily reversed to the present research. Also, the eyes and frons of Noto- ﬁliform state in Melanosoma. The prementum is conops are sexually dimorphic, a condition not longer than the width of the head in all members of observed in any members of Conopidae. Furthermore,

Conopidae (43), but has been secondarily shortened in Notoconops lacks all but one (78 – vein CuA2 + A1 Myopa occulta, Abrachyglossum, Baruerizodion, Het- elongate) of the apomorphic characters of Conopidae. eroconops, Leopoldius, Pseudomyopa, Tanyconops and For these reasons, Notoconops cannot be included

© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167, 43–81 62 J. F. GIBSON AND J. H. SKEVINGTON in Conopidae. Its proper family classiﬁcation is Hendel (1936) suggests that the absence of ocelli is unknown and it should be left as incertae sedis within characteristic of all Conopinae, but Hennig (1966) Schizophora pending further evidence of its proper notes that this is not the case. The presence of three placement. ocelli on a well-developed ocellar tubercle is the plesiomorphic state within Conopidae. A reduction of the ocellar tubercle is apomorphic to Conopinae, but CONOPINAE LATREILLE, 1802 the loss of the central and lateral ocelli and the Type genus Conops Linnaeus, 1758: 604. Included ocellar tubercle has occurred more than once within tribes: Asiconopini trib. nov., Brachyceraeini, Caeno- Conopinae (2, 3, 4). In a clade including all members conopini trib. nov., Conopini, Gyroconopini trib. nov., of Conopinae (except Neoconopini, Gyroconopini, Microconopini trib. nov., Neoconopini trib. nov., and Pleurocerinellini), the central ocellus is absent. Physocephalini, Pleurocerinellini, Siniconopini trib. In a clade including the Conopini + Asiconopini + nov., Tropidomyiini. Genera incertae sedis: Euconops Euconops + Mallachoconops + Brachyceraeini + Caeno- and Mallachoconops. Uncontroverted morphological conopini + Physocephalini + Tropidomyiini the lateral autapomorphies: apical, stylate arista (33); and epan- ocelli have been lost as well. Some reversal events drium fused beyond cerci (115). The vertex is com- have also occurred with regard to ocelli. All members plete and undivided in Conopinae (5), but this of Tropidomyiini have regained the lateral ocelli. character state is also observed in Pyrgotidae. Ocellar Within Tropidomyiini, all members of Physoconops bristles are absent in all members of Conopinae (s.l.) have regained the central ocellus as well, (14); this character state is also observed in Pyrgoti- although it is often very small. Parallel to these dae, Syrphidae, Dalmanniinae, and some species character changes, the central ocellus has been of Stylogaster (S. biannulata + S. stylata + S. frauci + lost in members of Heteroconops + Setosiconops + S. pauliani + S. westwoodi + S. sp.). Postocellar bris- Tanyconops, and all three ocelli have been lost tles are absent in all members of Conopinae (15); this in Heteroconops gracilis + Setosiconops + Tanyconops. character state is also observed in Pyrgotidae, Syr- This clade has also lost the ocellar tubercle com- phidae, and Dalmanniinae. The scape is elongate in pletely. Delkeskampomyia and Stenoconops are each all members of Conopinae (27); this character state described as having two ocelli, placing them within has been reversed in Brachyceraeini and is also the Heteroconops + Setosiconops + Tanyconops clade. observed in Pseudoconops. The second aristomere is A parallel loss of the ocellar tubercle has occurred expanded ventrally in all members of Conopinae and in Physocephalini and Asiconopini + Euconops + has been reversed in Asiconops (Sphenoconops) (36). Mallachoconops. The anterior margin of the subcranial cavity is A well-developed lunule (21) and the presence of a projects forward to a narrow point (37) and the max- posterior hump on the epandrium (117) are autapo- illary palpi are reduced or absent (40) in all members morphies of Conopinae excluding Neoconopini. The of Conopinae; these character states are also observed lunule has been secondarily reduced in Euconops + in Stylogastrinae. The posterolateral extensions of Mallachoconops. the basisternum have been reversed to blunt points in In addition to being an apomorphy of Stylogastri- all members of Conopinae (49); a parallel reversal is nae, the loss of the maxillary palpi (39) has occurred observed in Sicinae. The petiole of vein CuA2 + A1 is three times within Conopinae. This character state shorter than crossvein dm-cu (78) in all members of is an apomorphy of Asiconopini, Physocephalini + Conopinae (except Atrichoparia, Heteroconops, Neo- Tropidomyiini, and Neoconopini. The maxillary conops, Setosiconops, Smartiomyia, and Tanyconops). palpi have been regained in Camrasiconops. The A vena spuria is present in all members of Conopinae state of the maxillary palpi is unknown for (81), although sometimes only as a fold in the wing Delkeskampomyia. membrane; this character state is also observed in In addition to being an apomorphy of Pleuroceri- Pyrgotidae and Syrphidae. nellini, the reduction of the scutellum (65) is also Gibson et al. (2012) recover Conopinae as mono- an apomorphy of Asiconopini + Euconops + Mallacho- phyletic with the same set of apomorphic charac- conops + Brachyceraeini + Caenoconopini + Physoce- ter states, but only include representatives of phalini + Tropidomyiini. This character has been Asiconopini, Conopini, Microconopini, Physocephalini, reversed in Euconops, in which the scutellum is greatly Neoconopini, Tropidomyiini, and Euconops. They enlarged with many bristles. include the following additional apomorphic character The petiole of CuA2 + A1 (78) has been secondarily states: heavily sclerotized dorsal bridge of hypan- shortened in Conopinae. This character state has drium forming nearly complete tube around phallus; been reversed to the long state in Atrichoparia + single anterior arm on the hypandrium (absent in Neoconops + Smartiomyia + Heteroconops + Setosico- Atrichoparia sp. B); and absence of posterior surstyli. nops + Tanyconops.

The distinctly petiolate shape to the male abdomen Smithiconops guineensis comb. nov. (Conops); Smith- (105) is an apomorphy of Brachyceraeini + Caeno- iconops rondanii comb. nov. (Conops). conopini + Physocephalini + Tropidomyiini that has arisen independently in Siniconops microvalvus. Other characteristic states are observed in BRACHYCERAEINI ZIMINA, 1960 members of Conopinae but are not included in the Type genus Brachyceraea Röder, 1892: 366. Genera present analysis. A prominent fronto-facial spot is included: Brachyceraea, Microbrachyceraea Kröber, present in many Afrotropical species of various tribes. 1940b: 217, Neobrachyceraea Szilady, 1926: 587. A greatly elongate ventral genital plate is observed in Uncontroverted morphological autapomorphy: scape many species, but the size and shape of the ventral quadrate (27). The single species of Microbrachycer- genital plate is highly variable within genera. Both aea possesses a quadrate scape and petiolate male of these characters prove to be too homoplasious to abdomen, the two key characteristics of Brachycer- contribute to higher-level classiﬁcation. aeini, thereby warranting its inclusion. The presently proposed membership of Brachycer- aeini matches the previously proposed deﬁnitions for ASICONOPINI GIBSON TRIB. NOV. the group. Zimina’s (1960) original description of Type genus Asiconops Chen, 1939: 171. Genera Brachyceraeini includes the following diagnostic included: Anticonops Kröber, 1936: 285, Archiconops characters: antennae shortened; basisternum broad

Smith 1975: 375, Asiconops, Smithiconops Camras, with very small posterolateral extensions; vein R2+3

2000: 221. Uncontroverted morphological autapomor- parallel to R1 and meeting the costa midway between phy: angle between vein CuA1 and crossvein dm-cu the end of R1 and R4+5. Hennig (1966) also suggests obtuse (77). Setae on the maxillary palpi are com- that the reduced antenna is probably a shared, pletely absent in Asiconopini (39); this character state derived apomorphy of the Brachyceraea. The present is also observed in Physocephalini + Tropidomyiini analysis confirms that the characteristic antennal and most Neoconopini. shape is due to a reduction in the scape. The other The genus Anticonops is described based on one characters included by Zimina are plesiomorphic to female specimen from the Congo. Kröber’s (1936) Conopinae. original description suggests that it is similar to Species examined – Brachyceraea brevicornis; Conops, but does not specify a species or subgenus. Microbrachyceraea pendleburyi Neobrachyceraea The extremely long prementum suggests a similarity elongata; Neobrachyceraea obscuripennis. to Asiconops (s.l.), but with a distinct condition of the female terminalia. All of the genera currently placed in Asiconopini CAENOCONOPINI GIBSON TRIB. NOV. were previously placed in Conopini, with two of the Type genus Caenoconops Anonymous in Imperial genera being subgenera of Conops. The absence Institute of Entomology, 1940: 365. Genus included: of an ocellar tubercle, maxillary palpi, scutellar bris- Caenoconops. Uncontroverted morphological autapo- tles, and broad male abdominal tergites 3 and 4, as morphy: vertex triangular in shape with longitudinal well as the presence of a characteristic acute angle grooves (8). between vein CuA1 and crossvein dm-cu, separate Previous catalogues included Caenoconops in members of Asiconopini from Conopini as presently Conopini. The condition of the vertex, the absence of defined. scutellar bristles, the petiolate male abdomen, the Species examined – Anticonops abdominalis; notched posterior margin of the eye, and the presence Archiconops insularis; Archiconops pseudoerythro- of a characteristic dark anterior/hyaline posterior cephalus; Asiconops (Aegloconops) quadripunctatus wing pattern separate members of Caenoconopini comb. nov. (Conops); Asiconops (Asiconops) ater comb. from Conopini as presently defined. nov. (Conops); Asiconops (Asiconops) aureomaculatus Species examined – Caenoconops bicolor; Caeno- comb. nov. (Conops); Asiconops (Asiconops) austral- conops claripennis; Caenoconops friedbergi; Caeno- ianus comb. nov. (Conops); Asiconops (Asiconops) conops rhodesiensis comb. rev. (Conops). chinensis comb. nov. (Conops); Asiconops (Asiconops) nubeculosus comb. nov. (Conops); Asiconops (Asi- conops) verus comb. nov. (Conops); Asiconops (Cerato- CONOPINI LATREILLE, 1802 conops) ornatus comb. nov. (Conops); Asiconops Type genus Conops Linnaeus, 1758: 604. Genera (Diconops) geminatus comb. nov. (Conops); Asiconops included: Abrachyglossum Kröber, 1919: 142, Conops, (Diconops) trichus comb. nov. (Conops); Asiconops Leopoldius Rondani, 1843: 35. Uncontroverted mor- (Sphenoconops) brunneosericeus comb. nov. (Conops); phological autapomorphy: male abdominal tergites 3 Asiconops (Sphenoconops) nobilis comb. nov. (Conops); and 4 broad and parallel-sided (104).

Previous classifications variously included Species examined – Australoconops perbellum; Aus- members of Asiconopini, Caenoconopini, Physocepha- traloconops phaeomeros; Australoconops unicinctus; lini, Siniconopini, and Tropidomyiini within Conopini. Microconops nigrithorax; Microconops ornatus; Micro- See diagnoses of each of those respective tribes for conops similis; Microconops tasmaniensis. means to distinguish them from Conopini. Zimina’s (1960) definition of Conopini includes the following key characteristics: facial foveae present; arista NEOCONOPINI GIBSON TRIB. NOV. three-segmented; vein R1 ending near the end of R2+3; Type genus Neoconops Kröber, 1915e: 75. Included basisternum quadrangular. In the present analysis, genera: Atrichoparia Schneider, 2010: 33, Callosi- each of these character states is plesiomorphic to conops Kröber, 1940a: 75, Camrasiconops Schneider, Conopinae (s.l.). Smith & Peterson (1987) include 2010: 78, Delkeskampomyia Kröber, 1940a: 71, no key characteristics of Conopini in their key to Heteroconops Kröber, 1915e: 80, Neoconops Kröber, Nearctic genera, except that the key characteristics of 1915e: 75, Pleurocerina, Setosiconops Schneider, Physocephala are absent. 2010: 157, Smartiomyia Kröber, 1940a: 72, Steno- Species examined – Abrachyglossum capitatum; conops Kröber, 1939b: 606, Tanyconops Schneider, Conops flavipes; Conops vesicularis; Leopoldius coro- 2010: 170. Uncontroverted morphological apomor- natus; Leopoldius punctatum comb. nov. (Neobrachy- phies: none. The frons has extensive lateral grooves glossum); Leopoldius signatus. (10) in all members of Neoconopini; there has been a parallel development of this character state in Asi- conops (sensu stricto, s.s.) + Ceratoconops + Diconops. GYROCONOPINI GIBSON TRIB. NOV. Setae on the maxillary palpi are absent (39) in all members of Neoconopini, except Camrasiconops; this Type genus Gyroconops Camras, 1955b: 174. Included character state is also observed in Asiconopini and genus: Gyroconops. Uncontroverted morphological Physocephalini + Tropidomyiini. The wings are com- autapomorphy: vertex large, square, and setose, with pletely hyaline (67) in all members of Neoconopini ocelli displaced forward (7). except Pleurocerina; this character state has arisen Previously placed as a subgenus of Physoconops, independently in Phoridae, Syrphidae, Strongyloph- members of Gyroconops have been variously placed thalmyiidae, and Microconops. Although not available within Conopini and Physocephalini. As presently for examination, the original, or subsequent, descrip- defined, members of Gyroconopini can be distin- tions of Delkeskampomyia and Stenoconops both guished from Physoconops by the characteristic shape include mention of these apomorphic character states. of the vertex and frons, the developed scutellum, the Their inclusion in Neoconopini is thus warranted. non-petiolate male abdomen, the presence of maxil- Gibson et al. (2012) also recover a monophyletic lary palpi, the absence of a notch in the posterior Neoconopini using molecular and morphological margin of the eye, and the absence of a characteristic analysis and including representatives of Atricho- dark anterior/hyaline posterior wing pattern. paria, Camrasiconops, Pleurocerina, Heteroconops, Species examined – Gyroconops abbreviatus and Smartiomyia. Members of Neoconopini have comb. nov. (Physoconops); Gyroconops parvus comb. not been placed within any tribe in previous nov. (Physoconops); Gyroconops sylvosus comb. nov. classifications. (Physoconops). The facial ridge meeting the subcranial cavity at a 90° angle (25) is an apomorphy of Atrichoparia + Hete- roconops + Neoconops + Setosiconops + Smartiomyia + MICROCONOPINI GIBSON TRIB. NOV. Tanyconops. This character cannot be assessed in Type genus Microconops Kröber, 1915e: 77. Included Delkeskampomyia. The same character state has genera: Australoconops Camras 1961: 64, Micro- arisen independently in Microconops. conops. Uncontroverted morphological autapomorphy: An extremely elongate first flagellomere (32) is an presence of a medial spine on the epandrium (116). apomorphy of Stylogaster inca + S. neglecta, and is Gibson et al. (2012) also recover a monophyletic observed in Lauxaniidae and Phoridae. This charac- Microconopini using molecular and morphological ter state is also an apomorphy of Heteroconops + analysis of species of both Australoconops and Micro- Neoconops + Setosiconops + Smartiomyia + Tanyconops. conops. Members of Microconopini have not been This latter clade also shares a stylate arista that placed within any tribe in previous classifications. is reduced and retracted into the first flagellomere Schneider, 2010 lists Microconops ornatus Kröber, (35) and a reduction of the spicules on the ventral 1915e as the type species for Microconops, as desig- genital plate to a few apical rows (92). The status of nated by Smith, 1989. This is incorrect as a type was these characters is unknown in Stenoconops and previously designated by Malloch. Delkeskampomyia.

While both species of Setosiconops and the one included members of the present Pleurocerinellini species of Tanyconops are recovered in a monophyletic and Tropidomyiini in past catalogues. In addition to clade with members of Heteroconops, the possible the apomorphies of the tribe, Physocephalini can be synonymy is considered here as unconfirmed. Some distinguished from Conopini by the absence of max- characters of Heteroconops, especially regarding illary palpi, the absence of scutellar bristles, the peti- number of ocelli, are highly variable. Revision of all olate male abdomen, the notched posterior margin species of Heteroconops will be necessary to confirm of the eye, and a characteristic wing pattern that is a possible taxonomic revision for Setosiconops and dark anteriorly and hyaline posteriorly. Differences Tanyconops. between Physocephalini, Pleurocerinellini, and Tropi- Species examined – Atrichoparia curticornis; Atri- domyiini are discussed below. choparia sp. A; Atrichoparia sp. B; Camrasiconops Species examined – Dacops abdominalis; Dacops ater; Callosiconops hirsutus comb. rev. (Chrysidi- kaplanae; Physocephala bimarginipennis; Physo- omyia); Callosiconops rufus comb. nov. (Chrysidiomyia cephala maculipes; Physocephala madagascarien- rufa); Callosiconops rugifrons comb. nov. (Chrysidi- sis; Physocephala marginata; Physocephala rufipes; omyia); Delkeskampomyia fasciata; Heteroconops Physocephala tibialis; Pseudophysocephala constricta; antennatus; Heteroconops gracilis; Heteroconops sp.; Pseudophysocephala platycephala. Neoconops brevistylus; Neoconops longicornis; Pleu- rocerina brevis; Pleurocerina longicornis; Pleurocerina luteiceps; Pleurocerina turneri; Pleurocerina vespi- PLEUROCERINELLINI ZIMINA, 1974 formis; Setosiconops robustus; Smartiomyia arena; Type genus Pleurocerinella Brunetti, 1923: 368. Smartiomyia obscura; Stenoconops niger; Tanyconops Included genera: Pleurocerinella, Tammo Stuke, 2008: longicaudus. 50. Uncontroverted morphological autapomorphies: none. In all members of Pleurocerinellini, the prementum is reduced from the elongate state plesiomorphic PHYSOCEPHALINI SMITH &PETERSON, 1987 to Conopidae (43); the reduced state is also observed in Type genus Physocephala Schiner, 1861: 137. Included Abrachyglossum + Leopoldius, Baruerizodion, Hetero- genera: Dacops Speiser, 1923: 99, Physocephala, Pseu- conops, Myopa occulta, Pseudomyopa, and Tanyconops. dophysocephala Kröber, 1939a: 374. Uncontroverted The scutellum is greatly reduced and lacking bristles morphological autapomorphies: prominent row of (65) in all members of Pleurocerinellini; this character setae on mesofemur absent (54). The ocellar tubercle is state is also observed in Asiconopini + Euconops + absent (4); this character state is also observed Mallachoconops + Brachyceraeini + Caenoconopini + in Asiconopini + Euconops + Mallachoconops and Physocephalini + Tropidomyiini. Heteroconops gracilis + Tanyconops + Setosiconops. Zimina (1974) first described this tribe and The ventral half of the proepisternum is bare in all included only the Palaearctic species, Pleurocerinella members of Physocephalini (58); this character state is tibialis. The original tribal description, translated by also observed in Baruerizodion and Scatoccemyia. Clements & Vincent (2001), includes the following Gibson et al. (2012) also recover a monophyletic diagnostic characters: pedicel elongate; three ocelli Physocephalini using molecular and morpholo- present; prementum short; labella ‘massive’; cross- gical analysis and including representatives of vein sc-r present; vein R4+5 + M present; pleura and only Physocephala. A characteristic shape of the coxae bare. While all of these character states are metafemur is listed as an autapomorphy of the observed in the present analysis, only the shortened clade, but this character is recovered in the present prementum is apomorphic to the tribe. Zimina analysis as an autapomorphy of Physocephala + (1974) suggests that Pleurocerinellini is intermediate Pseudophysocephala. between Conopinae and Myopinae, as it has the mor- The original usage of this tribal classification by phological characters of Conopinae, except ocelli are Camras (1965) was in a Nearctic catalogue without present. In the present analysis, the absence of ocelli diagnosis. Sabrosky (1999) notes that this makes the is not apomorphic to Conopinae. Members of Pleu- name unavailable. Smith & Peterson (1987) include rocerinellini display all character states apomorphic Physocephalini in their key to Nearctic genera and to Conopinae. this action makes them the authors of the name. They Previous classifications have included Pleuro- include only Physocephala in the tribe and list the cerinella within Physocephalini. Members of Pleu- following as key characteristics: crossvein r-m well rocerinellini can be distinguished from members beyond the middle of cell dm; prosepisternum bare; of Physocephalini by the presence of ocelli and an metafemur thickened in basal half; ocelli absent. ocellar tubercle, a shortened prementum, a non- While Physocephala has been included in Conopini petiolate male abdomen, the absence of a notch in in some classifications, the tribe Physocephalini has the posterior margin of the eye, the absence of a

© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167, 43–81 66 J. F. GIBSON AND J. H. SKEVINGTON characteristic dark anterior and hyaline posterior While usually placed within its own tribe, Tropid- wing pattern, the presence of maxillary palpi, the omyia has been placed in Physocephalini by some presence of a prominent row of setae on the mesofe- authors. Members of Tropidomyiini can be distin- mur, and the presence of setae on the ventral half of guished from Physocephalini by the presence of the proepisternum. lateral ocelli, an ocellar tubercle, a prominent row of Species examined – Pleurocerinella albohalterata; setae on the mesofemur, and setae or bristles on the Pleurocerinella copelandi; Pleurocerinella srilankai; ventral half of the proepisternum. Pleurocerinella tibialis; Pleurocerinella sp.; Tammo Species examined – Physoconops (Aconops) antenna- rufa. tus; Physoconops (Aconops) costatus; Physoconops (Aureoconops) aureolus; Physoconops (Jelte) neotropi- cum comb. nov. (Jelte neotropica); Physoconops (Kroe- SINICONOPINI GIBSON TRIB. NOV. beroconops) argentinus; Physoconops (Kroeberoconops) hermanni; Physoconops (Kroeberoconops) rufipennis; Type genus Siniconops Chen, 1939: 197. Included Physoconops (Pachyconops) brachyrhynchus; Physo- genera: Macroconops Kröber, 1927: 125, Siniconops. conops (Pachyconops) bulbirostris; Physoconops Uncontroverted morphological autapomorphies: patch (Pachyconops) guianicus; Physoconops (Physoconops) of bristles on female abdominal sternite 2 (86); apex discalis; Physoconops (Physoconops) fronto; Physo- of male abdomen square (110). While only a male is conops (Physoconops) obscuripennis; Physoconops described and the number of ocelli is not mentioned, (Shannonoconops) apicalis; Schedophysoconops the stated similarity to Siniconops maculifrons justi- notatifrons comb. nov. (Physoconops); Tropidomyia fies placement of Macroconops within Siniconopini. alexanderi; Tropidomyia aureifacies; Tropidomyia Members of Siniconopini have been previously bimaculata; Tropidomyia ornata; Tropidomyia sp. placed in Conopini. In addition to the two autapomorphic characters, members of Siniconopini can be distinguished from Conopini by the presence of two ocelli INCERTAE SEDIS WITHIN CONOPINAE and a narrow second male abdominal segment. Genera: Euconops Kröber, 1915e: 76, Mallachoconops Species examined – Macroconops helleri; Sini- Camras, 1955b: 160. Euconops and Mallachoconops conops elegans; Siniconops microvalvus comb. nov. are each monotypic and their respective species are (Conops); Siniconops nigripes comb. nov. (Conops); included in the present analysis. The present analysis Siniconops sepulchralis comb. nov. (Physoconops). recovers the two species in a monophyletic clade albeit it with little support. The only apomorphy for the group is a secondary reduction of the lunule TROPIDOMYIINI ZIMINA, 1960 (21). As they cannot with confidence be placed in Type genus Tropidomyia Williston, 1888: 11. Included any described tribe, these two genera remain genera genera: Physoconops Szilady, 1926: 588, Schedophyso- incertae sedis within Conopinae. Previous molecular conops Gibson gen. nov., Tropidomyia. Uncontroverted and morphological analysis (Gibson et al., 2012) failed morphological autapomorphies: none. Members of to place Euconops within any other clade of Conopi- Tropidomyiini are characterized by the presence of nae with any support. lateral ocelli (3); this character state is present Species examined – Euconops bellus; Mallachoc- in many other conopid taxa, but it represents an onops atratulus. autapomophic character reversal in Tropidomyiini. Gibson et al. (2012) also recover a monophyletic Tropidomyiini using molecular and morphological DALMANNIINAE HENDEL, 1916 analysis and including only representatives of Physo- Type genus Dalmannia Robineau-Desvoidy, 1830: conops. Zimina (1960) describes the tribe Tropidomy- 248. Included genera: Baruerizodion Papavero, iini, with the following characteristics: facial fovea 1970: 121, Dalmannia. Uncontroverted morphological absent; basisternum with anterolateral extensions; autapomorphies: female abdominal sternite and arista two-segmented; vein R1 not ending near end tergite 6 separate (88); female abdominal segment of R2+3. Hennig (1966) suggests that the absence of 7 compressed laterally along entire length (97). facial fovea is a secondary loss in Tropidomyia.Inthe Ocellar bristles are absent in all members of Dalman- present analysis, reduction of the facial fovea and niinae (14); this character state is also observed in facial ridge (20, 24) along with a strongly developed Pyrgotidae, Syrphidae, Conopinae, and some species facial carina (23) are apomorphies of Tropidomyia, not of Stylogaster. Postocellar bristles are absent in Tropidomyiini (s.l.). The condition of the basisternum, all members of Dalmanniinae (15); this character arista, and wing veins in Tropidomyiini is plesiomor- state is also observed in Pyrgotidae, Syrphidae, and phic to the Conopinae. Conopinae.

Gibson et al. (2012) recover Dalmanniinae as mono- two, separate, anterior arms on the hypandrium. Mac- phyletic with the same set of apomorphic character quart (1834) originally described Myopariae, including states, but only include representatives of Dalman- Myopa, Stachynia [now Dalmannia], and Zodion, with nia. They include the following additional apomor- the following diagnostic characters: veins R4+5 + M not phic character states: phallus coiled and setose; and petiolate; alula small; prementum elongate; labellum posterior surstyli absent. Hendel’s (1916) original sometimes elongate and folded back along prementum. mention of Dalmanniinae does not specify included In the present analysis, both petiolate and non- genera and chracter states. It only states that Dal- petiolate states of vein R4+5 + M are observed in genera manniinae possesses an unbroken costa and ‘[other of Myopinae. The prementum in Myopinae is in the primitive characteristics]’. state plesiomorphic to Conopidae. The elongate and Zimina (1960) describes the tribe Dalmanniini ventrally directed labellum is an autapomorphy of with the following key characteristics: female with Conopidae. long, curved ‘ovipositor’; phallus elongate and ‘band- In addition to the present members of Myopini shaped’; veins Sc and R1 ending not near one another; and Thecophorini, previous classifications also in- cell cup short; lower pleural sclerites bare. According cluded genera presently placed within Dalmanniinae, to the present analysis, the male and female termi- Sicinae, and Zodioninae within Myopinae. See discus- nalia characters are both autapomorphies of Dal- sions of those subfamilies for details in distinguishing manniinae. For the wing and pleural characters, the members of each respective subfamily. Dalmanniinae retains the condition plesiomorphic to Conopidae. The presence of an ‘ovipositor’ has previ- MYOPINI MACQUART, 1834 ously been used to define Dalmanniinae, and in the Type genus Myopa Fabricius, 1775: 798. Included present analysis, modification to female abdominal genera: Melanosoma Robineau-Desvoidy, 1853: 122, segment 7 is an autapomorphy of the subfamily. Myopa, Myopotta Zimina, 1969: 671, Paramyopa Superficially similar, yet distinct, modifications to the Kröber, 1916a: 91, Pseudomyopa Pearson, 1974: 148. female terminalia have occurred in other subfamilies Uncontroverted morphological autapomorphies: gena (e.g. Paramyopa, Parazodion, and Tanyconops). expanded to be over one-third of head height (16). While all other authors have afforded Dalmannii- Gibson et al. (2012) recover Myopini as mono- nae subfamily status, it was described by Zimina phyletic with the same apomorphic character state, (1960) as a tribe of Myopinae. According to the but include only representatives of Myopa. Zimina present analysis, in addition to its apomorphic char- (1960) described the tribe Myopini, including Melano- acter states, Dalmanniinae can be distinguished from soma, Myopa, and Thecophora, with the following key Myopinae by a broad basisternum, unfused veins Sc characteristics: veins Sc and R fused before reaching and R , absence of spicules on female abdominal ster- 1 1 costa; costa thickened at end of R ; lower pleural nites 5 and 6, absence of spicules on male abdominal 1 sclerites pilose. According to the present analysis, the sternite 5, narrow male syntergosternite 7 + 8, and first two characters are apomorphies of Myopinae and absence of male abdominal sternite 8. Thecophorini respectively and the third character Species examined – Baruerizodion steyskali; Dal- state is not observed. mannia aculeata; Dalmannia nigriceps; Dalmannia Pearson (1974) included Pseudomyopa in Dalman- vitiosa. niinae based on the presence of an elongate ‘append- age on the penis’. This structure is not present in the MYOPINAE MACQUART, 1834 specimens included in the present analysis. All other characters of Pseudomyopa are in agreement with the Type genus Myopa Fabricius, 1775: 798. Included present definition of Myopini. tribes: Myopini, Thecophorini trib. nov. Uncontro- Species examined – Melanosoma bicolor; Melano- verted morphological autapomorphies: basisternum soma hyalipenne; Myopa buccata; Myopa clausa; short, narrow, single sclerite (47); veins Sc and R fused 1 Myopa occulta; Myopa vesiculosa; Myopa sp.; Myo- before reaching costa (73). A double row of black spines potta pallipes; Myopotta rubripes; Paramyopa oestra- is present on the ventral surface of all femora (52) cea; Pseudomyopa camrasi. in all members of Myopinae; this character state is also observed in Baruerizodion and Robertsonomyia + Zodiomyia. THECOPHORINI GIBSON TRIB. NOV. Gibson et al. (2012) recover Myopinae as mono- Type genus Thecophora Rondani, 1845: 15. Included phyletic with the same set of apomorphic character genera: Pseudoconops Camras, 1962a: 183, Scatocce- states, but only include representatives of Myopa, myia Camras, 1957a: 13, Thecophora. Uncontroverted Thecophora, and Pseudoconops. They add the following morphological autapomorphies: costa thickened at additional apomorphic character state for those taxa: insertion of Sc + R1 (74).

Gibson et al. (2012) recover a monophyletic Theco- lata and S. stylata). The facial carina is strongly phorini using molecular and morphological analysis, developed (23) in all members of Stylogastrinae; this including representatives of Pseudoconops and character state has arisen independently in Tropid- Thecophora. Additional uncontroverted morphological omyia. The anterior margin of the subcranial cavity autapomorphies provided are: dorsal half of proepi- projects forward to a narrow point in Stylogastrinae sternum with patch of setae; hypandrium with broad, (37); this character state is also observed in Con- setose, anterior plate; and a repeated segment of DNA opinae. Maxillary palpi are absent (39, 40) in all sequence in the D2 expansion section of the 28S members of Stylogastrinae; this character state is rDNA region. The first character state is found by also observed in some members of Conopinae. Bristles the present analysis to be an autapomorphy of are present on the anepimeron (62) in all members of Pseudoconops + Thecophora. Stylogastrinae; this character state is also observed Species examined – Pseudoconops antennatus; in Strongylophthalmyiidae and Pyrgotidae. Crossvein Scatoccemyia plaumanni; Thecophora africana; sc-r is present (76) in all members of Stylogastrinae; Thecophora atra; Thecophora australiana; Theco- this character state is also present in Conopinae. The phora metallica; Thecophora modesta; Thecophora male cerci are elongate (113) in all members of Sty- occidensis. logastrinae; this character state is also observed in Pyrgotidae and Strongylophthalmyiidae. Gibson et al. (2012) recover Stylogastrinae as mono- SICINAE ZIMINA, 1960 phyletic with the same set of apomorphic character Type genus Sicus Scopoli, 1763: 1004. Included states. They add the following additional apomorphic genera: Carbonosicus Zimina, 1958: 933, Sicus. character states: bi- or tri-lobed posterior surstyli; Uncontroverted morphological autapomorphies: male male abdominal tergite 6 broad, hemispherical, inter- sternite 8 with dense, long, black setae (112). A sec- nal; phallapodeme and ejaculatory apodeme broad- ondary reversal to reduced posterolateral extensions ened into wide, thin lobe; sperm pump modified into of the basisternum is present in members of Sicinae a sclerotized sphere with lateral lobes and a long (49); a parallel reversal has occurred in members of sperm duct; and phallus with two distinct, sclerotized Conopinae. segments. See the note in Gibson et al. (2012) about Gibson et al. (2012) did not test the monophyly alternative spelling of this subfamily name and dis- of Sicinae, because only one species of Sicus was cussion of past status as a separate family. included in the analysis. Zimina (1958) first suggests Species examined – Stylogaster biannulata; Sty- the tribe Sicini, including the genera Carbonosicus logaster breviventris; Stylogaster decorata; Stylogaster and Sicus, but does not give diagnostic characters for frauci; Stylogaster inca; Stylogaster neglecta; Sty- the group. According to Sabrosky (1999), the lack of a logaster pauliani; Stylogaster rectinervis; Stylogaster description or diagnosis renders the genus-group stylata; Stylogaster westwoodi; Stylogaster sp. name invalid. In Zimina (1960), however, a diagnosis of the tribe is provided with the following key char- ZODIONINAE RONDANI, 1856 acteristics: basisternum broad; veins Sc and R1 parallel; cell cup elongate; long red bristles present on Type genus Zodion Latreille, 1797: 162. Included sides of mesonotum. According to the present analy- genera: Parazodion Kröber, 1927: 135, Robertsono- sis, the first three character states are plesiomorphic myia Malloch, 1919: 205, Zodiomyia Camras, 1957b: to Conopidae. 163, Zodion. Uncontroverted morphological autapo- Species examined – Carbonosicus carbonarius; morphy: basisternum reduced to a narrow sclerite Sicus abdominalis; Sicus ferrugineus. divided posteriorly (47). Gibson et al. (2012) recover Zodioninae as monophyletic with the same set of apomorphic character STYLOGASTRINAE WILLISTON, 1885 states, but include only representatives of Zodion and Type genus Stylogaster Macquart, 1835: 38. Included Parazodion. They add the following additional apo- genus: Stylogaster. Uncontroverted morphological morphic character states for those taxa: a developed, autapomorphies: ommatidia distinctly larger anteri- fleshy hypoproct in the male; a leaf-shaped phallus orly (18); facial fovea absent (19); protibial, apical with a single, central, sclerotized rod; and two, sepa- spurs present (50); katepisternum bare (64); alula rate, anterior arms on the hypandrium. narrow (82); female abdominal sternite 2 and 3 Rondani (1856) originally describes Zodionina, absent (85); female hypoproct sclerotized and elongate including only Zodion, based on the presence of a (99). The vertex is expanded to more than half the dorsal arista, an elongate prementum, and reduced length of the frons, sometimes reaching the ptilinum labellum. Zimina (1960) redefines the tribe Zodionini, (6) in all members of Stylogaster (except S. biannu- within Myopinae and containing only Zodion, with

© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167, 43–81 REVISION OF CONOPID GENERA 69 the following key characteristics: labella not elongate; Based on Hennig’s (1966) detailed discussion of the crossvein sc-r present; vein R4+5 + M usually petiolate. specimens, the following important character states Each of these characters are plesiomorphic, either to for †Palaeomyopa can be determined: arista thick- Conopidae or to Conopinae + Zodioninae. ened, mid-dorsal; second aristomere not expanded Robertsonomyia and Zodion have been placed ventrally; scape quadrate; pedicel elongate; facial within Myopinae in most previous classiﬁcations. fovea and medial carina present; ptilinum present; Members of Zodioninae can be distinguished from facial ridge present and not meeting subcranial cavity Myopinae by the short, broad labella, the shape of the at 90° angle; lunule absent; maxillary palpi elongate; basisternum, the presence of shiny patches near the three ocelli and ocellar tubercle present; ocellar and apex of the tibia, the extended vein R1, the unfused postocellar bristles present; vein M deﬂected towards veins Sc and R1, the presence of crossvein sc-r, the vein R4+5, but not petiolate; vein Sc ends at a point 0.4 ending of vein R2+3 near the end of vein R1, the curved times the length of the costa; vein R1 ends at a point vein CuA2, and the male cerci attached by a narrow, before 0.6 times the length of the costa; vein Sc and R1 sclerotized stalk. separate; crossvein sc-r absent; vein R2+3 ends well

Species examined – Parazodion schmidti; Parazo- beyond end of vein R1; vein CuA2 + A1 longer than dion sp.; Robertsonomyia mexicana; Robertsonomyia crossvein dm-cu; vein CuA2 straight; cell cup narrow; palpalis; Robertsonomyia parva; Robertsonomyia vena spuria absent; alula broad; female abdominal pearsoni; Zodiomyia sumbaensis; Zodion cinereum; sternite 6 with spicules. While Hennig describes Zodion erythrurum; Zodion fulvifrons; Zodion fronto-orbital bristles, it is difficult to determine if griseum; Zodion intermedium; Zodion pictum; Zodion these are true bristles or merely large setae. The pruinosum comb. nov. (Neozodion). prementum and labella are not clear on the specimens observed. Hennig also suggests that a ventral genital plate arises from female abdominal sternite 4. PLACEMENT OF EXTINCT TAXA This is probably a misinterpretation, as in many Five fossil species have been assigned to Conopidae. modern specimens, the ventral genital plate arises †Poliomyia recta Scudder, 1878 is described from the from the fifth segment but is displaced anteriorly so Green River shale. †Hoffeinsia baltica, †Palaeomyopa as to be ventral to the fourth segment. tertiaria, †Palaeomyopa hennigi Stuke, 2003, and Hennig (1966) offers one possible apomorphy for † † Palaeosicus loewi Meunier, 1916 are all described Palaeomyopa: a bend in vein CuA2. This is not to be from Baltic amber. In his analysis of the Baltic amber confused with the curved vein CuA2 observed in species, Hennig (1966) states that †Poliomyia prob- Conopinae + Zodioninae, which is in the opposite ably does not belong in Conopidae. He also notes that direction. Hennig also suggests that the antennae the type of †Palaeomyopa tertiaria is missing. Hennig of †Palaeomyopa are in a ‘primitive’ state. Camras examined Meunier’s specimens of †Palaeosicus and (1994) suggests that the elongate pedicel and ‘pecu- concluded that they are synonymous with the descrip- liar’ arista are apomorphies of †Palaeomyopa. The tion of †Palaeomyopa. Thus, two fossil genera of present analysis demonstrates that †Palaeomyopa Conopidae, †Hoffeinsia and †Palaeomyopa, are recog- retains the conopid groundplan state for all antennal nized at present. characters (27–36). Stuke (2005) places †Hoffeinsia in Conopidae due to Likewise, Hennig (1966) notes the presence of a its similarity to †Palaeomyopa. He lists the character- ‘short, thick proboscis’ with elongate maxillary palpi. istics of †Hoffeinsia as: absence of facial foveae; broad If this is so, then the palpi retain a condition plesio- parafacial; extremely short prementum; and distinct morphic to Conopidae, but the prementum and labella chaetotaxy of the head and thorax. The absence of would represent apomorphic states, possibly conver- facial foveae is found within Conopidae in Stylogaster. gent with the condition observed in Leopoldius. The size of the parafacial is variable within Conopi- Camras (1994) includes reduced mouthparts as a dae. Based on Stuke’s description and figures, the plesiomorphic condition in his description of †Palaeo- prementum does not appear to be fused and elongate; myopinae (containing only †Palaeomyopa). However, an autapomorphy of Conopidae. Additionally, the cha- Hennig (1966) also states that mouthparts are absent etotaxy of the head and thorax is unlike any other or otherwise very difficult to diagnose in the fossil examined member of Conopidae. Furthermore, †Hof- specimens of †Palaeomyopa. Further examination of feinsia lacks all but one (78 – vein CuA2 + A1 elongate) existing specimens will be necessary to assess these of the apomorphic characters of Conopidae. For these potential apomorphies. reasons, †Hoffeinsia cannot be included in Conopidae. Camras (1994) and Hennig (1966) both note that Its proper family classification is unknown and it †Palaeomyopa, Stylogaster, and Tropidomyia all should be left as incertae sedis within Schizophora possess a strong medial carina and weak, if not pending further evidence of its proper placement. absent, facial foveae. He suggests that this is a

© 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167, 43–81 70 J. F. GIBSON AND J. H. SKEVINGTON plesiomorphic condition in the prior two genera, but is additional apomorphic character states: male abdomi- secondarily derived in Tropidomyia. The illustration nal tergite 6 narrow, bare, and fused with syntergos- in Hennig (1966) clearly shows the presence of a ternite 7 + 8; and postgonites present. facial fovea and medial carina in †Palaeomyopa, sug- Elongate, narrow, posterolateral extensions of the gesting a condition plesiomorphic to Conopidae for basisternum (49) is an apomorphy of Conopidae these characters (19, 20, 22, 23). excluding Stylogastrinae and has been independently Hennig (1966) also notes the presence of fronto- reversed in Sicinae and Conopinae. An equally parsi- orbital bristles (12), a plesiomorphic condition in monious explanation is the independent evolution of †Palaeomyopa. Although the degree of setation varies this character state in three subfamilies: Dalmannii- throughout Conopidae, the presence of true bristles nae, Myopinae, and Zodioninae. Unfortunately, this on the frons is limited to some species of Stylogaster. character cannot be diagnosed in †Palaeomyopa. In the present analysis, the absence of fronto-orbital A clade including ((Conopinae + Zodioninae) + bristles is an apomorphy of Conopidae excluding Sty- Sicinae) + Myopinae is defined by a number of modi- logastrinae. If bristles are truly present in †Palaeo- fications to the female and male abdomen. These myopa, it would suggest excluding it from this clade, apomorphic character states include: presence of spi- but closer examination is necessary. cules on female abdominal sternites 5 and 6 (91, 90); Hennig’s (1966) and Camras’ (1994) interpretation the deflection of female abdominal segment 7 ventrally of the ventral genital plate as arising from the fourth (96); male abdominal sternite 5 broad and with spi- abdominal sternite in †Palaeomyopa leads each of cules (106); male abdominal sternite 8 present (111); them to conclude this is the groundplan state of the and male abdominal syntergosternite 7 + 8 broad and Conopidae excluding Stylogastrinae. However, if the hemispherical (109). The spicules on female sternite 6 ventral genital plate arises from the fifth sternite have been secondarily lost in Parazodion and the and has merely been displaced in the available deflection of female segment 7 has arisen independ- specimen, then there is nothing in the female abdomi- ently in Baruerizodion. The broad male syntergoster- nal characters to distinguish †Palaeomyopa from nite 7 + 8 has arisen independently in Old World Myopinae. Stylogaster (i.e. S. frauci + S. pauliani + S. westwoodi This leaves †Palaeomyopa without any clear apo- + S. sp.). While the male character states are morphic character states. This is unsurprising for unknown, Palaeomyopinae possesses all three of the a fossil species. Only further specimens and further female abdominal character states apomorphic to this examination will reveal possible apomorphies for clade. †Palaeomyopa. Gibson et al. (2012) recover ((Conopinae + Zodio- ninae) + Sicinae) + Myopinae as monophyletic with the same set of apomorphic character states. The EVOLUTION OF CHARACTER STATES WITHIN following additional apomorphic character state is CONOPIDAE added: anterior surstyli present. In addition to those characters previously discussed Hennig (1966) suggests †Palaeomyopa as a possible that are apomorphic to certain clades, the following sister group to Conopidae excluding Stylogastrinae. characters are present in varying states within He also suggests it as possibly an ancestral form of Conopidae. As such, these characters can help to either Conopinae or Myopinae (including Zodion and determine relationships between subfamily clades Dalmannia in his analysis). With respect to wing within Conopidae. venation, Hennig notes that the absence of crossvein

The absence of fronto-orbital bristles (12) is an sc-r and the equally spaced insertions of Sc, R1, and † apomorphy of Conopidae excluding Stylogastrinae. R2+3 along the costa suggest that Palaeomyopa has This character state, however, is also observed in retained the plesiomorphic wing condition. Camras Pyrgotidae, Psilidae, Syrphidae, Notoconops, and (1994) describes the wing venation of †Palaeomyopi- a clade of Old World Stylogaster (S. frauci + S. nae as plesiomorphic. In the present analysis, †Pal- pauliani + S. westwoodi + S. sp.). Other apomorphies aeomyopa lacks all of the wing characters apomorphic of Conopidae excluding Stylogastrinae include: the to (Conopinae + Zodioninae) + Sicinae (72, 75), Myopi- extension of vein Sc to a point beyond four-tenths the nae (73), and Stylogastrinae (76), respectively. There length of the costa (71); female sternite 8 divided into remains equal likelihood that †Palaeomyopa repre- two, sclerotized lobes (100); and presence of sclero- sents a sister group to (Conopinae + Zodioninae) + tized posteroventral hooks on female syntergite 8 + 9 Sicinae, or Myopinae, or is an ancestral form of either.

(102). The extension of vein R1 to a point beyond six-tenths

Gibson et al. (2012) recover Conopidae excluding of the length of the costa and near to the end of R2+3 (72, Stylogastrinae as monophyletic with the same set of 75) are apomorphies of (Conopinae + Zodioninae) + apomorphic character states. They add the following Sicinae. Gibson et al. (2012) recover (Conopinae +

Zodioninae) + Sicinae as monophyletic with the same Sicinae is limited in generic and species diversity. It set of apomorphic character states. is also exclusive to the Palaearctic Region and the Elongate, filiform, partly fused labella that fold northern reaches of the Oriental Region. back along the prementum are the groundplan Within Zodioninae, Zodion is global in distribution. conditions within Conopidae. The reduction of the Parazodion, Robertsonomyia, and Zodiomyia, how- labella to a shortened, separate, broad state (44, 45) ever, are endemic to the Neotropics, the New World, are autapomorphies of Conopinae + Zodioninae. Addi- and the Oriental Region, respectively. tional apomorphies of this clade include: the presence Conopinae is truly global in distribution, but closer of a shiny patch near the apex of the metatibia (53); analysis of its component tribes reveals some biogeo- the presence of crossvein sc-r (76); vein CuA2 curved graphical pattern. Neoconopini is the sister group (79); female abdominal tergite and sternite 5 fused of the remaining Conopinae and is one of two tribes (87); and male cerci attached by a narrow, sclerotized composed entirely of Australian endemic genera. The stalk (114). Gibson et al. (2012) recover Conopinae + remaining tribes of Conopinae are each restricted to a Zodioninae as monophyletic with the same set of specific area, with notable exceptions. Conopini and apomorphic character states. Pleurocerinellini are both exclusively Old World, but absent from Australia. A single species of Conops, pruinosus Bigot, 1887, is recorded from the New BIOGEOGRAPHY OF CONOPIDAE World, but this identification and/or locality informa- While an overall biogeographical pattern is difficult tion should probably be questioned. Gyroconopini is to determine for Conopidae, a number of intra- found only in the New World. The distribution of subfamilial observations can be made that may facili- Siniconopini is limited to the far eastern Palaearctic tate future investigation. The divisions between and Oriental Regions. Microconopini is exclusively biogeographical regions employed here follow those of Australian. Members of Brachyceraeini are found McAlpine et al. (1981) and Papp & Darvas (2000). only in the Palaearctic and Oriental Regions. Caeno- Stylogastrinae has a southern distribution. The conopini is the only tribe endemic to the Afrotropical subfamily is absent from the Palaearctic and only two Region. Asiconopini and Physocephalini are both species are found in the Nearctic Region. The bulk of global in distribution, with some genera and subgen- the diversity appears to be found in the Neotropics. era being endemic to certain regions. Furthermore, both Gibson et al. (2012) and the The remaining tribe, Tropidomyiini, presents an present analysis recover four Old World species of odd distribution amongst its three included genera. Stylogaster as a monophyletic clade within the Schedophysoconops is exclusively Afrotropical, but is subfamily. This phylogeny and distribution pattern monotypic. Tropidomyia is circumtropical in distribu- suggests a possible origin of Stylogaster in South tion. Physoconops is endemic to the New World. Four America with a single invasion and radiation in the species of Physoconops from the Afrotropical and Ori- Old World. ental Regions are reclassified here. The two remain- In the present analysis, Dalmanniinae is divided ing Oriental species (P. jutogensis Nayar, 1968 and into the Holarctic Dalmannia and the Neotropical P. borneensis Kröber, 1940a) need to be re-examined, Baruerizodion. Two species of Dalmannia are included but are probably not Physoconops. in Smith’s (1975) Oriental catalogue, but they are both While taxon representation of all global regions is from China’s Jiangsu province, which is not conven- certainly not equal in the present analysis, a general tionally considered part of the Oriental Region. pattern can be observed. The Stylogastrinae probably Myopinae and Thecophorini are each global in dis- originated in South America, with a subsequent radia- tribution with little pattern evident in the present tion into Africa and Australia. The remaining Conopi- analysis. Myopini, however, displays an interesting dae probably originated in the northern hemisphere. pattern. Three species of Myopa are included in This hypothesis is strengthened by the location of a Papavero’s (1971) Neotropical catalogue, but they fossil of †Palaeomyopinae in Baltic Amber. Some are all from the Chihuahua state of Mexico, which is clades (e.g. Dalmanniinae, Myopini, Sicinae) have not conventionally considered part of the Neotropical spread throughout the Holarctic, but little beyond. Region. The species of Myopa included in Smith’s Zodioninae appear to have spread to all geographical (1975) Oriental catalogue are all from northern China regions, but with little genus-level diversification. or northern India. A single species of Myopa is Finally, Conopinae represents a series of tribe-level recorded from Australia. Melanosoma and Myopotta invasions and subsequent radiations within various are recorded only from the Palaearctic. Myopini can regions. This radiation has been accompanied by a thus be described as mainly Holarctic, with Paramy- great deal of genus- and species-level diversity, but opa and Pseudomyopa being the sole representatives little genitalic and molecular diversity (Gibson et al., of the tribe in Africa and South America, respectively. 2012).

CONCLUSIONS ACKNOWLEDGEMENTS The present analysis represents the most comprehen- We thank M. Mei, M. Irwin, S. Winterton, M. Hauser, sive phylogenetic analysis of relationships within K. Bayless, M. Locke, J. Cumming, S. Cumming, H. Conopidae to date. Parsimony analysis of a 117- Cumming, S. Brooks, J. Corrigan, M. Földvári, M. character morphological matrix recovers a phyloge- Jackson, C. Kehlmaier, O. Lonsdale, J. O’Hara, S. netic hypothesis for the family. A revised classification Marshall, M. Pollet, T. Wheeler, S. Gaimari, J. is proposed based on this phylogenetic hypothesis Savage, and B. Sinclair for providing specimens (see Appendix). All extant and two fossil genera are and/or identifications. Thanks, as well, to contacts at discussed. Only nine genera were unavailable for all insect collections who facilitated visits and/or observation and are included based on literature loans of materials. Special thanks to Dr S. Camras descriptions. (FMNH), one of the fathers of conopidology, who pro- All previously proposed subfamilies are recovered vided insight into conopid classification. Thanks as with numerous autapomorphic character states. well to M. Jackson (DEBU) and W. Knee (CNC) for Generic membership of most subfamilies (except assistance with analyses. Funding to J.H.S. was pro- Conopinae and Stylogastrinae) is revised. The previ- vided by Agriculture & Agri-Food Canada, an NSERC ously proposed tribe Sicini is elevated to subfamily Discovery Grant, and NSERC Canpolin. J.F.G. was rank. Relationships between subfamilies are also supported by an NSERC Postgraduate Scholarship recovered. Placement of the fossil subfamily †Palaeo- and an Ontario Graduate Scholarship. The manu- myopinae is ambiguous, but placement within the script was improved based on input from G. Capretta, ((Conopinae + Zodioninae) + Sicinae) + Myopinae M. Hauser, S. Peck, P. J. 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A new species of Neobrach- 54: 180–185. yceraea Szilady, 1926 (Diptera: Conopidae) from Thailand and a review of the genus. Tijdschrift voor Entomologie 148: 355–360. APPENDIX Szilady Z. 1926. Dipterenstudien. Annales Musei Nationlis Catalogue of genus-group names of Conopidae. Hungarici 24: 586–593. Villeneuve J. 1909. Contribution au catalogue des Diptères de France. La Feuille des Jeunes Naturalistes. Paris 39: FAMILY CONOPIDAE LATREILLE, 1802 153–155. Type genus Conops Linnaeus, 1758: 604.

SUBFAMILY CONOPINAE LATREILLE, 1802 Abrachyglossum Kröber, 1919:142; type – Conops Type genus Conops Linnaeus, 1758: 604. capitatus Loew, 1847, original designation. Conops Linnaeus, 1758: 604; type – Conops flavipes Linnaeus, 1758, designated by Curtis, 1831: TRIBE ASICONOPINI GIBSON TRIB. NOV. 377. Type genus Asiconops Chen, 1939: 171. Conopaejus Rondani, 1845: 7; type – Conops quad- Anticonops Kröber, 1936: 285; type – Anticonops rifasciatus De Geer, 1776, original designation [teste abdominalis Kröber, 1936, monotypy. Chvála & Smith, 1988: 248]. Archiconops Smith, 1975: 375; originally Conopoideus Rondani, 1845: 9; type – Conops fer- described by Kröber, 1939a: 381 without type; type – ruginea Macquart, 1834 [preoccupied by Conops Conops insularis Kröber, 1936, designated by Smith, ferruginea Linnaeus, 1761; = Conops vesicularis 1975. Linnaeus, 1761], monotypy [teste Coquillett, 1910: Asiconops Chen, 1939: 171; type – Conops aureo- 527]. maculatus Kröber, 1933, original designation, stat. Conopilla Rondani, 1845: 9; type – Conops ceriae- nov. formis Meigen, 1824, monotypy [teste Chvála & Asiconops (Aegloconops) Gibson subgen. nov.; Smith, 1988: 248]. type – Conops quadripunctatus Kröber, 1915d: 38, Sphyxosoma Rondani, 1856: 56; type – Conops present designation. quadrifasciatus De Geer, 1776, original designation Asiconops (Ceratoconops) Camras, 1955b: 159; [teste Coquillett, 1910: 527]. type – Conops ornatus Williston, 1892a, original des- Sphixosoma Rondani, 1857: 12 (unjustified emen- ignation, comb. nov. dation) [teste Chvála & Smith, 1988: 248]. Asiconops (Diconops) Camras, 1957a: 9; type – Spixosoma Rondani, 1857: 223 (unjustified emen- Conops trichus Camras, 1957a, original designation, dation) [teste O’Hara et al., 2011: 170]. comb. nov. Bombibia Lioy, 1864: 1326; type – Conops flavipes Asiconops (Sphenoconops) Camras, 1955b: 159; Linnaeus, 1758, monotypy [teste Coquillett, 1910: type – Conops nobilis Williston, 1892a, original des- 527]. ignation, comb. nov. Conopoejus Bigot, 1892: 164 (unjustified emenda- Smithiconops Camras, 2000: 221; type – Conops tion) [teste O’Hara et al., 2011: 63] rondanii Bezzi, 1901, original designation, stat. nov. Bombidia Aldrich, 1905: 407 (misspelling) syn. nov. Sphecosoma Bezzi, 1906: 51 (unjustified emendation) [teste O’Hara et al., 2011: 170]. TRIBE BRACHYCERAEINI ZIMINA, 1960 Conopejus Coquillett, 1910: 527 (misspelling) syn. Type genus Brachyceraea Röder, 1892: 366. nov. Brachyceraea Röder, 1892: 366; type – Conops Leopoldius Rondani, 1843: 35; type – Leopoldius brevicornis Loew, 1847, original designation. erostratus Rondani, 1843 = Conops signatus Wiede- Brachyceratias Röder, 1893: 62; unnecessary mann, 1824, original designation. replacement for Brachyceraea Röder, 1892 based on Brachiglossum Rondani, 1856: 56; type – Leopold- supposed preoccupation [teste Sabrosky, 1999: 70]. ius erostratus Rondani, 1843 = Conops signatus Wie- Microbrachyceraea Kröber, 1940b: 217; type – demann, 1824, original designation [teste Chvála & Microconops pendleburyi Brunetti, 1927, original des- Smith, 1988: 247]. ignation. Brachyglossum Bigot, 1858: 592 (unjustified emen- Neobrachyceraea Szilady, 1926: 587; type – dation) [teste O’Hara et al., 2011: 41]. Brachyceraea obscuripennis Kröber, 1913, original Neobrachyglossum Kröber, 1915e: 79; type designation. species, Neobrachyglossum punctatum Kröber, 1915e, monotypy, syn.nov. TRIBE CAENOCONOPINI GIBSON TRIB. NOV. Type genus Caenoconops Anonymous, 1940: 365. TRIBE GYROCONOPINI GIBSON TRIB. NOV. Caenoconops Anonymous in Imperial Institute of Entomology, 1940: 365; originally described by Type genus Gyroconops Camras, 1955b: 174. Kröber, 1939a: 373 without type; type – Caenoconops Gyroconops Camras, 1955b: 174; type – Conops subapicalis Kröber, 1939a (= Caenoconops bicolor sylvosus Williston, 1882, original designation, stat. (Kröber, 1931), teste Camras, 2000: 229), monotypy. nov.

TRIBE CONOPINI LATREILLE, 1802 TRIBE MICROCONOPINI GIBSON TRIB. NOV. Type genus Conops Linnaeus, 1758: 604. Type genus Microconops Kröber, 1915e: 77.

Australoconops Camras, 1961: 64; type – Conops bimarginipennis Karsch, 1887, monotypy [teste Smith splendidus Kröber, 1916b, original designation. & Cunningham-Van Someren, 1970: 439]. Microconops Kröber, 1915e: 77; type – Micro- Pseudophysocephala Kröber, 1940b: 221; type – conops fasciatus Kröber, 1915e, designated by Conops platycephala Loew, 1853, original designation. Malloch, 1933: 174.

TRIBE PLEUROCERINELLINI ZIMINA, 1974 TRIBE NEOCONOPINI GIBSON TRIB. NOV. Type genus Pleurocerinella Brunetti, 1923: 368. Type genus Neoconops Kröber, 1915e: 75. Pleurocerinella Brunetti, 1923: 368; type – Pleu- Atrichoparia Schneider, 2010: 33; type – rocerinella dioctriaeformis Brunetti, 1923, original Heteroconops curticornis Kröber, 1940a, original designation. designation. Tammo Stuke, 2008: 50: type – Euconops rufus Callosiconops Kröber, 1940a: 75; type – Callosi- Camras, 1955a, original designation. conops hirsutus Kröber, 1940a, monotypy, stat. rev. Chrysidiomyia Smith, 1989: 49 460; originally TRIBE SINICONOPINI GIBSON TRIB. NOV. described by Kröber, 1940a: 73 without type; type – Type genus Siniconops Chen, 1939: 197. Chrysidiomyia rufa Kröber, 1940a, original designa- Macroconops Kröber, 1927: 125; type – Macro- tion. syn. nov. conops helleri Kröber, 1927, monotypy. Camrasiconops Schneider, 2010: 78; type – Micro- Siniconops Chen, 1939: 197; type – Siniconops conops ater Camras, 1961, original designation. elegans Chen, 1939, original designation. Delkeskampomyia Kröber, 1940a: 71; type – Delkeskampomyia fasciata Kröber, 1940a, monotypy. Heteroconops Kröber, 1915e: 80; type – Hetero- TRIBE TROPIDOMYIINI ZIMINA, 1960 conops gracilis Kröber, 1915e, monotypy. Type genus: Tropidomyia Williston, 1888: 11. Neoconops Kröber, 1915e: 75; type – Neoconops Physoconops Szilady, 1926: 588; type – Conops longicornis Kröber, 1915e, monotypy. brachyrrhynchus Macquart, 1843 = Conops obscurip- Pleurocerina Macquart, 1851: 164; type – ennis Williston, 1882 [misidentiﬁed, see note in Pleurocerina fasciata Macquart, 1851, original Camras, 1955b], original designation. designation. Physoconops (Aconops) Kröber, 1919: 142; type Paraconops Kröber, 1915e: 74; type – Paraconops Conops antennatus Kröber, 1915f, designated by longicornis Kröber, 1915e, monotypy [teste Schneider, Malloch, 1933: 173 [see note in Camras, 1955b]. 2010: 133]. Physoconops (Aureoconops) Camras, 2004: 86; Setosiconops Schneider, 2010: 157; type – Neo- type – Physoconops aureolus Camras, 2004, original conops robustus Kröber, 1940a, original designation. designation. Smartiomyia Kröber, 1940a: 72; type – Smarti- Physoconops (Jelte) Stuke, 2008: 52; type – Pleu- omyia obscura Kröber, 1940a, monotypy. rocerinella neotropica Camras, 2008, original desig- Stenoconops Kröber, 1939b: 606; type – Steno- nation, comb. nov. conops niger Kröber, 1939b, monotypy. Physoconops (Kroeberoconops) Camras, 1955b: Tanyconops Schneider, 2010: 170; type – 175; type – Physoconops hermanni Kröber, 1915f, Tanyconops longicaudus Schneider, 2010, original original designation. designation. Physoconops (Pachyconops) Camras, 1955b: 161; type Conops bulbirostris Loew, 1853, original designation. TRIBE PHYSOCEPHALINI SMITH &PETERSON, 1987 Physoconops (Shannonoconops) Camras, 1955b: Type genus Physocephala Schiner, 1861: 137. 172; type Physoconops apicalis Camras, 1955b, origi- Dacops Speiser, 1923: 99; replacement name for nal designation. Pseudodacus Kröber, 1915e (preoccupied); type – Schedophysoconops Gibson, gen. nov.; type Pseudodacus abdominalis Kröber, 1915e, designated Physoconops notatifrons Camras, 1962b: 219, present by Smith, 1980: 513. designation. Pseudodacus Kröber, 1915e: 81; type Pseudodacus Tropidomyia Williston, 1888: 11; type – Tropid- abdominalis Kröber, 1915e, designated by Smith, omyia bimaculata Williston, 1888, monotypy. 1980: 513. Physocephala Schiner, 1861: 137; type – Conops ruﬁpes Fabricius, 1781, original designation. INCERTAE SEDIS WITHIN CONOPINAE Archiphysocephala Kröber, 1939a: 374, 392; type Euconops Kröber, 1915e: 76; type – Euconops bellus Physocephala gigantea Kröber, 1936 = Physocephala Kröber, 1915e, monotypy.

Mallochoconops Camras, 1955b: 160; type – Myopina Robineau-Desvoidy, 1853: 119; type – Conops atratula Malloch, 1933, original designation Myopa variegata Meigen, 1804, original designation [= Mallachoconops atratulus]. [teste Chvála & Smith, 1988: 262]. Haustellia Robineau-Desvoidy, 1853: 127; type – Myopa occulta Wiedemann, 1824, original designation SUBFAMILY DALMANNIINAE HENDEL, 1916 [teste Chvála & Smith, 1988: 262]. Type genus Dalmannia Robineau-Desvoidy, 1830. Heterellia Robineau-Desvoidy MS name; listed as Baruerizodion Papavero, 1970: 121; type – genus in Stuke & Clements, 2005, currently unavail- Baruerizodion steyskali Papavero, 1970, original des- able name, nomen dubium [teste Evenhuis et al., ignation. 2010: 89]. Dalmannia Robineau-Desvoidy, 1830: 248; type – Glossigona Rondani, 1856: 58; type – Myopa occulta Myopa gemina Wiedemann, 1824 [= Conops aculeatus Wiedemann 1824, original designation [teste Chvála Linnaeus, 1767], automatic [by designation of the & Smith, 1988: 262]. same species (by subsequent designation of Desmar- Gonirhyncus Rondani, 1856: 58; type – Gonirhyn- est in d’Orbigny, 1848: 798) for Stachynia Macquart, cus dispar Rondani, 1856 [= Myopa occulta Wiede- 1835] (see note in Evenhuis et al., 2010). mann, 1824], original designation [teste Chvála & Stachynia Macquart, 1835; type – Myopa gemina Smith, 1988: 262, but see note in O’Hara et al., 2011: Wiedemann, 1824 [= Conops aculeatus Linnaeus, 90]. 1767], designated by Desmarest in d’Orbigny, 1848: Arpagita Lioy, 1864: 1327; type – Stomoxys dorsalis 798 [teste Coquillett, 1910: 608]. Fabricius, 1794, original designation [teste Coquillett, Dalmania Robineau-Desvoidy, 1853: 92, 146 (unjus- 1910: 510]. tified emendation) Robineau-Desvoidy, 1830. Gonirhinchus Lioy, 1864: 1327 (unjustified emendation) [teste O’Hara et al., 2011: 90]. Gonirhynchus Marschall, 1873: 332 (unjustified emendation) [teste O’Hara et al., 2011: 197]. SUBFAMILY MYOPINAE MACQUART, 1834 Myiopa Giglio-Tos, 1893: 70, error [teste Sabrosky, Type genus Myopa, Fabricius, 1775: 798. 1999: 207]. Myopotta Zimina, 1969: 671; type – Myopa pallipes Wiedemann, 1824, original designation. TRIBE MYOPINI MACQUART, 1834 Paramyopa Kröber, 1916a: 91; type – Myopa insig- Type genus Myopa, Fabricius, 1775: 798. nis Jaennicke, 1867, monotypy. Melanosoma Robineau-Desvoidy, 1853: 122; type – Pseudomyopa Pearson, 1974: 148; type – Pseu- Myopa nigra Meigen, 1824, original designation. domyopa camrasi Pearson, 1974, original designation. Myopa Fabricius, 1775: 798; type – Conops buccata Linnaeus, 1758 (designated by Curtis, 1838: 677; I.C.Z.N., 1997: 133 [Opinion 1870]). TRIBE THECOPHORINI GIBSON TRIB. NOV. Pictinia Robineau-Desvoidy, 1853: 95; type – Myopa Type genus Thecophora Rondani, 1845: 15, present buccata Robineau-Desvoidy, 1830, original designa- designation. tion [teste Chvála & Smith, 1988: 261]. Pseudoconops Camras, 1962a: 183; type – Pseu- Myopella Robineau-Desvoidy, 1853: 98; type – doconops antennatus Camras, 1962a, original desig- Conops buccata Linnaeus, 1758, original designation nation. [teste I.C.Z.N., 1997: 133 [Opinion 1870]]. Scatoccemyia Camras, 1957a: 13; type Scatocce- Phorosia Robineau-Desvoidy, 1853: 109; type – myia plaumanni Camras, 1957a, original designation. Conops testaceus Linnaeus, 1767, original designation Thecophora Rondani, 1845: 15; type – Myopa atra [teste Chvála & Smith, 1988: 262]. Fabricius, 1775, monotypy. Fairmairia Robineau-Desvoidy, 1853: 111; type – Occemya Robineau-Desvoidy, 1853: 130; type – Myopa morio Meigen, 1804, original designation [teste Myopa atra Fabricius, 1775, original designation, Chvála & Smith, 1988: 262]. [teste Coquillett, 1910: 577] Fairemairia Robineau-Desvoidy, 1853: 111 (mis- Occemyia Schiner, 1861: 381 (unjustified emenda- spelling) [teste Evenhuis et al., 2010: 79]. tion) [teste Chvála & Smith, 1988: 267]. Lonchopalpus Robineau-Desvoidy, 1853: 115; type – Occemyia Marschall, 1873: 339 (unjustified emen- Myopa dorsalis Fabricius, 1794, original designation dation) [teste Chvála & Smith, 1988: 267]. [teste Chvála & Smith, 1988: 262]. Oncomyia Loew, 1866: 41 (unjustified emendation) Purpurella Robineau-Desvoidy, 1853: 117; [teste Chvála & Smith, 1988: 267]. type – Purpurella nobilis Robineau-Desvoidy, 1853, Eccemyia Graenicher, 1910: 44 (misspelling) [teste monotypy [teste Chvála & Smith, 1988: 262]. Chvála & Smith, 1988: 267].

Thecomyia Brues and Melander, 1932: 307 (mis- Stylogaster Macquart, 1835: 38; type – Conops spelling), syn. nov. stylatus Fabricius, 1805, monotypy. Stylomyia Westwood, 1851: 268; type – Stylomyia † SUBFAMILY PALAEOMYOPINAE CAMRAS, 1994 leonum Westwood, 1851, designated by Coquillet, Type genus †Palaeomyopa Meunier 1899: 145. 1910: 610, [teste Coquillett, 1910: 610]. †Palaeomyopa Meunier, 1899: 145; type †Palaeo- Ptychoproctus Bigot, 1859: 308; type – Ptychoproc- myopa tertiaria Meunier, 1912, subsequent monotypy. tus complexus Bigot, 1859, monotypy, [teste Coquillett, †Palaeosicus Meunier, 1916: 280; type †Palaeosicus 1910: 610]. loewi Meunier, 1916 = †Palaeomyopa tertiaria Meunier, 1912, monotypy [teste Hennig, 1966: 6]. SUBFAMILY ZODIONINAE RONDANI, 1856 SUBFAMILY SICINAE ZIMINA, 1960 Type genus Zodion Latreille, 1797: 162. Type genus Sicus Scopoli, 1763: 1004. Parazodion Kröber, 1927: 135; type – Parazodion Carbonosicus Zimina, 1958: 933; type – Melano- schmidti Kröber 1927, monotypy. soma carbonaria Kröber, 1915e, original designation. Robertsonomyia Malloch, 1919: 205; type – Zodion Sicus Scopoli 1763: 1004; type – Conops ferru- palpalis Robertson, 1901, original designation. gineus Linnaeus, 1761, designated by I.C.Z.N., 1997: Zodiomyia Camras, 1957b: 163; type – Zodiomyia 133, see note in Sabrosky, 1999: 282. sumbaensis Camras, 1957b, original designation. Cylindrogaster Lioy, 1864: 1327; type – Conops fer- Zodion Latreille, 1797: 162; type – Myopa cinerea rugineus Linnaeus, 1761 [teste Coquillett, 1910: 529]. Fabricius, 1794, subsequent monotypy by Latreille, 1802: 444. SUBFAMILY STYLOGASTRINAE WILLISTON, 1885 Neozodion Szilady, 1926: 590; type Neozodion pru- Type genus Stylogaster Macquart, 1835: 38. inosum Szilady, 1926, monotypy, syn. nov.

SUPPORTING INFORMATION Additional Supporting Information may be found in the online version of this article: Table S1. Character state matrix used for phylogenetic analysis.