An illustrated identification key to Nearctic genera of Empidoidea (exclusive of Dolichopodidae sensu stricto) (Diptera)
- CJAI #48
- September 27, 2023
Bradley J. Sinclair1,2*
,
Scott E. Brooks2
,
Jeffrey M. Cumming2
1 Canadian Food Inspection Agency, K.W. Neatby Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6; bradley.sinclair@inspection.gc.ca
2 Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6; scott.brooks@agr.gc.ca; jeff.cumming@agr.gc.ca
* Corresponding author
An illustrated identification key to Nearctic genera of Empidoidea (exclusive of Dolichopodidae sensu stricto) (Diptera)
An illustrated identification key to Nearctic genera of Empidoidea (exclusive of Dolichopodidae sensu stricto) (Diptera)
- CJAI #48
- September 27, 2023
Bradley J. Sinclair1,2*
,
Scott E. Brooks2
,
Jeffrey M. Cumming2
1 Canadian Food Inspection Agency, K.W. Neatby Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6; bradley.sinclair@inspection.gc.ca
2 Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, K.W. Neatby Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6; scott.brooks@agr.gc.ca; jeff.cumming@agr.gc.ca
* Corresponding author
Abstract
The Empidoidea (exclusive of Dolichopodidae sensu stricto), or “dance flies”, are a diverse group of Diptera that include six families and 72 genera in the Nearctic Region. An illustrated key to the dance fly genera is presented, updating the 1981 key from the “Empididae” chapter of the Manual of Nearctic Diptera, utilizing over 600 colour photographs displaying many unique characters as well as some rarely seen or photographed genera. Since publication of the Manual chapter, 14 generic names have been added (or updated), five generic names have been removed and two undescribed genera are now recognized. Currently, 953 Nearctic species of Empidoidea (exclusive of Dolichopodidae sensu stricto) have been described. A brief synopsis of each genus is included providing information on number of species, taxonomic literature, biology, defining morphological characters and Nearctic distribution.
Introduction
The superfamily Empidoidea has traditionally included two families: Empididae (dance flies) and Dolichopodidae (long-legged flies). Empidoids are very widespread from tundra to deserts and alpine streams to marine coastlines. Both adults and larvae are predaceous on a wide variety of soft-bodied arthropods (Sinclair & Cumming 2006). In addition, they are important pollinators, especially for early spring flowers and in colder ecological zones (Lefebvre et al. 2014). There are more than 13000 described species in the world and 2250 described species in Canada and the United States (Table 1).
The most recent keys to genera of the Empidoidea in the Manual of Nearctic Diptera recognized two families: Empididae (Steyskal & Knutson 1981) and Dolichopodidae (Robinson & Vockeroth 1981), but numerous taxonomic changes have been since proposed. As many as nine families are now recognized in the Empidoidea, although not all have been accepted by specialists. For the Nearctic Region, the former “Empididae” as treated in the Manual of Nearctic Diptera has been divided into five families: Atelestidae, Brachystomatidae, Empididae, Hybotidae and Iteaphilidae, as well as the subfamilies Microphorinae and Parathalassiinae which are now classified within the family Dolichopodidae sensu lato (Table 1) (Sinclair & Cumming 2006; Sinclair & Shamshev 2021). The remaining Dolichopodidae are now referred to as the Dolichopodidae sensu stricto and the Nearctic genera of that large group can be identified using the key in Bickel (2009). The main families of Empidoidea can be identified using the key to families and genus groups in Sinclair & Cumming (2006: 82) and using “Key Four” in Marshall (2012: 580).
There have also been changes in subfamily classification, with several newly proposed groups. In North America, these new or revised subfamilies include Bicellariinae, Brochellinae, Oedaleinae, Ragadinae, Trichininae and Trichopezinae (Table 1) (Sinclair & Cumming 2006; Yang et al. 2007; Sinclair 2016; Wahlberg & Johanson 2018).
In addition to changes in family status, there have been numerous taxonomic changes to the genera of “Empididae” in the Nearctic Region since Steyskal & Knutson (1981). These include 14 generic names added or updated, five generic names removed and two undescribed genera recognized (Table 2). These taxonomic changes support the urgent need to update the 1981 key to “Empididae” presented in the Manual of Nearctic Diptera to allow identification of the 72 genera currently recognized in the Region (Table 1).
Diagnosis of Empidoidea
Members of the superfamily Empidoidea can be characterized by the following combination of features:
- Antenna with apical flagellum consolidated into a compact, single-segmented postpedicel bearing a distinct robust stylus or arista-like stylus.
- Mouthparts with palpus one-segmented.
- Mouthparts with labrum usually armed at apex with epipharyngeal blades.
- Head without ptilinal fissure.
- Wing with vein CuA short, ending in vein CuA+CuP, usually near wing base.
- Wing with cell cua closed, never reaching wing margin.
- Wing with cell dm usually present, but if not, then R4+5 and M1+2 not forked.
- Legs often with one pair possessing raptorial modifications.
- Legs with apical tarsomere usually bearing two broad, flattened pads below tarsal claws.
- Male terminalia either unrotated, twisted 45º to 90º to right, or rotated 90º to 180º to right and lateroflexed; often asymmetrical.
- Female terminalia with or without acanthophorous spines.
- Female terminalia with one spermatheca internally.
Conveniently, Empidoidea (exclusive of Dolichopodidae sensu stricto) can generally be differentiated from similar Dolichopodidae (sensu stricto) (see Bickel 2009) by their lack of green metallic body colouration. However, some empidoids (e.g., most Lamprempis Wheeler & Melander and Porphyrochroa Melander) are metallic blue or blue-green and some dolichopodids (sensu stricto) are not metallic. A more precise separation of Dolichopodidae (sensu stricto) from the remaining Empidoidea is provided in our key, particularly in couplets 4 and 73, based on differences in wing venation. Some bombyliid and asiloid groups (e.g., Therevidae sensu lato), as well as Bolbomyia Loew (Rhagionidae sensu lato), can be confused with empidoids, but these flies generally have a larger wing cell cua that attains, or nearly attains, the wing margin and they lack predaceous modifications on either legs or mouthparts. A few platypezids (e.g., Microsania Zetterstedt) can also be confused with empidoids, but the antennal arista of these small flies is comprised of three articles, rather than two articles (as in the stylus of most empidoids), and the acrostichal setae of the thorax are uniserial, rather than biserial or absent.
Biology of the Empidoidea
The Empidoidea (exclusive of Dolichopodidae sensu stricto) represent a large group of mostly predaceous flies. Adults are often found in various riparian forest habitats, on leaves, tree trunks, aquatic vegetation, or in stream beds and seepage habitats, although some taxa are associated with more open areas such as fields, marshes, coastal zones and beaches (Collin 1961; Cumming & Sinclair 2009). Adults capture various arthropod prey, including small to medium sized Diptera, Hemiptera, Thysanoptera, Lepidoptera, Trichoptera, Hymenoptera, Neuroptera, Ephemeroptera, Plecoptera, Coleoptera, Collembola, and Acari (Tuomikoski 1952; Smith 1969; Cumming & Sinclair 2008).
A number of genera visit flowers as adults, presumably to obtain nectar, but at least a few groups (e.g., Anthalia Zetterstedt, Euthyneura Macquart, Iteaphila Zetterstedt, Schistostoma Becker) meet their protein requirements by feeding on pollen (Downes & Smith 1969; Chvála 1983; Sinclair & Shamshev 2012, 2021; Brooks & Cumming 2022). Many empidoids are important pollinators, especially during the spring, in high altitude ecological zones and higher latitudes (Kevan 1972; Lefebvre et al. 2014; Sinclair & Shamshev 2021). Larvae are generally found in moist soil, rotten wood, dung, or in aquatic habitats (Cumming & Sinclair 2009; Cumming et al. 2018) and appear to be predaceous on various arthropods, particularly other Diptera larvae (Cumming & Cooper 1993).
Many species mate on the ground or on vegetation, whereas others gather in mating swarms (Downes 1969; Chvála 1976, 1983). The synchronized movement of adult flies within these mating swarms is the basis for the common name “dance flies” that is sometimes used for the entire group. Members of one large subfamily, the Empidinae, transfer nuptial gifts from male to female during courtship and mating (Cumming 1994). Depending on the species, these nuptial gifts include prey, various types of inedible objects, or secreted balloons (Sinclair et al. 2013). Within the Empidinae, mate choice is generally performed by females that visit male-dominated swarms. However, many species exhibit sex-role reversed courtship behaviour where females gather in swarms to await males that choose mates. These species exhibit many female secondary sexual characters used in courting males, such as enlarged wings, pinnate leg scales, and eversible abdominal pleural sacs (Cumming 1994).
Material & Methods
Specimens and photography
Pinned specimens were selected from the Canadian National Collection of Insects, Arachnids and Nematodes (CNC), Ottawa, Canada and photographed with a Leica camera model DFC5400 using Leica Application Suite X, except as noted in the key and Acknowledgements. Additional photos of live empidoid flies were provided by the photographers and used with permission, as noted in the figure captions and Acknowledgements.
Specimen preparation
Adult empidoids (especially smaller specimens) are best collected directly into 75% ethanol and later critical-point-dried. This prevents shrinkage and distortion often seen in specimens that are mounted directly and allowed to air-dry. Larger specimens can also be mounted on pins directly or preserved in paper triangles, which helps to preserve flat and straight wings. Dried specimens are best glued directly to pins, which allows for easy manipulation and study. Terminalia can be cleared in hot 85% lactic acid, using either a hot plate or microwave oven.
Character selection
The text and character selection is modified from Steyskal & Knutson (1981) and Cumming & Sinclair (2009), with input from Collin (1961). Terms used in the key follow those described in Cumming & Wood (2017).
How to use the key
In the following key to genera, photos of each described character are provided below each couplet. Arrows and labels point out or highlight key features described in the couplet text. Hyperlinks advance the user to the next couplet and to each genus synopsis page. Each couplet has a “back button” (blue arrow at top left corner) which allows the user to return to the previous couplet. Hyperlinks are also included on each genus synopsis page to allow the user to return to the couplet where the genus is identified.
Checklist
Table 1. Classification and number of species of Nearctic Empidoidea.
| Family | Subfamily | Genus | Author, Year | No. of Species | |
|---|---|---|---|---|---|
| Iteaphilidae | |||||
| Iteaphila | Zetterstedt, 1838 |
35 |
|||
| Empididae | |||||
| Brochellinae | Brochella | Melander, 1928 | 1 | ||
| Clinocerinae | Asymphyloptera | Collin, 1933 | 2 | ||
| Clinocera | Meigen, 1803 | 42 | |||
| Dolichocephala | Macquart, 1823 | 7 | |||
| Oreothalia | Melander, 1902 | 5 | |||
| Proclinopyga | Melander, 1928 | 6 | |||
| Roederiodes | Coquillett, 1901 | 8 | |||
| Trichoclinocera | Collin, 1941 | 16 | |||
| Wiedemannia | Zetterstedt, 1838 | 7 | |||
| Empidinae | Empis | Linnaeus, 1758 | 91 | ||
| Hesperempis | Melander, 1906 | 5 | |||
| Hilara | Meigen, 1822 | 43 | |||
| Lamprempis | Wheeler & Melander, 1901 | undescribed (1) | |||
| Philetus | Melander, 1928 | 3 | |||
| Porphyrochroa | Melander, 1928 | undescribed (2) | |||
| Rhamphomyia | Meigen, 1822 | 205 | |||
| Hemerodromiinae | Chelifera | Macquart, 1823 | 22 | ||
| Chelipoda | Macquart, 1823 | 7 | |||
| Hemerodromia | Meigen, 1822 | 22 | |||
| Metachela | Coquillett, 1903 | 3 | |||
| Neoplasta | Coquillett, 1895 | 12 | |||
| Oreogetoninae | Oreogeton | Schiner, 1860 | 8 | ||
| Ragadinae | Hormopeza | Zetterstedt, 1838 | 7 | ||
| Ragas | Walker, 1837 | 4 | |||
| Zanclotus | Wilder, 1982 | 2 | |||
| Atelestidae | |||||
| Atelestinae | Meghyperus | Loew, 1850 |
2 |
||
| Brachystomatidae | |||||
| Brachystomatinae | Anomalempis | Melander, 1928 | 2 | ||
| Brachystoma | Meigen, 1822 | 4 | |||
| Trichopezinae | Apalocnemis | Philippi, 1865 | undescribed (1) | ||
| Boreodromia | Coquillett, 1903 | 1 | |||
| Ceratempis | Melander, 1928 | 1 | |||
| Gloma | Meigen, 1822 | 3 | |||
| Heleodromia | Haliday, 1833 | 6 | |||
| Niphogenia | Melander, 1928 | 2 | |||
| Sabroskyella | Wilder, 1982 | 1 | |||
| Saigusamyia | Sinclair, 2021 | 3 | |||
| Undescribed genus A | 2 | ||||
| Undescribed genus B | undescribed (1) | ||||
| Hybotidae | |||||
| Bicellariinae | Bicellaria | Macquart, 1823 |
12 |
||
| Hoplocyrtoma | Melander, 1928 | 2 | |||
| Hybotinae | Euhybus | Coquillett, 1895 | 13 | ||
| Hybos | Meigen, 1803 | 1 | |||
| Syndyas | Loew, 1857 | 6 | |||
| Syneches | Walker, 1852 | 11 | |||
| Ocydromiinae | Leptopeza | Macquart, 1828 | 5 | ||
| Ocydromia | Meigen, 1820 | 1 | |||
| Oedaleinae | Allanthalia | Melander, 1928 | undescribed (1) | ||
| Anthalia | Zetterstedt, 1838 | 12 | |||
| Euthyneura1 | Macquart, 1836 | 6 | |||
| Oedalea | Meigen, 1820 | 4 | |||
| Tachydromiinae | Allodromia | Smith, 1962 | 1 | ||
| Baeodromia | Cumming, 2007 | 1 | |||
| Chersodromia | Walker, 1851 | 8 | |||
| Crossopalpus | Bigot, 1857 | 15 | |||
| Drapetis | Meigen, 1822 | 18 | |||
| Elaphropeza | Macquart, 1828 | 2 | |||
| Megagrapha | Melander, 1928 | 3 | |||
| Micrempis | Melander, 1928 | 10 | |||
| Platypalpus | Macquart, 1828 | 117 | |||
| Stilpon | Loew, 1859 | 13 | |||
| Symballophthalmus | Becker, 1889 | 1 | |||
| Tachydromia | Meigen, 1803 | 15 | |||
| Tachyempis | Melander, 1928 | 6 | |||
| Tachypeza | Meigen, 1830 | 18 | |||
| Trichininae | Trichina | Meigen, 1830 | 1 | ||
| Trichinomyia | Tuomikoski, 1959 | 5 | |||
| Dolichopodidae sensu lato | |||||
| Microphorinae | Microphor | Macquart, 1828 | 5 | ||
| Schistostoma | Becker, 1902 | 28 | |||
| Parathalassiinae | Microphorella | Becker, 1909 | 10 | ||
| Parathalassius | Mik, 1891 | 12 | |||
| Thalassophorus | Saigusa, 1986 | 1 | |||
| Dolichopodidae sensu stricto | 1301 | ||||
| Total | 2254 | ||||
1 Of two original spellings (Euthyneura and Euthinevra) in Macquart (1836), Euthinevra was chosen by Macquart via Article 24.2.4 of the Code (International Commission on Zoological Nomenclature 1999) (see Evenhuis et al. 2016), but Euthyneura is used here as it is in prevailing usage in compliance with Article 33.3.1.
Taxonomic Changes
Table 2. Summary of taxonomic changes to the North American genera of Empidoidea since Steyskal and Knutson (1981).
| Genera added to North American fauna | |
| Allodromia Smith, 1962 | Micrempis testacea Melander, 1928 transferred by Chillcott & Teskey (1983) |
| Asymphyloptera Collin, 1933 | species collected in 1980s and 1990s and described (Sinclair 2015) |
| Baeodromia Cumming, 2007 | genus described for Drapetis pleuritica Melander, 1928 |
| Crossopalpus Bigot, 1857 | previously a subgenus of Drapetis Meigen, 1822 |
| Elaphropeza Macquart, 1828 | previously a subgenus of Drapetis Meigen, 1822 |
| Lamprempis Wheeler & Melander, 1901 | one undescribed species first collected in 1990s |
| Microphor Macquart, 1828 | change in spelling for Microphorus |
| Porphyrochroa Melander, 1928 | two undescribed species recently identified |
| Sabroskyella Wilder, 1982 | genus and species described |
| Saigusamyia Sinclair, 2021 | genus and species described |
| Schistostoma Becker 1902 | genus newly recorded in North America |
| Thalassophorus Saigusa, 1986 | species described (Brooks & Cumming 2011) |
| Trichinomyia Tuomikoski, 1959 | genus newly recorded in North America |
| Zanclotus Wilder, 1982 | genus and species described |
| Undescribed genus A | two described species currently in Apalocnemis |
| Undescribed genus B | one undescribed species identified |
| Genera removed from North American fauna | |
| Anthepiscopus Becker, 1891 | junior synonym of Iteaphila (Sinclair & Shamshev 2021) |
| Charadrodromia Melander, 1928 | junior synonym of Platypalpus (Shamshev & Grootaert 2012) |
| Phyllodromia Zetterstedt, 1837 | P. americana Melander, 1947 transferred to Chelipoda (MacDonald 1993) |
| Thanategia Melander, 1928 | junior synonym of Chelifera (MacDonald 1988) |
| Toreus Melander, 1906 | junior synonym of Hesperempis (Cumming et al. 2014) |
Key to Genera
Acknowledgements
This project would not have been possible without the vast Empidoidea holdings of the CNC. Past curators at the CNC, especially Jim Chillcott, Bill Mason, Guy Shewell, Dick Vockeroth and Monty Wood helped the collection grow to one of the largest in the world. In more recent years, the authors’ efforts to build the Empidoidea collection were assisted by Art Borkent (Salmon Arm, BC), Jeff Skevington (Ottawa, ON), Owen Lonsdale, Lubomir Masner and Jim O'Hara (CNC). We thank Michelle Locke and Gil Miranda (CNC) for comments on earlier drafts and for testing the key. Neal Evenhuis (Honolulu, Hawaii) provided nomenclatural advice. Alice Abela (Santa Maria, California), John Alcock (Tempe, Arizona), Christina Butler (Canton, Georgia), Allen Coovert (Laurellville, Ohio), Steve Marshall (Guelph, Ontario), Tom Murray (Groton, Massachusetts), John van der Linden (Decorah, Iowa), Gerrit Öhm (Göttingen, Germany), Dave Patton (Lafayette, Louisiana), Carlos Pradera (Barcelona, Spain), Masahiko Satô (Rishiri, Japan), Nikita Vikhrev (Moscow, Russia) and Nikolai Vladimirov (Moscow, Russia) kindly provided images of live, in situ flies; Justin Runyon (U.S. Forest Service, Bozeman, Montana, USA) provided images of a reduced-wing Tachyempis. Justin Runyon, Steve Paiero (University of Guelph), Karen Mclachlan Hamilton (Canadian Food Inspection Agency), Graham Thurston (CFIA), Brittany Wingert and Shawn Abraham (University of Alberta) and Owen Lonsdale kindly commented on the manuscript.
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