Eucoilinae, Thomson, 1862

Buffington, Matthew L., Forshage, Mattias, Liljeblad, Johan, Tang, Chang-Ti & Noort, Simon van, 2020, World Cynipoidea (Hymenoptera): A Key to Higher- Level Groups, Insect Systematics and Diversity 4 (2020), No. 1, pp. 1-69 : 56-59

Eucoilinae

Figs. 253–258

Within the Figitidae , the vast majority of both species diversity, and abundance, occurs within Eucoilinae . Eucoilines can be collected very easily on all continents (they are even found on Antarctic islands), and many species do very well in the suburban to urban environments, as well as around farms. Unlike most other cynipoids, the eucoilines are immediately recognizable by a single morphological feature: the scutellar plate. This feature is a structure holding up a glandular release pit the function of which is currently unknown. The feature is often referred to as a cup, a plate, a teardrop, or a disk. Because

of their commonness, and being immediately recognizable from all other cynipoids, many species have been more or less haphazardly described. As a result, this large group became an impenetrable taxonomic morass for decades until Nordlander’s work in the late 70s and early 80s began to make some sense of the diversity. Nordlander (1982b) summarized his work and generated generic groups that remained relevant well into the 2000s. Fontal-Cazalla et al. (2002) ignited renewed interest in the phylogeny of the group, and set the stage for an expanded analysis at the core of Buffington et al. (2007). The resulting phylogenies and recognition of phylogenetically informative characters have helped motivate addressing the taxonomy of larger groups of eucoilines, including the Diglyphosematini (Buffington 2011) , Zaeucoilini (Buffington 2009) and Eucoilini ( Forshage 2009) . The most comprehensive regional treatment of the Eucoilinae was published by van Noort et al. (2015) and establishes a format for future projects on eucoilines at other regional scales. In order to make sense of the genera within the group, tribes have recently been established. However, this is very much a work in progress and many genera currently lack tribal placement.

In all regions, the majority of species remain undescribed, and the described species are very often in completely wrong genera (due to the mentioned earlier lack of knowledge of phylogenetically informative characters). The latter problem (but not the former) has been addressed and largely rectified for some regions (Europe, North America, the Afrotropics) but remains at large elsewhere (the Oriental, Oceanic, and Neotropical regions all have a majority of described species still misplaced).

Eucoilines are parasitoids of cyclorraphous flies ( Buffington et al. 2012), with most host associations still unknown but spanning over a wide diversity of flies ( Ronquist 1999, Buffington 2007 , Buffington et al. 2012). Drosophila parasitoids in the genera Ganaspis and Leptopilina have been used in lab studies since the 1960s. Their biology has thus been studied in remarkable detail, and they are currently being considered for use in the biocontrol of Drosophila suzukii (“SWD”). Other eucoilines that have been used in the biological control of pest flies include: Aganaspis species on tephritids; Trybliographa species on onion maggot; Banacuniculus ; and Ganaspidium species on leafminers.

Biology. Koinobiont endoparasitoids of cyclorrhaphous flies. Early instar maggots are parasitized; and then after the host fly forms a puparium, the wasp kills the host, and completes its own pupation within the host puparium. Abe (2009) documented Gronotoma micromorpha as an egg-larval parasitoid of Liriomyza trifolii ; it is not know how widespread this type of biology is among Eucoilinae . Hosts are unknown for most species, and the records we have are very often anecdotal, but several preliminary patterns can be observed. First, that almost all reliable host records are indeed of muscomorphan (cyclorrhaphous) flies; some exceptional records of Kleidotoma on Sciaridae appears to us to be correct, whereas numerous, unisolated host records from Mycetophilidae are probably all erroneous. Second, that probably at least half of the Eucoilinae species attack saprophagous flies in more or less ephemeral habitats (dung, carrion, compost, debris, fermenting fruit and mushrooms) whereas another good portion attack phytophagous flies (leaf miners etc.). Thus, Diglyphosematini and Zaeucoilini are mostly but not exclusively on leafmining Agromyzidae , while Kleidotomini and Eucoilini are mostly but not exclusively on various saprophagous flies. Third, a “rule of thumb” that has been used among workers in the group for decades is to expect any genus of Eucoiline wasp to attack one particular family of flies. This is not valid in any strict sense but a mere pragmatic guidance, but with our limited data it works in a large number of cases. Large eucoiline genera tend to include exceptions (host switches), and two large genera ( Kleidotoma and Hexacola ) are known to have a wide range of hosts. The fly families attracting the largest number of eucoiline genera are Drosophilidae and Agromyzidae . Very little is known about host specificity of individual eucoiline species.

Distribution.Worldwide.Particularly speciose in the Neotropical Region.

Relevant literature. Weld (1952) remained dominant until the publications of Nordlander established a new standard of thoroughness and phylogenetic thinking in eucoiline research ( Nordlander, 1976, 1978, 1980, 1981, 1982a, summarized in Nordlander [1982b]). Van Lenteran et al. (1998) and van Alphen et al. (1991) investigated biology and host use. Forshage and Nordlander (2008) provided basic circumscription of tribes and keyed western Palearctic genera, Buffington revised Diglyphosematini (Buffington 2011) and the new tribe Zaeucoilini (Buffington 2009) . Forshage (2009) summarized global overview of the subfamily and especially Eucoilini . Van Noort et al. (2015) provided a substantial overview of theAfrotropical fauna,and Forshage et al.(2013) cataloged Nearctic taxa.A combination of the Afrotropical key and the European key ( Van Noort et al. 2015 and Forshage and Nordlander 2008) will allow generic recognition of most Eucoilinae worldwide, except in the utterly diverse Neotropics (cf Buffington et al. 2006) and highly aberrant Pacific islands (cf Beardsley 1989). Species-level identification is very often not possible, but many common European species can still be keyed with Quinlan (1978) even though taxonomy is obsolete, and odd taxa globally can be recognized using Weld (1952). Useful generic treatments are available for Ganaspidium ( Buffington 2010a) , Banacuniculus ( Buffington 2010b) , Zaeucoila ( Buffington et al. 2018) , European Rhoptromeris ( Nordlander 1978, Costa Baião and Forshage 2018), Leptopilina in different regions ( Nordlander 1980, Allemand et al. 2002, Novkovic et al. 2011, Lue et al. 2016), and several genera in Taiwan ( Lin 1987, 1988), as well as for several lesser, recently described genera or regional assemblies thereof, while many recent studies still await publication. Fontal-Cazalla et al. (2002) and Buffington et al. (2007) provided phylogenies.

Classification.

Eucoilinae Thomson, 1862

Diglyphosematini Belizin, 1961

Afrostilba Benoit, 1956 ; 18 species AT

Banacuniculus Buffington, 2010 ; 8 species NA, NT plus Hawaii Diglyphosema Förster, 1869 ; 7 species PA

Disorygma Förster, 1869 ; 7 species PA, OR

Ealata Quinlan, 1986 ; 5 species AT, OR

Ganaspidium Weld, 1955 ; 6 species NA, NT, AT but mainly arid North American Southwest

Gronotoma Förster, 1869 ; currently 35 species but a few more described species belong here, worldwide

Microstilba Förster, 1869 ; 6 species, wPA

Nordlanderia Quinlan, 1986 ; 4 species AT, PA Paradiglyphosema Lin, 1988 ; 3 species but at least 1 more described and some undescribed belong here, mostly OR but also AT

Sinatra Buffington, 2011 ; 1 species around the Pacific Tobiasiana Kovalev, 1979 ; 4 species arid southern Palearctic

Eucoilini Thomson, 1862

Afrodontaspis Weld, 1962 ; 2 species AT

Bothrochacis Cameron, 1904 ; 8 species currently in genus but a few more belong here, mostly AT but also OR and Hawaii Eucoila Westwood, 1833 ; only 3 described species currently are classified as Eucoila in a meaningful sense, while many need to be removed elsewhere and yet a few others need to be moved in or described as new, PA, NAPA, NA

Leptopilina Förster, 1869 ; 41 described species currently in the genus in a meaningful sense but more are currently being described and still ca 12 need to be moved in from other genera, worldwide

Linaspis Lin, 1988 ; 1 species ePA

Linoeucoila Lin, 1988 ; 11 species, OR but undescribed species also AT

Maacynips Yoshimoto, 1963 ; 3 described species and numerous undescribed in Australia and throughout the Pacific and East Asia

Quasimodoana Forshage, Nordlander & Ronquist, 2008 ; 2 species PA, NA

Trybliographa Förster, 1869 ; 43 described species currently in the genus in a meaningful sense but some 20 more need to be moved in from other genera and far more described as new, worldwide but mainly Holarctic

Ganaspini Belizin, 1961

Acantheucoela Ashmead, 1900 ; 6 species NT

Aganaspis Lin, 1987 ; 7 described species currently in the genus in a meaningful sense but ca 10 more need to be moved in and more described as new; worldwide but mainly Oriental and Neotropic

Areaspis Lin, 1988; 2 species but 2 more need to be moved in and additional ones described as new, OR, AT

Aspidogyrus Yoshimoto, 1962 ; 4 species Hawaii

Caleucoela Kieffer, 1909 ; 1 species NT

Chrestosema Förster, 1869 ; 3 described species currently in the genus in a meaningful sense but more will soon be moved in, and described as new, while remaining others will be moved out; mainly OR, PA

Coneucoela Kieffer, 1909 ; 1 species NT

Didyctium Riley, 1879 ; 12 described species currently in the genus in a meaningful sense but ca 10 need to be moved in and many more described as new; worldwide

Dieucoila Ashmead, 1903 ; 7 described species currently in the genus in a meaningful sense but ca 10 need to be moved in and more described as new; NT, NA

Discaspis Lin, 1988 ; 1 species OR

Ditanyomeria Yoshimoto, 1963 ; 4 nominal species AU, to be synonymized

Endecameris Yoshimoto, 1963 ; currently 2 species but many undescribed, PA, OR, AT, AU

Epicoela Borgmeier, 1935 ; 2 species NT

Epochresta Lin, 1988 ; 1 species OR

Euxestophaga Gallardo, 2017 ; 1 species NT

Fontaliella Pujade-Villar, 2013 ; 1 species NT

Ganaspis Förster, 1869 ; 25 described species currently in the genus in a meaningful sense, but ca. 40 more need to be moved in and yet more described as new; worldwide Gastraspis Lin, 1988 ; 2 species OR, AT

Glauraspidia Thomson, 1862 ; 3 described species currently in the genus in a meaningful sense, but a few more are being moved in or described as new; PA, rare in NA, NT Hexacola Förster, 1869 ; 43 described species currently in the genus in a meaningful sense but ca 25 more need to be moved in and many more described as new; worldwide Humboldteria Buffington 2017 ; 4 species NT Hydrelliaeucoila Díaz & Gallardo, 2009 ; 1 species NT Hypodiranchis Ashmead, 1901 ; 9 described species currently in the genus in a meaningful sense but a few more need to be moved in or described as new: Pacific and East Asian Lispothyreus Yoshimoto, 1962 ; 2 species Hawaii Mirandicola Belizin, 1968 ; 8 described species currently in the genus in a meaningful sense but some more are currently being described and many remain undescribed, OR, PA Nesodiranchis Perkins, 1910 ; 6 species Hawaii Nordlandiella Díaz, 1982 ; 2 species but 3 more need to be moved in and some described as new; NT, NA Odonteucoila Ashmead, 1903 ; 8 species NT

Odontosema Kieffer, 1909 ; 1 species NT

Paraganaspis Díaz & Gallardo, 1996 ; 2 species but 6 more need to be moved in and some described as new; NT, NA Pentamerocera Ashmead, 1896 ; 1 species but very poorly known; NT

Perischus Weld, 1931 ; 2 species NT

Promiomera Ashmead, 1903 ; 1 species NT

Pressia Belizin, 1968 ; 1 nominal species PA, to be synonymized Pseudodiranchis Yoshimoto, 1962 ; 10 nominal species, but poorly known, real species number might be smaller or larger; Hawaii

Sinochresta Lin, 1988 ; 2 species OR

Steleucoela Kieffer, 1908 ; 2 species NT

Striatovertex Schick, Forshage & Nordlander, 2011 ; 13 species but some are synonyms NT, NA and Hawaii Trissodontaspis Ashmead, 1903 ; 1 species NT

Weldia Yoshimoto, 1962 ; 6 species, but poorly known, real species number might be smaller or larger; Hawaii Zamischus Ashmead, 1903 ; 3 species NT

Kleidotomini Hellén, 1960

Cothonaspis Hartig, 1840 ; 7 species, mainly Holarctic and AT, as well as a cosmopolitan species

Eutrias Förster, 1869 ; 1 species, Holarctic

Garudella Buffington & Forshage, 2014 ; 4 species, OR, AT Kleidotoma Westwood, 1833 ; 137 species worldwide, but by far most are still undescribed

Muhaka Buffington & Copeland, 2015 ; 1 species AT Triplasta Kieffer, 1901 ; 4 species NT

Trichoplastini Kovalev, 1989

Angustocorpa Quinlan, 1988; 4 species AT

Nanocthulhu Buffington, 2012 ; 1 species AT Nordlanderiana Kovalev, 1989 ; 1 species PA

Rhoptromeris Förster, 1869 ; 46 described species currently in the genus in a meaningful sense, but ca. 10 more need to be moved in and several more described as new, worldwide but mainly AT & PA

Stentorceps Quinlan, 1984 ; 6 species AT

Trichoplasta Benoit, 1956 ; 27 described species currently in the genus in a meaningful sense, but a few need to be moved in and several more described as new, worldwide but mainly AT, PA, NA

Zaeucoilini Buffington, 2009

Dettmeria Borgmeier, 1935 ; 2 species NT

Dicerataspis Ashmead, 1896 ; 2 species NT

Lopheucoila Weld, 1951 ; 2 species NT

Marthiella Buffington, 2009 ; 2 species NT

Moneucoela Kieffer, 1907 ; 2 species NT

Moritiella Buffington, 2006 ; 2 species NT

Paradettmeria Gallardo & Díaz, 2011 ; 1 species NT Penteucoila Weld, 1951 ; 1 species NT

Preseucoela Buffington, 2004 ; 3 species NT, NA Rhabdeucoela Kieffer, 1907 ; 7 species NT

Tropideucoila Ashmead, 1903 ; 9 species NT

Zaeucoila Ashmead, 1903 ; 14 species NT, NA

‘ Leptolamina group’, not currently assigned to a tribe Leptolamina Yoshimoto ; 16 species and more undescribed; OR, AU, AT, PA

Micreriodes Yoshimoto, 1962 ; 1 species currently in genus, a few need to be moved in and some more described as new; mainly PA, OR, AU but single specimens found in AT, NA

Unplaced Nomina inquierenda

Delomeris Diaz & Gallardo, 1996 ; 1 species NT Unplacable Nomina dubia

Macrocereucoila Ashmead, 1887 ; 1 species

Tetraplasta Ashmead, 1903 ; 1 species

Figures

Plate 21. Eucoilinae. Fig. 253, Ganaspis brasiliensis (Ihering, 1905), USNMENT01520001. Fig. 254, Odonteucoila sp., USNMENT01231882. Fig. 255, Kleidotoma sp., USNMENT01525865. Fig. 256, Gronotoma sp., USNMENT01231843. Fig. 257, Trybliographa melanoptera, USNMENT01231838. Fig. 258, Striatovertex sp., USNMENT01231830.