ISSN 0013-8738, Entomological Review, 2014, Vol. 94, No. 8, pp. 1131–1190. © Pleiades Publishing, Inc., 2014. Original Russian Text © Z.A. Fedotova, 2014, published in Entomologicheskoe Obozrenie, 2014, Vol. 93, No. 3, pp. 666–739.
Classification of Gall Midges of the Supertribe Brachineuridi (Diptera, Cecidomyiidae: Lasiopterinae) with Description of New Genera and New Species from the Kurile Islands Z. A. Fedotova All-Russia Scientific-Research Institute of Plant Protection, Pushkin, St. Petersburg, Russia e-mail:
[email protected],
[email protected] Received March 8, 2014
Abstract—A new classification is proposed for the gall-midge supertribe Brachineuridi including 4 tribes, 9 subtribes, 24 genera, and 156 species: Brachineurini (Brachineurina, 4 genera, 23 species; Epimyiina, 4 genera, 9 species); Undoneurini trib. n. (Undoneurina subtrib. n., 5 genera, 8 species; Coccidomyiina subtrib. n., 2 genera, 8 species); Rhizomyiini stat. n. (Rhizomyiina, 4 genera, 33 species; Stellaserenina subtrib. n., 3 genera, 3 species); Ledomyiini (Ledomyiina, 8 genera, 61 species; Acinacistylina subtrib. n., 2 genera, 5 species; Kovaleviolina subtrib. n., 2 genera, 3 species), and also 2 genera Palaeospaniocera Meunier, 1901 (1 species) and Ledomyiella Meunier, 1904 (4) with an unclear systematic position. One new tribe, 5 subtribes, 3 genera, and 7 species (Stellaserena vaga gen. et sp. n., Stelladiurna flustra gen. et sp. n., Kuriloneura multifida gen. et sp. n., K. trifida sp. n., Cornistyla tufa sp. n., Rhizomyia flammula sp. n., Acinacistyla sundukovi sp. n.) are described. Keys to the tribes, subtribes, genera, and species of Brachineuridi are given. The genera Ramineura stat. n. and Cornistyla stat. n. are promoted from subgenera of Brachineura. A new synonymy is established: the genus Pararhizomyia Mamaev, 1998, stat. n. (= Effusomyia Fedotova, 2004, syn. n.). New combinations are formed: Ramineura ussurica (Fedotova, 2004), comb. n. (Brachineura), R. musiva (Fedotova et Sidorenko, 2006a), comb. n. (Brachineura), R. subulata (Fedotova et Sidorenko), comb. n. (Brachineura), and R. alia (Mamaev, 1994), comb. n. (Brachineura); Cornistyla nijveldti (Fedotova, 2004), comb. n. (Brachineura), C. flabellata Fedotova et Sidorenko, 2006a, comb. n. (Brachineura), C. infirma (Fedotova et Sidorenko, 2006) (Brachineura); Cingola pleiomorpha Mamaev, 1998, comb. n. (Brachyneurina), Stabiliola arsenjevi (Fedotova, 2004), comb. n. (Brachyneurina); S. umbra (Fedotova et Sidorenko, 2006b), comb. n. (Rhizomyia), Alatostyla binaria (Mamaev, 1998), comb. n. (Rhizomyia), Ledomyia eocenica (Meunier, 1904), comb. n. (Ledomyiella), Kovaleviola pyxidiiformis (Fedotova, 2005), comb. n. (Brachyneurina). DOI: 10.1134/S0013873814080107
The present paper continues the series of publications dealing with the study of the gall-midge supertribe Brachineuridi of the subfamily Lasiopterinae (Fedotova, 2004; Fedotova and Sidorenko, 2005, 2006a, 2006b, 2007). Mamaev (1968) considered the subtribes Brachineurina, Rhizomyiina, and Epimyiina within the tribe Oligotrophini of the subfamily Cecidomyiinae. Later (Fedotova, 2000), these subtribes were raised to tribes and, together with the closely related tribe Ledomyiini within the supertribe Oligotrophidi, were transferred to the subfamily Lasiopterinae. Gagné (2004, 2010) did not divide the supertribe Brachineuridi into tribes and placed part of its genera in the tribe Ledomyiini of the supertribe Lasiopteridi which, together with Brachineurini, was attributed later to the supertribe Brachineuridi of the subfamily Cecidomyiinae (Gagné and Jaschhof, 2014).
Some years after the publication by Krivosheina and Zaitsev (1989), in which the families Cecidomyiidae and Lestremiidae were combined again in the superfamily Cecidomyioidea, the author proposed a supplemented classification of the gall midges (Fedotova, 2000, 2013), where Cecidomyiidae was subdivided into the subfamilies Porricondylinae, Lasiopterinae, and Cecidomyiinae; the former tribes were raised to supertribes, and the supertribes, to subfamilies. The supertribes Lasiopteridi and Oligotrophidi were combined into the subfamily Lasiopterinae, and the supertribe Cecidomyiidi became a subfamily. The establishment of the subfamily Lasiopterinae including two supertribes was caused by the presence of the plesiomorphic characters common to these supertribes (a very dense cover of scales over the body, legs, and occasionally antennae; a compact body with short
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legs; the flagellar segments without necks or with basal enlargements; an unequal number of the flagellar segments in the male and the female; a poor development of loop-shaped threads on the flagellar segments of the male and female; the presence of a basal projection on the gonocoxite; and completely developed tergites and abdominal sternites), in contrast to the other plesiomorphic states of the characters of Cecidomyiinae (an equal number of the flagellar segments in both sexes; vein R4+5 running into or at the apex of the wing; the gonostylus almost entirely covered with microtrichiae; and the usually short non-telescopic ovipositor bearing large unfused apical plates) (Fedotova, 2000). Following Mamaev’s (1968) classification (with little changes), the author included the closely related tribes Brachineurini, Ledomyiini, Rhizomyiini, and Epimyiini in the supertribe Oligotrophidi of the subfamily Lasiopterinae, together with the 8 other tribes. It was found later that only these 4 tribes conformed to the characters of Brachineuridi sensu lato (Fedotova, 2014), and preliminary classification of the supertribe Brachineuridi of the world fauna was given, according to which Brachineuridi included the subtribes considered by Mamaev (1968) as tribes. However, this number of suprageneric taxa was insufficient to reflect the whole diversity of the genera included now in the supertribe Brachineuridi (Gagné and Jaschhof, 2014; Fedotova and Perkovsky, 2014; Fedotova, 2014) for which the classification remained undeveloped below the rank of a tribe. After the publication of a number of revisions of separate gall-midge groups of the supertribe Oligotrophidi, the taxonomic position of some supertribes was clarified. The previously established relationship between the supertribes Stomatosematidi (Gagné, 1975) and Cecidomyiidi (now the subfamily Cecidomyiinae) was confirmed by the results of the revision of the world fauna (Fedotova, 2011a, 2011b). The supertribes Lasiopteridi, Oligotrophidi, and Brachineuridi (selected from Oligotrophidi) were attributed to the subfamily Lasiopterinae. In the present publication, the classification of the supertribe Brachineuridi, implying its division into 4 tribes, 9 subtribes, 24 genera, and 156 species, is proposed for the first time. The taxonomic positions of two genera, Palaeospaniocera Meunier, 1901 (1 species) and Ledomyiella Meunier, 1904 (4 species), remain unclear.
MATERIALS AND METHODS The study is based on examination of the material collected using various methods in the south of the Russian Far East by V.S. Sidorenko (1965–2010) from the Lazovskii and Ussuri Nature Reserves in 2001– 2007, by the present author, near the Ussuri Nature Reserve (near Kamenushka Vill.) in 2001, and by Yu.N. Sundukov, from the southern Kuriles (Shikotan Island) in 2012. The adults were collected using traps (mainly light and Malaise traps) and fixed in 70% ethyl alcohol. The constant slides were made after keeping of the adults in 96% alcohol and then in oil cloves for removal of the residual water from the body and for clarification of the cuticle. The specimens were placed in Canadian balsam dissolved in xylene on a subject glass and covered with a glass. Some adult specimens were dissected; their separated heads, wings, and genitalia were placed in the same slide under separate small covering glasses. In addition to the standard measurements of the length of the body, antennae, wings, and all the legs, the characters usually used for a species diagnostics (the length to width ratio of the wing, the 2nd to 1st tarsal segment length ratio, the length to width ratio of the 5th flagellar segment, and the 1st to 2nd flagellar segment length ratio) are included in the descriptions of the new taxa; in addition, the antenna to wing length ratio, the wing to vein R1+2 length ratio, the body to femur length ratio, etc. (see table) are also used. For comparison of the biometric parameters of the adults belonging to different subtribes of Brachineuridi, the table is supplemented with the measurements of some species described earlier. The holotypes and paratypes of the new species are deposited in the collections of the Zoological Institute, the Russian Academy of Sciences, St. Petersburg (ZIN). The material from Rovno amber, including the types, is deposited in the collections of Schmalhauzen Institute of Zoology, the National Academy of Sciences of Ukraine, Kiev. The History of the Study of Brachineuridi Based on an integrated assessment of the characters (the structure of the legs, the venation of the wing, and the morphology of the antennal segments), Kieffer (1913) proposed the classification of the gall-midge family, according to which the subfamily CecidomyiiENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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nae included a combined group in the rank of the tribe Brachyneurariae with 22 genera. Felt (1925) did not accept this tribe and included its genera mainly in the tribe Oligotrophariae, in the additionally distinguished tribe Dasineurariae, and partly in the subfamily Heteropezinae. The grounding of the modern classification of the gall midges was laid by Rübsaamen and Hedicke (1926–1939) who, sharing on the whole Kieffer’s (1913) viewpoint on the composition of the tribe, distinguished 4 supertribes, Cecidomyiidi, Asphondyliidi, Oligotrophidi, and Lasiopteridi, and also included a tribe with the changed name Brachyneurini in Oligotrophidi. This changed name was first mentioned by Enderlein (1920) who later established within the supertribe Oligotrophidi the accessory tribe Ledomyiini Enderlein, 1936, including there 8 genera (Cryptolauthia Kieffer, 1912; Coccomorpha Rübsaamen, 1899; Lauthia Kieffer, 1904; Ledomyia Kieffer, 1895; Mikiola Kieffer, 1896; Phaenolauthia Kieffer, 1904; Rhizomyia Kieffer, 1898; and Stomatosema Kieffer, 1904) and attributing 4 genera to the tribe Brachineurini: Brachineura Rondani, 1840; Brachyneurella Kieffer, 1913; Acriectasis Rübsaamen, 1910; and Spaniocera Winnertz, 1853. The three latter names were soon (Edwards, 1937) reduced to the synonyms of Brachineura. Further Mamaev (1967) regarded the Rübsaamen and Hedicke’s supertribes Cecidomyiidi, Asphondyliidi, Oligotrophidi, and Lasiopteridi as tribes and combined Brachineurini with the other Oligotrophini, including the genera of the tribe Ledomyiini. In this and the next paper (Mamaev, 1968), this author did not mention the tribe Ledomyiini not did he refer to Enderlein’s (1936) publication, in which Ledomyiini and Brachineurini were attributed to the supertribe Oligotrophidi. Mamaev (1967) initially considered artificial the erection of the tribe Brachineurini differing from the typical Oligotrophini only in characters of the structure of the ovipositor plates. Later, Mamaev (1968) proposed the advanced classification of the gall midges, in which he distinguished within the tribe Oligotrophini the subtribe Brachineurina (with the only genus Brachineura) and the subtribes Rhizomyiina, Stomatosemina, and Epimyiina first established by him; he also gave diagnoses of the subtribes Brachineurina and Rhizomyiina and included only one genus in each subtribe, without listing the closely related ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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genera considered by him earlier in the review of the tribe Oligotrophini (Mamaev, 1967). The absence of phytophages among the representatives of these subtribes, along with similarity of their morphological characters, was considered an indirect evidence of the existence of relationships between these subtribes (Mamaev, 1967, 1968). Soon Gagné (1976) compiled a list of 16 Nearctic species of the genus Ledomyia, mentioned Phenolauthia and Neuromyia Felt, 1911 among its synonyms, and reduced the supertribes Oligotrophidi and Lasiopteridi to tribes. Then he analyzed the phylogenetic relationships between the tribes Ledomyiini, Oligotrophini, and Lasiopterini and comprised them in the supertribe Lasiopteridi. According to this classification, the tribe Brachineurini was not accepted, and its genera (Brachineura, Epimyia Felt, 1911, and Rhizomyia) were placed in the subfamily Cecidomyiinae as genera with an unclear taxonomic position. Later, Gagné (1985) came back to the discussion of the tribe Ledomyiini. One of the subtribes distinguished earlier within the tribe Oligotrophini and considered at present, similarly to Brachineuridi, a distinct supertribe, Stomatosematidi (the name is based on a corrected name of the subtribe Stomatosemina, originated from a Greek word; Gagné, 1975), belongs to the subfamily Cecidomyiinae and comprises 2 tribes, 3 subtribes, 7 genera, and 49 species, including 3 genera and 11 species from the Late Eocene Rovno and Baltic amber (Fedotova, 2011). The taxa Epimyiina and Rhizomyiina have not been mentioned in the catalogs of the gall-midge world fauna (Gagné, 2004, 2010; Gagné and Jaschhof, 2014), data on these taxa can be found only in the bibliographical tool of Sabrosky (1999). The independence of these taxa accepted here as the tribe Rhizomyiini stat. n. and the subtribe Epimyiina in the supertribe Brachineuridi is discussed below. Gagné (1989) was the first to mention Brachineuridi in the rank of a supertribe within the subfamily Cecidomyiinae, together with Cecidomyiidi, Stomatosematidi, and Lasiopteridi (also including Oligotrophini and Ledomyiini among the other tribes). The tribe Ledomyiini was included in the supertribe Lasiopteridi based on a similarity between the representatives of Lasiopterini and Oligotrophini in the structure of the genitalia characterized by the presence of a basal projection of the gonocoxite, which in Brachineuridi can
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be reduced, lost, or replaced with a sclerotized paramere. These concepts of the gall-midge classification were reflected in the first and second catalogs of the gall midges of the world fauna (Gagné, 2004, 2010). Skuhravá followed earlier classifications of the gall midges (Rübsaamen and Hedicke, 1926; Mamaev, 1968) and, in the catalog of and a key to the Palaearctic Cecidomyiidae (Skuhravá, 1986, 1997), she placed the tribe Brachineurini in the supertribe Oligotrophidi and included there 11 genera: Mikiola, Rhizomyia, Epimyiella, Lasiopteryx, Lauthia, Plesiolauthia, Ledomyia, Prolauthia, Spaniocera, Brachineura, and Brachyneurina. All the 4 species of the genus Mikiola Kieffer, 1896 are phytophagous insects causing formation of galls on the beech (Fagus spp.), which distinguishes them from the mainly mycetophagous species of the other genera (Mamaev, 1967). The tribe Mikiolini Skuhravá, 2006 attributed to the supertribe Oligotrophidi was established for the genus Mikiola. In the catalogs published later (Gagné, 2010; Gagné and Jaschhof, 2014), this taxon was not included, and the genus Mikiola was placed in the supertribe Lasiopteridi without indication of its position in any of the tribes. In the first catalog of the gall-midge world fauna (Gagné, 2004), the supertribe Brachineuridi included only 7 genera (Brachineura Rondani, 1840; Brachyneurina Mamaev, 1967; Chrybaneura Gagné, 1968; Coccidomyia Felt, 1911; Epimyia Felt, 1911; Epimyiella Mamaev, 1967; and Rhizomyia Kieffer, 1898) with 49 species. The tribe Ledomyiini, closely related to Brachineuridi, included 6 genera (Isogynandromyia Spungis, 1981; Lauthia Kieffer, 1912; Ledomyia Kieffer, 1895; Ledomyiella Meunier, 1904; Plesiolauthia Mamaev, 1967; Prolauthia Rübsaamen, 1916) with 53 species and was attributed to the supertribe Lasiopteridi (Gagné, 2004, 2010). Later, Skuhravá (2006) shared on the whole Gagné’s (2004) concept of the scope of the tribe Ledomyiini, but included it in the supertribe Oligotrophidi, according to the conventional classification of the gall midges (Rübsaamen and Hedicke, 1926). In the classification proposed by Mamaev (1968), the combined group in the rank of the subtribe Brachineurina sensu lato, with inclusion of the genera combined at present to the tribe Ledomyiini, also belonged to Oligotrophini (now Oligotrophidi). The list of the members of Brachineuridi of the world fauna was supplemented after description of
9 new genera, 2 subgenera, and 38 species from the Russian Far East (Fedotova, 2004; Fedotova and Sidorenko, 2005, 2006a, 2006b, 2007), and also 5 new genera and 10 new species from the Late Eocene Rovno amber (Fedotova and Perkovsky, 2005, 2008; Perkovsky and Fedotova, 2008), which was reflected in the next catalog (Gagné, 2010) listing 18 genera and 82 species within Brachineuridi and 6 genera and 64 species within Ledomyiini. In the third catalog of the gall midges of the world fauna (Gagné and Jaschhof, 2014), Brachineurini and Ledomyiini were combined to form the supertribe Brachineuridi, but the volume of this group (Brachineurini + Ledomyiini) almost did not change (24 genera with 143 species). The names of 3 species, Lauthia aurofulgens Kieffer, 1911, L. monilicornis Kieffer, 1911, and Ledomyia styloptera Kieffer, 1911, were recognized as nomina dubia, since the preserved types, mainly females, were badly damaged and could not be reliably identified with any taxon (Harris and Dorchin, 2012). The combination of these two tribes into the supertribe Brachineuridi was argued by the authors of the catalog by the presence of a short primitive ovipositor and also by the arrangement of abdominal sternite VIII behind tergite VIII. A list of the genera attributed to the tribes Brachineurini (15 genera and 79 species) and Ledomyiini (7 genera and 62 species) in this catalog requires specification because each taxon is characterized by a complex aggregate of the characters. After the publication of this catalog, 2 species of the genus Cingola Fedotova et Sidorenko, 2006 from Southern China (Jiao and Bu, 2014) and also 3 genera and 6 species from the Late Eocene Rovno amber (Fedotova and Perkovsky, 2014), belonging to Brachineuridi sensu lato, were described. This catalog, as well as the previous ones, does not mention the subtribes Rhizomyiina and Epimyiina established by Mamaev (1967, 1968). The Monophyletic Origin of Brachineuridi The monophyletic origin of the supertribe is substantiated by the distinctive morphological characters which are mosaically manifested in various groups of Brachineuridi taxa and, as a whole, clearly distinguish this supertribe from Oligotrophidi and Lasiopteridi closely related to it. Another distinctive character of the supertribe is the peculiarity of the food specialization of the mycetophagous, xylophagous species feeding on remains of insects, on films of eggs, and on exuvia or on coccid adults and eggs. Such feeding ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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allows Brachineuridi to be considered an archaic group which has been separated before the passage of the gall midges to the feeding on plants and to gallforming. The archaism of Brachineuridi, very sparsely occurring in the recent fauna, is corroborated by numerous records of these insects from the Late Eocene Baltic and Rovno amber (the age is 34 million years), where they show a wide generic and specific diversity. The fraction of Brachineuridi constitutes 11.9% of the number of the species of the superfamily Cecidomyioidea (27 out of 227) in Rovno amber and 9.0% (6 out of 67 species) in Baltic amber. Among the higher gall midges, the Cecidomyiidae proper, their fraction constitutes 20.1 (27 out of 132) and 12.5% (6 out of 48 species), respectively (Fedotova and Perkovsky, 2009). No phytophagous or gall-forming gall midges have been found in the amber examined. Comparison of Ledomyiini with Brachineurini and with the other Lasiopterinae shows that, in addition to the body entirely covered with scales, reduced antennal segments, palpus, and wing veins, and also a welldeveloped basal projection of the gonocoxite, which are characteristic of many genera of Lasiopterinae, the number of the features (mainly plesiomorphic) relating Brachineuridi to Oligotrophidi is greater than that relating it to Lasiopteridi. In Lasiopteridi, the flagellar antennal segments are secondarily deprived of necks (as the result of their considerable reduction or loss) and consist of a greater and various number of segments; this number is greater in the female than in the male; the apical flagellar segments are frequently fused (Fedotova, 2000), which is not observed in Brachineurini and Ledomyiini. The gall midges of the tribes Brachineurini and Ledomyiini are similar to those of the supertribe Oligotrophidi in reduction of the segments of the palpus and flagellum, in a shorter vein R4+5, in an equally widely varying degree of development of the basal projection of the gonocoxite: from totally absent (replaced by a small lobe at the base of the gonocoxite) to rather well developed or modified, and also in incompletely fused or unfused cerci of the ovipositor in part of the genera. The tribe Ledomyiini, judging by the plesiomorphies present in some genera (a long and fine basal projection of the gonocoxite adjoining the aedeagus, a 4-segmented palpus, an entirely developed furcation of the cubital vein, unfused apical flagellar segments, and unfused apical plates of the ovipositor), is less advanced in comparison with Brachineurini demonstrating characteristic synapomorphies (a short ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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8–12-segmented antenna, reduced palpus, and shortened vein R4+5). The genera belonging to Brachineurini and Ledomyiini frequently share some characters occurring mosaically in various genera and tribes. The structure of the male flagellar segments in the genera of these tribes usually exhibits secondary sexual characters (absent in Brachineura and Isogynandromyia). The wing venation is similar in the shape and arrangement of some veins: R4+5 is short (Chrybaneura, Ledomyia) or long (Coccidomyia, Acinacistyla), reaches the apex of the wig; vein Cu is bifurcate (Rhizomyia, Plesiolauthia) or simple (Cingola, Alatostyla); Rs is usually absent but rarely present (Acinacistyla, Rabindrodiplosis). The claws of all the tarsi are simple or each bearing a tooth at the base, or the claws of only the fore and hind tarsi are simple or each bearing a tooth at the base. The basal projection of the gonocoxite is present, absent, or appears as a sclerotized paramere (Epimyia, Volsatiola). The hypoproct is complete, concave, or emarginate. The ovipositor is short and non-telescopic nearly in all the genera, or telescopic (Ledomyia, Isogynandromyia), but always bears two cerci. The plesiomorphic state of some characters common for Ledomyiini and Brachineurini and also the abundance of fossil records of genera and species of these tribes suggest that Brachineuridi has been separated before the formation of the actually higher gall midges of the supertribe Lasiopteridi showing mainly apomorphic characters linked with the development of reduction in the structure of the antenna, palpus, and wing venation and with an extreme specialization of the structure of the ovipositor which was caused by the passage to phytophagy. The monophyletic origin of the supertribe is confirmed by evolutionary development of its characters among which we can trace a descent from the genera of the archaic tribe Ledomyiini to the more advanced Brachineurini. SUPERFAMILY CECIDOMYIOIDEA NEWMAN, 1834 FAMILY CECIDOMYIIDAE NEWMAN, 1834 Subfamily LASIOPTERINAE Rondani, 1856 Supertribe Brachineuridi Kieffer, 1913 Brachyneurariae Kieffer, 1913 : 17, 101. Brachyneurini: Enderlein, 1920 : 271. Brachineurini: Rübsaamen and Hedicke, 1926; Foote, 1965; Skuhravá, 1986, 2006; Fedotova, 2000, 2004; Fedotova and Sidorenko, 2006a, 2006b.
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Brachyneurinae: Edwards, 1937 : 149. Brachyneurina: Mamaev, 1968 : 16, 24. Brachineuridi: Gagné, 1989, 1994, 2004, 2010; Fedotova and Perkovsky, 2013; Gagné and Jaschhof, 2014. Diagnosis. Body densely covered with scales. Antenna 2+9–12-segmented. Pattern of sensory threads covering basal enlargement widely varying from one or two rings connected by longitudinal crosspieces to nets of serpentine or loop-shaped threads densely covering segment (Figs. 1, 3, 11, 18; 15, 7–9; 16, 6–9). Palpus usually reduced, 1–3-segmented. Wing venation widely varying. Vein R4+5 varying from very short, running into margin of wing far before its apex (Fig. 2, 1–9) to long, running into or at wing apex (Fig. 2, 10–12), in combination with simple cubital vein. Occasionally vein Cu forming furcation Cu1 and Cu2 (Fig. 15, 1–10); R4+5 running into margin of wing at various distance from its apex. Veins Rs (Fig. 11, 6, 7), M3+4, Cup (Fig. 11, 6), and also accessory vein originating from confluence of R1+2 into costal vein (Fig. 2, 3), which not occurring in higher gall midges of family Cecidomyiidae, rarely additionally developed. Gonocoxites separated (Fig. 1, 1, 7, 16) or fused near bases on ventral side (Fig. 11, 14, 15). Basal projection developed to various extent (Figs. 1, 7; 3, 17, 20; 15, 10) or absent (Figs. 1, 1, 16; 5, 1, 3, 12). Abdominal segments VII and VIII with very fine reduced tergites (Fig. 4, 7, 8). Ovipositor non-telescopic, with large apical cerci (Fig. 4, 1–13) and large ventral lobe on short abdominal segment VIII, or telescopic, with long abdominal segment VIII and with small cerci which always separated, without traces of fusion (Fig. 12, 25) (Ledomyia and Isogynandromyia, Fig. 17, 3). The proposed classification of the gall midges of the supertribe Brachineuridi includes 4 tribes, 9 subtribes, 24 genera, 156 species, and also 2 genera and 5 species with an unclear taxonomic position. The number of the species included in the genus is given in parentheses. Brachineurini Kieffer, 1913. Brachineurina: Brachineura Rondani, 1840 (17); Ramineura Fedotova et Sidorenko, 2006a (4); Spaniocera Winnertz, 1853 (1); Nascensluna Fedotova et Perkovsky, 2014 (1); Epimyiina Mamaev, 1968 : Epimyia Felt, 1911 (2); Epimyiella Mamaev, 1967 (1); Cornistyla Fedotova et Sidorenko, 2006a (4); Nodalistyla Fedotova et Sidorenko, 2006b (2).
Undoneurini trib. n. Undoneurina subtrib. n.: Brachyneurina Mamaev, 1967 (3); Chrybaneura Gagné, 1968 (1); Cingola Fedotova et Sidorenko, 2006a (4); Popovineura Fedotova et Perkovsky, 2014 (1); Undoneura Fedotova et Sidorenko, 2006a (2); Coccidomyiina subtrib. n.: Coccidomyia Felt, 1911 (2); Megommata Barnes, 1939 (6). Rhizomyiini Mamaev, 1967. Rhizomyiina: Rhizomyia Kieffer, 1898 (26), Stabiliola Fedotova et Sidorenko, 2006a (3); Alatostyla Fedotova et Sidorenko, 2006b (2); Pararhizomyia Mamaev, 1998 (2); Stellaserenina subtrib. n.: Stellaserena gen. n. (1); Stelladiurna gen. n. (1); Rovnobrachineura Fedotova et Perkovsky, 2014 (1). Ledomyiini Enderlein, 1936. Ledomyiina: Compositola Fedotova et Sidorenko, 2006a (1); Isogynandromyia Spungis, 1981 (1); Lauthia Kieffer, 1912 (29); Ledomyia Kieffer, 1895 (27); Plesiolauthia Mamaev, 1967 (1); Prolauthia Rübsaamen, 1916 (1); Rabindrodiplosis Grover, 1964 (2); Volsatiola Fedotova et Sidorenko, 2006a (2); Acinacistylina subtrib. n.: Acinacistyla Fedotova et Sidorenko, 2006b (3); Kuriloneura gen. n. (2); Kovaleviolina subtrib. n.: Kovaleviola Fedotova et Perkovsky, 2005 (2); Spungisiola Fedotova et Perkovsky, 2005 (1). (Fedotova, 2005, 2006a, 2006b—in papers by Fedotova and Sidorenko, 2005, 2006a, 2006b). The genera with an unclear taxonomic position: Palaeospaniocera Meunier, 1901 (1); Ledomyiella Meunier, 1904 (4). The above list includes genera and species of the gall midges occurring in the Late Miocene Mexican and Dominican amber—Lauthia (Gagné, 1973), in the Late Eocene Baltic amber—Ledomyia, Ledomyiella*, in African copal—Palaeospaniocera* (Meunier, 1901), and 8 genera occurring only in the Late Eocene Rovno amber: Brachineura, Brachyneurina, Kovaleviola*, Ledomyia, Nascensluna*, Popovineura*, Rovnobrachineura*, Spungisiola* (Fedotova and Perkovsky, 2005, 2008; Perkovsky and Fedotova, 2008; Fedotova and Perkovsky, 2014). Among these, 7 genera (noted with an asterisk*) have not been found yet in the recent fauna. The species of the supertribe Brachineuridi, mainly mycetophagous and xylophagous, do not cause formation of galls even when their larvae contact with living vegetative tissues. The larvae were found to develop on remains of insects and spiders, in the forest litter; ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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Fig. 1. Undoneurini (1–5, 17–20) and Brachineurini (6–16), males: (1–5) Cingola certa Fedotova et Sidorenko; (6–11) Nodalistyla undata Fedotova et Sidorenko; (12) N. aberrata (Mamaev); (13–16) Ramineura subulata (Fedotova et Sidorenko); (17–20) Chrybaneura harrisoni Gagné; (1, 7, 16, 19) genitalia; (2, 8) claw; (3, 10, 14, 17) 5th flagellar segment, dorsal view; (18) 5th flagellar segment, ventral view; (4) 1st flagellar segment; (5, 6, 13) mouthparts; (9) pedicel and 1st and 2nd flagellar segments; (11) 8th flagellar segment; (12) gonostylus; (15) scape, pedicel, and 1st and 2nd flagellar segments; (20) aedeagus and basal projection of gonocoxite (after: Mamaev, 1967; Gagné, 1968, 1994; Fedotova and Sidorenko, 2006a).
the adult gall midges were reared from a decaying wood. Some species of Ledomyiini develop in the fruit bodies of fungi; some Nearctic species were reared from vessels of the tree xylem. The species of the ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
genera Chrybaneura, Coccidomyia, and Megommata are associated with coccids, other insects, and spiders; some species of these genera feed on adults and eggs of coccids; Prolauthia circumdata Winnertz is an in-
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Fig. 2. Brachineurini (1, 3, 7–9), Undoneurini (2, 4–6, 12), and Rhizomyiini (10, 11), wing [(1–8, 10–12) males; (9) female]: (1) Brachineura impleta Fedotova et Sidorenko; (2) Undoneura phaseoliformis (Fedotova); (3) Ramineura subulata (Fedotova et Sidorenko); (4) Undoneura fasciculata Fedotova et Sidorenko; (5) Chrybaneura harrisoni Gagné; (6) Cingola certa Fedotova et Sidorenko; (7) Epimyia calcarata Fedotova et Sidorenko; (8) Nodalistyla undata Fedotova et Sidorenko; (9) Spaniocera squamigera Winnertz; (10) Coccidomyia pennsylvanica Felt; (11) Alatostyla velutina Fedotova et Sidorenko; (12) Brachyneurina peniophorae Harris (after: Nijveldt, 1963; Gagné, 1968; Harris, 1968, 1979; Fedotova, 2004; Fedotova and Sidorenko, 2006a, 2006b).
quiline in galls of Dasineura crataegi Winnertz. The majority of the species of Brachineuridi were collected with traps (Mamaev, 1967; Harris, 1968; Rock and Jackson, 1985; Gagné, 2004). The larva is known for Isogynandromyia terricola Spungis (Spungis, 1981), Brachyneurina peniophorae
Harris (Harris and Evans, 1979), and Chrybaneura harrisoni Gagné (Gagné and Ėtienne, 2009). The head capsule is hemispherical or triangular. The exocephalic rods are long, well developed. The antenna are 1- or 2-segmented, cylindrical or pear-shaped, directed laterally. The spatula is bidentate; the denticles ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
are triangular, dark. The posterior segment of the body bears 6 or 8 setae varying in length. The anal opening is narrow, situated ventrocaudally. A Key to the Tribes and Subtribes of the Supertribe Brachineuridi 1. Flagellar segments of antenna entirely densely covered with scales, without horseshoe-shaped lunulae (peritremes) (Figs. 1, 10, 11, 14, 15; 3, 19), with basal whorl of setae. Basal enlargement bearing serpentine (Fig. 5, 15) or alveolate sensory threads situated longitudinally, or threads under scales indistinct. Neck in male and female slightly longer or shorter than wide, occasionally absent in female (Fig. 5, 17). Cercus much wider than gonocoxite (Figs. 1, 1, 7, 16; 3, 13, 16, 17). Vein R4+5 very strongly approximate to costal vein, far not reaching wing apex (Fig. 2, 1, 3, 7, 9), straight at base. Vein Cu without furcation. Labrum wide, rounded, entirely concealing labellum or leaving lateral sides slightly open (Fig. 1, 6, 13) (Brachineurini) ........ 2. —Flagellar
segments of antenna not covered with scales, with large peritremes (Figs. 1, 3, 4, 8; 3, 2, 3, 8). Basal enlargement bearing sensory threads in shape of one or two rings connected by crosspieces, rarely threads absent. Length of neck of male always distinctly exceeding its width and usually exceeding width of basal enlargement (Figs. 5, 2, 10; 8, 4, 6); neck of female very short (Fig. 4, 15–21). Cercus slightly wider than gonocoxite (Fig. 5, 5, 9). Labrum triangular, not concealing labellum (Figs. 3, 1, 15; 5, 6; 8, 15) ............... 3.
2. Gonostylus well developed, widened at base and gradually narrowed toward apex (Fig. 1, 16), longer than width of gonocoxite, without pubescent dorsal lobe. Gonocoxite without basal projection. Aedeagus unsclerotized or weakly sclerotized, straight. Vein R4+5 zigzag-like curved at base (Fig. 2, 2, 4, 6), running into margin of wing far not reaching its apex ............................... Brachineurina. —Gonostylus
very short, rudimentary (Fig. 3, 6, 12, 13, 17, 20) or with elongate pubescent dorsal lobe (Fig. 1, 7, 12); its length less than width of gonocoxite. Aedeagus occasionally heavily sclerotized, claw-shaped (Figs. 1, 7; 3, 13, 18), curved near middle, strongly swollen at base (Fig. 3, 13, 18). Vein R4+5 straight at base, running into margin of wing before its apex (Fig. 2, 7, 8) .......... Epimyiina.
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3. Gonocoxite without basal projection. Genitalia very wide. Gonostylus large, even, gradually narrowed toward apex or with lobiform processes (Figs. 5, 5; 8, 1; 9, 1; 11, 14, 15, 17), or narrow and swollen at base (Figs. 1, 19; 5, 1, 3, 9). Vein Rs not developed. Tegmen oval (Fig. 8, 1), with apical processes (Fig. 9, 1) or triangular (Fig. 11, 14, 15) .............. 4. —Basal
projection of gonocoxite developed, long, fine, closely adjoining aedeagus (Figs. 10, 11; 12, 21; 14, 1, 3, 7; 15, 10; 16, 1, 3). Genitalia usually rounded or slightly elongate. Vein R4+5 running into margin of wing far before its apex; furcation of Cu always distinct (Figs. 11, 5, 6; 13, 8; 14, 8); when Rs developed, furcation of Cu occasionally absent (in the Late Eocene genera from amber). Tegmen wide (Fig. 10, 3, 7) or with apical processes and lateral branches (Figs. 10, 11; 12, 21) (Ledomyiini) ......................................................... 7.
4. Necks of middle flagellar segments nearly as long as or slightly shorter than basal enlargements (Figs. 5, 7; 8, 6; 9, 5); apical whorl of setae welldefined; peritremes forming middle whorl. Vein R4+5 straight, running into wing margin near its apex (Figs. 2, 11; 11, 1–5; 13, 8), or beyond apex (Figs. 8, 1; 9, 7); Cu bifurcate; when Cu not bifurcate, gonostylus with lobe (Fig. 5, 5), or aedeagal complex (Figs. 8, 7; 9, 1) with large tegmen and with very wide hypoproct forming semicircular emargination between pointed lobes. Gonostylus usually wide, even, without distinct swelling, slightly shorter than gonocoxite bearing no basomedial pubescent lobe at base (Rhizomyiini) ............ 5. —Necks of middle flagellar segments short or slightly longer than half of basal enlargements (Figs. 1, 3, 4, 17, 18; 4, 18, 20, 21; 5, 2, 10); apical whorl of setae absent; peritremes densely covering entire basal enlargement. Vein R4+5 running into margin of wing far before its apex; Cu simple (Fig. 2, 2, 4, 6, 12); when Cu bifurcate or indistinct, R4+5 running into or beyond wing apex (Fig. 2, 10). Gonostylus narrow, swollen at base (Figs. 1, 19; 5, 1, 3, 9), shorter than gonocoxite usually bearing basal pubescent lobe or swelling with setae (Fig. 1, 20) (Undoneurini trib. n.) .................................................................. 6. 5. Hypoproct with elongate oval emargination and narrow lateral lobes (Fig. 12, 6, 14, 18), slightly widened or narrowed toward base. Gonostylus large, swollen at base, occasionally with lobe or bifurcate (Figs. 5, 5; 11, 14, 15). Vein R4+5 running
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into margin of wing before its apex, or running nearly into wing apex (Figs. 2, 11; 11, 10; 13, 8); Cu usually with furcation; when Cu without furcation, it reaching margin of wing; gonostylus bifurcate (Fig. 5, 5) .................................... Rhizomyiina. —Hypoproct
with semicircular emargination and pointed lobes, very strongly widened toward base (Figs. 8, 1; 9, 1). Gonostylus long, wide. Vein R4+5 running beyond wing apex (Figs. 8, 7; 9, 7); Cu simple, not reaching margin of wing ............................. ........................................... Stellaserenina subtrib. n.
6. Vein R4+5 arcuately curved at base, running into margin of wing far before its apex. Cu arcuate or straight (Fig. 2, 4–6). Gonocoxite with large pubescent mediobasal lobe (Figs. 1, 1, 19; 5, 1, 3) or with projection bearing setae (Fig. 1, 20). Gonostylus very narrow, glabrous or covered with microtrichiae only dorsobasally. Hypoproct much narrower than cercus, parallel-sided or slightly widened toward base, complete (Fig. 5, 1, 3) or emarginate apically (Fig. 1, 19), rarely V-shaped (Fig. 1, 1). Gonocoxite without basal projection ................................ ............................................ Undoneurina subtrib. n. —Vein
R4+5 straight at base, running nearly into or beyond wing apex (Figs. 2, 10; 17, 19). Vein Cu straight or bifurcate. Gonocoxite without pubescent lobe (Fig. 17, 10, 12), rarely with projection bearing setae (Fig. 5, 9). Gonostylus swollen at base, entirely covered with microtrichiae (Fig. 17, 9, 11). Hypoproct nearly as wide and long as, or slightly narrower and longer than cercus, with shallowly emarginate or complete apex. Roots of genitalia usually bifurcate ............. Coccidomyiina subtrib. n.
7. Vein Cu2 reduced, not bifurcate; Cu1 and M3+4 well developed; Rs approximate to place of fusion of R1+2 with costal vein. Vein R4+5 running into margin of wing far from its apex. Flagellar segments with short necks in male, without necks in female .......... ........................................... Kovaleviolina subtrib. n. —Veins
Cu1 and Cu2 always developed, forming furcation; M3+4 usually developed; Rs distant from place of fusion of R1+2 with costal vein. R4+5 running into wing margin near its apex. Necks of flagellar segments of male nearly as long as basal enlargements ............................................................................... 8.
8. Hypoproct with shallow semicircular or triangular emargination (Figs. 10, 1, 3, 7; 12, 1, 23), rarely complete (Figs. 10, 19; 17, 2, 7). Gonostylus short,
wide or clearly widened at base. Vein Cu2 forming smooth continuation of basal stem of this vein; Cu1 arcuately originating from it, forming acute angle (Fig. 11, 8, 9). Necks of middle flagellar segments about 0.7 times as long as basal enlargements (Fig. 10, 5, 9) ....................................... Ledomyiina. —Hypoproct
complete or with lateral processes (Figs. 10, 11, 24; 12, 21; 14, 1–3; 15, 10; 16, 1, 3). Gonostylus very long and fine, almost not widened at base. Veins Cu1 and Cu2 not forming smooth continuation of basal stem of this vein; Cu2 forming obtuse angle with basal stem (Figs. 11, 5, 6; 14, 8; 15, 11; 16, 11). Necks of middle flagellar segments nearly as long as basal enlargements (Figs. 10, 10; 12, 20; 15, 2), or distinctly shorter than the long nodose basal enlargements (Figs. 15, 7; 16, 9) ......... .......................................... Acinacistylina subtrib. n. Tribe BRACHINEURINI Kieffer, 1913
Diagnosis. Body small, dark, densely covered with scales. Antenna 2+9–11-segmented; segments always entirely densely covered with scales. Flagellar segments varying in shape and length, each with very short neck length of which not exceeding width of segment [rarely (in Epimyiella), neck slightly shorter than basal enlargement]. Basal enlargements frequently covered with serpentine, vertically situated threads or with loop-shaped threads forming net. Palpus strongly reduced, 2- or 3-segmented. Wing widest in distal part, with 3 longitudinal veins: R1+2, R4+5, and Cu. Vein R4+5 very strongly approximate to costal vein, far not reaching wing apex; Cu without furcation, frequently disappearing before its confluence with margin of wing. Palpus 1–3-segmented. Fore femur and tibia shorter than hind ones. Claws of tarsi narrow, curved near middle, occasionally with basal tooth hardly visible because of scales densely covering ends of tarsi. Male genitalia always with very large wide cerci (tergite IX) separated by deep emargination (Figs. 1, 16; 3, 13, 16; 5, 12, 16, 18; 6, 1, 10, 12, 13, 15; 7, 1). Cercus overlapping half width of gonocoxite on inner side. Hypoproct narrow, also with deep emargination. Ovipositor short, non-telescopic, with large ventral lobe (abdominal segment VIII); its end with 2 large dorsal plates (cerci or tergite IX) and with small ventral plate (tergite X) (Figs. 4, 12, 14; 6, 18). The tribe comprises 2 subtribes: Brachineurina (4 genera, 23 species) and Epimyiina (4 genera, 9 species). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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Fig. 3. Undoneurina (1–9) and Epimyiina (10–20), male (1–8, 10–20) and female (9): (1–5) Undoneura fasciculata Fedotova et Sidorenko; (6–9) U. phaseoliformis (Fedotova); (10–15) Epimyia calcarata Fedotova et Sidorenko; (16–19) Epimyia sp.; (20) Epimyiella palustris Mamaev [(1, 15) mouthparts; (2) 1st and 2nd flagellar segments of antenna; (3) 4th and 5th flagellar segments; (4, 6, 13, 17) genitalia, dorsal view; (16) genitalia, ventral view; (5, 14) claw; (7, 19) 5th flagellar segment of antenna; (8) 8–10th flagellar segments; (9) ovipositor; (10) 8th flagellar segment; (11) 3rd flagellar segment; (12) gonocoxite and gonostylus; (18) aedeagus, lateral view; (20) gonocoxite, gonostylus, and paramere (after: Mamaev, 1967; Gagné, 1994; Fedotova, 2004; Fedotova and Sidorenko, 2006a).
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Fig. 4. Rhizomyiini (1, 8–11, 18), Undoneurini (2–4, 6, 15–17, 20, 21), Brachineurini (5, 7, 12–14, 19), females: (1, 18) Coccidomyia pennsylvanica Felt; (2, 3, 20) Brachyneurina peniophora Harris; (4, 15–17) Cingola certa Fedotova et Sidorenko; (5) Spaniocera squamigera Winnertz; (6, 21) Chrybaneura harrisoni Gagné; (7) Epimyia sp.; (8) Rhizomyia turriformis Fedotova et Sidorenko; (9–11) Rhizomyia operculata Fedotova et Sidorenko; (12, 13, 19) Brachineura enigmatosa Fedotova et Sidorenko; (14) B. transversa Fedotova et Sidorenko; (1–6, 9–14) ovipositor [(1, 2, 5, 6, 10, 12, 14) lateral view; (3, 4, 11, 13) dorsal view; (9) ventral view]; (7, 8) abdominal segments VI–VIII; (15–21) flagellar segments of antenna [(15, 18) 1st and 2nd, (16) 4th and 5th, (17) 9th and 10th, (19, 21) 5th, (20) 3rd segments] (after: Nijveldt, 1963; Gagné, 1968, 1994; Harris, 1968, 1979; Fedotova and Sidorenko, 2005, 2006a, 2006b).
Subtribe Brachineurina Kieffer, 1913 Description. Body, including antennae, palpus, tarsi, and genitalia, very densely covered with scales. Antenna 2+10-segmented. Flagellar segments of male with short apical necks and with cylindrical, urceolate or distally widened basal enlargements (Figs. 5, 17;
6, 9). Neck usually not longer than wide, less frequently 1/4 as long as basal enlargement. As exception, neck subequal in length to basal enlargement (Epimyiella), absent in all segments (Nascensluna) or in segments of apical part of antenna (Fig. 6, 8). In female, flagellar segments with short necks (Fig. 4, 19) ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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Fig. 5. Undoneurini (1–3), Rhizomyiini (4–11), Brachineurini (12–18), males: (1, 2) Brachyneurina peniophorae Harris; (3) B. xylophila Mamaev; (4–7) Alatostyla velutina Fedotova et Sidorenko; (8–11) Coccidomyia pennsylvanica Felt; (12, 13) Brachineura impleta Fedotova et Sidorenko; (14, 15) B. fuscogrisea Rondani; (16) Spaniocera squamigera Winnertz; (17, 18) Brachineura transversa Fedotova et Sidorenko; (1, 3, 5, 9, 12, 14, 16, 18) genitalia [(14) ventral view; others, dorsal view]; (2, 7, 10) 3rd flagellar segment of antenna; (4) claw; (6, 8, 13) mouthparts; (11) gonostylus; (15) 5th flagellar segment; (17) 1st flagellar segment (after: Nijveldt, 1963; Mamaev, 1967; Harris, 1968, 1979; Gagné and Solinas, 1996; Fedotova and Sidorenko, 2006a, 2006b).
or without necks (Spaniocera). Basal enlargement not bearing swollen horseshoe-shaped pores (peritreme)— places of attachment of long setae. Flagellar segments bearing sensory threads which occasionally invisible because of scales and indistinct without scales (Fig. 1, 14, 15) or varying in shape: vertical spiraliENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
form (Figs. 1, 11; 5, 15) or loop-shaped alveolate (Figs. 3, 19; 4, 15). Basal enlargements of middle flagellar segments in male usually more than 3.5–4.5 times as long as wide (Figs. 1, 11, 14; 6, 9), or, on contrary, short, more than 2.1–2.4 times as long as wide (Fig. 6, 2, 16). Necks always not longer than wide.
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Fig. 6. Brachineurina, males (1–17) and female (18): (1–5) Cornistyla infirma (Fedotova et Sidorenko); (6–10) C. flabellata (Fedotova et Sidorenko); (11, 12) C. nijveldti (Fedotova); (13, 14) Ramineura musiva (Fedotova et Sidorenko); (15–18) R. ussurica (Fedotova et Sidorenko); (1, 10, 12, 13, 15) genitalia; (2) 3rd and 4th flagellar segments of antenna; (3) 1st and 2nd flagellar segments; (4) mouthparts; (5, 11) claw; (6, 14) scape, pedicel, 1st and 2nd flagellar segments of antenna; (7) 10th flagellar segment; (8) 9th flagellar segment; (9, 16) 5th flagellar segment; (17) 8th flagellar segment; (18) ovipositor (after: Fedotova, 2004; Fedotova and Sidorenko, 2006a).
Proximal flagellar segments distinctly shorter than distal ones (Fig. 6, 6–9). Labrum complete, long and wide, concealing labellum and base of palpus (Fig. 1, 5, 13), usually rounded (Fig. 1, 6, 13); as exception (in Epimyia), labrum triangular, and labellum long and wide (Fig. 3, 15). Palpus 1–4-segmented (Fig. 1, 5, 6, 13). Basal projection of gonocoxite not developed; sclerotized paramere occasionally present (Figs. 1, 7; 3, 20).
The subtribe comprises 4 genera and 23 species, including the recent Brachineura Rondani, 1840 (a combined genus; 17 species), Ramineura Fedotova et Sidorenko, 2006b (as a subgenus of Brachineura), stat. n. (4), Spaniocera Winnertz, 1853 (1), and the fossil Nascensluna Fedotova et Perkovsky, 2014 (1 species, N. mellea Fedotova et Perkovsky, 2014 : 394) described from the Late Eocene Rovno amber. In Rovno amber, Brachineura polessica Fedotova ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
et Perkovsky, 2014 : 391 was also found (Fedotova and Perkovsky, 2014). A Key to the Genera of the Subtribe Brachyneurina 1. Vein R4+5 running into wing margin near its middle, fused with R1+2. Flagellar segments of female without necks. Apical plates of ovipositor small, pointed. A Late Eocene genus from Rovno amber ....................... Nascensluna Fedotova et Perkovsky. —Vein
R4 +5 running into margin of wing far before its apex but not fused with R1+2 at middle of wing (Fig. 2, 1–9). Flagellar segments of female with short necks (except in Spaniocera). Apical plates of ovipositor large, rounded. Recent and Late Eocene genera ................................................................... 2.
2.
Gonostylus short, strongly widened basally (Fig. 5, 12, 14, 18). Aedeagus thickened, weakly sclerotized, subparallel-sided distally, widely rounded apically (Fig. 5, 18) or with small rounded apical swelling (Fig. 5, 12, 14, 16). Cercus and hypoproct far not reaching apex of gonocoxite. Width of cell between veins C and R4+5 more than 3 times exceeding distance from C to R1+2 (Fig. 2, 1, 9). Small accessory vein beginning from place of fusion of veins C and R1+2 not developed ............. 3.
—Gonostylus
very long and fine, subparallel-sided or swollen basally (Figs. 1, 16; 6, 13, 15). Aedeagus fine, triangular, always with pointed apex and widened base, heavily sclerotized. Cercus and hypoproct far not reaching apex of gonocoxite or longer than it. Vein R4+5 strongly approximate to margin of wing (Fig. 2, 3); cell between veins C and R4+5 very narrow, its width less than 3 times distance from C to R1+2. Small accessory vein originating from fusion of veins C and R1+2 developed .......................... .......................... Ramineura Fedotova et Sidorenko.
3. Vein Rs present at base of wing (Fig. 2, 9); Cu straight, far not reaching wing margin. Gonocoxite without fine basal projection bearing setae (Fig. 1, 16). Palpus 4-segmented. Flagellar segments without sexual dimorphism, without necks in both sexes ..................................... Spaniocera Winnertz. —Vein
Rs absent at base of wing (Fig. 2, 1); Cu arcuate before apex, slightly not reaching margin of wing. Gonocoxite with fine glabrous basal projection bearing 1–3 setae (Fig. 5, 14), with lobe (Fig. 5, 18) or without lobe (Fig. 5, 12). Palpus 1–3-segmented. Structure of flagellar segments
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demonstrating sexual dimorphism: short necks present in male, absent in female ................................... .............................................. Brachineura Rondani. Genus Brachineura Rondani, 1840 Rondani, 1840 : 16. Brachyneura Anonymous, 1844 : 450 (misspelling). Brachynevra Agassiz, 1847 : 51, an unjustified correction. Acroectasis (as Acroëctasis) Rübsaamen, 1910 : 131. Type species Acroectasis maura Rübsaamen (by monotypy). Synonymy after: Gagné, 2010, with corrections. Mamaev, 1967 : 874; Gagné, Solinas, 1996 : 73; Fedotova, 2004 : 598; Fedotova and Sidorenko, 2006a : 78. Type species Brachineura fuscogrisea Rondani, 1840 : 17 (by monotypy). Description. Antenna 2+10-segmented; middle segments long, cylindrical, with short necks length of which in male almost not exceeding their diameter (Fig. 5, 15); in female, these necks even shorter (Fig. 5, 17). Labrum wide, semicircular; palpus 3-segmented (Fig. 5, 13). Vein R4+5 running into margin of wing far before its apex (Fig. 2, 1, 3, 9). Cu simple, without undulate curvature at base, curved distally, disappearing before margin of wing. Claws in all tarsi or only in fore tarsus with tooth at base. Gonocoxite slender, occasionally with small basal projection bearing setae (Fig. 5, 14). Gonostylus short or very long, swollen at base (Fig. 5, 12, 14, 18). Cercus very wide, cordate, frequently concealing more than middle of gonocoxite on inner side (Fig. 5, 12, 16, 18). Hypoproct narrow, with deep emargination. Aedeagus cylindrical, occasionally widened at base and rounded at end. Mamaev (1967) marked that “since the description of the genus Brachineura was incomplete and that of the genus Spaniocera included errors, the species belonging to the genus Brachineura were repeatedly described in the genera Brachyneurella Kieffer and Acroectasis Rübsaamen. After Edwards’s (1937) correction of these inaccuracies in the descriptions, there are few doubts in the synonymy of the genera given above [Spaniocera, Brachyneurella and Acroectasis].”
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FEDOTOVA
This cosmopolitan genus comprises 17 species, among which 13 are Palaearctic, 3 Nearctic, and 1 Australian. In Russia, 9 species were found, including 5 from the Far East. Biology not studied. Most adults were collected in nature. Food specialization is known only for several species. The larvae develop in decaying vegetative tissues (Gagné, 2004). The larvae of Brachineura fungicola (Mamaev, 1967) inhabit fungi on the bark of alder knots. The adults of B. squamigera were reared from fungi of the genera Russula and Rhizopogon (Basidiomycetes) (Bayram and Skuhravá, 2004; Skuhravá et al., 2005). Genus Spaniocera Winnertz, 1853 Winnertz, 1853 : 190. Spaniocera: Skuhravá, 1997 : 154. Brachineura (= Spaniocera): Gagné, 2004, 2010; Skuhravá, 2006. Type species Spaniocera squamigera Winnertz, 1853 : 377 (by monotypy). Brachyneura squamigera (Winnertz, 1853): Edwards, 1937; Milne, 1960; Nijveldt, 1963 (figure); Skuhravá, 1997 (keys). Diagnosis. Antenna 2+10-segmented in male and female; flagellar segments cylindrical, with short necks, covered with scales (Fig. 5, 15). Sensory thread developed in male, but hardly visible. Male genitalia with characteristic bilobed dorsal plate (cerci) and with hypoproct divided into two linear lobes (Fig. 5, 16). Ovipositor short, consisting of two large dorsal plates, small ventral plate, and large ventral lobe (Fig. 4, 5). Differential diagnosis. The genus Spaniocera is closely related to Brachineura, but differs in a 4-segmented palpus, in the absence of necks of the flagellar antennal segments in both sexes, and in the presence of cross-vein Rs at the base of the wing. Skuhravá (1997) accepts the independence of this genus, based on its description which nearly fits the characters of Brachineura. Later (Gagné, 2004, 2010; Skuhravá, 2006), Spaniocera was considered a synonym of Brachineura. The question of the independence of the genus Spaniocera requires further investigation. The genus includes only the type species (Figs. 2, 9; 5, 15, 16), the description of which (Winnertz, 1853)
was supplemented and clarified later (Edwards, 1937; Barnes, 1954; Nijveldt, 1963). Biology unknown. The adults were reared from the white clover (Barnes, 1954). In England, this species is of no significant economic importance (Barnes, 1954; Nijveldt, 1963). Genus Ramineura Fedotova et Sidorenko, 2006, stat. n. Fedotova and Sidorenko, 2006a : 84 (as a subgenus of Brachineura). Type species Brachineura ussurica Fedotova, 2004 : 601 (by monotypy). Diagnosis. Necks of flagellar segments short in male, inconspicuous in female. Basal enlargements long, parallel-sided (Fig. 1, 14, 15) or slightly swollen laterally (Fig. 6, 16, 17), densely covered with scales; sensory threads indistinct. 1st flagellar segment nearly without neck (Figs. 1, 15; 6, 14). 10th segment with rounded apical swelling (Fig. 6, 7, 17). Labrum complete, rounded (Fig. 1, 13) or with small triangular projection (R. ussurica). Palpus short, 2- or 3-segmented. Wing 2.4–2.6 times as long as wide (Fig. 2, 3). Width of cell between veins C and R4+5 less than 3 times distance from C to R1+2. Small accessory vein originating from fusion of R1+2 and C developed. Vein R4+5 straight or slightly curved, together with costal vein forming very narrow cell, occasionally with pore at place of their fusion. Femora strongly swollen, especially hind one. Gonostylus very long and fine, constituting 3/4 of length of gonocoxite (Figs. 1, 16; 6, 13, 15), very fine along entire length of distal half. Aedeagus triangular, or wide at base, or narrow, nearly needle-shaped, or wide and subparallel-sided, heavily sclerotized and pointed at end (Fig. 1, 16). Hypoproct with long and fine lobes, parallel-sided, horseshoe-shaped or slightly widened basally. Ovipositor with elongate apical lobes; ventral lobe of segment VIII large, rounded, with sclerotized inner structure (Fig. 6, 18). Differential diagnosis. The genus Ramineura is closely related to Brachineura, but differs in a very fine distal half of the gonostylus, in a sclerotized aedeagus, in the presence of an accessory vein at the place of fusion of veins R1+2 and R4+5, and in a narrow cell between the costal vein and R4+5. The genus comprises 4 species: Ramineura ussurica (Fedotova, 2004), comb. n. (Brachineura) (Fig. 6, 15– ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
18), R. musiva (Fedotova et Sidorenko, 2006a), comb. n. (Brachineura, subgen. Ramineura) (Fig. 6, 13, 14), and R. subulata (Fedotova et Sidorenko, 2006a), comb. n. (Brachineura, subgen. Ramineura) (Figs. 1, 13–16; 2, 3) all described from Primorskii Territory (the type species, from the environs of the Ussuri Nature Reserve, the others, from the Lazovskii Nature Reserve), and also R. alia (Mamaev, 1994), comb. n. (Brachineura) (Fig. 17, 8) known from Kamchatka (Mamaev, 1994). Biology unknown. In Primorskii Territory, all the species were collected using Malaise, yellow, and light traps, respectively. R. alia was noted to be mycetophagous (Mamaev, 1994). A Key to the Species of the Genus Ramineura 1. Gonocoxite very fine, apically nearly as wide as gonostylus. Gonostylus more strongly curved in distal 1/3 (Fig. 17, 8). Palpus 3-segmented ................... ..................................................... R. alia (Mamaev). —Gonocoxite
apically much wider than base of gonostylus (Figs. 1, 16; 6, 13, 15). Gonostylus straight or regularly arcuate. Palpus 1 or 2-segmented ......................................................... 2.
2. Gonostylus very fine from base to apex, 0.7 times as long as gonocoxite (Fig. 6, 13). Aedeagus strongly widened toward base. Labrum complete, with widely rounded margin. 1st flagellar segment 2.6 times as long as wide (Fig. 6, 14). Body length 0.83 mm, length of wing 0.95 mm ........................... .......................... R. musiva (Fedotova et Sidorenko). —Gonostylus
swollen at base (Figs. 1, 16; 6, 15), 0.8–0.9 times as long as gonocoxites. Aedeagus nearly needle-shaped, weakly widened toward base, or short, wide, subparallel-sided .......................... 3.
3. Aedeagus short, subparallel-sided, pointed near apex (Fig. 1, 15). Hypoproct heavily sclerotized, horseshoe-shaped. Labrum complete, with widely rounded margin. 1st flagellar segment 2.9 times as long as wide (Fig. 1, 15). Body length 0.88 mm, length of wing 0.9 mm ............................................. ....................... R. subulata (Fedotova et Sidorenko). —Aedeagus
very fine, nearly needle-shaped. Hypoproct slightly sclerotized, with constriction in proximal half (Fig. 6, 15). Labrum with small triangular projection. 1st flagellar segment 1.9 times as long as wide. Body length 0.51–0.64 mm, length of
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wing 0.94 mm ........................................................... ....................... R. ussurica (Fedotova et Sidorenko). Genus Nascensluna Fedotova et Perkovsky, 2014 Fedotova and Perkovsky, 2014 : 394. Type species Nascensluna mellea et Perkovsky, 2014 : 394 (by monotypy).
Fedotova
Diagnosis. Antenna 2+10-segmented. All flagellar segments without necks, slightly rounded laterally, gradually narrowed and shortened toward apex. Sensory threads not visible. Veins R1+2 and R4+5 nearly fused with costal vein. Cu regularly arcuately curved, far distant from margin of wing. Fore femur and tibia slightly longer than middle ones and shorter than hind ones. Wing 2.8 times as long as wide. 1st segment of hind tarsus 0.2 times as long as 2nd. Apical plates of ovipositor with pointed apices. Differential diagnosis. The genus Nascensluna is closely related to Brachineura, but differs in the absence of necks of the flagellar segments, in characters of the wing venation, and in pointed apical plates of the ovipositor. The genus includes only the type species described from the Late Eocene Rovno amber. Subtribe Epimyiina Mamaev, 1968 Mamaev, 1968 : 16, 24. Diagnosis. Antenna 2+9 or 10-segmented. 1st flagellar segment with nearly rounded basal enlargement, 0.58–0.66 times as long as 2nd segment (Fig. 1, 9). Flagellar segments with short necks not longer than wide; as exception, neck slightly shorter than basal enlargement in Epimyiella. Middle flagellar segments long, cylindrical. Basal enlargements with longitudinal serpentine sensory threads (Nodalistyla, Fig. 1, 11); or threads forming meshes in distal half (Epimyia, Fig. 3, 19) or net (Fig. 3, 10), or hardly visible under dense scales on segment. Proximal flagellar segments shorter than distal ones (Fig. 1, 9–11). Palpus 3-segmented (Fig. 1, 6). Labrum wide, triangular, not concealing dorsally base of palpus, projecting far beyond labellum (Fig. 3, 1, 15). Vein R4+5 straight at base, running into wing margin near its apex (Epimyiella) or far before apex (Epimyia) (Fig. 2, 7). Vein Rs in Nodalistyla poorly developed, not fused with R4+5 (Fig. 2, 8). Vein Cu simple, sharply curved in distal 1/4 before confluence with margin of wing. Tarsal claw simple or with tooth at base. Gonocoxite
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large, with apical or apicolateral lobes (Fig. 3, 12, 13, 16, 17, 20) or without lobes; sides of gonocoxite rounded or straight, its middle with wide basomedial emargination. Gonostylus fine, rudimentary. Cercus with emargination; lobes narrow and hooked outward, or wide and cordate (Figs. 1, 7; 3, 13, 16, 17). Hypoproct narrow, emarginate or cordate. Tegmen complete, in shape of distinctly sclerotized plate. Basal projection of gonocoxite absent, or developed in shape of sclerotized paramere which adjoining wide sclerotized aedeagus or distant from it (Fig. 3, 20). Base of gonocoxite always without rounded lobe. Aedeagus usually arcuately curved (Figs. 1, 7; 3, 13, 18), occasionally heavily sclerotized, claw-like curved near middle and directed dorsally, or straight and equal in length to gonocoxite. Differential diagnosis. The subtribe Epimyiina differs from Brachineurina in rudimentary gonostyli, in a complicated aedeagal complex with the aedeagus widely varying in structure between representatives of different genera, and in slightly longer flagellar antennal segments. The subtribe comprises 4 genera and 9 species: Epimyia Felt, 1911 (2 species), Epimyiella Mamaev, 1967 (1), Nodalistyla Fedotova et Sidorenko, 2006a (2) and Cornistyla Fedotova et Sidorenko, 2006a (4) (Mamaev, 1967; Gagné, 1994; Fedotova, 2004; Fedotova and Sidorenko, 2006a). A Key to the Genera of the Subtribe Epimyiina 1. Necks of middle flagellar segments slightly shorter than basal enlargements. Basal enlargement bearing 2 rings of sensory threads connected by crosspieces; dorsal ring forming small loops. Gonocoxite with long apical lobe (Fig. 3, 20). Gonostylus straight, subequal in width to gonocoxites. Cercus cordate, weakly sclerotized. Paramere spiniform, sclerotized, curved at right angle, shorter than gonocoxite. Aedeagus straight, weakly sclerotized. Vein Cu sharply curved in distal 1/4. Body length 1.5 mm ................................................. Epimyiella Mamaev. —Necks
of middle flagellar segments not longer than wide and distinctly shorter than basal enlargements (Figs. 1, 9–11; 3, 19; 6, 2, 9; 7, 7, 8). Basal enlargement covered with longitudinal serpentine threads (Fig. 1, 11), or its distal half covered with net of sensory threads (Fig. 3, 19), or threads not visible (Fig. 6, 2, 9). Gonocoxite without apical lobe, with wide median emargination (Figs. 1, 7;
3, 13, 17; 6, 1, 10, 12; 7, 1). Gonostylus with projection (Figs. 1, 7, 12; 7, 4, 5) or slightly curved (Figs. 3, 13, 17; 6, 1, 10, 12), length of gonostylus less than width of gonocoxite. Paramere not developed. Cercus cordate (Fig. 1, 7) or narrow, hookshaped at end (Fig. 3, 13, 16) ................................ 2. 2.
Gonostylus attached to gonocoxite apically (Figs. 1, 7; 3, 12, 13, 17). Basal projection of gonocoxite or paramere developed, very large sclerotized tegmen present (Fig. 3, 13, 17). Aedeagus always curved and widened at base .................................. 3.
—Gonostylus
attached to gonocoxites subapically (Figs. 6, 1, 10, 12; 7, 1). Basal projection of gonocoxite not developed. Aedeagus straight, parallelsided. Cercus cordate. Vein Cu arcuately curved before confluence with margin of wing (Fig. 7, 17). 1st flagellar segment elongate, distinctly shorter than 2nd one (Figs. 6, 3, 6; 7, 7, 8) ........................... ........................... Cornistyla Fedotova et Sidorenko.
3. Gonocoxite with short apicomedial pubescent lobe (Fig. 3, 12, 13, 17). Gonostylus without dorsal lobe. Cercus narrow, hook-shaped at end. Tegmen in shape of clearly sclerotized plate. Paramere not developed. Basal projection of gonocoxite not developed. Aedeagus weakly sclerotized, smoothly curved medially in lateral view, slightly thickened near base (Fig. 3, 13) or strongly curved medially and very strongly thickened proximally (Fig. 3, 18). Vein Cu arcuately curved (Fig. 2, 8). Flagellar egments elongate, slightly differing in shape (Fig. 3, 10, 11, 19) ............................. Epimyia Felt. —Gonocoxite
without lobe (Fig. 1, 7). Gonostylus with pubescent dorsal lobe (Fig. 1, 7, 12). Cercus cordate. Basal projection of gonocoxite fine, sclerotized. Aedeagus heavily sclerotized, subulate, sharply curved dorsocaudally or straight. Vein Cu weakly curved (Fig. 2, 8). Flagellar segments strongly rounded laterally; 1st segment with nearly rounded basal enlargement (Fig. 1, 9–11); other segments very strongly differing in shape and size ......................... Nodalistyla Fedotova et Sidorenko. Genus Epimyia Felt, 1911 Felt, 1911 : 38.
Gagné, 1994 : 58; Fedotova and Sidorenko, 2006a : 91. Type species Epimyia carolina Felt, 1911 : 38 (by monotypy). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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1151
Fig. 7. Cornistyla tufa sp. n., male (1, 2, 4–11, 13, 15, 17) and female (3, 12, 14, 16): (1) genitalia; (2, 11) claw of hind tarsus; (3) ovipositor; (4, 5) gonostylus (variants of shape); (6) junction of 2nd and 3rd segments of tarsus; (7, 8) scape, pedicel, and 1st and 2nd flagellar segments; (9) claw of middle tarsus; (10) claw of fore tarsus; (12) pedicel, 1st–3rd flagellar segments; (13) wing venation, dorsal view; (14) hind trochanter, femur, and tibia; (15, 16) mouthparts; (17) wing. Scale 0.1 mm [(a) to 1–3; (b) to 4, 5; (c) to 6–12; (d) to 13, 14; (e) to 15, 16; (f) to 17].
Diagnosis. Antenna 2+10-segmented; flagellar segments with basal enlargements slightly widened distally or cylindrical, strongly narrowed toward apically (Fig. 3, 10, 11, 19). Basal enlargement densely ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
covered with scales, with meshes of sensory threads in distal half. Short necks developed in all segments. Palpus 3-segmented (Fig. 3, 15). Claw with tooth, empodium shorter than claw (Fig. 3, 14). Vein R4+5
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Fig. 8. Stellaserena vaga gen. et sp. n., male: (1) genitalia; (2) mouthparts; (3) facets of eye bridge on vertex side of head; (4) 1st–3rd flagellar segments of antenna; (5) scape and pedicel; (6) 6th flagellar segment; (7) wing. Scale 0.1 mm [(a) to 1; (b) to 2, 3; (c) to 4; (d) to 5, 6; (e) to 7].
arcuately curved distally, running into margin of wing far before its apex; Cu strongly curved distally, reaching margins of wing (Fig. 2, 7). Gonocoxite strongly widened at base and at apex, with wide median emargination (Figs. 1, 7; 3, 12, 13, 16, 17). Gonostylus very small, rudimentary, curved, shorter than width of gonocoxite. Cercus with narrow lobes, hook-shaped at end. Hypoproct heavily sclerotized, with apical emargination and with short rounded lobes. Aedeagus wide, heavily sclerotized, widened at base, with distal half bent dorsally (Figs. 1, 7; 3, 13, 18).
This Holarctic genus comprises 2 species: Epimyia carolina Felt described from the USA, North Carolina (Felt, 1911) (Fig. 3, 16–19) and E. calcarata Fedotova et Sidorenko from Russia, Primorskii Territory, the Lazovskii Nature Reserve (Fedotova and Sidorenko, 2006a) (Fig. 3, 10–15). Gagné (1994) reported that he knew several additional undescribed species from the Nearctic and Neotropical regions. Biology. Epimyia carolina develops in fungi of the genera Boletus, Leccinum, Paxillus, Suillus, and Strobilomyces (Boletales) (Gagné, 1994, 2004, 2010); ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
E. calcarata was collected with Malaise and light traps (Fedotova and Sidorenko, 2006a). The species of the genus Epimyia differ in the following characters. 1. Aedeagus nearly twice folded in middle (Fig. 3, 18), its base very strongly swollen. Hypoproct short, not narrowed near middle. Middle flagellar segments with subcylindrical basal enlargements (Fig. 3, 19). Claw simple. Body length 1.0 mm ........................... ........................................................ E. carolina Felt. —Aedeagus
very weakly curved (Fig. 3, 13), regularly widened toward base. Hypoproct narrowed in middle. Middle flagellar segments with basal enlargements widened distally (Fig. 3, 10, 11). Claw with tooth at base (Fig. 3, 14). Body length 0.70–0.85 mm, length of wing 0.83–1.0 mm ........... ........................ E. calcarata Fedotova et Sidorenko. Genus Epimyiella Mamaev, 1967
Mamaev, 1967 : 879. Type species Epimyiella palustris Mamaev, 1967 : 880 (by monotypy). Diagnosis. Antenna 2+10-segmented. Middle flagellar segments of male antenna with long necks slightly shorter than basal enlargements. Circular thread on basal enlargement well-developed, consisting of two rings connected by crosspieces. Dorsal ring of circular thread forming small loops. Vein Cu sharply curved in distal 1/4. Tarsi densely covered with scales. Gonocoxite thick, with long wide apical lobe (Fig. 3, 20). Gonostylus situated at base of this lobe, thickened in middle part, emarginate at end. Parameres forming 2 spines curved at right angle. Aedeagus weakly sclerotized. Differential diagnosis. The genus Epimyiella is closely related to Epimyia, but differs in a long apical lobe of the gonocoxite, in the long necks of the middle flagellar segments of the male, and in sclerotized parameres. This Palaearctic genus includes only the type species described from Moscow Province, Lake Glubokoe (Mamaev, 1967) (Fig. 3, 20). Genus Nodalistyla Fedotova et Sidorenko, 2006 Fedotova and Sidorenko, 2006a : 91. Type species N. undata Fedotova et Sidorenko, 2006a : 91 (by monotypy). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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Diagnosis. Antenna 2+9 or 10-segmented; 2nd flagellar segment 1.5–1.7 times as long as 2nd one (Fig. 1, 9). Middle flagellar segments cylindrical, with short necks; basal enlargements covered with spiralshaped sensory threads (Fig. 1, 11). Claw simple (Fig. 1, 8) or with tooth at base; empodium half as long as claw or equal to it in length. Gonocoxite large, heavily sclerotized. Gonostylus very short, its dorsal part forming long pubescent lobe (Fig. 1, 7, 12). Cercus cordate, with large rounded lobes. Hypoproct parallel-sided or cordate. Aedeagus heavily sclerotized, subulate, distal half sharply bent dorsally or straight. Differential diagnosis. The genus Nodalistyla is closely related to Epimyia Felt in the presence of rudimentary gonostyli, but differs in the absence of an apical lobe of the gonocoxite and in the presence of a sclerotized projection at the base of the gonocoxites and of serpentine sensory vertical threads on the flagellar antennal segments. This Palaearctic genus includes the type species N. undata known from Primorskii Territory (the Lazovskii Nature Reserve) (Fedotova and Sidorenko, 2006a) (Figs. 1, 6–11; 2, 8) and N. aberrata (Mamaev, 1967), comb. n. (Brachineura) from Moscow Prov., Danki Vill. (Mamaev, 1967) (Fig. 1, 12). The species of the genus Nodalistyla differ in the following characters. 1. Antenna 2+9-segmented; 2nd flagellar segment 1.7 times as long as 1st one (Fig. 1, 9). Tarsal claw simple, empodium as long as claw (Fig. 1, 8). Hypoproct cordate (Fig. 1, 7). Aedeagus spiniform, sharply curved dorsocaudally. Distal margin of dorsal lobe of gonostylus forming right angle with its other part. Body length 0.98 mm .............................. ............................ N. undata Fedotova et Sidorenko. —Antenna
2+10-segmented; 2nd flagellar segment 1.5 times as long as 1st one. Claw with tooth at base; empodium half as long as claw. Hypoproct parallel-sided. Aedeagus straight, equal in length to gonocoxite. Dorsal lobe of gonostylus hanging over its other part (Fig. 1, 12). Body length 1.7 mm ........ ............................................. N. aberrata (Mamaev). Genus Cornistyla Fedotova et Sidorenko, 2006, stat. n.
Fedotova and Sidorenko, 2006a : 87 (as a subgenus of Brachineura). Type species Brachineura nijveldti Fedotova, 2004 : 601 (by monotypy).
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Description. Sensory threads on segments not visible under dense scales. Necks of flagellar segments in male longer than in female. Basal enlargements of flagellar segments parallel-sided (Fig. 6, 8, 9) or swollen (Fig. 7, 7, 8, 12) in both sexes. 2nd flagellar segment 1.5–twice as long as 1st one. Mouthparts very dark, palpus paler. Labrum wide, rounded, entirely concealing labellum, complete (Fig. 7, 16) or slightly split in middle (Fig. 6, 4). Palpus 3-segmented. Wing 2.4–2.6 times as long as wide. Vein R4+5 not undulate proximally, arcuate distally; R1+2 fused with costal vein up to middle of wing; cell between them indistinct; veins R1+2 or R4+5 with pores (Fig. 7, 13, 17). Femora arcuately curved (Fig. 7, 14), all femora shorter than tibiae and 2nd tarsal segments. Claw simple in all tarsi (Fig. 6, 11), or with tooth in fore tarsus (Fig. 7, 10). Gonocoxite apically rounded (Figs. 6, 1; 7, 1), or truncate (Fig. 6, 10) or bevelled inwards (Fig. 6, 12), with group of long scales along apical margin. Occasionally gonocoxite very strongly widened distally (Figs. 6, 10; 7, 11). Gonostylus attached to gonocoxite subapically, fine, horn-shaped, curved near middle, swollen at base (Fig. 6, 1, 10) or at middle (Figs. 6, 12; 7, 1, 4, 5); its distal half straight; apical claw-shaped process very small, black. Cercus very wide, cordate, with deep wide emargination. Hypoproct narrow, parallel-sided distally, widened proximally or basally, with fine parallel-sided or triangular lobes separated by oval emargination. Aedeagus wide, parallel-sided, weakly sclerotized, with small rounded apical process or rounded. Cercus of ovipositor rounded, situated dorsocaudally. Differential diagnosis. The genus Cornistyla is similar to Epimyia Felt in elongate genitalia, rudimentary gonostyli, and in a narrow hypoproct, but differs in the subapical attachment of the gonostyli to the gonocoxite, in wide cerci with rounded lateral lobes, in a straight aedeagus far not reaching the apex of the gonocoxite, and in a rounded, instead of triangular, labrum. The genus comprises 4 species: Cornistyla nijveldti (Fedotova, 2004), comb. n. (Brachineura) (Fig. 6, 11, 12), C. flabellata (Fedotova et Sidorenko, 2006a), comb. n. (Brachineura, subgen. Cornistyla) (Fig. 6, 6–10), C. infirma (Fedotova et Sidorenko, 2006), comb. n. (Brachineura, subgen. Cornistyla) (Fig. 6, 1–5), and C. tufa Fedotova, sp. n. (Fig. 7, 1– 17).
Biology unknown. All the species were collected at light or with Malaise traps. A Key to the Species of the Genus Cornistyla 1. Aedeagus with smoothly rounded apex (Figs. 6, 10; 7, 1). Middle flagellar segments long, parallel-sided (Figs. 6, 8, 9; 7, 8). Gonocoxite widest distally (Figs. 6, 10; 7, 1). Hypoproct weakly widened near base, with triangular lobe. 2nd flagellar segment 3.1–3.3 times as long as wide (Figs. 6, 6; 7, 8). Length of wing 1.13–1.20 mm .............................. 2. —Aedeagus
with small rounded apical projection (Fig. 6, 1, 12). Middle flagellar segments short, swollen (Fig. 6, 2, 3). Gonocoxite widest near base or at middle (Fig. 6, 1, 12). Hypoproct sharply widened proximally, with fine parallel-sided lobe. 2nd flagellar segment 2.6 times as long as wide (Fig. 6, 3). Vein R4+5 arcuate. Length of wing 0.95–0.97 mm ....................................................... 3.
2. Gonostylus widened at base; gonocoxite straightly truncate at apex (Fig. 6, 10). Vein R4+5 straight. Gonocoxite 2.4–2.6 times as long as wide. Body length 0.95 mm, length of wing 1.20 mm ................. ..................... C. flabellata (Fedotova et Sidorenko). —Gonostylus
widened near middle, gonocoxite rounded at apex (Fig. 7, 1). Vein R4+5 arcuate (Fig. 7, 13, 17). Gonocoxite 1.8 times as long as wide. Body length 0.94 mm, length of wing 1.13 mm .............................................. C. tufa sp. n.
3. Gonostylus strongly curved in basal 1/3 (Fig. 6, 12); gonocoxite bevelled from apex to inner margin. Length of wing 0.97 mm .... C. nijveldti (Fedotova). —Gonostylus
nearly straight (Fig. 6, 1); gonocoxite widely rounded apically. Body length 0.75 mm, length of wing 0.95 mm ............................................ ........................ C. infirma (Fedotova et Sidorenko). Cornistyla tufa Fedotova, sp. n. (Fig. 7, 1–17; see also table)
Material. Holotype, ♂ (slide 542/K-1/1): Russia, Southern Kuriles, Shikotan Island, Tserkovnaya Bay, Malaise trap, 9–15.IX.2012 (Yu.N. Sundukov). Diagnosis. Male. Body dark brown. Body length 0.94 mm; length of wing 1.13 mm, width of wing 0.46 mm. Head with widely rounded occiput. Eyes entirely well visible in view from facial side of head. Eye bridge narrow, consisting of 3 facets. Antennae ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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lost; 2 remained flagellar segments more heavily sclerotized than legs under scales. Scape, pedicel, and flagellar segments similarly uniformly sclerotized. Pedicel wider than long. Scape strongly widened distally; pedicel 0.77 times as long as scape and subequal to it in width. 1st flagellar segment of antenna uniformly sclerotized, without pale stripe at apex of distal neck, 0.6–0.7 times as long as 2nd segment. 2nd segment 3.3 times as long as wide; basal enlargement widely oval, 7.4 times as long as neck. Palpus 2-segmented, without palpiger, slightly swollen; length ratio of palpal segments 1 : 1.8 : 2.2; 3rd segment slightly swollen proximally, pointed apically. Thorax uniformly sclerotized, dark yellow, 1.3 times as long as wide. Anepimeron with 6 pores. Notum more heavily sclerotized than thorax. Haltere very densely covered with dark scales, pale under scales. Wing 2.5 times as long as wide. Vein R1+2 0.43 times as long as wing. Margin of wing not broken at place of fusion of costal vein and R4+5 (Fig. 7, 13). Vein Cu sharply curved in distal half, running into margin of wing. Legs dark because of dense scales. Hind leg 1.1 times as long as fore one. All femora slightly swollen and curved, 0.7–0.8 times as long as tibiae. Fore femur 0.62 times as long as hind one. Fore and middle femora nearly as long as 2nd segment of tarsus; hind femur 0.83 times as long as 2nd segment of tarsus. Hind tarsus 1.7 times as long as fore one. 2nd segment of fore tarsus 1.9 times as long as 2nd segment of hind tarsus and 4.0 times as long as 1st segment. 2nd segment of hind tarsus 6.4 times as long as 1st and in 3 times as long as 5th segment. In all legs, 3rd–5th tarsal segments combined always 1.3–1.5 times as long as 2nd segment. Claws of middle and hind tarsi simple, semicircular; fore tarsus with hook-shaped tooth at base; empodium shorter than claw. 2nd segment of tarsus with long apical projection fitting into emargination at proximal end of 3rd segment (Fig. 7, 6). Wing widest near middle, 2.5 times as long as wide; vein R1+2 0.43 times as long as wing. Genitalia elongate. Gonocoxite 1.8 times as long as wide, strongly widened in distal half, with wide and deep median emargination and with nearly semicircular wide subapical emargination to which very fine small gonostylus attached. Sclerotized stripe originating from this emargination and extending in shape of long chord toward base of gonocoxite. Gonostylus very narrow in distal half, slightly swollen proximally or near middle, 4.5–4.6 times as long as wide and ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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0.37 times as long as gonocoxite; length of gonostylus subequal to width of gonocoxite. Cercus far not reaching apex of gonocoxite, cordate, with deep triangular emargination between wide lobes. Hypoproct nearly as long and 0.47 times as wide as cercus, weakly widened toward base, with elongate apical lobes separated by rounded emargination. Aedeagus narrow, subparallelsided, heavily sclerotized, with rounded apex, as long as cerci and hypoproct. Female. Body dark brown. Body length 1.28 mm; length of wing 1.36 mm, width 0.54 mm. Length of antenna 0.90 mm. Occiput widely rounded. Eye bridge narrow, consisting of 3 facets. Antennae lost, only 3 flagellar segments remained. Pedicel rounded. 1st and 2nd flagellar segments subequal in length. 3rd segment 2.5 times as long as wide; basal enlargement widely oval, 7.7 times as long as neck. Palpus 3-segmented, without palpiger, slightly swollen; length ratio of palpal segments 1 : 0.7 : 0.6; 3rd segment slightly swollen distally, rounded apically. Hind leg 1.5 times as long as fore one. All femora curved (Fig. 7, 14), 0.7–0.9 times as long as tibiae. Fore femur 0.59 times as long as hind one. Fore and middle femora nearly as long as 2nd segment of tarsus; hind femur 0.91 times as long as 2nd segment of tarsus. Hind tarsus 1.7 times as long as fore one. 2nd segment of fore tarsus nearly twice as long as 2nd segment of hind tarsus and 4.3 times as long as 1st segment. 2nd segment of hind tarsus 7.4 times as long as 1st and 3.4 times as long as 5th segment. Combined length of 3rd–5th segments of all tarsi always 1.2– 1.4 times length of 2nd segment. Ovipositor short, with large rounded apical lobes 1.1 times as long as wide. Lobes densely covered with scales and with elongate setae bent apically. Ventral plate slightly elongate. Differential diagnosis. The new species is closely related to Cornistyla flabellata (Fedotova et Sidorenko, 2006), comb. n. (Brachineura, subgenus Cornistyla) described from Primorskii Territory (the Lazovskii Nature Reserve) (Fedotova and Sidorenko, 2006a) (Fig. 6, 6–10), but differs in a much longer 1st flagellar segment, in shorter, swollen, instead of parallel-sided, middle flagellar segments, in a shorter and narrower gonostylus with a wider and longer hypoproct, in a shorter vein R1+2, and in the wing not widened distally. Etymology. The name of the species is a Latin noun of feminine gender, means “a small flame, spark.”
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Tribe UNDONEURINI Fedotova, trib. n. Diagnosis. Flagellar antennal segments not covered with scales; middle segments with oval or rounded basal enlargements bearing horseshoe-shaped lunulae—peritremes (Figs. 1, 3, 4; 3, 2, 3) and with distinct necks. Basal enlargement bearing sensory threads constituting net or formed by 1 or 2 rings connected by longitudinal crosspieces; rarely threads absent. Basal enlargement in male and female with middle whorl of setae projecting from large peritremes (Fig. 17, 18, 20). Neck of flagellar segment in male at least slightly longer than wide (Figs. 1, 3, 4, 17, 18; 17, 20) or longer (Brachyneurina, Coccidomyia) (Fig. 5, 2, 10); that in female very short (Fig. 4, 20, 21). In male, apical whorl of setae not developed. Proximal flagellar segments longer than distal ones (Fig. 3, 2, 3, 8). Labrum triangular, short, not concealing labellum and base of palpus. Labellum wide, elongate, projecting far beyond labrum (Fig. 3, 1, 15). Palpus 3- or 4-segmented (Figs. 1, 5; 3, 1). Vein R4+5 running into wing margin near or far before its apex (Fig. 2, 4, 5, 6, 12). Gonocoxite without basal projection (Figs. 1, 1; 3, 4, 6; 5, 1, 3). Hypoproct complete or emarginate; mediobasal pubescent lobes or processes bearing setae present (Figs. 1, 1, 19, 20; 5, 1, 3, 9; 17, 21, 22) or absent (Fig. 17, 10, 12). Cerci rather small, situated between gonocoxites, not overlapping half width of gonocoxites on inner side. Differential diagnosis. The new tribe differs from the nominotypical one in wider flagellar antennal segments characterized by a long neck and a swollen basal enlargement not covered with scales but bearing swollen peritremes, in the absence of an apical whorl of setae on the basal enlargement, in an obvious undulate curvature of vein R4+5 at the base of the wing, in a short triangular labrum which does not conceal the labellum and palpal bases, in narrow cerci which do not extend onto the inner side of the gonocoxites, and in a usually shallowly emarginate or complete hypoproct being shorter than the cerci. The tribe comprises 2 subtribes: Undoneurina (5 genera and 11 species) and Coccidomyiina (2 genera and 8 species). Subtribe Undoneurina Fedotova, subtrib. n. Diagnosis. Antenna 2+10 or 11-segmented. Flagellar segments with short (Fig. 1, 3, 4, 17, 18) or long necks (Fig. 5, 2). Basal enlargement densely covered with strongly swollen peritremes bearing long or short
setae and rings of sensory threads connected by longitudinal crosspieces (Fig. 1, 3, 4). Apical flagellar segment oviform (Fig. 3, 8). Wing widest near middle or in distal part, entirely covered with scales densest on veins. Vein R4+5 running into margin of wing far before or near its apex, undulate at base (Fig. 2, 2, 4, 6, 12). Claw with large tooth at base, empodium as long as or shorter than claw. Vein Cu simple, without furcation, arcuately curved in distal 1/3 before confluence with margin of wing. Gonocoxite densely covered apicolaterally with very long setae or without setae. Hypoproct complete or emarginate; medial pubescent rounded or triangular lobes developed (Figs. 1, 1; 3, 4, 6; 5, 1, 3). Ovipositor non-telescopic, with large ventral projection and with small rounded and pointed apical plates directed dorsocaudally (Fig. 4, 1–4, 6). The subtribe comprises 5 genera and 11 species: the recent Brachyneurina Mamaev, 1967 (3 species), Chrybaneura Gagné, 1968 (1), Cingola Fedotova et Sidorenko, 2006a (4), and Undoneura Fedotova et Sidorenko, 2006a (2) and the Late Eocene Popovineura Fedotova et Perkovsky, 2014 (1) from Rovno amber. A Key to the Genera of the Subtribe Undoneurina subtrib. n. 1. Hypoproct complete, narrow. Flagellar segments with long necks 1/3–1/2 as long as basal enlargements (Fig. 5, 2) .................................................... 2. —Hypoproct
emarginate. Vein R4+5 running into margin of wing far from its apex (Fig. 1, 5, 6). Flagellar segments with short necks (Fig. 1, 3, 4, 17, 18) ... 4.
2. Gonocoxite narrow, without long setae apicolaterally (Fig. 5, 1, 3). Gonostylus arcuately curved near base. Hypoproct and cerci subequal in length; aedeagus much longer than cerci. Palpus 2- or 3-segmented. Middle flagellar segments with long necks more than half as long as basal enlargements (Fig. 5, 2) .............................................................. 3. —Gonocoxite
wide, densely covered with very long setae apicolaterally, without rounded lobe at base (Fig. 3, 4, 6). Gonostylus nearly straight, slightly thickened at base or regularly roundly thickened. Hypoproct longer than cerci; aedeagus distinctly shorter than cerci. Palpus 4-segmented (Fig. 3, 1). Middle flagellar segments with short necks slightly longer than wide (Fig. 3, 2, 3) .................................. ......................... Undoneura Fedotova et Sidorenko. ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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3. Gonocoxite with rounded basal lobe. Gonostylus strongly swollen and curved near base. Vein R4+5 straight, running into wing margin near its apex (Fig. 2, 12). Palpus 3-segmented. Antenna 2+10-segmented. Vein Cu straight ........................... ........................................... Brachyneurina Mamaev.
The genus comprises 2 species: the type species U. fasciculata (Fig. 3, 1–5) and U. phaseoliformis (Fedotova, 2004) (Brachineura), comb. n. (Fig. 3, 6–9) both described from Primorskii Territory, from the Lazovskii Nature Reserve and from the environs of the Ussuri Nature Reserve, respectively.
—Gonocoxite
The species of the genus Undoneura differ in the following characters.
without basal lobe. Gonostylus not swollen at base, arcuately curved. Vein R4+5 curved near middle, running into margin of wing far not reaching its apex. Palpus 2-segmented. Antenna 2+11-segmented. Vein Cu arcuate ........................... ....................... Popovineura Fedotova et Perkovsky.
4. Aedeagus long, projecting far beyond apices of cerci and hypoproct. Gonostylus very long and fine, weakly swollen at base (Fig. 1, 1). Gonocoxite with pubescent basal lobe (Fig. 17, 21, 22) ...................... ............................... Cingola Fedotova et Sidorenko. —Aedeagus
short, reaching apices of cerci and gonocoxites (Fig. 1, 19). Gonostylus short, strongly swollen at base. Gonocoxite basally with fine projection bearing long setae (Fig. 1, 20) ...................... ................................................. Chrybaneura Gagné. Genus Undoneura Fedotova et Sidorenko, 2006
Fedotova and Sidorenko, 2006a : 94. Type species Undoneura fasciculata Fedotova et Sidorenko, 2006a : 94 (by monotypy). Diagnosis. Antenna 2+10-segmented. Flagellar segments in male with strongly swollen basal enlargements covered with numerous horseshoeshaped lunulae of elongate setae forming middle whorl (Fig. 3, 2, 3). Base of basal enlargement with row of short setae of basal whorl. Basal flagellar segments longer than wide, middle ones shortened and more strongly swollen, apical segments subparallel-sided (Fig. 3, 8). Middle flagellar segments with short necks slightly longer than wide. Palpus 4-segmented (Fig. 3, 1). Claw with tooth, as long as empodium (Fig. 3, 5). Vein R4+5 strongly undulate in proximal 1/3, then straight, running into margin of wing far before its apex (Fig. 2, 2, 4). Gonocoxite wide, subparallel-sided, densely covered with scales, apicomedially covered with long black hairs. Gonostylus straight, slightly widened at base or roundly thickened, bean-shaped. Cercus cordate. Hypoproct complete, slightly sclerotized. Aedeagus short, sclerotized, triangular or narrow and parallel-sided. Base of genitalia with wide sclerotized plate. ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
1. Gonostylus straight, slightly widened at base (Fig. 3, 4). Hypoproct widened toward base. Aedeagus triangular. 4th palpal segment twice as long as 3rd one (Fig. 3, 1). Body length 0.95 mm, length of wing 1.0 mm .............................................. ..................... U. fasciculata Fedotova et Sidorenko. —Gonostylus
bean-shaped (Fig. 3, 6). Hypoproct narrowed toward base. Aedeagus parallel-sided. 4th palpal segment slightly longer than 3rd one. Body length 0.54 mm, length of wing 0.77 mm ................. .................................... U. phaseoliformis Fedotova. Genus Brachyneurina Mamaev, 1967
Mamaev, 1967 : 876. Harris and Evans, 1979 : 23. Type species Brachyneurina xylophila Mamaev, 1967 : 876 (by monotypy). Diagnosis. Antenna 2+10-segmented. Flagellar segments cylindrical, with short necks less than 1/3 as long as basal enlargements (Fig. 5, 2). In female, flagellar segments with shorter necks (Fig. 4, 20). Gonocoxite slender or wide (Fig. 5, 1, 3), with pubescent triangular mediobasal lobe. Gonostylus long, widened at base. Cercus widened toward base, with triangular emargination between rounded lobes. Hypoproct with rounded apex, short, slightly widened toward base, as long as cercus. Aedeagus longer than cercus, narrow, widened toward base, with rounded or truncate apex. Ovipositor with rounded apical plates (Fig. 4, 1, 2). Larva. Body strongly flattened dorsoventrally. Head long, rounded at end; exocephal rods long. Palpus 1-segmented, as long as wide. Each side of ultimate palpal segment with 2 large and 2 small papillae each bearing seta. Differential diagnosis. The genus Brachyneurina is similar to Undoneura Fedotova et Sidorenko in the structure of the complete hypoproct, but differs in longer necks of the flagellar segments, in the presence
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of denser swollen peritremes on the basal enlargements, in the presence of a basal projection at the base of the gonocoxite, in the absence of sclerotized stripes on the inner side of the gonocoxite, in the aedeagus considerably elongate in comparison with the cerci, in the gonostylus arcuately curved at the base, and in vein R4+5 more closely situated to the apex of the wing. This Palaearctic genus comprises 3 species, among which 2 occur in Russia. The larvae of Brachyneurina peniophorae Harris, 1979 are associated with the fungi Peniophora cinerea, and the adults of B. xylophila were reared from the flat larvae found on the surface of decaying alder knots (Mamaev, 1967; Harris and Evans, 1979). The larvae of Brachyneurina are rather abundant in the litter, but they prefer small knots and bark fragments decaying in the litter rather than decaying leaves; the larvae were also found in tree stubs split with an axe (Mamaev and Krivosheina, 1965). Genus Chrybaneura Gagné, 1968 Gagné, 1968 : 33. Gagné, 1994 : 57 (clarified diagnosis); Gagné and Ėtienne, 2009 : 342 (description of larva and biology).
Differential diagnosis. The genus Chrybaneura is closely related to Brachyneurina Mamaev, but differs in an emarginate hypoproct with pointed lateral lobes, in vein R4+5 running into the costal vein farther from the wing apex, and in the absence of a rounded pubescent lobe at the base of the gonocoxite. This Neotropical genus includes one species, Chrybaneura harrisoni Harris described from Panama and found later in Guadeloupe, Costa Rica, and Ecuador (Gagné and Ėtienne, 2009). Biology poorly studied. This genus is associated with the films which remained from spider eggs and also with spider egg batches (Gagnė, 1968). The adults were reared from the larvae found among the remains of old dead colonies of Iceria setchellarum (Westwood) (Hemiptera : Monophlebidae); they are also associated with the coccids infecting Pinus taeda L. (Pinaceae). The exuvia were associated with Orgyia sp. (Lepidoptera : Lymantriidae) (Gagné and Ėtienne, 2009). Genus Cingola Fedotova et Sidorenko, 2006 Fedotova and Sidorenko, 2006a : 96. Jiao and Bu, 2014 : 155.
Type species Chrybaneura harrisoni Harris, 1968 : 34 (by monotypy).
Type species Cingola certa Fedotova et Sidorenko, 2006 : 96 (by monotypy).
Diagnosis. Antenna 2+10-segmented. Flagellar segments with short necks; basal enlargement covered with short setae originating from numerous swollen peritremes (Fig. 1, 18), its base and apex bearing sensory threads in shape of rings connected by 2 longitudinal crosspieces. Necks shorter in female than in male (Fig. 4, 21). Vein Cu straight. Anepimeron with 7–9 setae. Gonocoxite short and strong, with short basal projection bearing setae (Fig. 1, 20). Gonostylus short, sharply narrowed distally. Cercus as long as hypoproct, with triangular narrow lobes separated by deep emargination (Fig. 1, 19). Ovipositor nontelescopic, with 2 dorsolateral cerci and with long ventral plate. Ventral to apical plates, 2 large triangular lobes (divided segment VIII) projecting beyond them (Fig. 4, 6).
Diagnosis. Antenna 2+10-segmented. Necks of flagellar segments subsquare in male (Fig. 1, 3, 4), very short in female (Fig. 4, 15–17). Labrum long and wide, with straight margin (Fig. 1, 5) or triangular; palpus 3-segmented. Vein R4+5 running into margin of wing far before its apex (Fig. 2, 6); Cu arcuately curved distally. Hypoproct more heavily sclerotized than cerci, with V- or U-shaped emargination (Fig. 1, 1). Base of gonocoxite with pubescent lobe; chrysanthemum-shaped sclerotized structure occasionally present dorsal to aedeagus (Fig. 17, 21, 22), or absent (Fig. 1, 1). Ovipositor with rounded apical lobes (Fig. 4, 4).
Larva and pupa recently described (Gagné and Ėtienne, 2009). Body flattened dorsoventrally. Head capsule conical, with long exocephalic rods. Length of antenna twice its diameter. Spatula bidentate. Terminal segment with 6 papillae: 2 papillae with long setae, and 4, with short setae.
Differential diagnosis. The genus Cingola is closely related to Chrybaneura Gagné, but differs in a very long fine gonostylus, cordate cercus, long aedeagus much longer than the cercus, and in the presence of a large pubescent basal lobe of the gonocoxite. After the description of two new species from Southern China (Jiao and Bu, 2014), the diagnosis of the genus was clarified owing to inclusion of the new ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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characters clearly distinguishing these species from the type species from Primorskii Territory; a key to 3 species was also given (Jiao and Bu, 2014). The structures with an unclear origin and position were listed. C. certa possesses well-defined sclerotized lateral stripes at the sides of the aedeagus, which resemble a hypoproct but entirely adjoin the aedeagus. In the description of C. circularis and C. ceratodicrata, a lateral sclerotized stripe on the aedeagus is not mentioned. The aedeagus bears fine, non-branched lateral processes at the apex (C. certa) (Fig. 1, 1) or branched processes at the middle (C. circularis, C. ceratodicrata). A sclerotized chrysanthemum-shaped structure situated dorsally to the aedeagus at the base of the gonocoxite is indicated to be present in C. ceratodicrana and C. circularis (Fig. 17, 21, 22); C. certa does not possess this structure. Jiao and Bu (2014) link its origin with the tegmen. The basal lobe of the gonocoxite is rounded and pubescent in C. certa, and is slightly curved and pointed in C. ceratodicrata and C. circularis. The genus comprises 4 species: Cingola certa Fedotova et Sidorenko, 2006a known from Primorskii Territory (the Lazovskii Nature Reserve) (Fig. 1, 1–5), C. circularis Jiao et Bu, 2014 : 157 (Fig. 17, 20, 21) and C. ceratodicrata Jiao et Bu, 2014 : 160 (Fig. 17, 22) both recently described from Southern China, and C. pleiomorpha Mamaev, 1998 : 6, comb. n. (Brachyneurina) described from Primorskii Territory, in the Kedrovaya Pad Nature Reserve (Mamaev, 1998; Fedotova and Sidorenko, 2006a; Jiao and Bu, 2014). The latter species was attributed to the genus Cingola according to the following characters: baculiform appendages are present at the sides of a needle-shaped pointed aedeagus, the basal triangular lobe of the gonocoxite is rounded at the end, the gonostylus is long and fine in the distal half, the basal enlargements of the antennal segments are long, cylindrical, with very short necks, and vein R4+5 runs into the wing margin at the level of its distal 1/4. A key to the species was compiled by Jiao and Bu (2014). Genus Popovineura Fedotova et Perkovsky, 2014 Fedotova and Perkovsky, 2014 : 392. Type species—Popovineura nacta et Perkovsky, 2014 : 393 (by monotypy).
Fedotova
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Diagnosis. Antenna 2+11-segmented. Middle flagellar segments with elongate necks more than half as long as basal enlargements. Palpus 2-segmented. Claw simple. Vein R4+5 running into margin of wing far from its apex. Vein Cu arcuate, situated far from margin of wing and running into its margin. Gonocoxite slender, without rounded basal lobe. Gonostylus not widened at base, fine, arcuately curved. Cercus cordate. Hypoproct complete, nearly as long as cercus. Aedeagus much longer than cercus and hypoproct, projecting beyond apex of gonocoxite. Differential diagnosis. The genus Popovineura is closely related to Brachineura, but differs in some characters of the wing venation, in a narrow gonostylus, 2+11-segmented antenna, 2-segmented palpus, and simple claw. The genus includes only the type species described from the Late Eocene Rovno amber (Ukraine, amberbearing deposits near Rovno) (Fedotova and Perkovsky, 2014). Subtribe Coccidomyiina Fedotova, subtrib. n. Diagnosis. Antenna 2+10–12-segmented. Flagellar segments with long cylindrical basal enlargements and shorter necks (Fig. 5, 10) in male, with longer basal enlargements and shorter necks in female (Fig. 4, 18). Vein R4+5 straight, fused with costal vein beyond wing apex; Cu simple, with indistinct furcation, straight, and not reaching margin of wing, or bifurcate. Palpus 2–4-segmented (Figs. 5, 8; 17, 15, 16). Claw with tooth in fore tarsus, simple in middle and hind tarsi. Gonocoxite large. Gonostylus short and narrow (Figs. 5, 9; 17, 9–12). Cercus and hypoproct bilobed, or hypoproct complete or shallowly emarginate. Ovipositor non-telescopic. Apical lobes triangular, situated dorsoapically, directed caudally, covered with numerous short setae, occasionally with apical spines (Fig. 17, 17). Differential diagnosis. The new subtribe differs from Undoneurina in vein R4+5 being straight at the base and running into the wing apex, in the base of the gonocoxite bearing neither a pubescent lobe nor a projection with setae, in an usually bifurcate vein Cu (Cu2 is indistinct or absent in Coccidomyia), in the gonostylus entirely covered with microtrichiae, in a very wide hypoproct only slightly narrower than the cerci, and in a very short aedeagus far from reaching the cerci.
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Two genera are included in the subtribe: Coccidomyia Felt, 1911 (1 species) and Megommata Barnes, 1939 (6).
This Nearctic genus includes only the type species known from the type series. It was redescribed and illustrated based on the type material (Harris, 1968).
The genera of the subtribe Coccidomyiina differ in the following characters.
Biology unknown. The species was reared from beech (probably Fagus grandifolia) leaves intensely infected with young coccids of the genus Lecanium (Hemiptera: Coccidae). The presence of exuvia of dipterans, possibly gall midges, projecting from under the coccids, suggest that this species is a predator or parasite.
1. Vein Cu simple, disappearing far from running into wing margin (Fig. 2, 10). Antenna 2+10-segmented. Palpus 2- or 3-segmented (Fig. 5, 8), with very short segments. Hypoproct clearly emarginate at apex (Fig. 5, 9). Roots of aedeagus not developed. Gonocoxite with narrow basal projection bearing setae ..................................................... Coccidomyia Felt. —Vein
Cu with furcation (Fig. 17, 19). Antenna 2+12-segmented. Palpus 4-segmented, with long segments (Fig. 17, 15, 16). Hypoproct with rounded apex (Fig. 17, 10, 12). Roots of aedeagus in shape of sclerotized furcation. Gonocoxite without basal projection ............................... Megommata Barnes. Genus Coccidomyia Felt, 1911 Felt, 1911 : 45. Harris, 1968 : 411; Gagné, 1994 : 57.
Type species Coccidomyia pennsylvanica Felt, 1911 : 45 (by monotypy). Diagnosis. Antenna 2+10-segmented. Palpus very short, with short segments, usually 2-segmented, but with very small rudimentary 3rd segment in some specimens (Fig. 5, 8). Vein Cu simple (furcation indistinct or absent: Fig. 2, 10). Cercus with deep oval emargination, hypoproct with small, nearly semicircular emargination or shallowly emarginate (Fig. 5, 9). Aedeagus weakly gradually widened toward base, rounded at end, slightly shorter than hypoproct and distinctly shorter than cerci. Abdominal segment IX (base of ovipositor) very wide, straightly truncate dorsoventrally, densely covered with short setae. Cercus very small; its length less than half width of abdominal segment IX in place of their joint. Differential diagnosis. The genus Coccidomyia is closely related to Chrybaneura Gagné, but differs in very short palpal segments, elongate necks of the flagellar segments, in a wide hypoproct rounded laterally and shallowly emarginate apically (in Chrybaneura, the hypoproct is narrow, widened toward the base, deeply emarginate apically: Fig. 5, 9), and in vein R4+5 running into the wing apex.
Genus Megommata Barnes, 1939 Barnes, 1939 : 333. Harris, 1968 : 455. Type species Megommata seychelli Barnes, 1939 : 334 (by monotypy). Diagnosis. Antenna 2+12-segmented. Eye bridge shifted onto facial side of head, with non-faceted area in middle. Palpus 4-segmented (Fig. 17, 15, 16). Flagellar segments of female with long cylindrical basal enlargements and with very short necks (Fig. 17, 18). Circular sensory thread simple, situated near bases of basal enlargements. Vein R4+5 fused with costal vein slightly beyond wing apex; Cu bifurcate (Fig. 17, 19). Claw with tooth in fore tarsus, simple in middle and hind tarsi. Gonocoxite widely oval; gonostylus swollen at base, very fine distally (Fig. 17, 9–12). Cercus elongate, not cordate, widened or slightly narrowed toward base, with deep emargination between triangular or pointed lobes. Hypoproct very wide, widely rounded or slightly bent at apex, slightly narrower and frequently longer than cercus. Aedeagus short, slightly widened toward base, with paired sclerotized roots and with rounded or pointed apex, frequently with pores in distal half. Ovipositor with narrow cerci bearing several terminal spines (Fig. 17, 17). Differential diagnosis. The genus Megommata is closely related to Coccidomyia, but differs in a very well-developed hypoproct occasionally slightly narrower but longer than the cerci, in shorter necks of the flagellar antennal segments, and in a distinct furcation of the cubital vein. The genus comprises 3 Afrotropical and 3 Oriental species developing on coccids (Coccoidea : Pulvinaria spp., Planococcus sp., Pseudococcus sp.). In Indonesia, M. leefmansi (Nijveldt, 1964) was reared from the ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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Fig. 9. Stelladiurna flustra gen. et sp. n., male: (1) genitalia; (2) 1st segment of tarsus; (3) mouthparts; (4) 1st and 2nd flagellar segments; (5) 3rd flagellar segment; (6) scape and pedicel; (7) wing. Scale 0.1 mm [(a) to 1–6, (b) to 7].
larvae feeding on eggs of Pulvinaria polygonata (Cockerell) (Harris, 1968). Tribe RHIZOMYIINI Mamaev, 1968 Diagnosis. Antenna 2+10–12-segmented. Flagellar segments without scales, with oval or rounded basal enlargements bearing horseshoe-shaped lunulae, and with distinct elongate necks. Basal enlargement bearing sensory threads in shape of two rings connected by longitudinal crosspieces; rarely only one ring present, or threads absent. Proximal flagellar segments shorter than middle ones and slightly longer than apical segENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
ments. Palpus 2–4-segmented Labrum triangular; labellum usually strongly swollen. Vein R4+5 running into margin of wing before or nearly in wing apex (Figs. 2, 11; 11, 10; 13, 1); Cu usually bifurcate, when it non-bifurcate, then it reaching margins of wing. Gonocoxite wide or swollen laterally, usually with sclerotized stripe along inner margin or near middle (Figs. 5, 5; 10, 14; 11, 14, 15; 12, 6, 11, 14, 18; 13, 1). Gonostylus large, swollen at base, occasionally with lobe or bifurcate, with sclerotized stripe at basal margin (Figs. 5, 5; 11, 14, 15). Base of gonocoxite without pubescent mediobasal lobe, except in Alatostyla bi-
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naria (Mamaev) (Fig. 17, 1): its aedeagus with 2 densely pubescent appendages at base. Cercus and hypoproct emarginate. Hypoproct with elongate oval emargination and with narrow lateral lobes (Figs. 5, 5; 11, 14, 15). Gonocoxite without basal projection. Aedeagus small, fine, heavily sclerotized or with fine apex (Figs. 5, 5; 8, 1; 17, 1); when aedeagus large, aedeagal complex strongly complicate (Fig. 9, 1). Differential diagnosis. The tribe Rhizomyiini differs from Undoneurini in longer flagellar segments with very long necks and with usually cylindrical basal enlargements, in the presence of an apical whorl of setae on the basal enlargements, in vein R4+5 forming no undulate curvature at the base and running into the wing margin near its apex, in a usually strongly complicate aedeagal complex, or in the shape of the gonostylus and in the absence of a mediobasal projection of the gonocoxite. The tribe comprises 2 subtribes, 7 genera, and 36 species: Rhizomyiina (4 genera and 33 species), Stellaserenina (3 genera and 3 species). Subtribe Stellaserenina Fedotova, subtrib. n. Diagnosis. Necks of flagellar segments (Figs. 8, 4, 6; 9, 4, 5) slightly shorter than or as long as (in recent genera) basal enlargements; or neck very short but basal enlargement long and varying in shape on different segments. Labrum triangular; labellum long, occasionally swollen laterally, subequal in length to labrum (Figs. 8, 2; 9, 3). Palpus 3-segmented, with elongate segments (Figs. 8, 2; 9, 3). Vein R4+5 running into margin of wing beyond wing apex (Figs. 8, 7; 9, 7); Cu simple, not reaching margin of wing. Genitalia with complicate aedeagal complex. Gonocoxite elongate (Figs. 8, 1; 9, 1). Gonostylus large, straight, regularly narrowed toward apex. Cercus very wide, cordate (Fig. 8, 1) or widened distally (Fig. 9, 1), with lateral lobes. Hypoproct with parallel narrow lateral lobes separated by oval emargination. Ventral plate with pointed lobes separated by semicircular emargination. Tegmen in shape of wide complete sclerotized plate concealing genitalia dorsally. Aedeagus small, with narrow pointed apex. Bifurcate sclerotized roots of aedeagus developed. Differential diagnosis. The subtribe Stellaserenina differs from Rhizomyiina in vein R4+5 running beyond the wing apex, in the absence of sclerotized chords near the middle of gonocoxite, in the absence of a furcation of vein Cu, in a very large, even, nearly
straight gonostylus, and in a well-developed complicate aedeagal complex, including a large tegmen and also a ventral plate with the pointed lobes separated by a semicircular emargination. The subtribe comprises 3 genera and 3 species: the recent Stellaserena gen. n. (1) and Stelladiurna gen. n. (1) and the Late Eocene Rovnobrachineura Fedotova et Perkovsky, 2014 (1) from Rovno amber. Biology unknown. The species of the recent genera were collected with traps. A Key to the Genera and Species of the Subtribe Stellaserenina 1. Flagellar segments very strongly differing in shape. Necks very short. Basal enlargements of proximal segments short, constricted near middle; distal segments with strongly swollen rounded basal necks and with narrow continuations of basal enlargements. Aedeagus large, wide, conical, projecting very far beyond apex of gonocoxite. Cercus cordate. Palpus 2-segmented, with long segments. From the Late Eocene Rovno amber ........................................ ............. Rovnobrachineura Fedotova et Perkovsky. —Flagellar
segments almost not differing in shape (Figs. 8, 4, 6; 9, 4, 5). Aedeagus narrow, not projecting beyond apex of gonocoxite, rarely slightly longer than it (Fig. 9, 1). Palpus 3-segmented. Gonostylus wide, long, gradually narrowed toward apex (Figs. 8, 1; 9, 1). Ventral plate strongly, almost trapeziformly widened from apex to base, with pointed apex and with wide semicircular emargination. Basal enlargements and necks of middle flagellar segments of antenna subequal in length (Figs. 8, 6; 9, 5). Recent genera ............................ 2.
2. Cercus widened toward apex, concave, passing into lateral processes, strongly narrowed proximally (Fig. 9, 1). Tegmen with long filiform processes at apex. Gonocoxite without pubescent apicomedial projection. Vein Cu far distant from margin of wing (Fig. 9, 7). Labellum strongly swollen (Fig. 9, 1). Eye bridge not broken on vertex ............................... .................................................. Stelladiurna gen. n. —Cercus
cordate, with rounded emargination between large wide rounded lobes (Fig. 8, 1). Gonocoxite with apicomedial pubescent projection. Tegmen in shape of sclerotized oval plate. Vein Cu very approximate to margin of wing (Fig. 8, 7). Labellum narrow, triangular (Fig. 8, 2). Eye bridge broken ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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off on vertex by sclerotized non-faceted stripe (Fig. 8, 3) .................................. Stellaserena gen. n. Genus Stellaserena Fedotova, gen. n. Type species Stellaserena vaga sp. n. Diagnosis. Male. Eye bridge in shape of narrow sclerotized stripe equal in width to 4 facets, non-faceted, bearing small triangular accessory lunulae (Fig. 8, 3). Scape and pedicel less heavily sclerotized than flagellar segments which equally uniformly sclerotized. 1st flagellar segment without pale stripe at apex of neck, slightly shorter than 2nd segment. Middle flagellar segments with long necks 0.8–0.9 times as long as widely oval basal enlargements. Palpus 3-segmented, slightly widened laterally (Fig. 8, 2), without palpiger. Haltere very densely covered with dark scales, pale under scales. Legs dark because of dense cover of scales. Fore and middle femora 0.7–0.8 times as long as tibiae. Fore femur half as long as 2nd segment of tarsus. 2nd segment of fore tarsus nearly 15 times as long as 1st one. Claw of fore tarsus with tooth, empodium shorter than claw. Vein R4+5 running into wing apex; margin of wing broken at place of fusion of R4+5 and costal vein (Fig. 8, 7). Vein Cu nearly straight, extending near margin of wing but disappearing before margin. Genitalia elongate (Fig. 8, 1). Gonocoxite strongly widened in distal half, with apicomedial hairy projection, with wide and deep median emargination, and with nearly semicircular wide emargination to which long and thick gonostylus attached. Cercus reaching apex of gonocoxite, cordate, with deep and wide emargination between wide rounded lobes. Hypoproct nearly as long as cercus, with narrow parallel lobes. Ventral plate with sclerotized pointed lobes, slightly narrower than cercus. Aedeagus narrow, with sclerotized rod, narrowed before apex, with fine apical process, distinctly shorter than cercus and hypoproct. Roots of aedeagus bifurcate, sclerotized. Sclerotized tegmen on dorsal side of genitalia occasionally widely oval (Fig. 8, 1). Female unknown. Differential diagnosis. The new genus is closely related to Alatostyla Fedotova et Sidorenko, 2006b, but differs in a simple, instead of bifurcate, gonostylus, in the presence of a large hairy apicomedial projection of the gonocoxite, in a nearly trapeziform, instead of parallel-sided, hypoproct, in a concave and not cordate cercus, and in the presence of a tegmen and a ventral plate. ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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The genus includes only the type species Stellaserena vaga sp. n. Etymology. The name of the genus is a noun of feminine gender, originates from the Latin expression “Stella serena” which means the morning sun. Stellaserena vaga Fedotova, sp. n. (Fig. 8, 1–7; see also table) Material. Holotype, ♂ (slide 543/K-2/1): Russia, Southern Kuriles, Shikotan Island, Krabozavodskoe Vill., Malaise trap, 18–20.VIII.2012 (Yu.N. Sundukov). Description. Male. Body dark brown. Body length 1.06 mm; length of wing 1.76 mm, width 0.65 mm. Head with widely rounded occiput. Eyes entirely well visible from facial side of head. Antennae lost; 6 remained flagellar segments more heavily sclerotized than legs (with fell scales). Pedicel nearly square, 0.91 times as long as and narrower than scape. 1st flagellar segment 3.3 times as long as wide, neck 0.62 times as long as basal enlargement which twice as long as wide. 2nd segment 3.3 times as long as wide; neck 0.83 times as long as basal enlargement. 5th flagellar segment 3.4 times as long as wide; basal enlargement 1.2 times as long as neck, 1.9 times as long as wide. Length ratio of palpal segments 1 : 1.7 : 3.1; 3rd segment uniformly thickened, pointed at end. Thorax uniformly sclerotized, dark yellow, 2.1 times as long as wide. Anepimeron without pores. Notum sclerotized as thorax. Only fore leg remained; it 3.9 times as long as body and 2.4 times as long as wing. Fore femur 0.91 times as long as middle one. 2nd segment of fore tarsus 14.9 times as long as 1st, and 6.9 times as long as 5th segment. Combined length of 3rd–5th segments of fore tarsus 0.9 times length of 2nd segment. Wing 2.7 times as long as wide. Vein R1+2 0.42 times as long as wing. Gonocoxite twice as long as wide. Gonostylus widened at base and regularly narrowed toward apex, 4.6 times as long as wide, 0.77 times as long as gonocoxite, very narrow in distal 1/4. Hypoproct weakly widened toward base, with elongate parallel-sided apical lobes separated by rounded emargination. Etymology. The name of the species is a Latin adjective of feminine gender, which means “wandering.”
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Etymology. The name of the genus is a noun of feminine gender and originates from the Latin expression “Stella diurna”—the morning star.
Genus Stelladiurna Fedotova, gen. n. Type species Stelladiurna flustra sp. n. Diagnosis. Eye bridge continuous, not broken off by sclerotized non-faceted plate. Head with widely rounded sclerotized projection. Scape and pedicel paler than heavily sclerotized flagellar segments. 1st flagellar segment of antenna uniformly sclerotized, without pale stripe. Basal enlargements of flagellar segments widely rounded (Fig. 9, 4, 5). Palpus 2- or 3-segmented, without palpiger (Fig. 9, 3). All femora 0.8 times as long as tibiae, shorter than 2nd tarsal segment. Fore femur 0.59 times as long as 2nd segment of tarsus. 2nd segment of fore tarsus more than 12 times as long as 1st one. Claw of fore tarsus with hook-shaped tooth at base, empodium subequal in length to claw. Wing widest in proximal half. Haltere very densely covered with dark scales. Vein R4+5 running into wing apex, not bearing pores. Vein Cu straight, not reaching margin of wing, far distant from its margin (Fig. 9, 7). Gonocoxite widely ovoid, with sclerotized areas at base and on inner side (Fig. 9, 1). Cercus complete, distally with long lateral processes and with arcuate emargination along margin. Hypoproct nearly as long as and 0.77 times as wide as cercus, slightly widened distally, with rounded emargination between rounded lobes. Tegmen narrow, with long filiform processes at rounded apex. Aedeagus conical, with lateral setae and pores, much longer than gonocoxite, cercus, and hypoproct. Ventral side of genitalia bearing trapeziform plate with pointed apices separated by small rounded emargination. Differential diagnosis. The new genus is closely related to Stellaserena gen. n. in the elongate shape of the gonocoxite, gonostylus, hypoproct, and ventral plate, but differs in a continuous eye bridge, in the absence of a medioapical projection on the gonocoxite, in the presence of lateral lobes on the distally widened cerci (not cordate as in Stellaserena), in the distinctive shape of both the tegmen with an apical pubescence of threads and a very large triangular aedeagus, in the absence of roots of the aedeagus, and also in the distinctive shape of the larger mouthparts, in the shape of the flagellar segments with their basal enlargements strongly rounded laterally, and in the cubital vein more distant from the wing margin. The genus includes Stelladiurna flustra sp. n.
only
the
type
species
Stelladiurna flustra Fedotova, sp. n. (Fig. 9, 1–7; see also table) Material. Holotype, ♂ (slide 544/K-2/1): Russia, Southern Kuriles, Shikotan Island, Krabozavodskoe Vill., Malaise trap, 18–20.VIII.2012 (Yu.N. Sundukov). Diagnosis. Male. Body dark yellow. Body length 1.17 mm; length of wing 1.64 mm, width 0.65 mm. Scape slightly elongate; pedicel rounded, as wide as and 0.91 times as long as scape. 1st flagellar segment of antenna 0.91 times as long as 2nd one. 1st segment 3.0 times as long as wide; basal enlargement twice as long as wide, neck 0.53 times as long as basal enlargement. 3rd flagellar segment 2.9 times as long as wide; basal enlargement 1.5 times as long as wide, as long as neck. Segments strongly swollen; 2 apical segments fused, their length ratio 1 : 1.1; 2nd segment with constriction, rounded at end. Gonostylus long, strongly swollen at base. Thorax uniformly sclerotized, dark yellow, 1.9 times as long as wide. Legs dark because of dense cover of scales. Hind leg 2.9 times as long as body, 2.1 times as long as wing. Hind femur 0.62 times as long as 2nd segment of tarsus. Hind tarsus 1.1 times as long as fore one. 2nd segment of fore tarsus 12.3 times as long as 1st and 5.7 times as long as 5th segment. Combined length of 3rd–5th segments of fore tarsus 0.9 times length of 2nd segment. Wing 2.5 times as long as wide. Vein R1+2 0.38 times as long as wing. Gonocoxite 2.2–2.5 times as long as wide, 1.1– 1.3 times as long as gonostylus. Gonostylus 4.2– 4.5 times as long as wide. Cerci with lateral lobes entirely overlapping distance between gonocoxites. Hypoproct nearly half as wide as cerci. Female unknown. Etymology. The name of the species is a Latin noun of feminine gender meaning “a calm.” Genus Rovnobrachineura Fedotova et Perkovsky, 2014 Fedotova and Perkovsky, 2014 : 388. Type species Rovnobrachineura kirieyevi Fedotova et Perkovsky, 2014 : 389 (by monotypy). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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Diagnosis. Male. Body 1.6 times as long as antenna. Antenna 2+10-segmented, with segments becoming distinctly longer toward apex of antenna. Flagellar segments long; middle segments 2.2 times as long as wide. Basal enlargements on different segments strongly differing in shape; necks very short. Each of middle antennal segments consisting of long basal enlargement with nearly spherical base and long fine continuation; neck of segment very short. Palpus 2-segmented, with long narrow segments. Wing 2.7 times as long as wide. Vein R4+5 running into wing apex, Cu extending closely to wing margin but disappearing before confluence with margin. Gonostylus long and slender, subparallel-sided. Aedeagus conical. Cercus short, cordate; hypoproct narrow. Fore femur shorter than tibia and hind femur. Differential diagnosis. The genus Rovnobrachineura is closely related to Alatostyla, but differs in the antennal segments varying in shape, in a 2-segmented palpus, in the straight cubital vein not reaching the wing apex, in a very long aedeagus, and in the absence of a dorsal lobe of the gonostylus. The genus was described from the Late Eocene Rovno amber (Fedotova and Perkovsky, 2014). Subtribe Rhizomyiina Mamaev, 1968 Mamaev, 1968 : 24. Diagnosis. Necks of flagellar segments long, about as long as basal enlargements in male (Figs. 11, 11; 12, 9) and short and widened in female (Fig. 12, 10, 16). Labrum triangular, palpus 3-segmented (Fig. 12, 7). Vein R4+5 running into wing margin near its apex (Figs. 10, 10; 13, 8) or into wing apex; Cu with furcation; Cup present. Genitalia widely transverse; gonostylus long, nearly as long as or slightly shorter than gonocoxite. Gonocoxite without basal projection (Figs. 12, 6, 11, 18; 13, 1). When inner side of gonocoxite with sclerotized spine, then sclerotized tegmen also present. Hypoproct with lateral lobes separated by emargination. Gonostylus long, with narrow claw-shaped process. When gonostylus wide or with projection, then genitalia of complex structure (Fig. 5, 5; 11, 14, 15) and vein Cu2 reduced (Fig. 2, 11). The subtribe comprises 4 genera and 33 species: the combined genus Rhizomyia Kieffer, 1898 (26 species), Stabiliola Fedotova et Sidorenko, 2006b (3 species), Alatostyla Grover, 1964 (2 species), and Pararhizomyia Mamaev, 2004, stat. n. (2). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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A Key to the Genera of the Subtribe Rhizomyiina 1. Gonostylus bifurcate (Fig. 5, 5) or with long triangular projection ventrally (Fig. 11, 14, 15, 17). Basal enlargements of middle flagellar antennal segments distinctly longer than necks (Figs. 5, 7; 11, 11). Hypoproct narrow, heavily sclerotized, with narrow deep emargination between fine lobes. Cercus cordate, with triangular emargination, as long as hypoproct. Roots of aedeagus bifurcate, projecting beyond base of genitalia ....................................... 2. —Gonostylus
slightly swollen near base (Figs. 10, 14; 12, 14; 13, 11) or narrow, nearly straight (Fig. 12, 6, 11). Basal enlargements of middle flagellar segments nearly as long as or longer than basal enlargements (Figs. 10, 15, 18; 12, 9; 13, 3). Aedeagus very narrow, its proximal part frequently invisible, apex usually sclerotized (Figs. 10, 14; 12, 6, 11, 14, 18; 13, 1). Roots of aedeagus indistinct, not projecting beyond base of genitalia ....... 3.
2. Vein Cu simple, weakly curved before confluence with margin of wing (Fig. 2, 11). Aedeagus long, unsclerotized, with strongly pointed apex, reaching apex of gonocoxite (Figs. 5, 5; 17, 1). Tegmen not developed .......... Alatostyla Fedotova et Sidorenko. —Vein
Cu bifurcate (Fig. 11, 10). Aedeagus short, heavily sclerotized, not reaching apex of gonocoxite (Fig. 11, 14, 15). Tegmen heavily sclerotized, with pointed apex ..................... Pararhizomyia Mamaev.
3. Base of gonocoxite with tegmen. Medial side of gonocoxite heavily sclerotized, with fine spine or with triangular projection near middle (Figs. 10, 14; 12, 14). Labrum short; mentum rudimentary, horseshoe-shaped (Figs. 10, 17; 12, 22). Basal enlargement of 5th flagellar segment urceolate (Fig. 10, 15, 18) ...................... Stabiliola Fedotova et Sidorenko. —Base of gonocoxite without tegmen. Medial side of gonocoxite unsclerotized (Figs. 12, 6, 11; 13, 1), without spine. Labrum wide; mentum as wide as labrum, shallowly emarginate, nearly as long as labellum (Fig. 12, 12). Basal enlargement of 5th flagellar segment regularly cylindrical (Figs. 12, 9; 13, 3) ..... ................................................... Rhizomyia Kieffer. Genus Rhizomyia Kieffer, 1898 Kieffer, 1898 : 56. Mamaev and Zaitsev, 2002 : 3 (diagnosis of the genus and a revision of the Palaearctic species).
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Fig. 10. Ledomyiini (1–19) and Rhizomyiini (20–25), males: (1, 2) Volsatiola aequa Fedotova et Sidorenko; (3–6) Ledomyia lignosa Fedotova et Sidorenko; (7–9) L. rustica Fedotova et Sidorenko; (10–13) Acinacistyla papposa Fedotova et Sidorenko; (14–17) Stabiliola serrata Fedotova et Sidorenko; (18) S. umbra (Fedotova et Sidorenko); (19) Lauthia spinigerella Mamaev; (20–25) Rabindrodiplosis orientalis Sharma et Rao; (1, 3, 7, 11, 14, 19, 24) genitalia; (2, 23) palpus; (4) aedeagus and basal projection of gonocoxite; (5, 9, 10, 15, 18) 5th flagellar segment of antenna; (6, 13, 17) mouthparts; (8, 12, 16, 25) claw; (20) cerci, hypoproct, and aedeagus; (21) 10–12th flagellar segments; (22) 1st and 2nd flagellar segments (after: Mamaev, 1967; Sharma, Rao, 1980; Fedotova and Sidorenko, 2006b, 2007).
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Fig. 11. Ledomyiini (1–6, 8, 9) and Rhizomyiini (7, 10–17), males: (1) Volsatiola aequa Fedotova et Sidorenko; (2) Stabiliola umbra (Fedotova et Sidorenko); (3) S. serrata Fedotova et Sidorenko; (4) Compositola competiva Fedotova et Sidorenko; (5) Acinacistyla papposa Fedotova et Sidorenko; (6) A. applanata (Fedotova et Sidorenko); (7) Rabindrodiplosis orientalis Sharma et Rao; (8) Ledomyia rustica Fedotova et Sidorenko; (9) L. lignosa Fedotova et Sidorenko; (10–16) Pararhizomyia pusilla (Fedotova); (17) P. improbabilis Mamaev; (1–10) wing; (11) 5th flagellar segment; (12) mouthparts; (13) palpus; (14, 15) genitalia (variants of shape); (16) claw; (17) gonostylus and apex of gonocoxite (after: Sharma, Rao, 1980; Mamaev and Zaitsev, 2002; Fedotova, 2004; Fedotova and Sidorenko, 2005, 2006b, 2007).
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Fig. 12. Ledomyiini (1–5, 20–26) and Rhizomyiini (6–19), males (1–9, 11–15, 17–23) and females (10, 16, 24–26): (1–5) Compositola competiva Fedotova et Sidorenko; (6–10) Rhizomyia operculata Fedotova et Sidorenko; (11, 12) Rh. propensa Fedotova et Sidorenko; (13–15) Stabiliola umbra (Fedotova et Sidorenko); (16–19) S. arsenjevi (Fedotova); (20–22) Acinacistyla applanata (Fedotova et Sidorenko); (23) Plesiolauthia crassicornis Mamaev; (24–26) Ledomyia eminens Fedotova et Sidorenko; (1, 6, 11, 14, 18, 21, 23) genitalia; (2) 1st and 2nd flagellar segments of antenna; (3, 9, 10, 16, 17) 5th flagellar segment; (4, 8, 13) claw; (5, 15, 19, 24) palpus; (7, 12, 22) mouthparts; (20) pedicel, 1st and 2nd flagellar segments; (25) ovipositor; (26) scape, pedicel, 1st and 2nd flagellar segments.
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Type species Rhizomyia perplexa Kieffer, 1898 : 57 (by monotypy). Diagnosis. Antenna 2+9 or 10-segmented, flagellar segments not covered with scales. Basal enlargements and necks of antennal segments of male very long, slightly differing from each other in length; neck usually 0.83–0.91 times as long as basal enlargement. Basal enlargement subcylindrical, bearing basal whorl of short setae and numerous horseshoe-shaped swellings mainly in distal half; whorl of long setae projecting from these swellings. Basal and apical parts of basal enlargement with sensory rings occasionally connected by longitudinal crosspieces. Flagellar segments, especially their necks, gradually shortened toward apex of antenna. Apical segment smoothly rounded, without process. Flagellar segments of female with distinct short necks (Fig. 12, 10, 16, 26). Labrum transverse, with straight or emarginate edge, partly concealing mandibles (Fig. 13, 7), or triangular and wide; mentum as wide as labrum, shallowly emarginate, nearly as long as labellum (Fig. 12, 12). Palpus 3-segmented, ultimate segment longer than others. Vein R4+5 nearly straight, always slightly not reaching wing apex. Vein Cu with furcation; Rs absent or fragmentary. Coxae and femora covered ventrally with long hairs. Claws of all tarsi with large hook-shaped tooth, nearly as long as empodium. Abdominal tergites VII and VIII strongly reduced, in shape of narrow stripes.
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Holarctic (26 species) and Oriental (1 species) regions; 17 species are known in the Palaearctic Region, 12 in Russia, among which 7 are described from Primorskii Territory of the Far East: Rhizomyia furcata Fedotova et Sidorenko, 2005; Rh. operculata Fedotova et Sidorenko, 2006b (Fig. 12, 6–10); Rh. propensa Fedotova et Sidorenko, 2006b (Fig. 12, 11, 12); Rh. rossica Mamaev et Zaitzev, 1998 : 4; Rh. turriformis Fedotova et Sidorenko, 2006b; Rh. umbra Fedotova et Sidorenko, 2006b, and Rh. flammula Fedotova, sp. n. (Fig. 13, 1–8) (Mamaev and Zaitsev, 1998; Fedotova, 2004; Fedotova and Sidorenko, 2006b). Descriptions of two species and figures of the genitalia for 12 Palaearctic species were given by Mamaev and Zaitsev (1998). Biology. Rhizomyia perplexa Kieffer and Rh. circumspinosa (Rübsaamen) are known as the species in which the larvae develop on the roots and in the axillae of sedges (Carex spp., Cyperaceae). Many representatives of the genus Rhizomyia are mycetophagous and inhabit the litter (Mamaev and Krivosheina, 1965). The larvae were not described. Rhizomyia flammula Fedotova, sp. n. (Fig. 13, 1–8; see also table) Material. Holotype, ♂ (slide 542/K-2/1): Russia, Southern Kuriles, Shikotan Island, Krabozavodskoe Vill., Malaise trap, 18–20.VIII.2012 (Yu.N. Sundukov).
Genitalia usually strongly transverse (Fig. 12, 6) or subsquare (Fig. 12, 11). Gonocoxite wide, oval (Fig. 12, 6, 11), oviform (Fig. 13, 1), without sclerotized projection on inner side (Fig. 12, 18). Gonostylus very long, not less than 3/4 as long as gonocoxite, slightly curved near middle, swollen at base (Figs. 12, 6, 11, 18; 13, 1). Cercus cordate, with rounded or triangular emargination. Hypoproct narrow, slightly widened toward base, with rounded emargination between narrow lobes, usually as long as or longer than cercus. Frequently lobes of hypoproct densely covered with dark spiniform microtrichiae. Aedeagus very fine, usually shorter than cerci and hypoproct, very heavily sclerotized before apex, straightly truncate apically; rarely aedeagus visible along entire length (Figs. 12, 11; 13, 1). Roots of genitalia in shape of wide subsquare plate. Ovipositor short, non-telescopic, with very large and wide apical plates.
Diagnosis. Male. Body dark yellow. Body length 0.95 mm; length of wing 1.40 mm, width 0.57 mm; length of antenna 0.90 mm. Antenna 2+10-segmented. Head with widely rounded sclerotized projection (Fig. 13, 7). Scape and pedicel paler than heavily sclerotized flagellar segments. Scape strongly widened distally; pedicel 0.91 times as long as and narrower than scape. 1st segment 0.91 times as long as 2nd one, 3.1 times as long as wide, uniformly sclerotized, without pale stripe at apex of distal neck. Basal enlargement widely oval; neck 0.62 times as long as basal enlargement. 2nd segment 3.7 times as long as wide, with basal enlargement 1.2 times as long as neck. 5th segment 3.8 times as long as wide, with neck 0.83 times as long as basal enlargement. Palpus 3-segmented, with palpiger, slightly swollen; length ratio of palpal segments 1 : 1 : 1.4, 3rd segment slightly swollen distally, rounded at end (Fig. 13, 5, 7).
The genus comprises 27 species (after: Gagné, 2010, with changes). The genus is distributed in the
Thorax uniformly sclerotized, dark yellow, 1.5 times as long as wide. Haltere very densely cov-
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Fig. 13. Rhizomyia flammula sp. n., male: (1) genitalia; (2) scape, pedicel, 1st and 2nd flagellar segments of antenna; (3) 3rd flagellar segment; (4) 8–10th flagellar segments; (5) palpus; (6) claw of hind tarsus; (7) head, lateral view; (8) wing. Scale 0.1 mm [(a) to 1–5; (b) to 6; (c) to 7; (d) to 8].
ered with dark scales. Wing 2.5 times as long as wide (Fig. 13, 8). Vein R1+2 0.43 times as long as wing, Rs indistinct. Cu forming narrow furcation; Cup developed. Legs dark because of dense cover of scales. Hind leg 1.1 times as long as fore one. All femora 0.7–
0.9 times as long as tibiae. Fore femur 0.83 times as long as hind one. All femora shorter than 2nd segment of tarsus; hind femur 0.77 times as long as 2nd segment of tarsus. Hind tarsus 1.2 times as long as fore one. 2nd segment of fore tarsus 7.1 times as long as 1st one. 2nd segment of hind tarsus 8.7 times as long ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
as 1st and 5.5 times as long as 5th times segment. In all tarsi, 3rd–5th tarsal segments combined 0.8– 0.9 times as long as 2nd segment. Claws of all tarsi with hook-shaped tooth at base, empodium subequal in length to claw. Wing widest near middle. Wing 2.5 times as long as wide, vein R1+2 0.4 times as long as wing. Gonocoxite widely ovoid (Fig. 13, 1), its base with oval sclerotized plates connected by Y-shaped sclerotized root. Gonocoxite 1.8–2.1 times as long as wide, 1.4–1.5 times as long as gonostylus. Gonostylus long, strongly swollen at base, 2.9–3.3 times as long as wide. Distal half of gonostylus very narrow. Cercus reaching approximately middle of gonocoxite, with oviform lobes separated by deep triangular emargination. Hypoproct longer than and 0.77 times as wide as cercus, gradually widened toward base, apically with oval, densely pubescent dark spines separated by rounded emargination. Aedeagus slightly longer than hypoproct, very fine, straightly truncate at apex, heavily sclerotized before apex, distinctly less heavily sclerotized proximally, gradually narrowed from base to apex (Fig. 13, 1). Female unknown. Differential diagnosis. The new species is closely related to Rhizomyia arsenjevi (Fedotova) described from the environs of the Ussuri Nature Reserve in Primorskii Territory (Fedotova, 2004) (Fig. 12, 16– 19), but differs in the absence of a sclerotized angular projection on the inner side of the wider gonocoxite, in a long aedeagus projecting over the hypoproct but not at the level of its emargination, in the absence of a constriction near the middle of the hypoproct, in a longer, as compared with the neck, basal enlargement of the 5th flagellar antennal segment, and in a much longer 3rd palpal segment. The new species differs from the other closely related species Rh. operculata Fedotova et Sidorenko, 2006b described from the Lazovskii Nature Reserve (Fedotova and Sidorenko, 2006b) in a wider and longer gonocoxite and gonostylus, considerably shorter cercus and hypoproct, and longer flagellar segments.
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Diagnosis. Antenna 2+9 or 10-segmented, flagellar segments without cover of scales. Labrum short; mentum rudimentary, horseshoe-shaped (Figs. 10, 17; 12, 22). Labellum (Fig. 10, 17) large, widely rounded. Palpus 3-segmented, with palpiger. In middle flagellar segments, necks as long as basal enlargements (Fig. 10, 18), or longer (Fig. 10, 15). Basal enlargements covered with horseshoe-shaped lunulae bearing basal and middle whorls of setae and ring-shaped sensory threads with longitudinal crosspieces. Vein R4+5 nearly straight, always slightly not reaching wing apex. Vein Cu with furcation; Rs not developed. Wing 2.5 times as long as wide. Claws in all tarsi with large hook-shaped tooth, empodium subequal in length to claw (Figs. 10, 16; 12, 13). Genitalia usually widely rounded laterally (Figs. 10, 14; 12, 14). Gonocoxite widely rounded on outer lateral side; its inner side with spiniform projection situated on heavily sclerotized lateral area. Cercus and hypoproct with lateral lobes. Medial unpaired tegmen with straightly truncate (Fig. 12, 14) or pointed apex (Fig. 10, 14) sclerotized along margin. Gonostylus short or long, regularly curved near middle and very weakly swollen at base. Aedeagus very small, fine, sclerotized, spiniform or hook-shaped. Base of gonocoxite with conical sclerotized projection (Fig. 10, 14). Roots of genitalia in shape of wide subsquare plate. Ovipositor short, non-telescopic, with very large and wide apical plates. Differential diagnosis. The genus is closely related to Rhizomyia, but differs in the presence of inner sclerotized spines on the gonocoxite and long sclerotized chords originating from them, in the hypoproct long and pointed or straightly bevelled at the apex, and in a very fine aedeagus.
Etymology. The name of the species is a Latin noun of feminine gender meaning “a small flame, spark.”
The genus comprises 3 species: Stabiliola serrata Fedotova et Sidorenko, 2006b (Figs. 10, 14–17; 11, 3), and S. umbra (Fedotova et Sidorenko, 2006b) (Rhizomyia), comb. n. (Figs. 11, 2; 12, 13–15) described from the Lazovskii Nature Reserve (Fedotova et Sidorenko, 2006b) and S. arsenjevi (Fedotova, 2004), comb. n. (Brachyneurina) (Fig. 12, 16–19) from the environs of the Ussuri Nature Reserve.
Genus Stabiliola Fedotova et Sidorenko, 2006b
A Key to the Species of the Genus Stabiliola
Fedotova and Sidorenko, 2006b : 14. Type species Stabiliola serrata Fedotova et Sidorenko, 2006b : 14 (by monotypy). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
1. Tegmen with straightly truncate apex. Gonostylus short (Fig. 12, 14). Cercus widened distally. 5th flagellar antennal segment 3.5 times as long as
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wide, with basal enlargement 1.3 times as long as neck (Fig. 10, 18). Base of aedeagus without roots or projections. Body length 1.08 mm; length of wing 1.35 mm, width 0.58 mm ................................. ........................... S. umbra (Fedotova et Sidorenko). —Tegmen
with pointed apex. Gonostylus long (Figs. 10, 14; 12, 18), nearly as long as gonocoxite. Cercus nearly cordate ........................................... 2.
2. 5th flagellar segment 3.2 times as long as wide, with basal enlargement 0.83 times as long as neck (Fig. 10, 15). Medial side of gonocoxite with spine (Fig. 10, 14). Base of aedeagus with sclerotized projection with spine at end, but without sclerotized roots. Body length 1.25 mm; length of wing 1.55 mm, width 0.50 mm ......................................... ............................. S. serrata Fedotova et Sidorenko. —5th
flagellar segment 3.4 times as long as wide, with basal enlargement as long as neck (Fig. 12, 17). Medial side of gonocoxite with triangular projection (Fig. 12, 18). Base of aedeagus without sclerotized projection but with sclerotized lobiform roots. Body length 1.7 mm; length of wing 1.6 mm, width 0.64 mm ............................. S. arsenjevi (Fedotova). Genus Alatostyla Fedotova et Sidorenko, 2006
Fedotova and Sidorenko, 2006b : 2. Type species Alatostyla velutina Fedotova et Sidorenko, 2006b : 3 (by monotypy). Diagnosis. Necks of middle flagellar segments slightly shorter than basal enlargements. Palpus 3-segmented. Vein Cu simple, strongly curved before confluence with margin of wing. All claws with tooth at base. Gonocoxite parallel-sided or obovoid, with medial sclerotized chords (Figs. 5, 5; 17, 1). Gonostylus long, bifurcate. Cercus cordate; hypoproct sclerotized, narrow, with deep narrow emargination. Aedeagus sclerotized, wider than hypoproct, with elongate pointed spine. Roots of aedeagus and genitalia developed. The genus comprises 2 species: the type species Alatostyla velutina Fedotova et Sidorenko described from Primorskii Territory (the Lazovskii Nature Reserve) and collected with a Malaise trap (Fig. 5, 4–7) and A. binaria (Mamaev, 1998) (Rhizomyia), comb. n. (Fig. 17, 1) from the “Kedrovaya Pad” Nature Reserve (Mamaev, 1998; Mamaev and Zaitsev, 2002, figure). The species of the genus Alatostyla differ in the following characters.
1. Gonostylus bifurcate; gonocoxite parallel-sided (Fig. 5, 5), without densely pubescent appendage. Aedeagus subparallel-sided, slightly widened at base. Body length 1.3–1.5 mm; length of wing 1.5 mm ............. A. velutina Fedotova et Sidorenko. —Gonostylus
wide, widened near middle (Fig. 17, 1); gonocoxite obovoid, with long, densely pubescent appendage at base. Aedeagus conical, very strongly widened toward base. Body length 1.3 mm .............. ............................................... A. binaria (Mamaev). Genus Pararhizomyia Mamaev, 1998, stat. n.
Mamaev, 1998 : 6 (as a subgenus of Rhizomyia). Type species Rhizomyia (Pararhizomyia) improbabilis Mamaev, 1998 : 6 (by monotypy). Effusomyia Fedotova, 2004 : 596, syn. n. Type species Effusomyia pusilla Fedotova, 2004 : 596 (Fedotova, 2004) (by monotypy). In the catalogs (Gagné, 2004, 2010; Gagné and Jaschhof, 2014), the subgenus Pararhizomyia Mamaev, 1998 is indicated as nomen nudum. Diagnosis. Male (Fig. 11, 10–16). Eye bridge wide. Flagellar segments of male with long necks. Labrum small, triangular, not concealing base of palpus. Labellum large, wide, projecting forward far beyond labrum. Palpus 3-segmented. Claw with long tooth, empodium shorter than claw. Vein R4+5 running into wing apex; Cu bifurcate. Gonocoxite nearly hemispheric, with inner sclerotized chords. Gonostylus with subapical digitate process, heavily sclerotized ventrally, with tuft of setae near base, with small claw-shaped process at apex. Cercus cordate. Hypoproct with long lobes. Tegmen tent-shaped, heavily sclerotized. Aedeagus short, rod-shaped, heavily sclerotized. Sclerotized biramous structure situated dorsal to aedeagus. Roots of aedeagus bilobed, heavily sclerotized. Differential diagnosis. The genus Pararhizomyia is closely related to Alatostyla Fedotova et Sidorenko (Figs. 2, 11; 5, 4–7), but differs in the presence of a tegmen at the base of the gonocoxite, in a fine sclerotized aedeagus, and in a bifurcate cubital vein. The genus comprises 2 species. The type species Pararhizomyia improbabilis (Fig. 11, 17) was described from the Khekhtsirskii Nature Reserve; P. pusilla (Fedotova, 2004), comb. n. (Fig. 11, 10–16) was described from the environs of the Ussuri Nature Reserve, collected with Malaise and window traps (Fedotova, 2004). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
Biology unknown. The species of the genus Pararhizomyia differ in the following characters. 1. Neck of flagellar segment equal in length to basal enlargement. Empodium rudimentary. Ventral process of gonostylus bearing claw-shaped process, larger than the widest part of gonostylus, longer than its ventral side before place of origin of process (Fig. 15, 17). Gonocoxite subapically narrower than gonostylus. Length of wing 1.4 mm ......................... .......................................... P. improbabilis Mamaev. —Neck
of flagellar segment shorter than basal enlargement (Fig. 11, 11). Empodium only half as long as claw (Fig. 11, 16). Ventral process of gonostylus bearing claw-shaped process, smaller than the widest part of gonostylus and shorter than its ventral side before place of origin of process (Fig. 15, 14, 15). Gonocoxite subapically wider than gonostylus. Body length 1.0–1.3 mm; length of wing 1.4 mm ........... P. pusilla (Fedotova), comb. n. Tribe LEDOMYIINI Enderlein, 1936
Ledomyiini Enderlein, 1936 : 73. Ledomyiini: Gagné, 1976, 1985. Diagnosis. Vein R4+5 far distant from costal vein, running far from wing apex (Fig. 11, 7–9) or closely to it (Figs. 11, 5, 6; 14, 8; 15, 11; 16, 11). Cu bifurcate. Flagellum with segments slightly varying in shape and gradually becoming narrower and shorter toward apex of antenna (Figs. 13, 2–4; 14, 5, 6), with circular sensory thread; or segments varying in shape, with loopshaped sensory threads (Figs. 15, 7–9; 16, 6–9). Frequently length of neck of flagellar segment less than width of basal enlargement (Ledomyia, Fig. 10, 5, 9, 21, 22) or slightly exceeding it (Acinacistyla, Figs. 10, 10; 12, 20). Gonocoxite and gonostylus slender (Figs. 10, 11; 12, 21; 14, 1, 3, 7; 15, 10), except in Ledomyia possessing characteristic short gonostylus with very wide claw-shaped process extending onto dorsal part of gonostylus (Fig. 10, 3, 7, 24). Basal projection of gonocoxite developed (Fig. 10, 4, 11). Cercus with deep emargination. Hypoproct complete (Fig. 10, 19, 20), concave (Fig. 10, 1, 3), with small rounded emargination (Fig. 12, 23), or with processes (Figs. 10, 11; 12, 21; 14, 1, 3, 7), as exception (Compositola), with triangular emargination (Fig. 12, 1). The tribe comprises 3 subtribes: Ledomyiina (8 genera, 64 species), Acinacistylina subtrib. n. ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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(2 genera, 5 species), and Kovaleviolina subtrib. n. (2 genera, 3 species). Subtribe Ledomyiina Enderlein, 1936 Diagnosis. Antenna 2+8–12-segmented, necks of flagellar segments not less than 0.67 times as long as basal enlargements. Vein R4+5 running into margin of wing far from its apex (Fig. 11, 8, 9) or approximate to its apex (Fig. 11, 1). Vein Cu bifurcate; Cu1 forming continuation of common vein; Cu2 sharply arcuately originating from it near middle of wing at acute angle (Fig. 11, 1, 4–7, 8, 9). Palpus 3- or 4-segmented. Basal projection of gonocoxite developed. Gonocoxite slender (Figs. 10, 1, 7, 19; 12, 23) or wide (Figs. 10, 3; 12, 1). Gonostylus short (its length slightly exceeding width of gonocoxite (Fig. 10, 3, 7)), or long (Fig. 10, 19). Cercus cordate (rounded laterally and with triangular emargination between rounded lobes) or widened toward base (Fig. 10, 19). Hypoproct short, complete, with rounded apex (Fig. 10, 19), or with shallow (Fig. 10, 1, 3, 7), semicircular (Fig. 12, 23), or triangular emargination (Fig. 12, 1). Gonostylus short with wide claw-shaped process, or long and fine. The subtribe comprises 8 genera with 64 species: Compositola Fedotova et Sidorenko, 2006b (1 species), Isogynandromyia Spungis, 1981 (1), Lauthia Kieffer, 1912 (29) (Fig. 10, 19), Ledomyia Kieffer, 1894 (27) (Figs. 10, 3, 7; 11, 8, 9), Plesiolauthia Mamaev, 1967 (1) (Fig. 12, 23), Prolauthia Rübsaamen, 1916 (1); Rabindrodiplosis Grover, 1964 (2), and Volsatiola Fedotova et Sidorenko, 2006b (2) (Figs. 10, 1; 11, 1). A Key to the Genera of the Subtribe Ledomyiina 1. Vein Rs not developed; R4+5 straight, running into wing margin near its apex (Fig. 11, 1, 4–6); when R4+5 running far from apex, it arcuately curved distally (Fig. 11, 8) .................................................... 2. —Vein
Rs developed; R4+5 running into margin of wing far from its apex (Fig. 11, 7). Aedeagus very wide, slightly more than twice as long as wide, pubescent, weakly sclerotized (Fig. 10, 20). Hypoproct complete, with rounded apex. Apical tooth of gonostylus wide (Fig. 10, 24). Wing 2.2 times as long as wide (Fig. 11, 7). Antenna 2+12-segmented. 2nd segment of tarsus 7.4 times as long as wide. Flagellar segments of male antenna with short necks (Fig. 10, 21, 22). Claws of tarsi simple .................... ......................................... Rabindrodiplosis Grover.
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2. Hypoproct weakly concave (Fig. 10, 1, 3, 7), semicircular (Fig. 12, 23), or complete (Fig. 10, 19). Cercus widened toward base, elongate, or lateral lobes rounded. Gonostylus elongate, sharply narrowed distally, or straight, strongly emarginate ventrally (Fig. 10, 3, 7) .............................................. 3.
6. Eyes separated on vertex by sclerotized non-faceted stripe. Antenna 2+12-segmented. Flagellar segments of male with cylindrical basal enlargements and long necks. Hypoproct complete. Basal projection of gonocoxite and aedeagus unsclerotized (Figs. 10, 19; 17, 7) ........................ Lauthia Kieffer.
—Hypoproct
—Eyes
with deep triangular emargination (Fig. 12, 1). Cercus cordate, transverse, with rounded lobes. Gonostylus strongly widened at base, subtriangular. Basal projection of gonocoxite with small apical tubercle and with loop-shaped medial pattern. Flagellar segments with widely rounded basal enlargements; necks distinctly shorter than basal enlargements (Fig. 12, 2, 3); 1st segment shorter than 2nd one ................................................. ........................ Compositola Fedotova et Sidorenko.
3. Aedeagus fine and unsclerotized (Fig. 10, 4, 7), or nearly needle-shaped and sclerotized (Figs. 10, 19; 17, 2). Basal projection of gonocoxite very fine, closely adjoining aedeagus and nearly reaching its apex. Palpus 4-segmented .................................... 4. —Aedeagus
thickened, much longer than hypoproct which reaching middle (Fig. 10, 1) or apical 1/4 of aedeagus (Fig. 12, 23). Basal projection of gonocoxite shorter or as long as cercus and hypoproct, far not reaching apex of aedeagus (Figs. 10, 1; 12, 23). Gonostylus fine distally, with small claw-shaped process. Palpus 3-segmented (Fig. 10, 2) ............................................................ 7.
4. Vein R4+5 running into margin of wing distinctly not reaching its apex, slightly curved near fusion with costal vein (Fig. 11, 8). Tegmen developed. Basal projection of gonocoxite longer than cercus and hypoproct, reaching apex of aedeagus. Gonostylus long with small claw-shaped process, or short with wide claw-shaped process (Fig. 10, 3, 7) ............. 5. —Vein
R4+5 nearly straight, running into wing margin near its apex. Tegmen not developed ................... 6.
5. Vein R4+5 nearly straight. Tegmen transparent. Gonostylus long, strongly narrowed toward apex, with small claw-shaped process. Antenna 2+11 or 12-segmented .................... Prolauthia Rübsaamen. —Vein
R4+5 arcuately curved distally. Tegmen sclerotized, varying in shape. Gonostylus wide, short, weakly narrowed toward apex; claw-shaped process usually very wide, grasping dorsal side of gonostylus (Fig. 10, 3, 7). Antenna 2+8–11-segmented ....... ..................................................... Ledomyia Kieffer.
fused on occiput, forming eye bridge. Antenna 10-segmented. Flagellar segments in both sexes without necks, very fine, narrowed from base to distal part (Fig. 17, 4–6). Hypoproct slightly concave (Fig. 17, 2). Basal projection of gonocoxite and aedeagus sclerotized ..... Isogynandromyia Spungis.
7. Basal projection of gonocoxite narrow, fine, distinctly shorter than cercus (Fig. 12, 23). Hypoproct with semicircular emargination. Base of gonocoxite with rounded lobe. Aedeagus far not reaching base of gonostyli. Antenna 2+10-segmented .................... ............................................. Plesiolauthia Mamaev. —Basal
projection of gonocoxite wide, with medial constriction and widely rounded apex (Fig. 10, 1), as long as cercus. Hypoproct weakly concave. Base of gonocoxite without rounded mediobasal lobe (Fig. 10, 1). Aedeagus reaching base of gonostyli. (Antennae lost) .......................................................... ........................... Volsatiola Fedotova et Sidorenko. Genus Ledomyia Kieffer, 1895
Kieffer, 1894 : 201 (nom. praeocc., non Loew, 1894). Ledomyia Kieffer, 1895 : 320 (replacing name). Type species Lepidomyia lugens Kieffer, 1894 : 211 (by monotypy) Diagnosis. Antenna 2+8–11-segmented in recent species, 2+12-segmented in Late Eocene species; flagellar segments short, with basal enlargements rounded laterally (Fig. 10, 5, 9). Palpus 4-segmented (Fig. 10, 6). Vein R4+5 distally arcuately curved, running into margin of wing far from its apex. Gonocoxite from slender to subsquare (Fig. 10, 3, 7). Gonostylus wide, short, weakly narrowed toward apex, emarginate ventrally; claw-shaped process usually very wide, grasping dorsal side of gonostylus (Fig. 10, 3, 7). Tegmen sclerotized, varying in shape. Aedeagus fine, unsclerotized (Fig. 10, 4, 7). Basal projection of gonocoxite very fine, closely adjoining aedeagus and nearly reaching its apex. Ovipositor long, telescopic. Abdominal segments VII and VIII strongly reduced, ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
appearing as fine elongate sclerotized structures (Fig. 12, 25). The genus is nearly cosmopolitan, comprises 27 species: 18 from the Palaearctic, 1 from Afrotropical, 1 from Neotropical, and 7 from the Nearctic Region. The three Palaearctic species are known from the Late Eocene amber. One of them, Ledomyia eocenica (Meunier, 1904), comb. n. (Ledomyiella) from Baltic amber, is attributed to the recent genus owing to the absence of vein Rs and to arcuate vein R4+5 running into the wing margin very far from its apex. The other species, L. dextra Fedotova et Perkovsky, 2008 and L. olgae Fedotova et Perkovsky, 2014, were described from Rovno amber (Fedotova and Perkovsky, 2008, 2014). Ten new species of the genus Ledomyia were recently described from Primorskii Territory of the Russian Far East, and a key to them was given (Fedotova and Sidorenko, 2005, 2007). Ledomyia volucris Mamaev, 1972 described from Primorskii Territory (the Suputinskii, now the Ussuri, Nature Reserve) was attributed to the genus Lauthia in the catalogs of the gall-midge world fauna (Gagné, 2004, 2010; Gagné and Jaschhof, 2014), but its description fits the diagnosis of Ledomyia; in particular, a claw-shaped process constitutes the half length of the gonostylus on the dorsal side, and each plate of the ovipositor bears two long dark spines (Mamaev, 1972 : 113). Biology not studied. Most of the known species were reared from recently broken branches (Gagné, 1985, 2010; Rock and Jackson, 1985). Larvae of Ledomyia collarata Gagné, 1987 were found in galls on the fungus Xylaria enterogena Mont. (Ascomycetes, Xylariaceae) (Larew et al., 1987). Larvae of Ledomyia volucris were found in the wood of a dead elm Ulmus propinqua Koidz. (Mamaev, 1972). A key to the species of the genus Ledomyia of the fauna of the Russian Far East was published earlier (Fedotova and Sidorenko, 2007). Genus Rabindrodiplosis Grover, 1964 Grover, 1964 : 193. Type species Rabindrodiplosis champakii Grover, 1964 : 193 (by monotypy). Diagnosis. Antenna 2+12-segmented. Flagellar segments of male with short necks (Fig. 10, 21, 22). Palpus 4-segmented. 2nd segment of tarsus of R. orientalis 7.4 times as long as wide. Wing 2.2 times as ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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long as wide (Fig. 11, 7). Claw simple in all tarsi or with tooth in fore tarsus; empodium shorter than claw or rudimentary. Aedeagus pubescent, weakly sclerotized, very wide, slightly more than twice as long as wide (Fig. 10, 20). Hypoproct complete, with rounded apex. Apical tooth of gonostylus wide (Fig. 10, 24). Basal projection of gonocoxite wide, pointed distally and directed sideward from aedeagus. Ovipositor cylindrical, telescopic. Differential diagnosis. The genus Rabindrodiplosis is closely related to Ledomyia Kieffer in the shape and structure of the gonostylus with a very wide clawshaped process, in a cordate cercus, in short flagellar segments, and in a 4-segmented palpus, but differs in a 12-segmented antenna, complete hypoproct, a wide basal projection of the gonocoxite, in the presence of a basal projection of the gonocoxite, and in a short wide aedeagus. The genus comprises 2 species from India : Rabindrodiplosis champakii Grover, 1964 and R. orientalis Sharma et Rao, 1980 (Fig. 10, 20–25). Biology unknown. The genus was originally attributed to the subfamily Porricondylinae, but after specification of its diagnosis and taxonomic position (Grover, 1972), the genus was included in the tribe Oligotriphini of the subfamily Cecidomyiinae. A key to its species was published by Sharma and Rao (1980). Genus Lauthia Kieffer, 1912 Kieffer, 1912 : 2. Mamaev, 1967 : 880. A list of the synonyms of Lauthia includes 7 names (Gagné and Jaschhof, 2014). Type species Ledomyia divisa Kieffer, 1904 : 372 (by monotypy). Diagnosis. Eye bridge strongly reduced. Eyes on vertex connected only by non-faceted pigmented stripe, or eye bridge consisting of 1 row of facets. Male antenna 2+12-segmented. Flagellar segments cylindrical, with long necks and with distinct circular threads. Palpus 4-segmented. Vein R4+5 fused with costal vein before wing apex. Cu bifurcate; Cu1 forming smooth continuation of common stem of cubital veins. Claw with tooth at base. Hypoproct complete. Basal projections of gonocoxites in shape of two long fine plates (Fig. 10, 19).
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This is a combined cosmopolitan genus comprising 29 species: 13 Nearctic, 11 Palaearctic, and 5 Afrotropical. In Russia, 3 species were found: L. primigenia (Mamaev, 1994) (Ledomyia) described from the Kamchatka Peninsula (Kozyrevsk Vill.: Mamaev, 1994) and L. spilota Mamaev, 1967 and L. spinigerella Mamaev, 1967 (Mamaev, 1967) (Fig. 10, 19) from Yaroslavl Prov.; they were also found in Ukraine (Gagné, 2004). Biology unknown. Lauthia acariphaga (Marikovskij, 1968) and L. acarina (Giraud, 1863) were reared from galls of phytophagous mites (Eriophyidae); L. cardui (Kieffer, 1904) was reared from old galls of fruit-flies (Diptera, Tephritidae). Some species were reared from newly cut or partly decayed wood (Gagné, 2004). Genus Plesiolauthia Mamaev, 1967 Mamaev, 1967 : 880. Type species Plesiolauthia crassicornis Mamaev, 1967 : 881 (by monotypy). Diagnosis. Eye bridge consisting of 5 rows of facets. Antenna of male 2+10-segmented. Length of basal enlargements of middle flagellar segments 4 times their diameter. Necks 0.4 times as long as basal enlargements. Circular thread appressed, forming 2 rings connected by 2 crosspieces Palpus 3-segmented. Wing long, narrow; vein R4+5 running into costal vein at wing apex; Cu indistinct. Tarsi densely covered with scales. Claw of all pairs with tooth at base, empodium shorter than claw. Gonocoxite slender, with large rounded lobe at base (Fig. 12, 23). Gonostylus slender, curved, with black claw-shaped process at end. Cercus with wide rounded emargination. Hypoproct widened toward end, bilobed. Aedeagus as long as cerci, with 2 weakly sclerotized lateral plates—rudiments of gonosternum. Differential diagnosis. This genus is similar to Lauthia in the general plan of the structure of the genitalia and in a 3-segmented palpus, but differs in a distinct separated basal projection of the gonocoxite. A distinctive character of the genus is its long and narrow wing. The genus includes only the type species Pseudolauthia crassicornis (Fig. 12, 23) from Voronezh Prov. (the Tellermanovskoe forestry: Mamaev, 1967).
Genus Prolauthia Rübsaamen, 1916 Rübsaamen, 1916 : 505. Type species Cecidomyia circumdata Winnertz, 1853 : 226 (by monotypy). Diagnosis. Body, wings, and legs covered with scales. Antenna 2+11- or 12-segmented. Wing wide; vein R4+5 running into margin of wing before its apex. Gonocoxite and gonostylus elongate. Aedeagus reaching gonostyli, surrounded with fine membrane medioventrally. Ovipositor very short, with 2 apical rounded plates. This Western Palaearctic species was described from Germany. Biology. The larvae of Prolauthia circumdata (Winnertz, 1853) are known to develop (Rübsaamen, 1916) as inquilines in galls of Dasineura srataegi (Winnertz, 1853) on Crataegus oxyacantha. Genus Compositola Fedotova et Sidorenko, 2006 Fedotova and Sidorenko, 2006b : 18. Type species Compositola competiva Fedotova et Sidorenko, 2006b : 18 (by monotypy). Diagnosis. Eye bridge consisting of 5 or 6 facets. 1st flagellar segment slightly shorter than 2nd one (Fig. 12, 2). Middle flagellar segments with widely rounded basal enlargements and short necks distinctly shorter than basal enlargements (Fig. 12, 3). Palpus 4-segmented, with subparallel-sided segments (Fig. 12, 5). Wing widened in distal half. Vein R4+5 fused with costal vein before apex of wing and forming together with it narrow cell. Claw in all tarsi with large tooth at base, empodium shorter than claw (Fig. 12, 4). Gonocoxite and gonostylus with very wide bases (Fig. 12, 1). Cercus wide but very short, with wide rounded lateral lobes. Hypoproct more heavily sclerotized than cercus, with triangular lobes, shorter than gonocoxite. Basal projection of gonocoxite longer than cercus, slightly not reaching apex of gonocoxite, with sinuous pattern near middle and with small swelling at apex, entirely covering fine unsclerotized aedeagus. Gonocoxite without mediobasal projection. The genus includes only the type species Compositola sompetiva (Fig. 12, 1–5) described from Primorskii Territory (the Lazovskii Nature Reserve). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
Genus Isogynandromyia Spungis, 1981 Spungis, 1981 : 43. Type species Isogynandromyia terricola Spungis, 1981 : 44 (by monotypy). Diagnosis. Eye bridge long, consisting of 2 or 3 rows of facets. Antenna 2+10-segmented. Flagellar segments in male and female very long, strongly narrowed from base to distal part, each with short neck and 1 whorl of long deflexed hairs much longer than segment (Fig. 17, 4–6). Middle part of each segment with 2 ring sensory threads connected by longitudinal crosspieces. Apical segment rounded at end. Palpus 4-segmented. Vein R4+5 running into margin of wing immediately before its apex; Cu pronounced only in basal half, its branches poorly defined. Claw simple, empodium shorter than claws. Gonocoxite widely oval, narrowed toward apex (Fig. 17, 2). Gonostylus half as long as gonocoxite, fine, slightly widened at base. Cercus narrower than gonocoxite, with long lobes separated by deep narrow emargination. Hypoproct as long as cercus, shallowly emarginate at apex. Aedeagus slightly widened toward base, heavily sclerotized, reaching apex of gonocoxite. Basal projection of gonocoxite fine, sclerotized, closely adjoining aedeagus and slightly not reaching its apex. Ovipositor elongate (Fig. 17, 3), telescopic, with 2 small elongate apical plates situated dorsocaudally; ventral plates very fine. The genus includes only I. terricola (Fig. 17, 2–6) described from Latvia (Brotseny). It was also found in Poland (Bogdanowicz et al., 2007). Biology. Larvae develop in the upper layer of the forest soil. Subtribe Acinacistylina Fedotova, subtrib. n. Diagnosis. Hypoproct bent, shallowly emarginate or complete, occasionally marginate apically with long hairs (Fig. 12, 21), occasionally with accessory processes and with 2 long curved lateral branches attached subapically (Figs. 10, 11; 12, 21; 14, 1, 3, 7). Gonostylus usually fine and long. Basal projection of gonocoxite well developed, occasionally sclerotized. Antenna 2+10–12-segmented. Flagellar segments of male with long necks subequal in length to basal enlargements bearing circular sensory thread, or necks fine, shortened, and basal enlargement elongate, occasionally nodose, covered with very long sensory loops (Figs. 15, 7, 8; 16, 6–9). Claw with tooth or simple in ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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all tarsi, or simple in middle and hind tarsi. Vein R4+5 running into margin of wing slightly not reaching its apex. Vein Cu bifurcate; vein Cu2 forming obvious continuation of its common stem. Veins M3+4 and Cup indistinct in the species of Acinacistyla and Kuriloneura. Gonocoxite and gonostylus very slender and long. Cercus emarginate. Hypoproct complete or with long lateral processes (Fig. 10, 11), or without processes (Fig. 12, 21) (Acinacistyla), or in shape of plate strongly widened toward apex and bearing shallow emargination at apex and long lateral apical processes (Kuriloneura) which, apparently, similar to apical lateral processes of Acinacistyla. Differential diagnosis. The subtribe Acinacistylina differs from Ledomyiina in the aedeagus widened toward the base and surrounded with a loosely adjoining sclerotized basal projection of the gonocoxite, in a strongly modified hypoproct bearing various processes and lobes, in vein R4+5 running into the wing margin near its apex, in nearly always developed vein M3+4, and in very fine and long flagellar segments of the antenna. The subtribe comprises 2 genera and 5 species: Acinacistyla Fedotova et Sidorenko, 2006b (3 species) and Kuriloneura gen. n. (2). The genera of the subtribe Acinacistylina differ in the following characters. 1. Flagellar segments of male with elongate nodose basal enlargements covered with long loops of sensory threads (Figs. 15, 7–9; 16, 6–9). Hypoproct regularly widened toward apex (Fig. 16, 1, 3). Gonocoxite without pubescent mediobasal lobe ....... ................................................. Kuriloneura gen. n. —Flagellar
segments of male with long basal enlargements each bearing 1 ring thread at base (Figs. 10, 10; 12, 20). Hypoproct narrow, with apical margination of hairs (Fig. 12, 21) or with long lateral branches (Figs. 10, 11; 14, 1, 3, 7). Gonocoxite with pubescent mediobasal lobe .................... ........................ Acinacistyla Fedotova et Sidorenko. Genus Acinacistyla Fedotova et Sidorenko, 2006
Fedotova and Sidorenko, 2006b : 11. Type species Acinacistyla papposa Fedotova et Sidorenko, 2006b : 12 (by monotypy). Diagnosis. Antenna 2+11 or 12-segmented. Flagellar segments of antenna with long cylindrical basal
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Fig. 14. Acinacistyla sundukovi sp. n., male: (1, 3, 7) genitalia (variants of shape); (2) claw of middle tarsus; (4) mouthparts; (5) 11th and 12th flagellar segments of antenna; (6) 1st and 2nd flagellar segments; (8) wing. Scale 0.1 mm [(a) to 1; (b) to 2; (c) to 3–7; (d) to 8].
enlargements bearing numerous peritremes and circular sensory thread near bases; necks of middle segments longer than or nearly as long as basal enlargements (Figs. 10, 10; 12, 20; 15, 2). Palpus 3- or 4-segmented. Claw with tooth, empodium and claws subequal in length (Fig. 14, 4). Labrum wide, split in
middle or triangular (Fig. 12, 22). Vein R4+5 running into margin of wing near its apex (Figs. 11, 5, 6; 14, 8); vein Rs developed but occasionally hardly visible; veins M3+4 and Cup developed (Figs. 11, 6; 14, 8) but frequently hardly visible. Gonostylus very fine, weakly curved; gonocoxite slender, with pubescent ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
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Fig. 15. Acinacistyla sundukovi sp. n. (1–4) and Kuriloneura trifida sp. n. (5–12), males: (1) head, dorsal view; (2) 5th flagellar segment; (3, 5) palpus; (4) scape and pedicel; (6) mouthparts; (7) 8th flagellar segment; (8) pedicel and 1st flagellar segment; (9) scape, pedicel, and 1st flagellar segment; (10) genitalia; (11) wing; (12) head, lateral view. Scale 0.1 mm [(a) to 1; (b) to 2–9; (c) to 10; (d) to 11; (e) to 12].
lobe at base. Aedeagal complex short, not reaching apex of gonocoxite. Cercus usually cordate (Figs. 10, 11; 12, 21; 14, 1, 3, 7). Hypoproct apically without emargination, with long setae (Fig. 12, 21) or with rosette of fine processes and with long curved appendages originating near this rosette (Fig. 14, 1, ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
3, 7). Aedeagus widened toward base. Basal projection of gonocoxite fine, sclerotized, loosely adjoining aedeagus. The genus comprises 3 species: Acinacistyla papposa Fedotova et Sidorenko, 2006b (Figs. 10, 10–
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13; 11, 5) and A. applanata (Fedotova et Sidorenko, 2005) (Rhizomyia) (Figs. 12, 20–22; 11, 6) described from Primorskii Territory (Fedotova and Sidorenko, 2005, 2006b) and A. sundukovi sp. n. Biology unknown. The species was collected with Malaise and light traps. Acinacistyla sundukovi Fedotova, sp. n. (Figs. 14, 1–8; 15, 1–4; see also table) Material. Holotype, ♂ (slide 546/K-3/1): Russia, Southern Kuriles, south of Shikotan Island, Tserkovnaya Bay, Malaise trap, 12–18.VII.2012 (Yu.N. Sundukov). Paratypes 3 ♂ (slides 546/K-3/2–4), as holotype. Diagnosis. Male. Body pale yellow. Body length 1.07–1.24 mm; length of wing 1.43–1.72 mm, width 0.52–0.68 mm; length of antenna 1.60 mm. Antenna 2+12-segmented. Eyes shifted onto facial side of head, eye bridge formed by 2 or 3 facets (Fig. 15, 1). Scape and pedicel paler than the heavily sclerotized flagellar segments. Scape elongate and strongly widened distally; pedicel 0.67 times as long as and 0.77 times as wide as scape. 1st flagellar segment of antenna uniformly sclerotized, without pale stripe at apex of distal neck, 0.91 times as long as 2nd segment, 2.9 times as long as wide; its basal enlargement widely oval, 1.8 times as long as wide, neck 0.62 times as long as basal enlargement. 2nd flagellar segment 3.2 times as long as wide, with basal enlargement 0.91 times as long as neck. 5th flagellar segment 4.0– 4.6 times as long as, with basal enlargement 1.9 times as long as wide, and with neck 1.1–1.3 times as long as basal enlargement. 12th segment narrowed in distal half, without apical process, 0.62 times as long as 11th segment. Mouthparts very dark; palpus paler. Labrum wide, bilobed, entirely concealing labellum (Figs. 10, 13; 12, 22; 14, 4). Palpus 3-segmented, with palpiger, slightly swollen, densely covered with scales; length ratio of palpal segments 1 : 1 : 1.3; 3rd segment pointed at apex. Scale apically truncate, regularly dentate. Thorax uniformly sclerotized, dark yellow, 1.2– 1.6 times as long as wide. Notum white at base. Lower part of thorax, and also coxae, trochanters, and underside of femora densely covered with long golden hairs. Legs dark because of dense cover of scales. Hind leg about as long as fore one. All femora 0.7–0.8 times as long as tibiae, distinctly shorter than 2nd tarsal segment. Fore femur 1.1 times as long as hind one. Hind
femur 0.62–0.71 times as long as 2nd segment of tarsus. Hind tarsus slightly longer than fore one. 2nd segment of fore tarsus 10.3–12.3 times as long as 1st one. 2nd segment of hind tarsus 12.6–13.6 times as long as 1st and 5.4–5.9 times as long as 5th segment. Combined length of 3rd–5th segments of all tarsi 1.1– 1.3 times length of 2nd segment. Claw with hookshaped tooth at base in fore tarsus, simple in middle and hind tarsi; empodium shorter than claw. Wing widest in distal half. Haltere very densely covered with dark scales. Wing 2.6–2.9 times as long as wide. Vein R1+2 0.42–0.45 times as long as wing; vein Rs present but inconspicuous. Vein Cu forming nearly right-angled furcation; M3+4 and Cup developed. Anepimeron with 6 and 3 pores arranged in 2 rows. Genitalia slightly elongate. Gonocoxite narrow, subparallel-sided, with small rounded pubescent lobe at base. Gonocoxite 2.5–3.0 times as long as wide. Gonostylus very long, slightly swollen at base, 0.77– 0.83 times as long as gonocoxite. Gonostylus 6.5– 6.8 times as long as wide. Distal half of gonostylus very narrow, subparallel-sided. Cercus nearly reaching middle of gonocoxite, cordate, with rounded lobes separated by deep triangular emargination. Hypoproct slightly longer than cercus, weakly sclerotized, apically with group of fine transparent processes; long curved branch covered with microtrichiae originating from apex at each side. Aedeagus slightly longer than hypoproct, narrow, widely rounded or with rounded enlargement at apex, far not reaching apex of gonocoxite. Basal projection of gonocoxite in shape of fine, heavily sclerotized paramere adjoining aedeagus. Female unknown. Differential diagnosis. The new species is closely related to Acinacistyla papposa Fedotova et Sidorenko, 2006b described from the Lazovskii Nature Reserve in Primorskii Territory (Fedotova and Sidorenko, 2006b) (Fig. 10, 10–13), but differs in a longer gonostylus more strongly curved at the base, in a less heavily sclerotized hypoproct not swollen at the base and bearing distinctly shorter lateral processes originating from the apex, in longer basal enlargements of the middle antennal flagellar segments, in shorter segments of the palpus, and in vein R4+5 fused with the costal vein farther from the wing apex. Etymology. The species is named after Yu.N. Sundukov (the Kuril Nature Reserve) who collected the gall midges from Shikotan Island. ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
A Key to the Species of the Genus Acinacistyla 1. Hypoproct apically with group of transparent processes and with 2 long lateral branches covered with microtrichiae (Figs. 10, 11; 14, 1, 3, 7). Neck of 1st flagellar segment of antenna shorter than basal enlargement (Fig. 14, 6). Basal enlargements of flagellar segments densely covered with swollen peritremes (Figs. 10, 11; 14, 5, 6; 15, 2). Labrum widely rounded, bilobed (Figs. 10, 13; 14, 4). Palpus 3-segmented. Vein R4+5 running into margin of wing far from its apex (Figs. 11, 5; 14, 8) ....... 2. —Hypoproct
with elongate curved apex densely marginate with long hairs (Fig. 12, 21). Neck of 1st flagellar segment of antenna longer than basal enlargement (Fig. 12, 20). Basal enlargements of flagellar segments covered with sparse swollen peritremes (Fig. 12, 20). Labrum complete, triangular (Fig. 12, 22). Palpus 2-segmented. Vein R4+5 running into wing margin near its apex (Fig. 11, 6). Body length 2.0 mm; length of wing 1.94 mm ......... ........................ A. applanata Fedotova et Sidorenko.
2. Processes of lateral branches of hypoproct very long (Fig. 10, 11). Gonostylus weakly widened at base and slightly curved. Length of gonostylus 4.6– 4.9 times as long as wide. Antenna 2+11-segmented. Body length 1.4–1.6 mm; length of wing 1.8–2.0 mm ................................................. .......................... A. papposa Fedotova et Sidorenko. —Processes
of lateral branches of hypoproct short (Fig. 14, 1, 3, 7). Gonostylus strongly swollen at base and curved. Length of gonostylus 6.5–6.8 times as long as wide. Antenna 2+12-segmented. Body length 1.07–1.24 mm; length of wing 1.43–1.72 mm ... A. sundukovi sp. n. Genus Kuriloneura Fedotova, gen. n.
Type species Kuriloneura multifida sp. n. Diagnosis. Male. Eyes curved laterally (Fig. 15, 12), widely covering lateral and occipital parts of head; eye bridge formed by 3 or 4 facets (Fig. 15, 1). Antenna 2+12-segmented. Head with pointed projection. Scape and pedicel paler than flagellar segments which heavily sclerotized, especially on apical end of neck. Scape and pedicel subequally widened, subequal in length. 2nd flagellar segment uniformly sclerotized, with dark stripe at apex of neck, as long as 1st segment. Base of each flagellar segments very dark at place of attachment of precedENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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ing segment. Mouthparts very dark; palpus paler, densely covered with scales. Labrum narrow and short, triangular, not concealing labellum; ligula developed (Fig. 15, 6). Palpus 3-segmented, densely covered with scales. Each of flagellar segment with very long loops at base, near middle, and at apex of basal enlargement; these loops in shape of indistinct whorls also connected by long loops. Basal enlargement also with 3 indistinct nodes. Necks fine, uneven, long. Flagellar segments gradually becoming longer toward apex of antenna. Flagellar segments entirely irregularly covered with strong sharp dark spiniform microtrichiae. Wing widely oval, 2.5–2.6 times as long as wide; vein R4+5 running into margin of wing far not reaching its apex; at this place, wing broken and 1 pore present; R1+2 running into margin before middle of wing. Vein Rs not developed. Vein Cu bifurcate, furcation forming acute angle. Traces of vein M3+4 (Fig. 15, 11) occasionally marked. Lower part of thorax and also coxae, trochanters, and underside of femora densely covered with long golden hairs. Genitalia nearly as long as wide, covered with scales (Figs. 15, 10; 16, 1, 3) Gonocoxite wide, widely rounded laterally, covered with large pores, narrowed toward apex or strongly widened at apex in lateral view (Fig. 16, 3). Gonostylus very fine, nearly straight or arcuate, subparallel-sided, with small dark clawshaped process at apex. Cercus with pointed lateral lobes separated by triangular emargination (Fig. 16, 1), or cordate (Figs. 15, 10; 16, 1). Hypoproct weakly sclerotized, strongly widened toward apex, semicircularly edged along free margin. Basal projection of gonocoxite closely adjoining aedeagus, slightly widened distally or nearly parallel-sided, with straight or shallowly emarginate apex. Aedeagus conical, with rounded apex. Roots of genitalia weakly sclerotized, curved. Differential diagnosis. The new genus is closely related to Acinacistyla described from Primorskii Territory (Fedotova and Sidorenko, 2006b) in the shape of the gonocoxite and gonostylus, in the presence of a sclerotized basal projection of the gonocoxite, and in the complex hypoproct strongly widened distally and forming a shallow emargination between the apices resembling the anteriorly curved lateral processes of the hypoproct of Acinacistyla, and also in the shape and venation of the wing, but differs in the presence of loop-shaped sensory threads on the flagellar segments, in a very wide hypoproct and basal projection of the gonocoxite, in the absence of a pubescent lobe at the
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Fig. 16. Kuriloneura multifida gen. et sp. n. (1, 2, 5–9, 11) and K. trifida sp. n. (3, 4, 10), males: (1, 3) genitalia; (2) aedeagus and basal projection of gonocoxite; (4) palpus; (5) mouthparts; (6) 1st flagellar segment; (7) 12th flagellar segment; (8) 5th flagellar segment; (9) 2nd flagellar segment; (10) scape and pedicel; (11) wing. Scale 0.1 mm [(a) to 1, 2; (b) to 3; (c) to 4–10; (d) to 11].
base of the gonocoxite, in shorter palpal segments, in simple claws, and in a short narrow triangular labrum.
Kuriloneura multifida Fedotova, sp. n. (Fig. 16, 1, 2, 5–9, 11; see also table)
The genus comprises 2 species: Kuriloneura multifida sp. n. and K. trifida sp. n.
Material. Holotype, ♂ (slide 541/K-3/1): Russia, Southern Kuriles, south of Shikotan Island, TserkovENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
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Fig. 17. Rhizomyiini (1), Ledomyiini (2–7), Brachineurini (8), and Undoneurini (9–22), males (1, 2, 5–16, 19–22) and females (3, 4, 17, 18): (1) Alatostyla binaria (Mamaev); (2–6) Isogynandromyia terricola Spungis; (7) Lauthia primigenia Mamaev; (8) Ramineura alia (Mamaev); (9, 10, 13, 15, 17) Megommata leefmansi (Nijveldt); (11, 12) M. pulvinariae (Felt); (14, 16, 18, 19) M. seychelli Barnes; (20, 21) Cingola circularis Jiao et Bu; (22) C. ceratodicrana Jiao et Bu; (1, 2, 7, 10, 12, 21, 22) genitalia; (3, 17) ovipositor; (4, 5) 5th flagellar segment of antenna; (6) 10th flagellar segment; (8, 9, 11) gonostylus; (13, 14, 20) 3rd flagellar segment of antenna; (15, 16) palpus; (18) 1st and 2nd flagellar segments; (19) wing (after: Harris, 1968; Spungis, 1981; Mamaev and Zaitsev, 1994, 2002; Jiao and Bu, 2014).
naya Bay, Malaise trap, 12–18.VII.2012 (Yu.N. Sundukov). Diagnosis. Male. Body pale yellow. Body length 0.88 mm; length of wing 1.42, width 0.55 mm; length of antenna 1.46 mm. 1st flagellar segment as long as 2nd one, 3.9 times as long as wide; basal enlargement nodose, 3.0 times as long as wide; neck 0.3 times as long as basal enlargement. 2nd segment 4.2 times as long as wide; basal enlargement nodose, 3.0 times as long as wide; neck 0.4 times as long as basal enlargement. 5th segment 3.7 times as long as wide; basal ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
enlargement 2.5 times as long as wide, neck 0.43 times as long as basal enlargement. 11th segment 4.1 times as long as wide; its basal enlargement 2.9 times as long as wide, neck 0.42 times as long as basal enlargement. 12th segment narrowed in distal half, with long apical process, 0.83 times as long as 11th segment. Palpus 3-segmented, without palpiger, strongly irregularly swollen; ratio of length of segments 1 : 2.1 : 2.3; 3rd segment rounded at end. Thorax uniformly sclerotized, dark yellow, 1.5 times as long as wide, slightly darker dorsally than
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ventrally. Legs dark because of dense cover of scales. Hind leg slightly longer than fore one; middle leg distinctly shorter than fore and hind legs. All femora 0.8 times as long as tibiae; fore femur 1.1 times as long as hind one. All femora distinctly shorter than 2nd tarsal segment; hind femur 0.83 times as long as 2nd tarsal segment. Hind tarsus slightly longer than fore one. 2nd segment of fore tarsus 10.3–12.3 times as long as 1st one. 2nd segment of hind tarsus 7.5 times as long as 1st and 4.3 times as long as 5th segment. Combined length of 3rd–5th tarsal segments in all legs 1.1–1.2 times length of 2nd segment. Claw simple in all tarsi, empodium shorter than claw. Wing widest near middle, 2.6 times as long as wide. Vein R1+2 0.43 times as long as wing. Genitalia wider than long. Gonocoxite wide, rounded on outer side, without lobe at base, 2.1 times as long as wide. Gonostylus very long, slightly swollen at base, 0.91 times as long as gonocoxite, 6–8 times as long as wide. Distal half of gonostylus very narrow, subparallel-sided. Cerci in shape of 2 conical lobes separated by triangular emargination. Hypoproct as long as cerci, with long setae at apex. Basal projection of gonocoxite widened distally, more heavily sclerotized at base, emarginate apically (Fig. 16, 2). Aedeagus weakly sclerotized, slightly widened toward base. Female unknown. Etymology. The name of the species is a Latin adjective of feminine gender meaning “strongly split, divided, consisting of many parts” and emphasizing the complex structure of its genitalia. Kuriloneura trifida Fedotova, sp. n. (Figs. 15, 5–10; 16, 3, 4, 10; see also table) Material. Holotype, ♂ (slide 540/K-3/1): Russia, Southern Kuriles, south of Shikotan Island, Tserkovnaya Bay, Malaise trap, 12–18.VII.2012 (Yu.N. Sundukov). Paratype, ♂ (slide 540/K-3/2), same locality and date. Diagnosis. Male. Body pale yellow. Body length 0.84–0.88 mm; length of wing 1.36–1.44, width 0.54– 0.56 mm; length of antenna 1.37 mm. Scape 1.2 times as long as pedicel; pedicel wider than long. 1st flagellar segment slightly shorter than 2nd one, 3.7 times as long as wide; basal enlargement slightly narrowed between base and apex, 2.7 times as long as wide; neck 0.37 times as long as basal enlargement. 2nd flagellar segment 3.5 times as long as wide; basal
enlargement three-nodular, 2.3 times as long as wide; neck half as long as basal enlargement. 5th flagellar segment 3.7 times as long as wide; basal enlargement 2.5 times as long as wide; neck half as long as basal enlargement, heavily sclerotized at apex. 8th flagellar segment 3.9 times as long as wide; its basal enlargement 2.8 times as long as wide; neck 0.42 times as long as basal enlargement. 12th segment as long as 11th one, strongly swollen at base, narrowed in distal half, with rounded apical process. Palpus 3-segmented, without palpiger, weakly swollen; 2nd and 3rd segments subparallel-sided; length ratio of palpal segments 1 : 0.9 : 1.3 or 1 : 1.4 : 1.4; 3rd segment rounded at end. Thorax uniformly sclerotized, dark yellow, 1.6– 1.7 times as long as wide. Notum slightly darker than thorax. Legs dark because of scales densely covering them. Hind leg 1.1 times as long as fore one; middle leg distinctly shorter than fore and hind legs. All femora 0.7–0.8 times as long as tibiae. Fore femur 1.1 times as long as hind femur or nearly subequal to it in length. All femora distinctly shorter than 2nd tarsal segment; hind femur 0.83 times as long as 2nd tarsal segment. Hind tarsus 0.91 times as long as fore one. 2nd segment of fore tarsus 8.3–8.5 times as long as 1st one. 2nd segment of hind tarsus 8.1–8.5 times as long as 1st and 4.0–4.1 times as long as 5th segment. Combined length of 3rd–5th tarsal segments in all legs 0.9–1.2 times length of 2nd segment. Claw in all tarsi simple, setiform; empodium shorter than claw. Anepimeron with 5 pores. Wing widest near middle, 2.5–2.6 times as long as wide. Vein R1+2 0.43– 0.45 times as long as wing. Apex of wing widely rounded. Genitalia elongate. Gonocoxite narrow in dorsal view and wide in lateral view. Gonocoxite 1.8– 2.1 times as long as wide. Gonostylus very long, subparallel-sided, slightly swollen at base, 0.91 times as long as gonocoxite, 6.3–6.8 times as long as wide. Cercus with wide, oviform narrowed lobes separated by triangular emargination. Hypoproct longer than cercus, strongly widened distally, with pointed lobes and small triangular emargination between them, 0.83 times as wide as cercus. Basal projections of gonocoxites in shape of wide sclerotized processes surrounding weakly sclerotized aedeagus slightly widened toward base and nearly reaching apices of gonocoxites. Female unknown. ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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Differential diagnosis. The new species is closely related to Kuriloneura multifida sp. n., but differs in a finer gonostylus curved at the base, in a more widely rounded gonocoxite, in rounded, instead of pointed, lobes of the cercus, in a more widely rounded apex of the wing, in the gonostylus more strongly curved at the base, in a less heavily sclerotized hypoproct not swollen at the base and bearing distinctly shorter lateral processes originating from the apex, in longer basal enlargements of the middle flagellar segments, in shorter palpal segments, in a longer and less strongly curved vein R4+5 fused with the costal vein farther from the wing apex, in a short neck of the 1st flagellar segment, and in a sparser loop-shaped threads covering the segments. Etymology. The name of species is a Latin adjective of feminine gender meaning “divided into three parts.” The species of the genus Kuriloneura differ in the following characters. 1. Cercus with triangular lobes, widened toward base (Fig. 16, 1). Gonostylus fine, arcuately curved. Hind leg and tarsus longer than fore ones. Body length 0.88 mm; length of wing 1.42 mm ........................... ...................................................... K. multifida sp. n. —Cercus
with rounded lobes, cordate (Figs. 15, 10; 16, 3). Hind leg and tarsus shorter than fore ones. Body length 0.84–0.88 mm; length of wing 1.36–1.44 mm .................................. K. trifida sp. n. Subtribe Kovaleviolina Fedotova, subtrib. n.
Diagnosis. Antenna short, 2+11-segmented in male, 2+10-segmented in female, 2.4 and 2.6 times as long as head in male and female, respectively; pedicel swollen (Kovaleviola). Flagellar segments with necks shorter than half length of basal enlargements in male, without necks in female. Palpus 3- or 4-segmented. Wing 2.0–2.6 times as long as wide. Vein R4+5 running into margin of wing far from its apex. Veins M3+4 and Cu1 separate, distinct; Cu2 reduced; M1+2 present in Kovaleviola, absent in Spungisiola. Fore femur longer than tibia. Claws simple. Genitalia transverse; gonocoxite and gonostylus long and fine. Gonocoxite with pubescent lobe at base. Aedeagus longer than gonocoxite, occasionally hook-shaped at end (Spungisiola). Differential diagnosis. These genera described from the Late Eocene Rovno amber are attributed to the supertribe Brachineuridi according to the folENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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lowing characters: vein R4+5 running into the margin of the wing far from its apex, the presence of vein Rs, simple claws, short antennae, and long legs. The new subtribe is similar to Acinacistylina in the shape of long and slender gonocoxites and gonostyli, in the presence of a pubescent lobe at the base of the gonocoxite, in a relatively short antenna of the male (which is slightly shorter than the wing), in long legs with a subequal length of the fore and hind pairs, in a longer fore femur, as compared with the hind femur, in the femora longer than the tibiae in all the species, and in the presence of vein M3+4, but differs in the presence of a furcation of the cubital vein, in the arrangement of vein Rs approximate to the place of fusion of the costal vein and R4+5, in shorter flagellar antennal segments in both sexes, and also in the shape of the transverse, instead of elongate, genitalia. The subtribe comprises 2 genera: Kovaleviola Fedotova et Perkovsky, 2008 (1 species) and Spungisiola Fedotova et Perkovsky, 2008 (1) described from the Late Eocene Rovno amber (Perkovsky and Fedotova, 2008). The characters of these genera require further investigation. The genera of the subtribe Kovaleviolina differ in the following characters. 1. Palpus 4-segmented. Pedicel not swollen. Wing 2.6 times as long as wide. Fore femur half as long as tibia. 2nd segment 9.4 times as long as 1st segment in fore tarsus, 14.3 times as long as 1st segment in hind tarsus ...... Spungisiola Fedotova et Perkovsky. —Palpus
3-segmented. Pedicel swollen. Wing twice as long as wide. Fore femur 1.4 times as long as tibia. 2nd segment of only remained tarsus 4.6 times as long as 1st segment ................................................... ........................ Kovaleviola Fedotova et Perkovsky. Genus Kovaleviola Fedotova et Perkovsky, 2008
Fedotova and Perkovsky in Perkovsky and Fedotova 2008 : 65. Type species Kovaleviola injusta Fedotova et Perkovsky, 2008 : 66. Diagnosis. Female (Kovaleviola injusta). Antenna 2+10-segmented; flagellar segments urceolate, without necks, strongly narrowed toward apex of antenna. Pedicel considerably swollen, 1st flagellar segment longer than others. Palpus 3-segmented. Vein R4+5 strongly approximate to costal vein. Vein M1+2 present. Apical abdominal segments VIII–X strongly narrowed.
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2nd segment of tarsus 14.3 times as long as 1st one. Apical plate of ovipositor directed dorsocaudally. Femora much longer than tibiae. Tarsi more than twice as long as tibiae. Male of K. injusta unknown; diagnosis of this genus based on male of K. pyxidiiformis comb. n. Male. Antenna 2+12-segmented; flagellar segments with elongate necks longer than half of basal enlargements. Palpus 3-segmented, with palpiger. Veins M1+2 and M3+4 distinct when branching from vein R4+5, but gradually disappearing near middle of wing; Cu rudimentary. Femora longer than tibiae. Tarsi more than twice as long as femora and tibiae. 2nd segment of tarsus 5.3 times as long as 1st one. Gonocoxite and gonostylus slender. Aedeagus and basal projection of gonocoxite projecting far beyond apex of gonocoxite. Differential diagnosis. The genus Kovaleviola is closely related to Spungisiola, but differs in the presence of vein M1+2, in a 3-segmented palpus, a swollen pedicel, and in the femora longer than the tibiae. The genus comprises the type species and K pyxidiiformis (Fedotova, 2005 : 49, Brachyneurina), comb. n. described from the Late Eocene Rovno amber (Ukraine, amber-bearing deposits near Rovno City). The species of the genus Kovaleviola differ in the following characters. 1. 2nd segment of tarsus 5.3 times as long as 1st segment. Veins M1+2 and M3+4 poorly developed; Cu rudimentary. Palpus long, with palpiger .................. ..................................... K. pyxidiiformis (Fedotova). —2nd
segment of tarsus 14.3 times as long as 1st segment. Veins M1+2, M3+4, and Cu well developed. Palpus short, without palpiger .................................. ............................ K. injusta Fedotova et Perkovsky.
very slender, with rounded pubescent lobe at base. Aedeagus projecting beyond apex of gonocoxite, hooked at end. Differential diagnosis. The genus Spungisiola differs from Kovaleviola in the absence of vein M1+2, in a 4-segmented palpus, in a non-swollen pedicel, in the femora shorter than the tibiae, and in a very long 2nd segment of the tarsus (in comparison with the 1st one), which also distinguishes this genus from the other Ledomyiini. The genus includes only the type species described from the Late Eocene Rovno amber (Ukraine, amberbearing deposits near Rovno City). The Genera Requiring an Additional Study During their description, the genera Palaeospaniocera Meunier, 1901 (1 species) described from the African copal and Ledomyiella Meunier, 1904 (4 species) from the Late Eocene Baltic amber were compared respectively with the genera Spaniocera and Ledomyia belonging to Brachineuridi. According to the descriptions, these genera are similar first of all in the wing venation, but also in a 4-segmented tarsus, which is characteristic of Heteropezidi (Fedotova and Perkovsky, 2009). Judging by their habitus, structure of the antennal segments, and the presence of a simple cubital vein and Rs (Ledomyiella), these genera can be conditionally attributed to Brachineuridi. Genus Ledomyiella Meunier, 1904 Meunier, 1904 : 44. Felt, 1911 : 38 (redescription). Type species Ledomyiella succini Meunier, 1904 : 45 (by monotypy).
Type species Spungisiola insuperabilis Fedotova et Perkovsky, 2008 : 67 (by monotypy).
Description. Antenna of male and female usually 2+12-segmented; flagellar segments with necks in male, without necks in female. Palpus 3- or 4-segmented. Wing short, wide. Vein R4+5 running into margin of wing far from its apex; Rs developed. Tarsi 4-segmented; 1st segment shorter than 2nd, 3rd longer than 3rd and 4th segments combined.
Diagnosis. Antenna of male 2+11-segmented; flagellar segments with short necks length of which less than half length of basal enlargements. Palpus 4-segmented. Veins Cu1 and M3+4 widely spaced. Femora shorter than tibiae. 2nd tarsal segment 4.6 times as long as 1st one. Gonocoxite and gonostylus
The genus is closely related to the recent genus Ledomyia, but differs in a 2+12-segmented antenna in both sexes, in the presence of vein Rs, in a straight vein R4+5 situated more closely to the wing apex, and also in 4-segmented tarsi, which fits the characters of the subfamily Porricondylinae.
Genus Spungisiola Fedotova et Perkovsky, 2008 Fedotova and Perkovsky in Perkovsky and Fedotova 2008 : 67.
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CLASSIFICATION OF GALL MIDGES OF THE SUPERTRIBE BRACHINEURIDI
Earlier, 5 species described from the Late Eocene Rovno amber were attributed to this genus. Among these, Ledomyia eocenica (Meunier, 1904), comb. n., is transferred to the recent genus Ledomyia, according to the absence of vein Rs and to the arcuate shape of R4+5 running into the margin of the wing very far from its apex. Genus Palaeospaniocera Meunier, 1901 Meunier, 1901 : 192. Felt, 1911 : 36. Type species Palaeospaniocera palaeospaniocera Evenhuis, 1994 : 186 (by monotypy). Description. Body oblong-ovate. Thorax slightly convex. Antenna 2+11-segmented. Palpus 3-segmented. Wing with 3 simple veins (R1+2, R4+5, and Cu). Tarsi 4-segmented, 1st segment longer than 2nd. Ovipositor with distinct apical plate. The genus includes only the type species described from the female found in African copal (Meunier, 1901). Indian Species of the Genus Meinertomyia At present, the name Meinertomyia Felt, 1911 is held to be a synonym of Brephometra Strand, 1910 (this genus includes 1 phytophagous species belonging to the supertribe Cecidomyiidi). In the catalogs of the gall-midges of the world fauna (Gagné, 2004, 2010), all the 4 species of Meinertomyia are included in a group of species of Cecidomyiinae with an unclear taxonomic position (Meinertomyia aequipalpis Mani, 1935 : 426; M. inaequipalpis Mani, 1935 : 426; M. manii Rao, 1951 : 110; M. ovalis Grover, 1961 : 411); all were described from India (Uttar Pradesh State) based only on the females. I consider that at least M. ovalis belongs to the supertribe Brachineuridi, tribe Rhizomyiini, subtribe Stellaserenina, being closely related to the genus Stelladiurna in the following characters: vein R4+5 is zigzag-like curved at the base of the wing and runs into the margin of the wing far from its apex; vein Cu is simple, curved; the palpus is 3-segmented; the antenna is 2+12-segmented; the necks of the flagellar segments of the female are short; the claws bear a tooth at the base and a long empodium; and the apical plate of the ovipositor is very wide. A key to all the species of the genus was published by Grover (1961). ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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ACKNOWLEDGMENTS I thank Yu.N. Sundukov (the “Kurilskii” Nature Reserve) for granting the material collected by him. I remember with gratitude V.S. Sidorenko (1965– 2010), a co-author of my many publications on the gall midges of the Russian Far East, which were based on his collections, and I also thank E.E. Perkovsky (I.I. Schmalhauzen Institute of Zoology, the National Academy of Sciences of Ukraine) for the opportunity to examine gall midges from the Late Eocene Rovno and Baltic amber and for his help in examination of these material. REFERENCES 1. Barnes, H.F., “Gall Midges (Cecidomyidae) Associated with Coffee,” Rev. Zool. Bot. Afr. 32, 324–336 (1939). 2. Barnes, H.F., “Gall Midges Living in White Clover Flowers,” Entomologist 87, 258–264 (1954). 3. Bayram, Ş. and Skuhravá, M., “New Records of Cecidomyiidae (Diptera) from Turkey,” J. Entomol. Res. Soc. 6 (3), 9–13 (2004). 4. Bogdanowicz, W., Chudzicka, E., Pilipiuk, I., Skibińska, E., Eds., “Cecidomyiidae,” in Fauna Polski. Charakteristics and Checklist of Species (Muz. Inst. Zool. Pol. Acad. Sci., 2007), pp. 17–56. 5. Edwards, F.W., “New Records of British Cecidomyiidae (Diptera), with Taxonomic Notes on Certain Genera,” Entomol. Month. Mag. 73, 145–154 (1937). 6. Enderlein, G., “Ord. Diptera, Fliegen (Zweiflügler),” in Fauna von Deutschland, Ed. by Brohmer, P. (Quelle & Meyer, Leipzig, 1920), vol. 2, pp. 265–315, pl. 12. 7. Enderlein, G., “22. Ordnung: Zweiflügler, Diptera,” in Die Tierwelt Mitteleuropas. Vol. 6, No. 2. Insekten, Teil III, Ed. by Brohmer, P., Ehrmann, P., and Ulmer, G. (Quelle & Meyer, Leipzig, 1936), pp. 1–259. 8. Evenhuis, N.L., Catalogue of the Fossil Flies of the World (Insecta : Diptera) (Backhuys Publ., Leiden, 1994). 9. Fedotova, Z.A., Gall Midges (Diptera, Cecidomyiidae) of the Deserts and Mountains of Kazakhstan: Morphology, Biology, Distribution, Phylogeny, and Systematics (Samara State Agricult. Acad., Samara, 2000) [in Russian]. 10. Fedotova, Z.A., “Dopolnenie. 22. Family Cecidomyiidae—Gall Midges,” in A Key to the Insects of the Russian Far East in Six Volumes. Vol. VI. Diptera and Fleas. Part 3, Ed. by Lehr, P. (Dal’nauka, Vladivostok, 2004), pp. 565–629 [in Russian]. 11. Fedotova, Z.A., “Gall Midges of the Subtribe Didactylomyiina (Diptera, Cecidomyiidae, Stomatosematidi) with Descriptions of New Taxa from Russian Far East,” Internatn. J. Dipterol. Res. 22 (1), 11–53 (2011a).
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12. Fedotova, Z.A., “Gall Midges of the Supertribe Stomatosematidi (Diptera, Cecidomyiidae) with Descriptions of New Taxa from Russian Far East,” Internatn. J. Dipterol. Res. 22 (4), 149–203 (2011b). 13. Fedotova, Z.A., “Classification of the Gall Midge Tribe Aphidoletini (Diptera, Cecidomyiidae: Aphidoletidi) with Descriptions of a New Genus and a New Species from the Kurile Islands,” Entomol. Obozr. 92 (4), 823–848 (2013) [Entomol. Rev. 94 (7), 1031–1051 (2014)]. 14. Fedotova, Z.A., “Fauna, Systematics, and Biology of the Gall-midges Supertribe Brachineuridi (Diptera, Cecidomyioidea),” Trudy Stavropol. Otd. Russ. Entomol. O-va, No. 10, 18–25 (Proceedings of the VII International Scientific-practical Online Conference, January 31, 2014) (Paragraph, Stavropol, 2014). 15. Fedotova, Z.A. and Perkovsky, E.E., “New Gall Midges (Diptera, Cecidomyiidae) from Rovno Amber. Subfamily Porricondylinae, the Tribes Bryocryptini and Winnertziini; Subfamily Lasiopterinae, Tribes Brachineurini and Oligotrophini,” Paleontol. Zh., No. 1, 42–53 (2005). 16. Fedotova, Z.A. and Perkovsky, E.E., “New Gall-midge Taxa (Diptera, Cecidomyiidae) from Dubrovitsy (Rovno Amber),” Vestn. Zool. 42 (1), 69–82 (2008). 17. Fedotova, Z.A. and Perkovsky, E.E., “New Gall Midges of the Tribe Leptosynini (Diptera, Cecidomyiidae) from the Late Eocene Ambers and the Classification of the Supertribe Heteropezidi,” Paleontol. J. 43 (9), 1101–1179 (2009). 18. Fedotova, Z.A. and Perkovsky, E.E., “Gall Midges of the Supertribe Stomatosematidi (Diptera, Cecidomyiidae) in the Palaearctic Region, with Descriptions of New Taxa from the Late Eocene Rovno Amber: 1. Stomatosema, Vanchidiplosis, Clarumreddera gen. n.,” Zool. Zh. 90 (10), 1204–1215 (2011a) [Entomol. Rev. 92 (4), 427–439 (2012)]. 19. Fedotova, Z.A. and Perkovsky, E.E., “Gall Midges of the Supertribe Stomatosematidi (Diptera, Cecidomyiidae) in the Palaearctic Region, with Descriptions of New Taxa from the Late Eocene Rovno Amber. 2. The Genera Didactylomyia and Rovnodidactylomyia gen. n.,” Zool. Zh. 90 (11), 1374–1384 (2011b) [Entomol. Rev. 92 (5), 565–575 (2012)]. 20. Fedotova, Z.A. and Perkovsky, E.E., “Gall Midges of the Supertribe Stomatosematidi (Diptera, Cecidomyiidae) in the Palaearctic Region, with Description of New Taxa from the Late Eocene Rovno Amber: 3. Groveromyia gen. n. Analysis of Biometric Parameters of the Recent and Late Eocene Genera and Species,” Zool. Zh. 91 (1), 71–87 (2012) [Entomol. Rev. 92 (8), 932–948 (2012)]. 21. Fedotova, Z.A. and Perkovsky, E.E., “New Gall Midges (Diptera, Cecidomyiidae, Brachineurini, Ledomyiini) from the Late Eocene Rovno Amber,” Vestnik Zool. 48 (1), 387–393 (2014).
22. Fedotova, Z.A. and Sidorenko, V.S., “New Species of Gall Midges of the Supertribe Oligotrophidi (Diptera, Cecidomyiidae) from the Russian Far East,” Far Eastern Entomol. 146, 1–12 (2005). 23. Fedotova, Z.A. and Sidorenko, V.S., “New Species of Gall Midges from the Genus Brachineura Rondani, 1840 and New Related Genera (Diptera : Cecidomyiidae, Brachineurini) from the Russian Far East,” Internatn. J. Dipterol. Res. 17, 77–97 (2006a). 24. Fedotova, Z.A. and Sidorenko, V.S., “New Gall Midges of the Tribes Brachineurini Edwards, 1937 and Stomatosematini Mamaev, 1968 (Diptera : Cecidomyiidae) from the Russian Far East,” Far Eastern Entomol. 163, 1–28 (2006b). 25. Fedotova, Z.A. and Sidorenko, V.S., “Nine New Species of Genus Ledomyia Kieffer, 1895 (Diptera, Cecidomyiidae) from the Russian Far East,” Far Eastern Entomol. 169, 1–19 (2007). 26. Felt, E., “A Generic Synopsis of the Itonidae,” J. New York Entomol. Soc. 19, 31–62 (1911). 27. Felt, E., “Key to Gall Midges (A Resume of Studies I–VII, Itonididae),” New York State Mus. Bull. 257, 3–239 (1925). 28. Gagné, R.J., “Cecidomyiidae from Mexican Tertiary amber (Diptera),” Proc. Entomol. Soc. Washington 75, 169–171 (1973). 29. Gagné, R.J., “A Revision of the Nearctic Stomatosematidi (Diptera : Cecidomyiidae: Cecidomyiinae),” Ann. Entomol. Soc. America 68, 86–90 (1975). 30. Gagné, R.J., “New Nearctic Records and Taxonomic Changes in the Cecidomyiidae (Diptera),” Ann. Entomol. Soc. America 69, 26–28 (1976). 31. Gagné, R.J., “Descriptions of New Nearctic Cecidomyiidae (Diptera) That Live in Xylem Vessels of Fresh-cut Wood, and a Review of Ledomyia (s. str.),” Proc. Entomol. Soc. Washington 87, 116–134 (1985). 32. Gagné, R.J., The Plant Feeding Gall Midges of North America (Coms. Publ. Ass. Cornell Univ., Ithaca, London, 1989). 33. Gagné, R.J., The Gall Midges of the Neotropical Region. (Coms. Publ. Ass. Corn. Univ., London. 1994). 34. Gagné, R.J., “A Catalog of the Cecidomyiidae (Diptera) of the World,” Mem. Entomol. Soc. Washington 25 (2004). 35. Gagné, R.J., Update for a Catalog of the Cecidomyiidae (Diptera) of the World. Digital version 1, http:// www.ars.usda.gov/SP2UserFiles/Place/12754100/Gagné _2010_World_Catalog_Cecidomyiidae.pdf. 36. Gagné, R.J. and Étienne, J., “Note on the Cecidomyiidae from Guadeloupe (West Indies) with Description of a New Species of Paracalmonia (Diptera),” Bull. Soc. Entomol. France 114, 337–350 (2009). 37. Gagné, R.J. and Jaschhof, M., A Catalog of the Cecidomyiidae (Diptera) of the World. 3rd ENTOMOLOGICAL REVIEW Vol. 94 No. 8 2014
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38. 39. 40. 41.
42.
43.
44.
45. 46. 47. 48. 49. 50. 51. 52.
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