Revision of the genus Lepidopria Kieffer (Hymenoptera, Diapriidae, Diapriinae) of the world fauna
Abstract
All known species of the diapriid genus Lepidopria Kieffer, 1911 are revised, keyed and illustrated. A new species, L. bicarinata Chemyreva & Ku, sp. nov. is described from South Korea. The diagnosis of the genus Lepidopria, as well as that of all described species, has been clarified. New data on the distribution of the species L. masneri Notton, 1994 and L. pedestris Kieffer, 1911 are provided. The association of L. masneri with the nests of Solenopsis japonica Wheeler, 1928 is reported and the biology of Lepidopria species is discussed. Taxonomic status of the mysterious L. aberrans Brues, 1916 is considered and a new combination, Bruesopria aberrans (Brues), comb. nov., suggested.
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Revision of the genus Lepidopria Kieffer (Hymenoptera, Diapriidae, Diapriinae) of the world fauna Vasilisa G. Chemyreva1,2, Seonwoo Yoon3, Deok-Seo Ku4 1A.A. Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, Moscow, 117647, Russia 2Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St. Petersburg, 199034, Russia 3Ulsan Agricultural Technology Center, Ulsan, Republic of Korea 4The Science Museum of Natural Enemies, Geochang 50147, Republic of Korea Corresponding author: Vasilisa G. Chemyreva ([email protected]) Academic editor: Elijah Talamas|Received 31 August 2025|Accepted 4 November 2025|Published 18 November 2025 https://zoobank.org/D1292408-DD83-4D78-A557-F3A63128ACB4 Citation: Chemyreva VG, Yoon S, Ku D-S (2025) Revision of the genus Lepidopria Kieffer (Hymenoptera, Diapriidae, Diapriinae) of the world fauna. Journal of Hymenoptera Research 98: 1107–1126. https://doi.org/10.3897/jhr.98.169802 Abstract All known species of the diapriid genus Lepidopria Kieffer, 1911 are revised, keyed and illustrated. A new species, L. bicarinata Chemyreva & Ku, sp. nov. is described from South Korea. The diagnosis of the genus Lepidopria, as well as that of all described species, has been clarified. New data on the distribution of the species L. masneri Notton, 1994 and L. pedestris Kieffer, 1911 are provided. The association of L. masneri with the nests of Solenopsis japonica Wheeler, 1928 is reported and the biology of Lepidopria species is discussed. Taxonomic status of the mysterious L. aberrans Brues, 1916 is considered and a new combination, Bruesopria aberrans (Brues), comb. nov., suggested. Keywords Bruesopria, Diapriini, myrmecophilic wasps, new records, new species, symphiles, Solenopsis, taxonomy Introduction Lepidopria Kieffer, 1911 comprises a few species of tiny wasps belonging to the myrmecophilic diapriids (Huggert and Masner 1983). The genus currently includes the following four species: L. aberrans Brues, 1916, recorded in Maryland, USA; L. lloydi (Ferrière, 1935), recorded in Tanzania; L. masneri Notton, 1994, recorded in Japan JHR 98: 1107–1126 (2025) doi: 10.3897/jhr.98.169802 https://jhr.pensoft.net Copyright Vasilisa G. Chemyreva et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. RESEARCH ARTICLE
V. G. Chemyreva et al. / Journal of Hymenoptera Research 98: 1107–1126 (2025) 1108 and South Korea; and L. pedestris Kieffer, 1911, recorded in Europe. According to Masner and Muesebeck (1968), the L. aberrans is doubtfully placed in this genus and may belong to the Nearctic genus Bruesopria Wing, 1951 (Huggert and Masner 1983; Masner and García 2002); however, the reasons for the doubts and the morphological features of this species were not discussed. The specific characters of the Lepidopria species, such as reduced eye size and number of palpal segments, reduced pigmentation (especially in the female antenna, which has a yellow clava), the secondary apterism, abundant setation, and an enlarged petiole (similar to many ants), suggest that they all have myrmecophilic habits (Notton 1994). Lepidopria aberrans and L. pedestris were reared from the puparia of tachinid flies, which were found inside adult scarabaeid beetles (Brues 1916; Huggert and Masner 1983). It has also been confirmed that both species are symphilic in the nests of ants of the genus Solenopsis Westwood, 1840 (Brues 1916; Ferrière 1927; Hölldobler 1928; Wing 1951; Lachaud 1981b; Lachaud and Passera 1982; Masner and García 2002). The interaction with ants has been studied in detail for only one species, L. pedestris (Hölldobler 1928; Lachaud 1980, 1981a, b, c, 1982; Lachaud and Passera 1982). According these studies L. pedestris is a highly adapted myrmecophilous diapriid wasp that lives in close association with the ant Solenopsis fugax (Latreille, 1798). The wasp integrates into ant colonies primarily by mimicking or sharing tactile solicitation signals and by producing cuticular secretions that induce intense grooming or licking from ant workers. Adults (especially females) obtain food by trophallaxis from ants inside nests without aggression (Lachaud 1982). Morphological specializations like clusters of active secretory cells opening into dense tufts of hairs on the petiole and anterior part of S2 (these secretions are associated with the regions ants lick most intensively) underlie chemical integration of L. pedestris in the ant colony (Lachaud 1981a). The fore leg tapping patterns and antennal contacts are similar to those used by ants and underlie the behavioral integration of L. pedestris (Lachaud 1981c, 1982). The success of the wasp’s integration is most clearly demonstrated by the fact that workers of S. fugax may regurgitate food to newly introduced L. pedestris within ~10 minutes and show very little aggression (Lachaud 1981b). A detailed phenological study of L. pedestris was carried out by Lachaud and Passera (1982). Although the authors of this study were unable to establish the primary host of this species, they reached several significant conclusions. Firstly, L. pedestris is undoubtedly myrmecophilic; it uses Solenopsis fugax nests for protection, feeding purposes and hibernation in winter. Secondly, L. pedestris does not use S. fugax as a host and does not reproduce in its nest. Thirdly, L. pedestris specimens spend most of their life cycle outside the nest (from early spring to late autumn), where they mate, parasitize the primary host and develop into adults, and then, when winter arrives, they return to the S. fugax nests. Finally, L. pedestris is strictly associated with S. fugax nests, since it is not found in the nests of other ant species parasitized by the lestobiotic S. fugax and was even eliminated by two of these species (Plagiolepis pygmaea (Latreille, 1798) and Tetramorium semilaeve André, 1883) when experimentally introduced in their nest (Lachaud and Passera 1982). Only later the main host of L. pedestris was re-
Revision of Lepidopria (Hymenoptera, Diapriidae) 1109 ported as tachinid flies, which parasitise scarabaeid beetles that are not associated with ant nests (Huggert and Masner 1983). Unfortunately, we do not know much about the life histories of other Lepidopria species. The African species Lepidopria lloydi was reared from puparia of the Glossina Wiedemann, 1830 (Ferrière 1935; Notton 2014), whereas the biology of L. masneri remains unknown (Notton 1994). These two species have morphology that suggest myrmecophily, however, their connection with ants has not yet been documented. This work aims to review the morphology and biology of all described species of the genus Lepidopria, clarify the diagnostic characteristics of this genus and some closely related genera, and describe a new species of Lepidopria from the south-eastern Palaearctic region. Material and methods Material observed in this study originates from the several collections: MCSN − the Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy; NHML − the Natural History Museum, London, UK; NIBR − the National Institute of Biological Resources, Incheon, Republic of Korea; SMNE − the Science Museum of Natural Enemies, Geochang, Republic of Korea; NMNH − the National Museum of Natural History, Smithsonian Institution, Washington, USA; ZISP − the Zoological Institute of the Russian Academy of Sciences, St Petersburg, Russia; UATC – Ulsan Agricultural Technology Center, Republic of Korea. Most of images were taken with a Nikon stereo microscope SMZ25 equipped with objective lens Plan Apo 1.6× and digital camera DS-10. Some images were taken with a camera Olympus E-M5 Mark II and objective lens M.Zuiko 60 mm 1:2.8 combined with a Raynox MSN-505 (Fig. 7A, C). Final images were stacked using Helicon Focus 7.6. The morphological terminology, abbreviations and measurements follow Masner and García (2002), Yoder (2004) and the Hymenoptera Anatomy Ontology (Yoder et al. 2010). New regional and country records are indicated with an asterisk (*). Taxonomy Genus Bruesopria Wing, 1951 Note. A detailed diagnosis of this genus has been given by Masner and García (2002). Therefore, here we give only a short comparative diagnosis of this genus, emphasizing the diagnostic differences between it and the genus Lepidopria. Comparative diagnosis. Female antenna 11–12-merous [only 12-merous in Lepidopria], with abrupt 3-merous clava [nonabrupt or 4-merous clava in Lepidopria]; posterior margin of propodeum with deep semicircular excavation, median carina of propodeum rudimentary, posterolateral corner of propodeum angular and strongly
V. G. Chemyreva et al. / Journal of Hymenoptera Research 98: 1107–1126 (2025) 1110 projecting [same in Lepidopria to with median carina, without excavation of posterior margin and without posterolateral projection]; petiole strongly modified, remarkably higher than long in lateral view, with finger-like projection produced dorsally above anterior margin of T2 (Fig. 1C, D) [same in Lepidopria (Fig. 9E) to as long as high in lateral view and not produce dorsally above T2 (Fig. 2C, D)]; base of S2 bare, smooth, flat: not convex medially and not grooved laterally, with deep semicircular excavation of anterior margin (Fig. 1A) [densely setose, with lateral grooves and convex medially, excavation of anterior margin not deep in Lepidopria (Fig. 1B)]. Remarks. This genus comprises only two described species from the New World (Wing 1951; Johnson 1992). Masner and García (2002) reported an undescribed species associated with Solenopsis sp. in Arizona. An undetermined species of Bruesopria (possibly B. aberrans) was found by Dr Alexander L. Wild in a colony of Solenopsis molesta (Say, 1836) at Konza Prairie in Kansas, USA (see Fig. 1E, F). The biodiversity and morphological variability of the three closely related genera – Lepidopria, Bruesopria, and Solenopsia Wasmann, 1899 – have not yet been sufficiently studied and future discoveries may necessitate the synonymisation of these genera (Masner and García 2002). However, the diagnostic differences between Lepidopria and Bruesopria, as described above, are presently clear enough to maintain these taxa as valid. Furthermore, these genera have different distributions: all Lepidopria species are found in the Palaearctic region, whereas all Bruesopria species are found in the Nearctic region. Bruesopria aberrans (Brues, 1916), comb. nov. Fig. 1C, D Lepidopria aberrans Brues, 1916: 126. Lepidopria aberrans: Masner and Muesebeck 1968: 10. ?Bruesopria aberrans: Masner and García 2002: 73. Material examined. Holotype, • female: USNM no.20474, Hagerstown, Md., [Washington, Maryland, USA] Aug. 15, 1913, ex puparium of ?Cryptomeigenia theutis (Walker) in an adult of Phyllophaga inversa (Horn); stored in NMNH. The female of B. aberrans can easily be distinguished from other described Bruesopria species by its 12-merous antennae, whereas males of all known Bruesopria species have not been described or illustrated in detail (Wing 1951). Some details about Bruesopria male morphology are known only from the diagnosis of this genus given by Masner and García (2002). Biology. Bruesopria aberrans was reared from the puparium of a tachinid fly (possibly Cryptomeigenia theutis (Walker, 1849)), which was found inside an adult June beetle, Phyllophaga inversa (Horn, 1887). The association of this species with Solenopsis sp. was reported by Masner and García (2002), but without indication of the date or location of the observation. Distribution. Recorded only from the USA.
Revision of Lepidopria (Hymenoptera, Diapriidae) 1111 Figure 1. Bruesopria americana (A), Lepidopria masneri (B), B. aberrans, comb. nov., female, slide of holotype (C, D) and Bruesopria sp.(E, F): A, B metasoma, ventral view C base of metasoma D habitus, dorsolateral view E, F female of Bruesopria sp. in the colony of thief ant Solenopsis molesta (Say), image Alex Wild, used by permission. Remarks. Masner and Muesebeck (1968) reported on the type of Lepidopria aberrans Brues and reported that this species was doubtfully placed in this genus. Later, Huggert and Masner (1983) additionally noted that it may belong to the Nearctic genus Bruesopria Wing. This species was not mentioned again until 2002, when Masner and García wrote about it as a member of the genus Bruesopria, despite that no new combination had been suggested or justified. However, re-examination of the type spec-
V. G. Chemyreva et al. / Journal of Hymenoptera Research 98: 1107–1126 (2025) 1112 imen of this species and some other available Bruesopria specimens, as well as all known Lepidopria species, led us to the same conclusion: Lepidopria aberrans Brues should be considered a member of the genus Bruesopria, as it shares all the characters typical for Bruesopria species, as outlined in the comparative diagnosis of this genus above. Genus Lepidopria Kieffer, 1911 Type species. Lepidopria pedestris Kieffer, by monotypy. Genus diagnosis. Small sized insects (1.0–1.8 mm) with yellowish to almost black body, covered with short, erect, evenly distributed setae (usually females more pale and more densely setose than males). Head subglobular in dorsal and lateral views, subtriangular to globular in frontal view; eyes of female small, its largest diameter shorter than length of malar space; palpal formula 4–2. Male antenna 14-merous with only A4 to A3−A4 modified; female antennae 12-merous, with more or less abrupt 4-merous clava to clava indistinct; A1 about as long as four following antennomeres combined; A4−A8 from subquadrate to transverse. Propleuron, mesopleuron ventrally, metapleuron, lateral side of propodeum and petiole covered with dense setation or foamy structures; mesoscutum transverse, without notauli, almost flat but slightly convex antero-medially and depressed postero-medially; mesoscutellum usually with single transverse anterior scutellar pit or it subdivided into two pits; propodeum with more or less distinct median carina that is not elevated anteriorly or carina absent; plica and posterior plical process slightly to moderately projecting. Alate, fore wing as long as body or slightly shorter, with single costal vein developed in its basal third. All legs with cylindrical tarsi, tarsomeres about as long as wide. Petiole in lateral view humped dorsally, elevated above level of base of T2 and not broadened posteriorly; base of T2 distinctly wider than petiole, smooth and flat; base of S2 densely setose, with two groove-like invaginations laterally and median convexity between them; anterior margin of S2 without deep excavation; gaster depressed and ovoid. Lepidopria bicarinata sp. nov. https://zoobank.org/F9D56D76-04B2-4E9E-9D8E-EA21D8292FA0 Figs 2, 3 Type material. Holotype, • female: South Korea, Jeju-do (JJ), Jeju-si, Sanduk-ro, 593-40, near Temple Gwaneum rest area (Livestock Research Institute), MT, 1–24. IX.2022, Deokseo Ku & Jeongjun Ahn legs (NIBR). Paratypes: same locality as holotype, 1–16.VII.2022, 16–31.VII.2022, 1♀ and 2♂ (ZISP). Etymology. The name derives from Latin “bi-” and “carina” and refers to the welldeveloped two carinae on male A3 and A4. Diagnosis. Head globular in frontal view (Fig. 2E); female antennae stout, with A9−A11 transverse, A9 about as long as, and as wide as A10 (Fig. 2F); male A3 with carina and without emargination, male A4 slightly enlarged, slightly longer and slightly
Revision of Lepidopria (Hymenoptera, Diapriidae) 1113 Figure 2. Lepidopria bicarinata sp. nov., female. A habitus, lateral view B head, mesosoma and base of metasoma, dorsal view C metasoma, lateral view D head and mesosoma, lateral view E face F antenna, lateral view. Scale bars: 0.5 mm (A); 0.3 mm (B–E). wider than A3, with shallow emargination and membranous, pale carina (Fig. 3D, E); mesoscutellum with single transverse scutellar pit (Figs 2B, 3B); propodeum densely setose dorsally, with median carina; plica hardly indicated under setae (Figs 2B, D,
V. G. Chemyreva et al. / Journal of Hymenoptera Research 98: 1107–1126 (2025) 1114 3B); petiole weakly swollen dorsally and projecting upwards, dorsally covered with foamy structure in female and long pale setae in male (Figs 2B, D, 3B). Description. Female, holotype. Body length 1.5 mm, antennae length 0.85 mm; fore wing length 1.4 mm. Body dark brown except petiole and apex of metasoma brown; tegulae, antenna and legs brownish yellow. Head rounded, smooth, in dorsal view as long as wide, and 1.15 times as high as long in lateral view, covered with short, erect setae; postgenal cushion dense; ocelli small, inconspicuous, arranged in an almost equilateral triangle; LOL 2.5 times as long as diameter of frontal oculus; OOL large, 3 times as long as POL; compound eye small with fewer than 50 ommatidia; face smooth, epistomal sulcus indistinct; malar space large, 1.1 times as long as height of eye; tentorial pit small and distinct; malar distance about as long as shortest distance between eye in frontal view; mandibles bidentate, with apices slightly overlapping. Antenna 12-merous, covered with short, semidecumbent to erect setae; A1 slightly compressed and curved, about 3.7 times as long as its greatest width, smooth, shining, and bare ventrolaterally, coriaceous and setose otherwise; in dorsal view A2 cylindrical and 1.8 times as long as wide, in lateral view it tapering towards base; A3 slightly narrower than the A2, 1.5 times as long as wide; A4 and A5 about as wide as A3, subequal to each other and about 1.2 times as wide as long; A6−A8 distinctly transverse and becoming gradually wider; A9−A12 forming abrupt 4-segmented clava, its setation denser and slightly shorter than on other antennomeres; A10 and All subequal, slightly transverse; A9 transverse and slightly wider than A12; A12 ovoid, 1.2 times as long as wide. Mesosoma about as wide as head; pronotal collar weakly developed (Fig. 2B, D); pronotal shoulders prominent, crossed with sharply projecting pronotal suprahumeral sulcus (Fig. 2D); lateral side of pronotum smooth, with short scattered and erect setation; propleuron with setae and foamy structures on ventral 0.3 and smooth and bare dorsally; mesoscutum weakly convex anteriorly and laterally, and slightly depressed posteromedially, without notauli and covered with short, scattered erect setae; mesoscutellum with transverse and reniform anterior scutellar pit, pit smooth and bare inside; mesoscutellar disc with very weak median carina and elongate rugulae anteriorly, covered with several erect setae laterally; mesopleuron smooth, laterally covered with scattered, short, erect setae; mesopleuron ventrally with short recumbent setae and foamy structures anterior to mid-coxa; metanotum entirely setose, sculptured with scattered setigerous punctures; metascutellum short, with low median carina and weakly projecting lateral carina; ventral side of metapleuron with foamy structure; propodeum and metapleuron entirely densely setose, covered with short, adpressed white setae; metapleuron and lateral side of propodeum without visible carinae; in dorsal view posterior margin of propodeum deeply arcuate: propodeum short medially and significantly longer laterally; lateral and median carinae of propodeum almost parallel, narrow and hidden under setation; nuchal area smooth and bare. Legs moderately robust, covered with short, erect setae similar to those of body; fore tibia with apical spine, its length about half of apical width of tibia; all femora with short basal stalk; hind femur 2.3 times as long as its greatest width. Fore wing 0.9 times as long as body, with single costal vein reaching 0.3 of fore wing length.
Revision of Lepidopria (Hymenoptera, Diapriidae) 1115 Figure 3. Lepidopria bicarinata sp. nov., male. A habitus, lateral view B mesosoma and metasoma, dorsal view C antenna, lateral view D, E base of antenna in dorsal (D) and lateral (E) views. Scale bars: 0.5 mm (A); 0.3 mm (B, C). Metasoma. Petiole cylindrical in dorsal view, as wide as long, densely covered with foamy structure on dorsal surface; in lateral view, medial portion slightly swollen upwards; ventral side of petiole covered with short recumbent white setation, not projecting ventrally; gaster about 0.9 times as long as head and mesosoma combined, more than 1.7 times as long as its greatest width, dorsoventrally compressed; T2 dorsally with short, erect, evenly distributed setae, base of T2 flat, wider than petiole; T3−T5 smooth and scarcely setose; base of S2 densely setose, with deep lateral groove and median convexity; basal corners of S2 extending forwards either side of petiole beyond base of large tergite; ovipositor shortly exserted. Male. Body length 1.4 mm. Differing from female as follows: colour of body black, mandibles and tegula brown, antennae dark brown with A2 yellowish, legs yellowish brown with coxae, femoral and tibial clubs darkened; body less densely setose than in female, especially metasoma, foamy structures developed only on propleuron
V. G. Chemyreva et al. / Journal of Hymenoptera Research 98: 1107–1126 (2025) 1122 Figure 9. Lepidopria pedestris, male. A face and base of antennae B head, mesosoma and base of metasoma, dorsal view C habitus, dorsal view D flagellum, dorsal view E petiole, lateral view. Scale bars: 0.5 mm (C); 0.3 mm (A, B, D). Remarks. Huggert and Masner (1983) reported about “an interesting case of simultaneous parasitism by Lepidopria pedestris Kieffer and Trichopria inquilina Kieffer on a tachinid puparium in an adult may beetle, Melolontha melolontha Linnaeus is known to us from Yugoslavia (specimens in Nat. Museum, Praha)”. The male specimen from Bosnia and Herzegovina mentioned above most likely belongs to this group of specimens, and we are providing its label data for the first time here. Dr Jan Macek kindly presented it to us many years ago and it is now stored in ZISP.
Revision of Lepidopria (Hymenoptera, Diapriidae) 1123 Key to the all species of Lepidopria Kieffer 1 Female antennae slender with non-abrupt clava (Fig. 4D); male A1 swollen medially, A3 and A4 modified with deep excavation (Fig. 4A, B); pronotum with dense pronotal cushion; borders of anterior scutellar pit indistinct, smoothed (Fig. 4A, C) .................................................... L. lloydi (Ferrière) – Female antennae stout or/and with more or less distinct 4-segmented clava (Figs 1A, 2F, 5E, 8A); male A1 cylindrical, A3 not modified or, at least, without excavation; pronotal cushion reduced (Figs 2D, 5C); borders of anterior scutellar pit distinct (Figs 2B, 3B, 5D, 6B, 9B) ........................................... 2 2 Petiole strongly swollen dorsally and projecting upwards (Fig. 9E); pronotal shoulders not projecting in dorsal view, without pronotal suprahumeral sulcus (Figs 8A, 9B); anterior scutellar pit subdivided into two pits, with raised area in the middle (Figs 8A, C, 9B); head subtriangular in frontal view (Figs 8A, C, 9B); female A9−A11 as long as wide (Fig. 8A); male A4 strongly enlarged, distinctly longer and wider than A3, with deep emargination and strong, pigmented carina (Fig. 9A, D)............................ L. pedestris Kieffer – Petiole weakly swollen dorsally and projecting upwards (Figs 2D, 3A, 5F, 6A); pronotal shoulders distinctly projecting in dorsal view, with sharp pronotal suprahumeral sulcus (Figs 2D, 5C); anterior scutellar pit not subdivided into two pits (Figs 2B, 3B, 5D, 6B); head globular in frontal view (Figs 2E, 5A); female A9−A11 transverse (Figs 2F, 5E); male A4 slightly enlarged, slightly longer and almost not wider than A3, with moderately shallow emargination and membranous, pale carina (Figs 3D, E, 6D, E) ......................................3 3 Propodeum very finely setose dorsally (Figs 5D, 6B); female petiole partly bare to setose dorsally (Fig. 5D); female antennae with A9 distinctly shorter and narrower than A10 (Fig. 5E); male A3 not modified (Fig. 6D, E) .......... .......................................................................................L. masneri Notton – Propodeum densely setose dorsally (Fig. 2B); female petiole covered with foamy structure dorsally (Fig. 2B); female antennae with A9 about as long as A10 and as wide as A10 (Fig. 2F); male A3 with carina (Fig. 3D, E) ............. ..................................................................................L. bicarinata sp. nov. Discussion The generic diagnoses of Lepidopria given by Kieffer (1911, 1916) were based on a single species, Lepidopria pedestris. Subsequently, Huggert and Masner (1983) carried out a comparative analysis of myrmecophilous species of the Holarctic. Although they did not clarify the generic diagnosis, many diagnostic features were illustrated and used in their key. The description of the genus by Huggert and Masner (1983) is not entirely unambiguous. According to the key provided, L. pedestris has a propodeum that is “not
V. G. Chemyreva et al. / Journal of Hymenoptera Research 98: 1107–1126 (2025) 1124 excavated posteriorly, ... with distinct median keel”, which does not align with the morphology of the holotype (Fig. 8A, B) and Fig. 6 in Huggert and Masner (1983). In our opinion, the latter illustrated a Lepidopria sp. with a posteriorly excavated propodeum, whereas the holotype of L. pedestris does not have a median carina (keel). However, this discrepancy is not currently significant, as high variation in these two features has been reported in subsequently described species of the genus (Notton 1994; current study). The genus Lepidopria is most closely related to the Nearctic genus Bruesopria (the confused classification of L. aberrans may confirm this), and to the Palaearctic genus Solenopsia (Masner and García 2002). The differences between Lepidopria and Bruesopria are given in the comparative diagnosis above. Two described species of Solenopsia can be distinguished from Lepidopria and Bruesopria by having 11-merous antenna with a 2-merous clava in the females. They are also characterised by the presence of a distinct median propodeal carina which is slightly elevated anteriorly and forms a tooth-like, subtriangular structure. Other important diagnostic features of Solenopsia (most of which are mentioned in the comparative diagnosis given for Bruesopria and Lepidopria above) are similar to those of species Lepidopria or Bruesopria. Currently, we believe that it is not possible to reasonably discuss the synonymy of these genera, as additional morphological, molecular, and biological data on these wasps are required. It must be considered that all three genera may have arisen independently through convergent evolution. However, we would like to note that the life cycles of B. aberrans and L. pedestris most likely are remarkably similar. Photos taken by Dr Alexander Wild in Kansas (Fig. 1E, F) confirm once again the connection between Bruesopria species and the ant Solenopsis molesta. The Bruesopria in these photos either belongs to the B. aberrans species, as it has 12-merous antennae (two other described species of this genus have 11-merous antennae) and 3-segmented clava with slightly elongated segments that are nearly equal in width and about twice as wide as A4–A8 (clavameres are transverse and much wider in B. americana (Brues, 1936) and B. seeversi Wing, 1951), or it belongs to an undescribed species of this genus that is morphologically close to B. aberrans. Therefore, it is likely that all currently known species of Bruesopria are associated with S. molesta (Wing 1951). Unfortunately, the detailed biology of the other species Bruesopria, Lepidopria and Solenopsia is not well understood, but all species of these three genera, if known, are associated with the genus Solenopsis (Brues 1916; Lachaud 1980, 1981b, c, 1982; Lachaud and Passera 1982; Masner and García 2002; current study). At the same time, we can state with great confidence that the actual species diversity of these myrmecophilous wasps is significantly greater than is currently recognized. Thus, it cannot be discounted that additional discoveries will eliminate the tenuous boundaries between these myrmecophilous genera. Acknowledgments We are very thankful to Mr Joseph Monks (NHML) and Dr Ekaterina Tselikh (ZISP) for providing crucial information and images of the type material of Lepidopria lloydi. We also wish to thank Dr. Matt Buffington (NMNH) for the nice photos of B. aberrans slide. Our
Revision of Lepidopria (Hymenoptera, Diapriidae) 1125 appreciation goes to Dr. Maria Tavano (MCSN) for providing the opportunity to examine the types of L. pedestris. We thank Dr. Alexander Wild (The University of Texas at Austin, USA) for permission to use the photos of Bruesopria sp. Special thanks go to Dr Jean-Paul Lachaud (El Colegio de la Frontera Sur, Chetumal, Mexico) for reading the first draft of the manuscript and for his useful suggestions and additional literature, which enabled us to significantly improve this article. We would also like to thank Dr. Elijah Talamas (Florida Department of Agriculture and Consumer Services, USA) for his valuable comments on the first draft of the manuscript and his comprehensive support. This work was supported by a grant from the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR202502202) and partially supported by Russian State Research Project (No. 125012901042-9). References Brues CT (1916) A new species of Lepidopria from North America. Psyche 23(4): 126–127. https://doi.org/10.1155/1916/61831 Ferrière C (1927) Note sur un Diapriide (Hyménoptère), hôte de Solenopsis fugax Latr. Konowia 6: 282–286. Ferrière C (1935) Les Hyménoptères parasites des mouches Tsétsé. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 16: 328–340. https://doi.org/10.5169/seals-400820 Hölldobler K (1928) Zur Biologie der diebischen Zwergameise (Solenopsis fugax) und ihrer Gäste. Biologisches Zentralbtatt 48(3): 129–142. Huggert L, Masner L (1983) A review of myrmecophilic-symphilic diapriid wasps in the Holarctic realm, with descriptions of new taxa and a key to genera (Hymenoptera: Proctotrupoidea: Diapriidae). Contributions of the American Entomological Institute 20: 63–89. Johnson N (1992) Catalog of world species of Proctotrupoidea, exclusive of Platygastridae (Hymenoptera). Memoirs of the American Entomological Institute 51: 1–825. Kieffer JJ (1911) Proctotrypidæ. In: André E (Ed.) Species des Hyménoptères d’Europe et d’Algérie 10(6): 753–912. Kieffer JJ (1916) Diapriidae. Das Tierreich (Vol. 44). Walter de Gruyter & Co., Berlin, 627 pp. https://doi.org/10.5962/bhl.title.1109 Lachaud J-P (1980) Les communications tactiles interspécifique chez les Diapriides myrmécophiles Lepidopria pedestris Kieffer et Solenopsia imitatrix Wasmann et leur hôte Diplorhotrum fugax Latr. (Solenopsis fugax Latr.). Biologie-Écologie Méditerranéenne 7(3): 183–184. Lachaud J-P (1981a) Les glandes tégumentaires chez deux espèces de Diapriidae: Aspects structuraux et ultrastructuraux. Bulletin Intérieur de la Section Française de l’UIEIS, Toulouse, France, 83–85. Lachaud J-P (1981b) Étude des relations hôte-myrmécophile entre les Diapriidae Lepidopria pedestris Kieffer et Solenopsis imitatrix Wasmann et la fourmi Diplorhoptrum fugax Latreille. Bulletin Intérieur de la Section Française de l’UIEIS, Toulouse, France, 135–137. Lachaud J-P (1981c) Étude des relations hôte-myrmdcophile entre les Diapriidae Lepidopria pedestris Kieffer et Solenopsia imitatrix Wasmann et la fourmi Diplorhoptrum [ugax Latreille. PhD Thesis, Univeristé Paul-Sabatier, Toulouse, 139 pp.
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