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1 Annales de la Société Entomologique de France International Journal of Entomology ISSN: (Print) (Online) Journal homepage: Leiophron Nees (Hymenoptera, Braconidae: Euphorinae): a debated genus. Provisional key for West Palaearctic species, additional data on related genera and newly collected material from Corsica Giuseppina Simbolotti, Claire Villemant & Marie-Cécile AndreÏ-Ruiz To cite this article: Giuseppina Simbolotti, Claire Villemant & Marie-Cécile AndreÏ-Ruiz (2002) Leiophron Nees (Hymenoptera, Braconidae: Euphorinae): a debated genus. Provisional key for West Palaearctic species, additional data on related genera and newly collected material from Corsica, Annales de la Société Entomologique de France, 38:4, , DOI: / To link to this article: Published online: 31 May Submit your article to this journal Article views: 97 Citing articles: 1 View citing articles Full Terms & Conditions of access and use can be found at

2 Ann. Soc. entomol. Fr. (n.s.), 2002, 38 (4) : ARTICLE Leiophron Nees (Hymenoptera, Braconidae : Euphorinae): a debated genus. Provisional key for West Palaearctic species, additional data on related genera and newly collected material from Corsica Giuseppina SIMBOLOTTI (1), Claire VILLEMANT* (2) & Marie-Cécile ANDREÏ-RUIZ (2) (1) Università dell Aquila, Dipartimento di Scienze Ambientali, Via Vetoio, I Aquila, Italy. (2) Muséum national d Histoire naturelle, Laboratoire d Entomologie (ESA 8043 du CNRS), 45, rue Buffon, F Paris, France. Abstract Following a sampling of the hymenopteran parasitoid fauna in Corsica and identification of the collected specimens, the taxonomical aspects of the genus Leiophron Nees (Hymenopteran, Braconidae: Euphorinae) have been studied. Partial improving of the only existing key (Loan, 1974) to the European species of the genus is proposed based on insular collected new series, type-material specimens and evidence that the species Leiophron reclinator (Ruthe) belongs to the recently new designed Euphorinae genus Mama Belokobylskij (2000). A short re-description of Leiophron fulvipes Curtis is given, taking into account the variability observed in Corsican specimens.the diagnosis characters of the two closely related genera Leiophron and Peristenus, pointed out in the most recent taxonomical studies are discussed taking into account the above-mentioned material. Résumé Leiophron Nees (Braconidae, Euphorinae) : un genre controversé. Données nouvelles apportées par l étude de spécimens de Corse et clé proposée pour les espèces ouest-paléarctiques. La collecte en Corse de nombreux représentants du genre Leiophron Nees, l examen du matériel type des espèces européennes du genre et l appartenance évidente de Leiophron reclinator (Ruthe) au nouveau genre Mama Belokobylkij (2000), ont conduit à la révision de la seule clé disponible (Loan, 1974) pour l identification des espèces européennes de Leiophron. L espèce Leiophron fulvipes Curtis est re-décrite en prenant en compte la variabilité morphologique observée parmi les spécimens de Corse. Le matériel examiné au cours de cette étude sert de base à un examen critique des caractères de diagnose proposés dans les études taxonomiques récentes pour les deux genres voisins Leiophron et Peristenus. Euphorinae is a braconid subfamily containing about 500 species. Genera recognized in the group vary from 35 (Shenefelt 1969) to 54 (Chen & van Achterberg 1997). The group is characterized by great diversity in host association matched by a similarly great morphological diversity. In fact, most braconid subfamilies are adapted to parasitism of a single host order, whereas euphorines are associated with several: they have been recorded on Orthoptera, Hemiptera, Psocoptera, Neuroptera, Coleoptera, Lepidoptera and Hymenoptera. Adaptation to parasitism of adult insects, which is rather uncommon within hymenopteran parasitoids, is widely found within the euphorines. Complexity of Euphorinae taxonomy is well expressed by the variable number of recognized tribes in most recent taxonomical studies: Shaw (1985) recognises 9 tribes (Perilitini, Dinocampini, Euphorini, * Corresponding author. E mail: villeman@mnhn.fr Manuscrit accepté le Cosmophorini, Townesilitini, Microctonini, Centistini, Loxocephalini, Syntretini) and favours recognition of the subfamily Meteorinae Cresson for the genus Meteorus Haliday. Only 4 tribes, Euphorini, Cosmophorini, Centistini, Meteorini, are recognized by van Achterberg (1993). Within the Euphorinae, the genera Euphorus Nees, Leiophron Nees, Peristenus Foerster, Euphoriella Ashmead and Euphoriana Gahan are described as parasitic on Miridae and Psocoptera. The taxonomic position of these genera was studied by Loan (1974), Shaw (1985) and, most recently, by Chen & van Achterberg (1997), the latter, however, deals only with specimens from China. The opinions among authors of these studies vary greatly, but all of them agree on one main point: the removal of the genus Peristenus from synonymy with the genus Leiophron (Loan 1974). The synonymy between these two genera date back to Marshall (1887). We obtained a number of Euphorines specimens from a sampling of hymenopteran parasitoid fauna in 339 Published online 31 May 2013

3 G. SIMBOLOTTI, C. VILLEMANT & M.-C. ANDREÏ-RUIZ Fango Valley (Corsica) carried out in order to assess the biodiversity of this Man and Biosphere Reserve. Two Euphorinae species were mainly found in the caught material: Leiophron pallidistigma Curtis and Leiophron fulvipes Curtis. The genus and the species were identified according to Loan s diagnosis of this genus and his key for West Palaearctic species (Loan 1974). Furthermore, we compared our specimens with the lectotypes designed by this author for L. pallidistigma and L. fulvipes, as well as with the holotypes and lectotypes of other West Palaearctic Leiophron species. From this examination more detailed knowledge of the intraspecific Figure 1 The Fango Valley in Corsica and the distribution of the traps for sampling of hymenopteran parasitoids. 340

4 Leiophron: a debated genus of Braconidae morphological variability range within the species L. fulvipes has derived, thus suggesting the need for a short species re-description. Comparison with other West Palaearctic species of Leiophron, as well as specimens of the genus Mama (Belokolbylskij 2000), also drew attention to some new morphological diagnostic characters, thus suggesting additions and re-adaptation of the only existing key (Loan 1974) for West Palaearctic species of the genus. Finally, the study provided useful evidence that the diagnosis of the genera Leiophron Nees and Peristenus Foerster is still largely contested. Materials and methods The Fango Valley is situated in North-West Corsica and is scheduled as a Man and Biosphere Reserve owing to its outstanding evergreen oak (Quercus ilex) woodland, a typical sclerophyllous forest ecosystem in the Mediterranean Region. In the Valley all stages of the green oak succession sequence are present. Sampling of the Fango Valley s parasitoid hymenopteran fauna was done using yellow pan traps positioned at 1, 3.5 and 14 meters above the ground, as well as yellow pan traps and emerging traps placed at ground level. A total of 87 traps were spread in 24 micro-stations. Traps were left for two weeks, during the months of May and June The studied plot (0.5 ha) is situated at an altitude of about 180 m and includes three vegetation areas: oak grove, low maquis with oak, Cistaceae and brambles, and an ecotone grove-maquis (fig. 1). The majority of the euphorines was caught in the grove and the ecotone grovemaquis. Leiophron fulvipes specimens were mostly collected in the lowest traps (on the ground and at 1 m above the ground), while the majority of L. pallidistigma individuals were caught in the tree crowns (Villemant & Andreï-Ruiz 1999). Other specimens of L. fulvipes were also collected in spring 1997, at the same place, using a Malaise trap. A total of 21 females and 38 males of L. fulvipes and 21 females and 11 males of L. pallidistigma caught in the Reserve were studied. Male genitalia of both species were extracted, cleared with lactophenol and mounted on slide with Euparal. The material collected in Fango Valley was compared with the type specimens of the West-Palaearctic euphorines species diagnosed by Loan (1974) as belonging to the genus Leiophron. The examined material was : Leiophron fulvipes Curtis, 1833: Holotype (NMVM) male: Ent. 886; type of Leiophron fulvipes, G. Nixon det. 1948; type; holotypus Leiophron fulvipes Curtis det. B. C. Loan, Leiophron pallidistigma Curtis, 1833: Lectotype (NMVM), female: 16 (on specimen point): type of Leiophron pallidistigma Curtis, G. Nixon det. 1948; type; lectotypus Leiophron pallidistigma Curtis, det. B. C. Loan, Paralectotype (NMVM), female: 16; Ent. 886; paralectotype Leiophron pallidistigma Curtis, det. Loan Paralectotype (NMVM), female: 6; Ent. 886; paralectotype Leiophron pallidistigma Curtis, det. Loan Leiophron apicalis Haliday, 1833: Lectotype (NMVM), female: Ent. 886; Euphorus apicalis; type; type of Leiophron apicalis Curtis, G. Nixon det. 1948; lectotype of Leiophron apicalis Curtis, det. G. C. Loan Paralectotype (NMVM), male: 17. Paralectotype (NMVM), male: Ent. 886; 7; paralectotype Leiophron apicalis Curtis, det. G.C. Loan Paralectotype (NMVM), female: no data on label. Leiophron basalis Curtis, 1833: Holotype (NMVM), male: C Hym n. 7; 16; basalis B. E. (undersurface of point); type of Leiophron basalis Curt., G. Nixon det Leiophron similis Curtis, 1833: Lectotype (NMVM), male: 16; Ent. 886; lectotype Leiophron similis Curtis, det. C. Loan Paralectotype (NMVM), female: 16; not type of Leiophron similis Curt., G. Nixon det Leiophron ruthei Loan, 1974: Holotype (BMNH), male: holotype; Germany ; M. claviventris, Wsm.; Ruthe Coll ; B.M. type, Hym 3c 1913; holotype Leiophron ruthei, Loan. Leiophron truncator Ruthe, 1856: Holotype (BMNH), female: type H.T. B. M. type Hym. 3c 715; 16; Microctonus truncator Ruthe 1856; M. truncator m; Ruthe Coll Leiophron fascipennis (Ruthe), 1856: Lectotype (BMNH), male, designed Richards, 1967, lectotype; B. M. type Hym. 3 C 718; M. fascipennis Rut.; Ruthe Coll Leiophron heterocordyli Richards, 1967: Holotype (BMNH), female: type 72350; B. M. type Hym. 3C 1650; Leiophron heterocordyli Richards, female, holotype; Berks., Silkwood Park, ex nymph Heterocordylus tibialis. Leiophron deficiens (Ruthe), 1856: Lectotype (BMNH), female, designed Richards, 1967; Germany ; B.M. type Hym. 3C 716; Ruthe Coll ; Microctonus deficiens Ruthe 1856, female lectotype Richards 1966; M. deficiens Rut. Leiophron reclinator (Ruthe), 1856: Lectotype (BMNH), female; lectotype; B.M. Hym. type 3c 1943; lectotype Leiophron reclinator, design. G.C. Loan; M. reclinator (not clear) M.; Ruthe Coll ; Germany; M. reclinator Rut. Mama mariae Belokolbylskij, 2000: Paratypes, 2 females. Abbreviations. NMVM: National Museum of Victoria, Melbourne. BMNH: British Museum of Natural History. Figures 2-4 show how diagnostic characters of the Leiophron species were measured. Wing venation nomenclature refers to Loan (1974) and Shaw (1985). Redescription of Leiophron fulvipes Curtis (figs. 5-17) Leiophron fulvipes Curtis, 1833: 476. Euphorus fulvipes Curtis, 1837: 118. Redescription Female (fig. 17). Length of body: mm. Head. Head (figs. 5-7) rather rectangular, entirely smooth and polished, its height times the width. Antennae (fig. 11) comparatively less slender and much shorter than in L. pallidistigma (fig. 24): length of antenna times the length of the body. Number of flagellomeres: Length of flagellomeres times their width: 3.0 (1 st to 2 nd ), 2.0 (3 rd to 4 th ), 341

5 G. SIMBOLOTTI, C. VILLEMANT & M.-C. ANDREÏ-RUIZ Figures 2-4 Measurements. 2, frontal view of the head (a, malar space; b, distance between the lowest point of the eye and the clypeal pit; c, subantennal distance between the eyes; d, distance between the anterior articulations of the mandibles. 3, dorsal view of the head (e, width of temple; f, width of the eye). 4, spiracles position on first metasomal tergite (g, length of first tergite; h, distance between the spiracle and the tergite apical end. 1.6 (4 th to penultimate), penultimate distinctly quadrate. Eyes large and convergent (figs. 5-7): height of eye about times its width (fig. 7). Subantennal distance between eyes comparatively shorter than in male specimens: about 0.7 times the distance between the insertions of mandibles; malar space about as long as the distance between the lowest point of the eye and the clypeal pit (figs. 2c, d; 5). Width of the temple in dorsal view of the head, times the width of the eye (figs. 3e, f; 6). Anterior margins of posterior ocelli at the level of posterior margins of the eyes, occipital carina broadly broken behind vertex (fig. 6). Mesosoma. Largely glabrous all over with only a few long hairs medially, pronotal sides and mesopleurae sometimes weakly sculptured but generally smooth. Length of mesosoma times its width in lateral view. Notauli distinctly present on the basal 1/3 of mesoscutum (fig. 13) in the specimens from Fango Valley, but entirely absent in the holotype. Prescutellar sulcus with median carina. Prescutellar grooves mostly completely and distinctly sculptured (specimens from Fango Valley), but, also, at times smooth all over (type). Precoxal sulcus distinct, moderately deep, thin. Propodeum largely sculptured with various, mainly curved distinct carinae. Wing. Wing venation comparatively more defined and complete than in L. pallidistigma. Fore wing (fig. 16): radial cell short but comparatively less narrow than in L. pallidistigma (fig. 29); stigma about 2.0 times as long as wide; 2 nd abscissa of the discoideus and sub-discoideus distinctly present, although at times unsclerotized. Submediellan cell of hind wing open. Legs. Length of hind femur times its width. Length of hind tibia times its width. Length of hind basitarsus times its width. Metasoma. First tergite dorsally distinctly sculptured with fine rugulae (fig. 14) or reticulae (fig. 51) all over its apical half. Position of the 1 st tergite s spiracles rather low (figs. 4g, h; 14, 50): first tergite 2.0 times as long as the distance between spiracles and the tergite s apex. Ventral sides of the 1 st tergite (fig. 15) distinctly separated all along the tergite s length and distinctly diverging both basally and apically. Length of the 1 st tergite about 2.0 to 2.5 times its apical width. Remainder of metasoma smooth, glabrous and polished. Ovipositor very short and curved (fig. 17). Colour. Dark yellowish-brown. Male. Usually somewhat smaller than female. Head with eye comparatively much smaller (height about times its width), less convergent and more removed from clypeal pit (figs. 6-10). Antennae distinctly thinner and longer than in female (fig. 12). Subantennal distance between eyes as long as the distance between the anterior articulations of the mandibles; malar space times the distance between the lowest point of the eye and the clypeal pit (figs. 2, 8). Length of temple, in dorsal view of the head, times the width of the eye (figs. 3e, f; 9). Anterior margin of posterior ocelli behind posterior margin of the eyes. Mesoscutum more often lacking notauli and prescutellar grooves more often unsculptured than in the female. Remainder of the body largely similar to the female. Diagnostic characters for Leiophron fulvipes L. fulvipes is very close to L. pallidistigma with which it seems to share the host: in fact, while other Leiophron species are reported as parasitoids of Mirids, L. fulvipes and L. pallidistigma are said to be specialised on parasitation of Psocoptera (Loan 1974). External morphological differences between the two species are rather reduced: in general L. pallidistigma (figs ) can be distinguished from L. fulvipes only by its smoother and very polished aspect, its somewhat larger body-size, its elongated antennomeres and its comparatively reduced venation of fore wing. Loan (1974), however, has shown that the differences of male genitalia of the two species are clear cut (figs ), and our material from Corsica (figs ) confirm this: in fact, contrary to what clearly occurs in L. pallidistigma (figs. 31, 32), the ventral face of the parameres in males of L. fulvipes is not modified into median and apical hooks (figs. 33, 34). Discussion After Loan (1974) removed the genus Peristenus Foerster from synonymy with Leiophron Nees, Shaw (1985) and Chen & van Achterberg (1997) treated 342

6 Leiophron: a debated genus of Braconidae the taxonomy of these two genera. Table 1 shows the contrasting diagnosis of these two genera by all these authors. In addition, due to the fact that after Loan, no author used genitalia morphology, there is no objective way to estimate the validity of their diagnosis compared to Loan s. The separation of the genus Peristenus Foerster from Leiophron Nees was done by Loan based on the following characters: in Peristenus the sides of tergite 1 Figures 5-17 Leiophron fulvipes Curtis, specimens from Fango Valley (Corsica). 5, 6, 7, head, facial, dorsal and lateral view in female. 8, 9, 10, same in male. 11, female antenna. 12, male antenna. 13, female, mesoscutum and scutellum in dorsal view. 14, female first tergite in dorsal view. 15, same, ventral view. 16, female fore wing. 17, female habitus. 343

7 G. SIMBOLOTTI, C. VILLEMANT & M.-C. ANDREÏ-RUIZ Table 1 The way how different authors separate the genera Leiophron Nees and Peristenus Foerster. : Peristenus; : Leiophron; n.s. : not specified. AUTHOR OPINIONS CHARACTERS MORPHOLOGY Loan, Shaw, Chen & van Achterberg, HEAD Occipital carina - complete n.s. - dorsally and usually interrupted (fig. 6) n.s. Notauli - present (fig. 13) MESOSOMA - absent (fig, 26) Surface of mesonotum - smooth - punctured Sub-discoideus - present (fig. 36d) n.s. FRONT - absent (fig. 37) n.s. WING Discoidal cell - much more setose than median cell n.s. n.s. - as setose as median cell n.s. n.s. METASOMA Ventral aspect of the sides - parrallel or subparallel of the first tergite - diverging apically - fused at base - touching - separated (fig. 15) Ventral face of male paramers - anteriorly as long as dorsal face n.s. n.s. - anteriorly longer than dorsal face (fig. 34) n.s. n.s. MALE Aedeagus apex shape - rounded or squared off n.s. n.s. GENITALIA - pointed with protruding structures (fig. 34) n.s. n.s. Aedeagus length - longer posteriorly than paramers n.s. n.s. - shorter than paramers (fig. 34) n.s. n.s. are fused at the base and usually diverge so that apex of tergite 1 is petiolate and much wider than the base; the ventral face of male parameres is about the same length anteriorly as the dorsal face, the apex of the aedeagus is rounded or squared-off, the aedeagus itself is longer posteriorly than parameres and not at all enclosed by apices of parameres. Leiophron, on the contrary, would have sides of tergite 1 not fused basally and subparallel, the apex of this tergite a little wider than its base or at least subequal, the ventral face of parameres longer than dorsal face, the apex of the aedeagus pointed with protruding structures, and, finally, the aedeagus somewhat enclosed by the apices of the parameres. In Leiophron specimens from Corsica, the genitalia characters according to Loan s diagnosis are confirmed (figs ). Similarly, the ventral aspect of tergite 1 as described by this author was observed in all the Corsican specimens (figs. 15, 27) as well as in other types of Leiophron species. However we found little confirmation of Chen & van Achterberg s Leiophron diagnosis from our collected material, especially from L. fulvipes specimens: we cannot say that their notauli are entirely missing nor that their first discal cell of the fore wing is much more setose than the basal cell. All specimens from Corsica (both L. fulvipes and L. pallidistigma) show the first discal cell of the fore wing as setose as the basal cell. Moreover, at least three (i.e. L. basalis, L. similis and L. heterocordyli) of the Leiophron species-types we examined, showed distinct notauli (figs ). According to Shaw (1985) and Chen & van Achterberg (1997) the sub-discoideus (i.e. veins 3-CU1 and CU1a, in the nomenclatorial system used by these authors) is absent in the fore wing venation of Leiophron, while is clearly present in Peristenus (tab. 1). Status of this character is not reported in Loan (1974). Presence of the sub-discoideus is variable among our collected specimens of L. pallidistigma (but absent in the lectotype), while clearly distinct in all Corsican specimens and in the type of L. fulvipes, as well as in L. truncator and L. heterocordily types. In the remaining types of West-Palaearctic species of Leiophron, the character is variable in that the sub-discoideus is only partly present. Based on the pre-existing taxonomical situation, the study of the collected material from Fango Valley seems to support the distinctive male genitalia conformation 344

8 Leiophron: a debated genus of Braconidae attributed by Loan to the genus Leiophron as compared to the genus Peristenus. However, the fore wing venation of the specimens, as well as the venation of some of the types of the Palaearctic species of Leiophron, suggest a somewhat less clear-cut status of the genera Leiophron and Peristenus. Since wide specimens series as the ones we collected in Fango Valley are not yet available for all the various Palaearctic Leiophron and Peristenus described Figures Leiophron pallidistigma Curtis, specimens from Fango Valley (Corsica). 18, 19, 20, head, facial, dorsal and lateral view in female. 21, 22, 23, same in male. 24, female antenna. 25, male antenna. 26, female mesoscutum and scutellum in dorsal view. 27, female first tergite, dorsal view. 28, same, ventral view. 29, female fore wing. 30, female habitus. 345

9 G. SIMBOLOTTI, C. VILLEMANT & M.-C. ANDREÏ-RUIZ species, a final support in favour of Loan s distinctive characters for the two genera could only be provided by dissection and analysis of the genitalia morphology of the existing species types. However, the list of the examined type-material points out how often the types-series of Leiophron species only consist of one unique very old specimen. In this given situation, is not possible to carry out an extended genitalia analysis. In his key of the Leiophron species, Loan (1974) included Leiophron reclinator Ruthe, mentioning the hind wing venation characters and the antennal segments number. Presently, the holotype of this species a female in very bad condition is entirely lacking the head and has just enough remaining hind wing to show that the mediellan cell is closed (fig. 38); the remaining characters, due to the specimen s condition, are not clear. The recently, newly described Euphorinae genus, Mama (Belokolbylskij 2000), is characterised by the presence of spines on the scapus, a peculiar shape of the telotarsus (fig. 40), a straight ovipositor and a clearly closed mediellan cell on the hind wing. A specimen we found in the British Museum Collection, labelled L. reclinator and Germany Ruthe Coll., appears to be similar to the L. reclinator female holotype. Due to its fore telotarsus shape (fig. 41), it seems to be better related to the new genus Mama than to Leiophron. Evidence from the study of other possible Mama species (Simbolotti and Villemant, in prep.) have further convinced us, that L. reclinator could probably be moved to the genus Mama. The four characters on which this genus is designed seem very distinctive in relation to the characters currently defining the genus Leiophron. In addition, contrary to most of the species attributed to Leiophron, the only so far officially recognized species of the genus Mama, Mama mariae, is based on a typeseries of several specimens among which intraspecific variability seem scarce, thus giving someway additional support to the validity of this newly found genus. For all these reasons, therefore, we decided provisionally not to include the species L. reclinator in our key for West- Palaearctic Leiophron. Obviously, before really moving L. reclinator to the genus Mama it will be needed to have available, for study, more specimens of such a doubtful Leiophron species. Conclusions Emerging from the present study, the following problems seem to affect the taxonomy of the genus Leiophron in the West Palaearctic Region: a) to date, there is no clear agreement, as to how characterize this genus; Figures Male genitalia morphology in Leiophron, as found in Fango Valley specimens (figs. 32, 34) and as described by Loan (1974) (figs. 31, 33). 31, L. pallidistigma (redrawn after Loan, 1974). 32, idem (Fango Valley). 33, L. fulvipes (redrawn after Loan, 1974). 34, idem (Fango Valley). (a, ventral view; b, dorsal view) b) there are not sufficiently large type-series and available number of collected specimens to enable elucidation of the problem; 346

10 Leiophron: a debated genus of Braconidae c) there is evidence of characters shared by Leiophron/Mama as well as of characters shared by Peristenus/Leiophron. This the actual state of the facts, therefore it is clear that, in further study and/or interest about the genera Leiophron/Peristenus, priority must be given to large Figures Fore and hind wings in various species of Leiophron. 35, L. fascipennis, fore wing, male lectotype. 36, L. truncator, fore wing, female holotype (a, radius; b, cubitus; c, discoideus; d, subdiscoideus; A, radial cell; B, discoidal cell; C, brachial cell; D, median cell; E, submedian cell). 37, L. deficiens, fore wing, female lectotype. 38, L. reclinator and L. deficiens, submediellan cell (E ) of hind wing in female lectotypes. 39, L. pallidistigma, female from Corsica, hind wing with open submediellan cell. Figures Distinctive character of the genus Mama Belokolbylskij. 40, dorsal view of the fore tarsus of Mama mariae female paratype. 41, fore tarsus of the specimen reclinator, Germany Ruthe Coll : a, dorsal view; b, ventral view. 347

11 G. SIMBOLOTTI, C. VILLEMANT & M.-C. ANDREÏ-RUIZ collecting of specimen-series for the various species of the two genera (possibly, together with detailed data on the parasitized host-species). In the absence of abundant material and comprehensive ecological data, it would be highly unlikely to be able to formulate a stable taxonomy for these two or other closely related genera of Euphorinae. Actually, a state of synonymy between Leiophron and Peristenus would perhaps be more appropriate in view of the scarcity of the currently available material and data, as well as the contradictory opinions held by taxonomists who, recently or in the past, revised the two genera. Key to West-Palaearctic species of Leiophron Nees In the key, the genus Leiophron, and hence the included species, is provisionally interpreted according to LOAN (1974). With respect to this author s key, only additional characters for some species are introduced, while the number of overall treated species (except L. reclinator) remain unchanged. Therefore, for general description of any below-mentioned species the reader is referred to Loan s paper. 1. First metasomal tergite longer than 4.0 times its apical width (fig. 42); fore wing with band of yellowish colour distinctly running across the discoidal and brachial cell (fig. 35) L. fascipennis (Ruthe) Length of first metasomal tergite less than 4.0 times its apical width (figs ); discoidal and brachial cell of fore wing without band of colour (figs. 36, 37) Submediellan cell of hind wing closed (fig. 38); venation of fore wing largely incomplete (fig. 37), with cubitus, discoideus and sub-discoideus distinctly missing; first metasomal tergite about twice as long as its apical width (fig 43) L. deficiens (Ruthe) Submediellan cell of hind wing open (fig. 39); venation of fore wing complete (fig. 36); length of first tergite variable First metasomal tergite rather short and apically wide (fig. 44), its length less than or equal to 1.5 times its apical width; antennae comparatively short less than 0.5 times the length of the body L. heterocordyli Richards Figures Dorsal aspect of the first metasomal tergite in various Leiophron species. 42, L. fascipennis, male lectotype. 43, L. deficiens, male lectotype. 44, L. heterocordyli, female holotype. 45, L. truncator, female holotype. 46, L. apicalis, female lectotype. 47, L. similis, male lectotype. 48, L. basalis, male holotype. 49, L. ruthei, male holotype. 50, L. pallidistigma, female lectotype. 51, L. fulvipes, male holotype. 348

12 Leiophron: a debated genus of Braconidae First tergite comparatively longer than 1.5 times its apical width (figs ); antennae longer than 0.5 times the length of the body Propodeum unusually short (fig. 52) with posterior surface abruptly vertical; first metasomal tergite with distinctly knob-like spiracles (fig. 45) L. truncator (Ruthe) Propodeum of normal shape (figs ); spiracles of the first tergite comparatively smaller (figs ) Spiracles on first tergite positioned comparatively more basally (figs. 46, 47) with length of the tergite less than 1.7 times the distance between the spiracle position and the tergite s apex (fig. 4g, h); width of the temple in dorsal view of the head less than 0.4 times the width of the eye (figs. 3: e, f; 60-63) Spiracles on first tergite positioned rather apically (figs ), with the tergite distinctly longer than 1.7 times the distance between the spiracles position and the tergite s apex (fig. 4: g, h); width of the temple in dorsal view of the head (fig. 3: e, f) distinctly more than 0.4 times the width of the eye (figs ) Notauli absent; propodeum usually with distinct longitudinal carinae arranged in a sort of rectangular pattern L. apicalis Curtis Notauli present (fig. 55); propodeum largely foveolated L. similis Curtis 7. Notauli very well defined, complete, usually sculptured (figs ) Notauli entirely missing (fig. 58) or at least more scarcely defined and incomplete (fig. 59) Eyes distinctly not converging (figs ); notauli somewhat comparatively thinner (fig. 56); ventral margin of the clypeus not truncate (fig. 67)..... L. basalis Curtis Eyes comparatively more convergent (figs ); notauli very deep, complete, haevely sculptured (fig. 57); ventral margin of the clypeus medially truncate (fig. 69) L. ruthei Loan Figures Lateral view of the propodeum in various Leiophron species. 52, L. truncator, female holotype. 53, L. pallidistigma, female lectotype. 54, L. ruthei, male holotype. Figures Dorsal view of mesoscutum and scutellum. 55, L. similis, male lectotype. 56, L. basalis, male holotype. 57, L. ruthei, male holotype. 58, L. pallidistigma, female from Corsica. 59, L. fulvipes, male holotype. 349

13 G. SIMBOLOTTI, C. VILLEMANT & M.-C. ANDREÏ-RUIZ 9. Ventral face of parameres in male genitalia modified into median and apical hooks (fig. 32); antennae very long and thin (figs ) with most flagellomeres distinctly elongate and, usually, somewhat pale; prescutellar grooves entirely smooth (fig.58); fore wing venation usually lacking sub-discoideus and 2 nd abscissa of the discoideus (fig. 29); body-size large; mesoscutum usually smooth, polished, entirely lacking notauli (fig. 58) L. pallidistigma Curtis Ventral face of parameres in male genitalia without median and apical hooks (fig. 34); antennae comparatively short (figs. 11, 12) with much fewer elongate flagellomeres and always dusky; prescutellar grooves usually finely sculptured (fig. 59); fore wing venation always with sub-discoideus and 2 nd abscissa of the discoideus (fig. 16), although they can be, at times, unsclerotized; mesoscutum in females often with traces of notauli (fig. 59) L. fulvipes Curtis Figures The dorsal aspect of the head in various species of Leiophron. 60, L. apicalis, female lectotype; 61, L. apicalis, male paralectotype , L. similis, male lectotype. 63, L. similis, female paralectotype. 64, L. basalis, male holotype. 65, L. heterocordyli, female holotype. 66, L. pallidistigma, female lectotype. Figures Facial and frontal view of the head in some species of Leiophron. 67, 68, L. basalis, male holotype. 69, 70, L. ruthei, male holotype. Acknowledgements We are especially grateful to Miss Suzanne Lewis and Miss Christine Taylor of the British Museum (Natural History) for their helpful collaboration in providing Leiophron specimens present in the Museum collections, as well as several types. We are similarly indebted to Dr. Ken Walker of the National Museum of Victoria, Melbourne, for kindly providing types of the Curtis collection, to Dr. P. Belokolbylskij for sending us type material of the genus Mama and to Dr. Kees van Achterberg of the Museum of Natural History Naturalis in Leiden (The Netherlands) for his careful revision of the first draft of the paper and his fruitful suggestions in improving the final version of it. A special thanks goes to Mrs Denise Palczewski for her kind contribution in correcting and improving the English text. Finally, the senior author is deeply obliged to the Museum National d Histoire Naturelle for financial support during her working at the Laboratoire d Entomologie, where the major part of the research has been carried out. REFERENCES ACHTERBERG C. Van 1993 Illustrated key to the subfamilies of Braconidae (Hymenoptera: Braconidae). Zoologische Verhandelingen, 283 : BELOKOLBYLSKIJ S.A A new genus and subgenus of the subfamily Euphorinae (Hymenoptera: Braconidae) from East Asia. Zoologische Mededelingen, 73 : CHEN X., ACHTERBERG C. Van 1997 Euphorinae from China. Zoologische Verhandelingen, 313 : LOAN C.C The European species of Leiophron Nees and Peristenus Foerster (Hymenoptera: Braconidae, Euphorinae). Transaction of the Royal entomological Society of London, 126 : MARSHALL T.A Monograph of British Braconidae, 2. Transactions of the Royal entomological Society of London, 2 : SHAW S.R A phylogenetic study of the subfamilies Meteorinae and Euphorinae (Hymenoptera, Braconidae). Entomography, 3 : SHENEFELT R.D Braconidae 1:. in CH. Ferriere & J. Van Der Vecht (eds), Hymenopterorum Catalogus, Pars 4. W. Junk, s-gravenhage, 176 p. VILLEMANT C., ANDREI-RUIZ M.C Diversité et répartition spatiale des Hyménoptères parasitoïdes dans la chênaie verte du Fango (Haute Corse). Actes de la IV Conférence Internationale Francophone d Entomologie, Saint-Malo (France), 5-9 juillet, Annales de la Société entomologique de France (n.s.), 35 (suppl.) :