Quasimodoana, a new Holarctic genus of eucoiline wasps (Hymenoptera, Cynipoidea, Figitidae), with a phylogenetic analysis of related genera

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1 Systematic Entomology (2008), 33, Quasimodoana, a new Holarctic genus of eucoiline wasps (Hymenoptera, Cynipoidea, Figitidae), with a phylogenetic analysis of related genera M A T T I A S F O R S H A G E 1, G Ö R A N N O R D L A N D E R 2 and F R E D R I K R O N Q U I S T 3 1 Department of Systematic Zoology, Evolutionary Biology Centre, Uppsala, Sweden, 2 Department of Ecology, Swedish University of Agricultural Sciences, Uppsala, Sweden and 3 Department of Entomology, Swedish Museum of Natural History, Stockholm, Sweden Abstract. A new Holarctic genus, Quasimodoana gen.n. is described for the Palearctic eucoiline wasp Eucoila decipiens Fo rster, 1869, as Quasimodoana decipiens comb.n. A lectotype is designated for this species. A new North American species is described here as Quasimodoana gibba sp.n. As the new genus has similarities with several eucoiline genera, belonging to two related but distinct lineages, a phylogenetic analysis was carried out, based on 83 morphological characters. The two lineages included in the analysis are referred to here under the tribal names of Eucoilini and Trichoplastini stat.nov. We discuss reasons for adopting available tribal names (rather than informal genus groups ), and Trichoplastini is removed from synonymy with Eucoilini. The phylogenetic analysis places Quasimodoana unambiguously within Eucoilini, as a sistergroup to the Trybliographa complex. However, there is only weak support for the monophyly of Eucoilini and for the hypothesized sistergroup relationship between Eucoilini and Trichoplastini. Introduction Eucoilinae is a group of Diptera parasitic Cynipoidea, which has historically been treated as a distinct family but is now considered a subfamily of Figitidae (Ronquist, 1999). This poorly known group contains slightly less than 1000 nominal species (Va rdal, 2004), but the identities of many remain doubtful. Indeed, most Eucoilinae remain undescribed, even within the Holarctic region. With Eucoila Westwood, 1833 being the oldest genus in Eucoilinae, a number of species were originally described as being within that genus, but were then transferred to other genera of younger date, or simply forgotten, thus remaining within Eucoila as nomina dubia. Often taxa do not belong in Eucoila, but correct placement is denied by the loss of types and inadequate descriptions. Probably many nomina dubia refer to species of Trybliographa Fo rster, 1869, as this is the Correspondence: Mattias Forshage, Department of Systematic Zoology, Evolutionary Biology Centre, Norbyvägen 18D, SE Uppsala, Sweden. mattias.forshage@ebc.uu.se principal Holarctic genus of relatively large-sized eucoilines. However, some type specimens do exist, and it is only a task for the systematist to establish new combinations or describe the necessary new genera. The circumscription of Eucoila established by Weld (1952), and representing the long-held consensus view, actually grouped two quite distinct lineages. The Holarctic crassinerva group (Eucoila s.str., as crassinerva Westwood, 1833 is the type species) and the New World nudipennis group were separated first by Nordlander (1981). The nudipennis group was confirmed later to be distant to the true Eucoila, belonging to the neotropical grade and not to the higher eucoilines in the analysis of Fontal-Cazalla et al. (2002). The species group has been studied recently by Schick (1999), but the new genus name proposed in her unpublished doctoral dissertation is not available formally, and the common way of referring to this taxon remains Eucoila with quotation marks (Nordlander, 1982b; Fontal- Cazalla et al., 2002). The type material of one of the dubious Eucoila species, E. decipiens Fo rster, 1855 from Europe, studied by GN, is Journal compilation # 2008 The Royal Entomological Society 301

2 302 M. Forshage et al. not assignable to any valid genus and actually represents a distinct lineage, which also has Nearctic representatives. This lineage is described herein as a Quasimodoana gen.n. We describe a new species from North America; although other specimens examined probably represent additional new species, we refrain from describing these here because of the scarcity of specimens and the difficulties in associating the sexes. A phylogenetic analysis was carried out in order to investigate the phylogenetic position and monophyly of the new genus. Because Quasimodoana shares important characters both with Trybliographa and Leptopilina of Eucoilini and with Trichoplasta and Rhoptromeris of Trichoplastini, it is not immediately obvious to which of these lineages it belongs. The Eucoilini and Trichoplastini appeared (as the Trybliographa group and the Rhoptromeris group ) as sistergroups in Fontal-Cazalla et al. s (2002) analysis of higher-level eucoiline phylogeny, but support for this was not strong. Subsequently, in the analysis of Buffington et al. (2007) they grouped, but with Eucoilini found to be paraphyletic. In the phylogenetic analysis of the position of Quasimodoana we focused our attention on these two lineages, using a few other higher eucoiline representatives as outgroups, primarily because of the mosaic of Eucoilini and Trichoplastini characters in Quasimodoana and because we hoped that such an analysis might help to elucidate relationships. Tribal classification Nordlander made a tentative subdivision of Eucoilinae into informal genus groups (Nordlander, 1982b), and most workers have retained these informal group names. This has perhaps primarily been motivated by a relative lack of knowledge of the phylogeny of the group. The few authors utilizing family-level names within Eucoilinae usually applied the names to groupings that were not founded in careful hypotheses of phylogeny, and most available names were synonymized into Eucoilinae by Ronquist (1999). Taking the analyses of Fontal-Cazalla et al. (2002) and Buffington et al. (2007) into account, it would seem convenient to use available tribal names for those established genus groups having gained some support, in order to facilitate the discussion of phylogenetical relationships. Informal groups make sense on an informal level, or within the context of a single investigation. When informal groups become established they become similar to formal classificatory units and may even be mistaken for such, but without being subject to formal classificatory rules nor being available to formal classificatory actions. Thus, the primary purpose of erecting tribes here is to make conventional groups available formally as taxonomic units, and to encourage testing. The name Eucoilini is available with the same authorship as Eucoilinae, Thomson, The core of Eucoilini, as delineated here, consists of Eucoila (type genus), Trybliographa, and Bothrochacis Cameron, 1904, which are the traditional genera in the Trybliographa group (Nordlander, 1982a, b) [note that Trybliographa after the revision of Nordlander (1981) includes Psichacra Fo rster, 1869, Pseudeucoila Ashmead, 1903, Piezobria Fo rster, 1869, and Pilinothrix Fo rster, 1869]. Fontal-Cazalla et al. (1982) showed that Leptopilina belongs to the same lineage. Furthermore, Linaspis Lin, 1988 and Linoeucoila Lin, 1988 are included tentatively based on the original descriptions (Lin, 1988), although we cannot confirm this by examination of the type specimens. We refer Maacynips Yoshimoto, 1963 also to this group (Table 1). Morphologically, these taxa are recognized by the males having the second flagellomere modified (shared with Trichoplastini) and by having the posteroventral corner of the metapleuron raised, facing posterolaterally, entirely or almost glabrous. This latter structure has lacked a name, forcing authors to use long descriptions such as surface immediately posterior to (posteroventral corner of metapleuron) (probably a part of propodeum) is hairless (...), facing latero-posteriorly or, more rarely, directly posteriorly (Nordlander, 1981) and lateral part of metacoxal rim forming a smooth and nude triangular projection having its surface directed more laterally (Fontal-Cazalla et al., 2002). As the structure is often triangular in shape we refer to it as the metapleural triangle. Trichoplastini Kovalev, 1989 hereby is removed from synonymy with Eucoilidae Thomson, 1862, stat.n. It contains Trichoplasta (type genus) and Rhoptromeris from the Rhoptromeris group (note that Armigerina Belizin, 1968 is included in Trichoplasta). Daruna Benoit, 1956, Stentorceps Quinlan, 1984, and Angustacorpa Quinlan, 1988 were included in this group by Quinlan (1984, 1986, 1988), and Nordlanderiana was included by Kovalev (1989). Morphologically, these taxa are easily recognizable by having lateral bridges on the pronotal plate (closed lateral pits), and sharing the state of a modified second male flagellomere with Eucoilini (Table 2). Materials and methods Morphological terminology follows Nordlander (1982b), Ronquist & Nordlander (1989), Ronquist (1995), and Fontal-Cazalla et al. (2002). Descriptions are based on observations utilizing a Leica MZ16 stereo microscope and a Leo/Zeiss Supra 35VP scanning electron microscope (SEM). One drawing was made by hand, based on stereo microscope observations, but the rest were carried out in Adobe Photoshop CS2 based on images from various sources (digital images captured through the stereo microscope, SEM images, direct observations). All SEM images were captured with the Leo/Zeiss instrument mentioned, but only one of them with the usual process of goldcoating and high vacuum. All the images of Q. gibba (and one image of Q. decipiens to facilitate direct

3 Quasimodoana, new genus of eucoiline wasps 303 Table 1. Genera included in Eucoilini. The source given is for inclusion in this tribe (or in its equivalent the Trybliographa group ). Note that the number of nominal species is often substantially smaller than the real number. Some of these genera are poorly studied, and their robustness as distinct genera should not be considered as confirmed by the present authors simply by their inclusion in this table. Genus No. of species Hosts Habitats Geographical distribution Source Trybliographa Fo rster, Anthomyiidae Mushrooms, rootcrops, leaves of herbs, ferns, dung, conifer cones, etc. Eucoila Westwood, Muscidae, Calliphoridae, Sarcophagidae Leptopilina Förster, Drosophilidae Decaying fruit, mushrooms, and decaying debris in general Widespread but predominantly Holarctic Dung and carrion Widespread but predominantly Palearctic Nordlander (1982a, b); Fontal-Cazalla et al. (2002) Nordlander (1982a, b); Fontal-Cazalla et al. (2002) Cosmopolitan Fontal-Cazalla et al. (2002) Quasimodoana gen.n. 2?? Holarctic Present consideration Bothrochacis Cameron, ?? Afrotropical Nordlander (1982a, b) Linaspis Lin, ?? Taiwan only considered close to Leptopilina in Lin (1988) Linoeucoila Lin, ?? Taiwan only Lin (1988) Maacynips Yoshimoto, ?? Australopacific region Present consideration Table 2. Genera included in Trichoplastini. The source given is for inclusion in this tribe (or in its equivalent the Rhoptromeris group ). Note that the number of nominal species is often substantially smaller than the real number. Some of these genera are poorly studied, and their robustness as distinct genera should not be considered as confirmed by the present authors simply by their inclusion in this table. Genus No. of species Hosts Habitats Geographical distribution Source Rhoptromeris Fo rster, Chloropidae Grasses, conifer cones, bracket fungi, etc. Trichoplasta Benoit, Lonchaeidae and others? Widespread but predominantly Palearctic and Afrotropical Decaying wood Widespread but predominantly Holarctic and Afrotropical Nordlander (1982a, b); Kovalev (1989); Fontal-Cazalla et al. (2002) Nordlander (1982a); Kovalev (1989) Nordlanderiana Kovalev, Lonchaeidae? Decaying wood Palearctic Kovalev (1989) Daruna Benoit, ?? Afrotropical Tentatively suggested in Quinlan (1986) Angustacorpa Quinlan, ?? Afrotropical Quinlan (1988) Stentorceps Quinlan, ?? Afrotropical Quinlan (1984, 1986)

4 304 M. Forshage et al. comparison) were captured in low vacuum with uncoated specimens because of the rarity of specimens, which unfortunately resulted in poorer image quality. Character coding for the phylogenetic analyses was to a large extent based on SEM and compound microscopy images available from MorphBank ( but additional taxa and some of the additional characters were coded from specimens, utilizing SEM and stereo microscopy. The phylogenetic analysis was performed with PAUP 4.0B10 (Swofford, 2002), with parsimony as the optimality criterion, with equal as well as implied weights (Goloboff, 1993), the latter with a series of concavity functions (Goloboff weights k ¼ 1 6), various ordering schemes, full Branchand-Bound search, plus bootstrapping (100 replicates, 25 random addition sequences). The resulting trees were further investigated in MACCLADE 3.08A (Maddison & Maddison, 1999). Abbreviations used for specimen data are as follows: AEI, American Entomological Institute, Gainesville, U.S.A.; CNC, Canadian National Collection, Ottawa, Canada; coll GN, G. Nordlander s collection at the Swedish University of Agricultural Sciences, Uppsala, Sweden; LZM, Zoologiska Museet, Lund, Sweden; NHRS, Naturhistoriska Riksmuseet, Stockholm, Sweden; USNM, U.S. National Museum, Washington, U.S.A.; ZMHB, Museum fu r Naturkunde, Berlin, Germany. Included characters The analysis re-utilized (after scrupulous proofreading and some recoding) relevant parts of the data matrix of Fontal-Cazalla et al. (2002), and added some taxa and a number of characters (app. 1, 2). The new characters were coded mostly in the same manner as the available matrix, from direct comparison between prints of SEM pictures of exemplar specimens in standard views [available from MorphBank ( Fifty of the characters analysed here are entirely new (1, 2, 6, 11, 13, 14, 15, 17, 18, 19, 20, 24, 26, 29, 30, 32, 33, 34, 35, 39, 41, 42, 45, 46, 47, 49, 50, 54, 55, 56, 57, 58, 59, 60, 61, 63, 66, 67, 68, 69, 71, 73, 74, 75, 78, 79, 80, 81, 82, 83 in the present matrix), and 33 are based on characters taken from the matrix in Fontal-Cazalla et al. (2002). From that source all characters that were informative in the groups of genera studied here were taken. Fourteen characters were unchanged [4, 11, 28, 30, 36, 51, 53, 56, 62, 111, 114, 122, 128, 135 of Fontal-Cazalla et al. (2002)], and 17 were modified, either because it was possible to introduce gradations of the differences within this more narrowly circumscribed group, or because new scrutiny of the states of single taxa necessitated correction [5, 7, 9, 17, 23, 47, 48, 57, 71, 72, 75, 81, 86, 91, 107, 113, 136 of Fontal-Cazalla et al. (2002); 107 was divided into three distinct characters, in the present matrix]. Some problematic characters [17, 116, 120 of Fontal-Cazalla et al. (2002)] were removed, as were several uninformative characters or states. Multistate characters are either gradual characters, in which character states occur along a one-dimensional continuum, or complex characters, which may be grouped into a transformation series through careful interpretation of character evolution. Logically, the former would best be treated as ordered (Wagner parsimony) and the latter as unordered (Fitch parsimony). Different characters may of course evolve in different ways, some in a leap-wise manner without passing intermediate stages, but little is known about this, and ideally we should proceed in a manner chosen from a methodological standpoint as sound regardless of what might eventually be discovered. For characters that present an obvious grade of character states, to treat them as unordered, implying that it would be equally easy to go from one extreme to another as to an intermediate, would in fact represent quite a strong assumption, and, not only does that strategy fail to retain information on similarity present in the matrix, it is contrary to the general scientific principle of parsimony. However, a priori weighting schemes are not that important if we recognize that all transformation series are actually tested by the tree itself (Mickevich, 1982; Mickevich & Lipscomb, 1991; Wilkinson, 1992; Schulmeister, 2003). Checking the distribution of character states of a multistate character in the most parsimonous trees (starting out from the one obtained with unordered characters) empirically shows whether the character evolution necessarily passes through the intermediate stages or just as easily goes from one end to the other. A method of ordering that decides whether to treat each individual character as ordered or unordered based on how it is distributed in the tree is here called phylogenetic ordering as opposed to logical ordering. Analyses were made with characters unordered, a priori logically ordered, and a posteriori phylogenetically ordered. In the present matrix, only two (64 and 75) of the 25 multistate characters did not present an obvious order from the a priori perspective, and these were treated as unordered even in the logically ordered analysis. Ten of the 25 multistate characters showed an unordered distribution (including character 64 but not 75), and 15 characters retained their logical order after analysis. To investigate the effects of ordering on tree topology, the dataset was split into two-state characters (58 characters) and multistate characters (25 characters), which were analysed separately, the latter both as a priori logically ordered and as a posteriori phylogenetically ordered. Differential weighting was tried, based a posteriori on tree topology, with Goloboff s implied weighting (Goloboff, 1993) as implemented in PAUP. To investigate the consequences, concavity functions with k ¼ 1 6 were tried, as well as equal weights. Included taxa The taxon set chosen for the analysis includes representatives of all the Holarctic genera of the Eucoilini and Trichoplastini, with more than one species from the larger

5 Quasimodoana, new genus of eucoiline wasps 305 genera with which the new genus shares some similarities namely Leptopilina, Trybliographa and Trichoplasta (Table 3). The phylogeny among the higher eucoilines is not well resolved, and the best outgroup for this analysis is unclear. Several candidates were tried but performed poorly because of many outgroup autapomorphies, making polarity decisions impossible for many characters, and some specific similarities with ingroup taxa that appeared as potential synapomorphies in the context of our analysis but that were probably convergences based on the more comprehensive analysis of Fontal-Cazalla et al. (2002) and other background knowledge. The taxa chosen as outgroups are Chrestosema Fo rster, 1869 and Glauraspidia Thomson, Both were included in the Chrestosema group of Fontal-Cazalla et al. (2002), but that group was supported only weakly, characterized by a set of non-unique synapomorphies, none shared by all genera in the group. The non-monophyly of the Chrestosema group has been confirmed since by molecular and combined analyses (Buffington et al., 2007). The Glauraspidia species used here as an outgroup is actually a partly brachypterous and rather autapomorphic species, but its odd features appear to be of no importance within this analysis. Descriptions Quasimodoana gen.n. Type species. Eucoila decipiens Fo rster, 1855 (Fig. 1) The new genus has an unusual combination of characters. It has a metapleural triangle similar to that characteristic of genera of Eucoilini, whereas the general appearance, including the long and pointedly oval female metasoma and the posteriorly extended scutellum, is more similar to that of Trichoplasta and Nordlanderiana of Trichoplastini. Most representatives are relatively large (similar to Trybliographa), Fig. 1. Quasimodoana decipiens (Fo rster, 1869), male. have a distinctly humped mesoscutum (similar to Leptopilina) and a more or less strongly modified second flagellomere of male antenna (similar to Leptopilina and Trichoplastini). They lack the foremost diagnostic characters of all these genera (the closed lateral pits of the pronotal plate of Trichoplastini, the broad petiolar rim and often reduced hairy ring of syntergite of Leptopilina, and the subalar pits of Trybliographa). All species we have seen are characterized (uniquely among their close relatives) by the distinct subantennal striae and by the strong depressions in the metapleura, and usually by the shortened first flagellomere (third antennomere) of the female antenna, the latter feature being emphasized by Fo rster (1855) in his description of the type species. Table 3. Taxa included in the phylogenetic analysis. Presence in the analysis of Fontal-Cazalla et al. (2002) is indicated. Preparation is the source of coding (available ¼ matrix taken from Fontal-Cazalla et al. (2002), additional coding from images available on MorphBank; dissected ¼ fully dissected to provide most standard views in SEM images; not exhaustively dissected ¼ rare species, only one or two specimens dissected, not covering several standard views in SEM images; not dissected ¼ very rare species, no SEM images, only stereomicroscopy. NB for all taxa, some additional coding has been made from stereomicroscopy). Taxon Reason for inclusion Preparation Glauraspidia microptera (Hartig, 1840) Outgroup Type species of genus In FC Available Chrestosema erythropum Fo rster, 1869 Outgroup Type species of genus In FC Available Quasimodoana decipiens (Fo rster, 1869) Ingroup Type species of genus Not in FC Not exhaustively dissected Quasimodoana gibba sp.n. Ingroup Tew species Not in FC Not dissected Eucoila crassinerva Westwood, 1833 Ingroup Type species of genus In FC Available Trybliographa longicornis (Hartig, 1840) Ingroup Typical of species group Not in FC Dissected Trybliographa cubitalis (Hartig, 1841) Ingroup Typical of species group Not in FC Dissected Leptopilina longipes (Hartig, 1841) Ingroup Type species of genus In FC Available Leptopilina heterotoma (Thomson, 1862) Ingroup Typical of species group Not in FC Dissected Leptopilina boulardi (Barbotin et al., 1979) Ingroup Typical of species group Not in FC Dissected Rhoptromeris heptoma (Hartig, 1840) Ingroup Type species of genus In FC Available Trichoplasta (Armigerina) sp. Ingroup Typical of species group In FC Available Trichoplasta (s.str.) bouceki (P. Masner, 1960) Ingroup Typical of species group Not in FC Not dissected Nordlanderiana grunini (Belizin, 1968) Ingroup Type species of genus Not in FC Not exhaustively dissected

6 306 M. Forshage et al. Etymology. The name is derived from Quasimodo in Victor Hugo s Notre Dame de Paris, in reference to the hunchback appearance of these wasps. The gender is feminine. Fig. 3. Quasimodoana gibba sp.n., pronotal plate. Description. Compactly built eucoiline wasps, with brown or black body, length mm. Head intermediate in size, strongly reflexed, shape more or less broadly oval. Eyes slightly protruding, small or large. Frons broad with more or less protruding antennal sockets, with distinct lateral striae (Fig. 2). Vertex smooth, evenly rounded or more or less conical, with small protruding ocelli. Occiput with several hairs, without carina. Scattered hairs on head mostly short. Mandible basally dark and hairy, apically light and glabrous. Maxillary palpus four-segmented, terminal segment enlarged towards apex, with an erect seta. Labial palpus two-segmented. Female antenna with 13 articles, first flagellomere more or less shortened, apical seven to nine flagellomeres forming a rather indistinct club. Male antenna with 15 articles, second flagellomere more or less modified (asymmetric, elongated, curved), with placoid sensillae on all flagellomeres. Mesosoma high, usually humped (anterior end of mesoscutum abruptly rising). Entire mesosoma usually strongly sclerotised, black, with bulging sclerites and strong ridges. Pronotum with several hairs, without distinct ridges lateral to pronotal plate, but with strong wrinkles ventrally. Pronotal plate of varying shape, always with submedian depressions laterally open but narrow, the dorsal rims of these depressions weakly delineated (ventrolateral parts of dorsal area of pronotal plate sticking close to surface of pronotum without a distinct edge) (Fig. 3). Mesoscutum with four rows of hair punctures; two submedian ones representing rudimentary notauli, and two lateral ones in posterior half. Scutellum large, posterior part in lateral view protruding (over propodeum) in a subrectangular shape, in dorsal view more or less evenly rounded. Sculpture of scutellum punctate-reticulate, asymmetrical and with punctures of different sizes intermixed, but largely with finer punctures near scutellar plate and increasingly rough towards the lateral edge, and longitudinally reticulate on lateral surfaces. Lateral bars rather short, indistinctly or rather weakly striate. Scutellar plate raised, oval-shaped, with a distinct margin and slightly concave disc with irregularly distributed hairs in small pits, and a large posterior glandular release pit. Mesopleuron glabrous, with ridges along lower margin and across lower third (precoxal carina and mesopleural carina), and a diagonal groove from middle of anterior margin to middle of dorsal margin (by the forewing insertion). Subalar groove well developed but without subalar pit. Metapleuron with varying amount of hairs, with several ridges extending forwards and upwards from posterior margin, with depressed areas in between (Fig. 4). Anteroventral cavity round. Posteroventral corner largely glabrous Fig. 2. Quasimodoana decipiens, male, face and basal parts of antennae. Fig. 4. Quasimodoana decipiens, metapleuron, posterolateral view.

7 Quasimodoana, new genus of eucoiline wasps 307 and directed posterolaterally, with a concave surface, forming a well-developed metapleural triangle. Anterodorsally from the metapleural triangle runs a more or less developed carina (which is probably homologous with ridge 3 sensu Nordlander (1980) in Leptopilina, but which here is more dorsally oriented than in other eucoilines), running almost parallel to the posterior edge, and separated from it by a strongly depressed area, dorsally adjoining the median metapleural carina or disappearing by the metapleural groove. Area anterior to this subposterior carina also depressed but less sharply delimited. Median metapleural carina and metapleural groove varying. Subalar groove deep and wide, the calyptral carina delimiting it ventrally strong. Calyptra distinctly protruding. Propodeal complex, including smaller or larger parts of metapleura, with long hairs. Propodeal ridges glabrous, lyre-shaped (with an angle between the divergent lower thirds and the slightly concavely rounded upper twothirds), joined by nucha ventrally but separate dorsally, only weakly protruding posteriorly, almost vertical in lateral view. Legs rather long, light-coloured. Middle and hind coxae convex, with a few parallel rows of hairs laterally and scattered hairs apically. Hind coxae with an additional distinct small dorsolateral tuft of hairs; middle coxae lacking such a tuft. Claws small and simple. Wings well developed, with short pubescence over entire surface. Ciliation along fringe well developed (intermediate between the short pubescence of Trybliographa and the long of Leptopilina). Forewings large and rather long, apically rounded, with ordinary figitid nervature (lacking costal vein, with distinct R þ Sc and marginal cell forming a W-like shape). Marginal cell closed, neither narrowed nor deepened (similar to Trybliographa, probably plesiomorphic), curvature of 2r and Rs varying, M þ Cu1, Cu1, Rs þ M and M reduced, nebulous or indistinct, 1A and Cu1a entirely indistinct. Hindwings narrow with a single vein, R þ Sc. Petiolus short, striate, posteriorly slightly enlarged. Hair ring on large tergite (fused terga of abdominal segments 3 to 5 (females) or 3 to 4 (males)) varying, weak or rather dense but never felt-like, complete except for a very short interruption dorsally, with very thin second row of long hairs. Female metasoma large, often strikingly long, usually with an outstretched, pointedly oval form. Hypopygium protruding, with a ventral row of setae. Ovipositor (terebra) long and strong, protruding. Male metasoma small, usually more or less quadrangular, suggesting diamond-shape, length approximately the same as mesosoma but height distinctly less. (Note, however, that overall metasomal shapes in eucoilines may vary substantially between specimens, depending on the degree and pattern of telescoping of the posterior tergites.) Distribution. Holarctic, with the single previously described species European, and one herein described species (plus some undescribed species) Nearctic. Biology. The single European species was collected in open deciduous forest. The Nearctic specimens usually lack such information. Hosts are unknown. All specimens studied were collected between June and October. Quasimodoana decipiens (Fo rster, 1855) comb.n. Originally described as Eucoila decipiens by Fo rster (1855: 255f) Redescription mm long, dark brown with lighter brown antennae and light brown legs and mandibles. Compound eye relatively small (in males approximately 1.5 wider than genae, with longest diameter distinctly less than twice the distance from upper rim to vertex, in female somewhat larger) and moderately protruding. Antennal sockets more or less protruding between eyes, face below them with faint medial vertical carina, and broad, distinct subantennal striae (Fig. 2). Vertex somewhat conical. Scattered hairs on head short around ocelli, compound eyes and antennal sockets, longer on the upper rim of compound eye, on occiput and on mandibular bases. Female antenna shorter than body, club formed by eight to nine rounded flagellomeres with placoid sensillae and denser setae; third flagellomere (fifth antennomere) forming transitory article, as long as club articles (longer than the preceding one) but not or only slightly rounded, with a few placoid sensillae and varying density of setae. Flagellomeres subequal in length, except for first flagellomere (third antennomere) more or less distinctly shorter than others (Fig. 5). Male antenna longer than body, with a moderately modified second flagellomere (fourth antennomere) inner edge weakly bicurved, outer edge distinctly but not strongly outward-curved, making the article asymmetrical and apically enlarged not or very slightly longer than the neighbouring articles (Fig. 2). Fig. 5. Quasimodoana decipiens, female antenna.

8 308 M. Forshage et al. Pronotal plate wide, with narrow but laterally open submedian depressions, very weakly striate anterior flange, dorsal area with an irregular pattern of hair punctures, dorsal margin bilobed, not projecting above pronotal margin. Lateral parts of pronotal plate usually distinctly elevated above surface of pronotum, with distinct but very short traces of ridges discernible at junction. Pronotum uneven with scattered hairs, anterolateral parts with long, more or less dense hairs, ventral parts usually strongly wrinkled. Mesoscutum markedly humped. Scutellum rather large, produced posteriorly, roundedly rectangular in lateral view, with an irregular punctatereticulate sculpture. Lateral bars basally more or less weakly wrinkled but not striate, with enlarged ventral lobes. Scutellar plate of intermediate size, ovoid shape, with distinct margin and flat or slightly concave surface, posterior third with a large glandular release pit, other parts with an irregular pattern of several minor pits with a hair in each. Lateral foveae large, shallow, slightly oblique (Fig. 7). Metapleuron with well-developed narrow metapleural triangle, and well-developed subposterior carina. Ventral third of the metapleuron, anterior to the metapleural triangle, more or less entirely depressed. Median metapleural carina rather weak, not reaching anterior margin, angled in the posterior part that forms the dorsal limit of the subposterior depression. Posterior parts dorsal to that carina only weakly depressed. Metapleural groove usually indistinct. One small tuft of hairs in posteroventral corner (anterior to calyptral base) consisiting of only a few single hairs, and one slightly larger tuft in anteroventral concavity. Propodeal complex covered with long hairs and rough puncture, propodeal carinae broadly lyre-shaped. Legs yellow-brown or rarely chestnut-brown; femur, coxa and apical tarsal joints darker. Marginal cell of forewing closed (R1 entire but distally indistinct) with intermediate length (Rs often less than 1.5 longer than 2r). 2r evenly curved, Rs almost straight, distinctly curved only in apical end. Rs þ M and M nebulous and rather short, M þ Cu1, Cu1 and Cu1a not clearly visible. Female metasoma large, outstretched, pointed oval. Male metasoma remarkably small. Surface mostly diaphanic, with very few and weak punctures, slightly more distinct on posterior tergites. Type specimen. A female, glued to a triangle, with left antenna broken after sixth article, carrying a label Eucoila decipiens M. apparently by Fo rster, a red Typus label, and a lectotype label added by GN 1978 (with unique signature Em 385). As no lectotype designation was published at the time, the specimen lacked lectotype status. The specimen is the only decipiens in the Fo rster collection, but that may be insufficient reason to assume that there are no syntypes [in accordance with Recommendation 73F of the International Code of Zoological Nomenclature (ICZN, 1999)]. Thus, in order to prevent any ambiguity in the interpretation of the species should any such specimens surface, we designate a lectotype in conformity with Recommendation 74G (ICZN, 2003). Distribution. Europe. Specimens studied by the authors (listed below) from south and middle Sweden, from Germany (type locality) and from France. Also recorded from the Czech Republic (Mikula, 1989), but the identity of these specimens remains to be confirmed. Habitat. Open deciduous forest, often oak, cattle-grazed. Phenology. Specimens caught mostly in late summer (July August) but also in autumn (September October). Host. Unknown. Specimens studied. Lectotype, female, GERMANY, Aachen, before 1855 (Fo rster) (ZMHB). Additional material: FRANCE: male, Mertzwiller, 9.ix.1980 (Huggert) (coll. GN); SWEDEN: female Småland, Odensviholm, 23.vii.1947 (Jansson) (coll. GN); female, Gotska Sando n 7.vii.1946 (Jansson) (coll. GN); one female, one male, Gotska Sando n, Lilla lo vskogen, 8.vii.1946 (Jansson) (coll. GN); female, Gotska Sando n, Stora lo vskogen, 9.vii.1946 (Jansson) (coll. GN); three males, same locality, 12.vii.1946 (Jansson) (coll. GN); male, same locality, 17.vii (Jansson) (coll. GN); two males, same locality, 15.viii (Jansson) (coll. GN); female, Gotska Sando n without date (Jansson) (NHRS); three males, one female, Gotska Sando n 11.viii.1963 (Sundholm) (coll. GN); male, O stergo t- land, S:t Anna, Svensmaro, Sanningholmen, 7.viii.1976 (Wa ngsjo ) (NHRS); female, Na rke, Oset, 14.viii.1939 (Jansson) (coll. GN); female, Närke, probably O rebro, Sommarbo ( O r.somm. ), 3.vii.1943 (Jansson) (coll. GN); male, So dermanland Väsbyo n (or similar), 22.vii.1949 (Jansson) (LZM); male, So dermanland, Tockeno n, 20.viii (Jansson) (coll. GN); male, So dermanland, Sandemar deciduous forest, 18.viii.1976 (Nordlander) (coll. GN); male, So dermanland, Ga lo, Stegsholm, Olsholmen oak forest, 12.vii.1975 (Nordlander) (coll. GN); male, So dermanland, Ga lo, Stegsholm grazed mixed oak forest, 29.vii.1975 (Nordlander) (coll. GN); male Södermanland, Dalarömalmen, Sjo täppan oak grove, 3.x.1976 (Nordlander) (coll. GN); male, So dermanland, Ornö, Sundby, Näset damp mixed forest, 13.viii.1980 (Nordlander) (coll. GN); male, Södermanland, A va, Stavholmen mixed oak forest, 19.ix.1976 (Nordlander) (coll. GN); male, same locality, 2.x.1976 (Nordlander) (coll. GN); female, labelled Ent. Anst. Mjöberg (collected by Eric Mjo berg early 20th century; indicating either a collecting site near Entomological Institute, Uppland or merely the fact that the specimen was integrated into the collections of the Entomological Institute); male, Västmanland, Strömsholm mixed oak forest (Huggert) (coll. GN); male, Västmanland, Kärrbo, Solbacken (RN 1552/6600) Malaise trap, vii.1991 (Nilsson) (LZM). Quasimodoana gibba sp.n. Etymology. Gibba is the feminine form of gibbus (Latin) ¼ humped. Diagnostic characters. The first distinguishing character from Q. decipiens is that the fourth flagellomere in

9 Quasimodoana, new genus of eucoiline wasps 309 the female antenna is the transitory joint to the club, not the third as in Q. decipiens. Furthermore, Q. gibba is distinguished by its larger eye size (distance from ventral rim of compound eye to top of head less than one-half the longest diameter of eye), and the denser pubescence on the metapleuron, often obscuring other metapleural characters (see below). Often the scutellum is smaller with distinctly striate lateral bars (at most wrinkled in Q. decipiens) and a comparatively larger scutellar plate (surface of scutellum protruding beneath scutellar plate is less than one-half the width of the scutellar plate to the sides, and often less than one-half the length of the scutellar plate to the posterior), and the marginal cell of the forewing is often longer than in Q. decipiens. These additional characters are all rather variable, and it can be difficult safely to assign some male specimens to species without taking the geographical origin into account. Description mm, chestnut brown to dark brown (Figs 9, 10). Compound eye large, particularly in females, in both sexes length of eye (longest diameter) distinctly more than twice the distance between upper rim and top of vertex. Antennal sockets more or less protruding between eyes, face below them with faint medial vertical carina, and broad, distinct subantennal striae, like in Q. decipiens. Vertex very slightly conical. Scattered hairs on head like in Q. decipiens. Female antenna shorter than body, club formed by (seven-)eight rounded flagellomeres with placoid sensillae and denser setae; fourth flagellomere (sixth antennomere) is a clear transitory segment, rounded like the true club articles but longer, with placoid sensilla only in distal three-quarters or so. Flagellomeres subequal in length, first often somewhat shorter, fourth and 11th (first and last club article) somewhat longer than others (Fig. 6). Male antenna longer than body with flagellomeres subequal in length, second flagellomere (fourth antennomere) moderately modified, distinctly curved especially along outer edge, longer than other articles. Pronotal plate wide with narrow, laterally open submedian depressions. Anterior flange with distinct striae, dorsal area with an irregular pattern of hair punctures, dorsal margin bilobed, not projecting above pronotal margin Fig. 7. Quasimodoana decipiens, scutellum. (Fig. 3). Lateral parts of pronotal plate weakly elevated above surface of pronotum, any ridges very indistinct. Pronotum usually uneven with scattered hairs, anterolateral parts more or less hairy, ventral parts usually strongly wrinkled. Mesoscutum more or less strongly humped. Scutellum usually smaller and narrower than in Q. decipiens, roundedly rectangular in lateral view, not protruding very far posteriorly, with an irregular punctate-reticulate sculpture. Lateral bars smooth dorsally, weakly striate laterally. Scutellar plate large, covering most of scutellar surface (dorsal area exposed beneath it in dorsal view usually less than one-half the width of the plate on the sides and up to one-half the length of the plate to the posterior) (Fig. 8). Metapleuron with well-developed narrow metapleural triangle. Subposterior carina less well developed than in Q. decipiens. The whole posterior end of metapleuron heavily depressed, making less contrast between ventral and dorsal parts than in Q. decipiens. Median metapleural carina indistinct, but metapleural groove very distinct. Hairs more abundant than in Q. decipiens, covering much of ventral parts and scattered along posterior parts. Fig. 6. Quasimodoana gibba sp.n., female antenna.

10 310 M. Forshage et al. Fig. 10. Quasimodoana gibba sp.n., habitus, male. Fig. 8. Quasimodoana gibba sp.n., scutellum. Propodeal complex covered with long hairs and rough puncture, propodeal carinae lyre-shaped, usually narrower than in Q. decipiens. Legs yellow-brown or chestnut-brown; femur, coxa and apical tarsal joints slightly darker. Marginal cell of forewing closed (R1 entire but distally indistinct) and rather long (Rs often more than 1.5 longer than 2r). 2r rather evenly curved, Rs almost straight, distinctly curved only in apical end. Rs þ M and M nebulous and rather short, M þ Cu1, Cu1 and Cu1a not clearly visible. Female metasoma large, outstretched, pointedly oval. Male metasoma remarkably small. Surface mostly diaphanic, with very few and weak punctures, slightly more distinct on posterior tergites. Type specimens. Holotype, female, U.S.A., Maryland, Bowie 9.v.1945, DDT Expedition (USNM) pinpoint-glued on left side, upward-bent head but otherwise in good condition (Fig. 9); Paratypes, U.S.A.: female identical data as holotype (USNM); male (damaged, lacking antennae and Fig. 9. Quasimodoana gibba sp.n., habitus, female. most of legs) identical data as holotype (USNM); male, North Carolina, Mt Pisgah, ft., 24.vii.1957 (Townes & Townes) (AEI); male, Washington, Ashford, 10.vii.1940 (Townes & Townes) (AEI); male (damaged, lacking head and fore legs), Tennessee, Gatlinburg 3Gst4/ 4c//3G Oak-hickory//GMNSF 1500, NE/Sweeps (Whittaker) (USNM). Distribution. North America (see above). Additional Nearctic specimens of Quasimodoana have been studied by the authors. These specimens probably represent a few new species, but the scarcity of specimens and the difficulties in associating the sexes mean that we prefer to refrain from describing them. U.S.A.: female, Georgia, Warwoman Creek, 26.vii.1957 (Richards) (CNC); male, South Carolina, Greenville, 26.vi.1955 (L. & G. Townes) (AEI); CANADA: male, Quebec, Cap Rouge, 4.vii.1957 (Peck) (CNC); female, Quebec, Gatineau Park, 23.viii.1981 (Masner) (CNC); male, British Columbia, Lake Erroch near Deroche, 2.vi.1953 (Mason) (CNC). Results All analyses under equal weights ( unweighted ) resulted in two shortest trees with identical topologies (consensus: Fig. 11). The ingroup was not retained as monophyletic; instead, Chrestosema becomes the sistergroup of Eucoilini. Eucoilini and Trichoplastini are monophyletic, as are the genera Quasimodoana, Trybliographa and Leptopilina. The only conflict in the tree is within Trichoplastini, but no tree retains a monophyletic Trichoplasta. Of these analyses, treating the characters as unordered gave a tree-length of 236 steps, a consistency index (CI) of 0.42, and a retention index (RI) of Ordering the characters a priori (logical ordering) resulted in a tree-length of 246 steps, a CI of 0.44 and a RI of Re-ordering the characters a posteriori (phylogenetic ordering) as expected retained the shortest possible tree-length, 236 steps, but with higher consistency

11 Quasimodoana, new genus of eucoiline wasps 311 and retention, namely CI ¼ 0.46, RI ¼ Support values differed somewhat between these trees, for most groups identical or slightly higher with phylogenetically ordered characters, for Leptopilina substantially higher (0.79 unordered, 0.95 reordered), whereas Trichoplastini decreased somewhat in support. Analyses under implied weights (with phylogenetically ordered characters) all resulted in one single shortest trees, each with an identical topology for most concavity functions tried (k ¼ 2 6) (Fig. 12), but with a minor difference for k ¼ 1(Eucoila crassinerva grouping with Trybliographa longicornis, rendering Trybliographa non-monophyletic). Compared with the topologies from analyses under equal weights, the ingroup is retained as monophyletic, and one of the two alternative topologies inside Trichoplastini is favoured. Except for the fact that Trybliographa is not monophyletic under k ¼ 1, the topology is identical between different concavity functions. These trees had CI ¼ 0.46 and RI ¼ 0.63, and their parsimony score ranged from (k ¼ 1) to (k ¼ 6). Support values differed between trees, usually in an apparently random way, but for two groups of interest (Leptopilina and Eucoila þ Q decipiens Q gibba T rapae T cubitalis T longicornis E crassinerva L longipes L heterotoma L boulardi N grunini T(A) sp T(T) bouceki Q decipiens Q gibba T rapae T cubitalis T longicornis R heptoma Chrestosema Glauraspidia Fig. 12. Single most parsimonous tree of analyses under implied weights (characters phylogenetically ordered) for k ¼ 2 6. Thick branches have bootstrap support values over 0.90 in all analyses and over 0.95 in some. E crassinerva L longipes L heterotoma L boulardi Chrestosema N grunini T(A) sp T(T) bouceki R heptoma Glauraspidia Fig. 11. Consensus tree of the two most parsimonous trees of analyses under equal weights. Thick branches have bootstrap support values over 0.95 in all analyses (characters unordered, logically ordered, phylogenetically ordered). Trybliographa þ Quasimodoana) support had smooth peaks at intermediate k values. When the dataset was split into two partitions, one with the 58 two-state characters, and the other with the 25 multistate characters, support values diminished for some groups, and some differences in topology were produced, but almost all groups of interest were still present for both datasets. The multistate character dataset was analysed both as logically ordered and as phylogenetically ordered, with the latter producing a clearly shorter tree (96 steps vs. 107) with one remarkable difference, namely that Quasimodoana grouped with Leptopilina rather than with Trybliographa þ Eucoila in the logical-order tree (Figs 14, 15). The tree with only two-state characters did not retain a monophyletic ingroup, Chrestosema grouping with Leptopilina (Fig. 13). In all analyses, Quasimodoana come out as monophyletic and (in all analyses of the entire dataset) well supported (bootstrap ). A non-monophyletic Quasimodoana required eight extra steps, and a Quasimodoana within Trichoplastini required ten extra steps. Quasimodoana has four unique synapomorphies, namely the distinct subantennal striae, the shortened first flagellomere, and two concerning the metapleuron; and four uniting it with

12 312 M. Forshage et al. Q decipiens Q decipiens Q gibba Q gibba T rapae L longipes T cubitalis L heterotoma T longicornis L boulardi E crassinerva T rapae L longipes T longicornis L heterotoma E crassinerva L boulardi Chrestosema T cubitalis N grunini N grunini T(A) sp T(T) bouceki R heptoma Glauraspidia Fig. 13. Single most parsimonous tree of analysis with two-state character partition, equal weights. T(A) sp T(T) bouceki R heptoma Chrestosema Glauraspidia Fig. 14. Single most parsimonous tree of analysis with multistate character partition, logically ordered, equal weights. Trybliographa þ Eucoila; four other characters are shared with parts or the whole of Leptopilina or Trichoplastini and result as homoplasies in this analysis (Table 3). Trybliographa is monophyletic in many trees but always with low support (often collapsed into a polytomy with Eucoila in bootstrap trees), but Eucoila þ Trybliographa is retained as monophyletic and well supported in all trees (1.0 in all analyses of the entire dataset); and all analyses of the entire dataset have Quasimodoana as the sister of Eucoila þ Trybliographa (mostly intermediate support around 0.85). The internal topology in this group varies, with different attachments of the root in different analyses. Leptopilina is monophyletic in all trees, but with varying support (bootstrap ). Inside Leptopilina, L. heterotoma and L. boulardi constantly group together with good support. In all trees except the two-state-only tree, Eucoilini comes out as monophyletic, but with very little support (usually around 0.6, but even lower with unordered-only or logically ordered characters), being dependent on a small number of characters (only three morphological synapomorphies, all in the same body part: characters 52, 53 and 56). A tree with Trichoplastini within Eucoilini (as a sistergroup of Quasimodoana þ Trybliographa þ Eucoila), thus rendering Eucoilini paraphyletic as in Buffington s analysis (2007), required only four extra steps. Trichoplastini is well supported as monophyletic in all trees (bootstrap ), but its internal topology varies and is never well supported; however, Trichoplasta (Trichoplasta s.str. þ Armigerina) is never retained as monophyletic without Nordlanderiana. Discussion The support found for the monophyly of the new genus seems sufficient, as does that for its inclusion in the Eucoilini. The support for the monophyly of Eucoilini remains weak. Possibly Trichoplastini is nested within Eucoilini, as suggested by other analyses, for example Buffington et al. (2007). In that analysis, too, morphology favoured inclusion of Leptopilina in a monophyletic group with Eucoila and Trybliographa, but the 28S molecular data strongly favoured Leptopilina as paraphyletic vis-a-vis the Trichoplastini genera, and made the combined data analysis favour that topology. Members of the ingroup (Eucoilini þ Trichoplastini) share characters, the most important of which may be the modified second flagellomere in males

13 Quasimodoana, new genus of eucoiline wasps 313 Q decipiens Q gibba T rapae T longicornis E crassinerva T cubitalis L longipes L heterotoma L boulardi N grunini T(A) sp T(T) bouceki R heptoma Chrestosema Glauraspidia Fig. 15. Single most parsimonous tree of analysis with multistate character partition, phylogenetically ordered, equal weights. (character 16 in this matrix), which is an almost unique, unreversed synapomorphy of great diagnostic importance: all other eucoilines except Cothonaspis have the first male flagellomere modified, not the second. However, there is much homoplasy between genera of higher eucoilines (as documented in Fontal-Cazalla et al., 2002), so relationships at that level can be convincingly resolved probably only in a comprehensive, combined analysis of morphological and preferably multigene molecular data, and including other taxa. Optimizing biological traits on the tree makes it probable that Quasimodoana are parasitoids of some calyptrate Diptera larvae living in decaying or fresh plant or mushroom matter (as in most species of Trybliographa, Eucoila and Leptopilina with known life histories). The usual pattern in Eucoilinae is one wasp genus for one host fly family, and thus the host choices of the relatives of Quasimodoana (Drosophilidae in Lepotopilina, Anthomyiidae in Trybliographa, but several families in Eucoila) provide no ground for speculation. The feeding habits of these flies (herbivorous, mycophagous or saprophagous, including coprophagous and necrophagous) are diverse enough, and the host records few enough, to provide little ground for speculation on where the host shift occurred, whether in mushrooms (many Trybliographa, some Leptopilina), animal dung (Eucoila, some Trybliographa), carcasses (Eucoila), roots or vegetables (some Trybliographa), fruit (some Leptopilina), leaves of herbs and ferns (some Trybliographa), conifer cones (some Trybliographa) or some other microhabitat. Thus, in this analysis, further support has been provided for the monophyly of the Trichoplastini Kovalev, 1989, although the situation is still uncertain for the Eucoilini Thomson, 1862, despite the fact that our analysis is consistent with the hypothesis of monophyly. Morphological data at this point argue unequivocally for the inclusion of Leptopilina into Eucoilini, but further research may necessitate a reclassification of Eucoilini that either excludes Leptopilina or includes the whole Trichoplastini. Some critical issues regarding the phylogeny of this group are touched upon but not solved by this analysis. Is Trybliographa actually a monophyletic group without Eucoila s.str. (Nordlander, 1981)? Trybliographa is a monophyletic sistergroup to Eucoila in most parsimonous trees, but bootstrap trees collapsed Trybliographa þ Eucoila into Table 4. Some synapomorphies of Quasimodoana. Character no. State Description Shared with 15 2 Subantennal striae distinct Unique 20 2 First flagellomere of female antenna shorter than other flagellomeres Unique 56 2 Lower metapleural carina pointing high, almost vertically Unique 57 2 Surface of metapleuron anterior to posteroventral corner strongly depressed Unique 7 1 Hair punctures on lateral part of vertex distinct Trybliographa, Eucoila, Nordlanderiana 23 0 Erect seta medially on apical segment of maxillary palp present Trybliographa, Eucoila 25 1 First segment of labial palp longer than apical segment Trybliographa, Eucoila 67 1 Dorseroventral hairpatch of metacoxa present and small Trybliographa, Eucoila 6 1 Compound eye large, twice as long as gena Leptopilina, Trichoplasta bouceki 28 1 Submedian pronotal depressions intermedially deep, with ventral edge distinct but dorsal edge indistinct Leptopilina boulardi (Trichoplastini have entirely shallow depressions) 33 1 Mesoscutum humped Leptopilina 44 2 Posterior part of scutellum in lateral view distinctly produced posteriorly Trichoplastini