NRM - Forshage et al systent 2008Correspondence: Mattias Forshage, Department of Systematic Zoology, Evolutionary Biology Centre, Norbyva¨gen 18D, SE-75236 Uppsala, Sweden. E-mail:

Systematic Entomology (2008), 33, 301–318

Quasimodoana, a new Holarctic genus of eucoilinewasps (Hymenoptera, Cynipoidea, Figitidae), witha phylogenetic analysis of related genera

MATT I A S FOR SHAGE 1 , G O RAN NORDLANDER 2 andFREDR IK RONQU I S T 3

1Department of Systematic Zoology, Evolutionary Biology Centre, Uppsala, Sweden, 2Department of Ecology, SwedishUniversity of Agricultural Sciences, Uppsala, Sweden and 3Department of Entomology, Swedish Museum of NaturalHistory, Stockholm, Sweden

Abstract. A new Holarctic genus, Quasimodoana gen.n. is described for thePalearctic eucoiline wasp Eucoila decipiens Forster, 1869, as Quasimodoanadecipiens comb.n.A lectotype is designated for this species. A new North Americanspecies is described here as Quasimodoana gibba sp.n. As the new genus hassimilarities with several eucoiline genera, belonging to two related but distinctlineages, a phylogenetic analysis was carried out, based on 83 morphologicalcharacters. The two lineages included in the analysis are referred to here under thetribal names of Eucoilini and Trichoplastini stat.nov. We discuss reasons foradopting available tribal names (rather than ‘informal genus groups’), andTrichoplastini is removed from synonymy with Eucoilini. The phylogeneticanalysis places Quasimodoana unambiguously within Eucoilini, as a sistergroupto the Trybliographa complex. However, there is only weak support for themonophyly of Eucoilini and for the hypothesized sistergroup relationship betweenEucoilini and Trichoplastini.

Introduction

Eucoilinae is a group of Diptera parasitic Cynipoidea,which has historically been treated as a distinct family butis now considered a subfamily of Figitidae (Ronquist, 1999).This poorly known group contains slightly less than 1000nominal species (Vardal, 2004), but the identities of manyremain doubtful. Indeed, most Eucoilinae remain unde-scribed, even within the Holarctic region.With Eucoila Westwood, 1833 being the oldest genus in

Eucoilinae, a number of species were originally described asbeing within that genus, but were then transferred to othergenera of younger date, or simply forgotten, thus remainingwithin Eucoila as nomina dubia. Often taxa do not belong inEucoila, but correct placement is denied by the loss of typesand inadequate descriptions. Probably many nomina dubiarefer to species of Trybliographa Forster, 1869, as this is the

principal Holarctic genus of relatively large-sized eucoilines.However, some type specimens do exist, and it is only a taskfor the systematist to establish new combinations or de-scribe 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 ‘nudipennisgroup’ were separated first by Nordlander (1981). Thenudipennis group was confirmed later to be distant to thetrue Eucoila, belonging to the ‘neotropical grade’ and not tothe ‘higher eucoilines’ in the analysis of Fontal-Cazalla et al.(2002). The species group has been studied recently bySchick (1999), but the new genus name proposed in herunpublished 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 Forster, 1855 from Europe, studied by GN, is

Correspondence: Mattias Forshage, Department of SystematicZoology, Evolutionary Biology Centre, Norbyvagen 18D,SE-75236 Uppsala, Sweden. E-mail: [email protected]

# 2008 The AuthorsJournal compilation # 2008 The Royal Entomological Society 301

not assignable to any valid genus and actually representsa 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; althoughother specimens examined probably represent additionalnew species, we refrain from describing these here because ofthe scarcity of specimens and the difficulties in associatingthe sexes.A phylogenetic analysis was carried out in order to

investigate the phylogenetic position and monophyly ofthe new genus. Because Quasimodoana shares importantcharacters both with Trybliographa and Leptopilina ofEucoilini and with Trichoplasta and Rhoptromeris of Tri-choplastini, it is not immediately obvious to which of theselineages it belongs. The Eucoilini and Trichoplastini ap-peared (as the ‘Trybliographa group’ and the ‘Rhoptromerisgroup’) as sistergroups in Fontal-Cazalla et al.’s (2002)analysis of higher-level eucoiline phylogeny, but supportfor this was not strong. Subsequently, in the analysis ofBuffington et al. (2007) they grouped, but with Eucoilinifound to be paraphyletic. In the phylogenetic analysis of theposition ofQuasimodoana we focused our attention on thesetwo lineages, using a few other higher eucoiline representa-tives as outgroups, primarily because of the mosaic ofEucoilini and Trichoplastini characters in Quasimodoanaand because we hoped that such an analysis might help toelucidate relationships.

Tribal classification

Nordlander made a tentative subdivision of Eucoilinae intoinformal genus groups (Nordlander, 1982b), and mostworkers have retained these informal group names. Thishas perhaps primarily been motivated by a relative lack ofknowledge of the phylogeny of the group. The few authorsutilizing family-level names within Eucoilinae usuallyapplied the names to groupings that were not founded incareful hypotheses of phylogeny, and most available nameswere synonymized into Eucoilinae by Ronquist (1999).Taking the analyses of Fontal-Cazalla et al. (2002) andBuffington et al. (2007) into account, it would seemconvenient to use available tribal names for those estab-lished genus groups having gained some support, in order tofacilitate the discussion of phylogenetical relationships.Informal groups make sense on an informal level, or withinthe context of a single investigation. When informal groupsbecome ‘established’ they become similar to formal classi-ficatory units and may even be mistaken for such, butwithout being subject to formal classificatory rules norbeing available to formal classificatory actions. Thus, theprimary purpose of erecting tribes here is to make conven-tional groups available formally as taxonomic units, andto encourage testing.The name Eucoilini is available with the same authorship

as Eucoilinae, Thomson, 1862. The core of Eucoilini, asdelineated here, consists of Eucoila (type genus), Trybliog-

rapha, and Bothrochacis Cameron, 1904, which are thetraditional genera in the ‘Trybliographa group’ (Nord-lander, 1982a, b) [note that Trybliographa after the revisionof Nordlander (1981) includes Psichacra Forster, 1869,Pseudeucoila Ashmead, 1903, Piezobria Forster, 1869, andPilinothrix Forster, 1869]. Fontal-Cazalla et al. (1982)showed that Leptopilina belongs to the same lineage.Furthermore, Linaspis Lin, 1988 and Linoeucoila Lin, 1988are included tentatively based on the original descriptions(Lin, 1988), although we cannot confirm this by examina-tion of the type specimens. We refer Maacynips Yoshimoto,1963 also to this group (Table 1). Morphologically, thesetaxa are recognized by the males having the second flag-ellomere modified (shared with Trichoplastini) and byhaving the posteroventral corner of the metapleuron raised,facing posterolaterally, entirely or almost glabrous. Thislatter structure has lacked a name, forcing authors to uselong descriptions such as ‘surface immediately posterior to(posteroventral corner of metapleuron) (probably a part ofpropodeum) is hairless (. . .), facing latero-posteriorly or,more rarely, directly posteriorly’ (Nordlander, 1981) and‘lateral part of metacoxal rim forming a smooth and nudetriangular projection having its surface directed morelaterally’ (Fontal-Cazalla et al., 2002). As the structure isoften triangular in shape we refer to it as the metapleuraltriangle.Trichoplastini Kovalev, 1989 hereby is removed from

synonymy with Eucoilidae Thomson, 1862, stat.n. It con-tains Trichoplasta (type genus) and Rhoptromeris from the‘Rhoptromeris group’ (note that Armigerina Belizin, 1968 isincluded in Trichoplasta). Daruna Benoit, 1956, StentorcepsQuinlan, 1984, and Angustacorpa Quinlan, 1988 wereincluded in this group by Quinlan (1984, 1986, 1988), andNordlanderiana was included by Kovalev (1989). Morpho-logically, these taxa are easily recognizable by having lateralbridges on the pronotal plate (closed lateral pits), andsharing the state of a modified second male flagellomerewith Eucoilini (Table 2).

Materials and methods

Morphological terminology follows Nordlander (1982b),Ronquist & Nordlander (1989), Ronquist (1995), andFontal-Cazalla et al. (2002).Descriptions are based on observations utilizing a Leica

MZ16 stereo microscope and a Leo/Zeiss Supra 35VPscanning electron microscope (SEM).One drawing was made by hand, based on stereo micro-

scope observations, but the rest were carried out in AdobePhotoshop 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 usualprocess of goldcoating and high vacuum. All the images ofQ. gibba (and one image of Q. decipiens to facilitate direct

302 M. Forshage et al.

# 2008 The AuthorsJournal compilation # 2008 The Royal Entomological Society, Systematic Entomology, 33, 301–318

Table

1.Generaincluded

inEucoilini.Thesourcegivenisforinclusionin

thistribe(orin

itsequivalentthe‘Trybliograp

hagroup’).Note

that

thenumber

ofnominal

speciesisoften

substan

tially

smallerthan

thereal

number.Someofthesegeneraarepoorlystudied,an

dtheirrobustnessas

distinct

generashould

notbeconsidered

asconfirm

edbythepresentau

thors

simply

bytheirinclusionin

thistable.

Genus

No.of

species

Hosts

Hab

itats

Geograp

hical

distribution

Source

Trybliograp

haForster,18

6915

1Anthomyiidae

Mushrooms,rootcrops,leav

esofherbs,

ferns,dung,

conifer

cones,etc.

Widespread

butpredominan

tly

Holarctic

Nordlander

(198

2a,b);

Fontal-Cazalla

etal.(200

2)Eucoila

Westw

ood,18

3310

Muscidae,Calliphoridae,

Sarcophag

idae

Dungan

dcarrion

Widespread

butpredominan

tly

Palearctic

Nordlander

(198

2a,b);

Fontal-Cazalla

etal.(200

2)Leptopilina

Forster,18

6927

Drosophilidae

Decay

ingfruit,mushrooms,an

ddecayingdebrisin

general

Cosm

opolitan

Fontal-Cazalla

etal.(200

2)

Qua

simodoa

nagen.n.

2?

?Holarctic

Presentconsideration

Bothrocha

cisCam

eron,19

044

??

Afrotropical

Nordlander

(198

2a,b)

LinaspisLin,19

881

??

Taiwan

only

considered

close

toLeptopilina

inLin

(198

8)Linoeucoila

Lin,19

8813

??

Taiwan

only

Lin

(198

8)Maa

cynips

Yoshim

oto,19

633

??

Australopacific

region

Presentconsideration

Table

2.Generaincluded

inTrichoplastini.Thesourcegivenisforinclusionin

thistribe(orin

itsequivalentthe‘Rho

ptromerisgroup’).Note

that

thenumber

ofnominal

speciesisoften

substan

tially

smallerthan

thereal

number.Someofthesegeneraarepoorlystudied,an

dtheirrobustnessas

distinct

generashould

notbeconsidered

asconfirm

edbythepresentau

thors

simply

bytheirinclusionin

thistable.

Genus

No.of

species

Hosts

Hab

itats

Geograp

hical

distribution

Source

Rho

ptromerisForster,18

6940

Chloropidae

Grasses,conifer

cones,

bracket

fungi,etc.

Widespread

butpredominan

tly

Palearctican

dAfrotropical

Nordlander

(198

2a,b);

Kova

lev(198

9);Fontal-Cazalla

etal.(200

2)Trichop

lastaBenoit,19

5627

Lonchaeidae

andothers?

Decay

ingwood

Widespread

butpredominan

tly

Holarctican

dAfrotropical

Nordlander

(198

2a);Kova

lev(198

9)

Nordlan

derian

aKova

lev,

1989

1Lonchaeidae?

Decay

ingwood

Palearctic

Kova

lev(198

9)DarunaBenoit,19

561

??

Afrotropical

Tentatively

suggestedin

Quinlan(198

6)Ang

ustacorpaQuinlan,19

884

??

Afrotropical

Quinlan(198

8)Stentorceps

Quinlan,19

841

??

Afrotropical

Quinlan(198

4,19

86)

Quasimodoana, new genus of eucoiline wasps 303


comparison) were captured in low vacuum with uncoatedspecimens because of the rarity of specimens, which unfor-tunately resulted in poorer image quality.Character coding for the phylogenetic analyses was to

a large extent based on SEM and compound microscopyimages available from MorphBank (www.morphbank.net),but additional taxa and some of the additional characterswere coded from specimens, utilizing SEM and stereomicroscopy.The phylogenetic analysis was performed with PAUP

4.0B10 (Swofford, 2002), with parsimony as the optimalitycriterion, with equal as well as implied weights (Goloboff,1993), the latter with a series of concavity functions (Goloboffweights k ¼ 1–6), various ordering schemes, full Branch-and-Bound search, plus bootstrapping (100 replicates, 25random addition sequences). The resulting trees were furtherinvestigated 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; collGN, G. Nordlander’s collection at the Swedish Universityof Agricultural Sciences, Uppsala, Sweden; LZM, Zoolog-iska Museet, Lund, Sweden; NHRS, Naturhistoriska Riks-museet, Stockholm, Sweden; USNM, U.S. NationalMuseum, Washington, U.S.A.; ZMHB, Museum fur Na-turkunde, Berlin, Germany.

Included characters

The analysis re-utilized (after scrupulous proofreadingand some recoding) relevant parts of the data matrix ofFontal-Cazalla et al. (2002), and added some taxa and anumber of characters (app. 1, 2). The new characters werecoded mostly in the same manner as the available matrix,from direct comparison between prints of SEM picturesof exemplar specimens in standard views [available fromMorphBank (www.morphbank.net)].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 thepresent matrix), and 33 are based on characters taken fromthe matrix in Fontal-Cazalla et al. (2002). From that sourceall characters that were informative in the groups of generastudied here were taken. Fourteen characters wereunchanged [4, 11, 28, 30, 36, 51, 53, 56, 62, 111, 114, 122,128, 135 of Fontal-Cazalla et al. (2002)], and 17 weremodified, either because it was possible to introduce grada-tions of the differences within this more narrowly circum-scribed group, or because new scrutiny of the states of singletaxa 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, 50–52 in thepresent matrix]. Some problematic characters [17, 116, 120of Fontal-Cazalla et al. (2002)] were removed, as wereseveral uninformative characters or states.

Multistate characters are either gradual characters, inwhich character states occur along a one-dimensional con-tinuum, or complex characters, which may be grouped intoa transformation series through careful interpretation ofcharacter evolution. Logically, the former would best betreated as ordered (Wagner parsimony) and the latter asunordered (Fitch parsimony). Different characters may ofcourse evolve in different ways, some in a leap-wise mannerwithout passing intermediate stages, but little is knownabout this, and ideally we should proceed in a mannerchosen from a methodological standpoint as sound regard-less of what might eventually be discovered. For charactersthat present an obvious grade of character states, to treatthem as unordered, implying that it would be equally easy togo from one extreme to another as to an intermediate, wouldin fact represent quite a strong assumption, and, not onlydoes that strategy fail to retain information on similaritypresent in the matrix, it is contrary to the general scientificprinciple of parsimony. However, a priori weightingschemes are not that important if we recognize that alltransformation series are actually tested by the tree itself(Mickevich, 1982; Mickevich & Lipscomb, 1991; Wilkinson,1992; Schulmeister, 2003). Checking the distribution ofcharacter states of a multistate character in the mostparsimonous trees (starting out from the one obtained withunordered characters) empirically shows whether the char-acter evolution necessarily passes through the intermediatestages or just as easily goes from one end to the other. Amethod of ordering that decides whether to treat eachindividual character as ordered or unordered based onhow it is distributed in the tree is here called phylogeneticordering 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 25multistate characters did not present an obvious order fromthe a priori perspective, and these were treated as unorderedeven in the ‘logically ordered’ analysis. Ten of the 25multistate characters showed an unordered distribution(including character 64 but not 75), and 15 charactersretained their logical order after analysis. To investigatethe effects of ordering on tree topology, the dataset was splitinto two-state characters (58 characters) and multistatecharacters (25 characters), which were analysed separately,the latter both as a priori logically ordered and as a poste-riori 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 conse-quences, concavity functions with k ¼ 1–6 were tried, aswell as equal weights.

Included taxa

The taxon set chosen for the analysis includes representa-tives of all the Holarctic genera of the Eucoilini andTrichoplastini, with more than one species from the larger



genera with which the new genus shares some similarities –namelyLeptopilina,Trybliographa andTrichoplasta (Table 3).The phylogeny among the ‘higher eucoilines’ is not wellresolved, and the best outgroup for this analysis is unclear.Several candidates were tried but performed poorly because ofmany outgroup autapomorphies, making polarity decisionsimpossible for many characters, and some specific similaritieswith ingroup taxa that appeared as potential synapomorphiesin the context of our analysis but that were probablyconvergences based on the more comprehensive analysis ofFontal-Cazalla et al. (2002) and other background knowledge.The taxa chosen as outgroups are Chrestosema Forster, 1869and Glauraspidia Thomson, 1862. Both were included in the‘Chrestosema group’ of Fontal-Cazalla et al. (2002), but thatgroup was supported only weakly, characterized by a set ofnon-unique synapomorphies, none shared by all genera in thegroup. The non-monophyly of the ‘Chrestosema group’ hasbeen confirmed since by molecular and combined analyses(Buffington et al., 2007). TheGlauraspidia species used here asan outgroup is actually a partly brachypterous and ratherautapomorphic species, but its odd features appear to be of noimportance within this analysis.

Descriptions

Quasimodoana gen.n.

Type species. Eucoila decipiens Forster, 1855 (Fig. 1)The new genus has an unusual combination of characters.

It has a metapleural triangle similar to that characteristic ofgenera of Eucoilini, whereas the general appearance, includ-ing the long and pointedly oval female metasoma and theposteriorly extended scutellum, is more similar to that ofTrichoplasta andNordlanderiana of Trichoplastini. Most rep-resentatives are relatively large (similar to Trybliographa),

have a distinctly humped mesoscutum (similar to Leptopi-lina) and a more or less strongly modified second flagello-mere of male antenna (similar to Leptopilina andTrichoplastini). They lack the foremost diagnostic charac-ters of all these genera (the closed lateral pits of the pronotalplate of Trichoplastini, the broad petiolar rim and oftenreduced hairy ring of syntergite of Leptopilina, and thesubalar pits of Trybliographa). All species we have seen arecharacterized (uniquely among their close relatives) by thedistinct subantennal striae and by the strong depressions inthe metapleura, and usually by the shortened first flagello-mere (third antennomere) of the female antenna, the latterfeature being emphasized by Forster (1855) in his descrip-tion 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 thesource 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 specimensdissected, not covering several standard views in SEM images; not dissected ¼ very rare species, no SEM images, only stereomicroscopy. NBfor 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 AvailableChrestosema erythropum Forster, 1869 Outgroup Type species of genus In FC AvailableQuasimodoana decipiens (Forster, 1869) Ingroup Type species of genus Not in FC Not exhaustively dissectedQuasimodoana gibba sp.n. Ingroup Tew species Not in FC Not dissectedEucoila crassinerva Westwood, 1833 Ingroup Type species of genus In FC AvailableTrybliographa longicornis (Hartig, 1840) Ingroup Typical of species group Not in FC DissectedTrybliographa cubitalis (Hartig, 1841) Ingroup Typical of species group Not in FC DissectedLeptopilina longipes (Hartig, 1841) Ingroup Type species of genus In FC AvailableLeptopilina heterotoma (Thomson, 1862) Ingroup Typical of species group Not in FC DissectedLeptopilina boulardi (Barbotin et al., 1979) Ingroup Typical of species group Not in FC DissectedRhoptromeris heptoma (Hartig, 1840) Ingroup Type species of genus In FC AvailableTrichoplasta (Armigerina) sp. Ingroup Typical of species group In FC AvailableTrichoplasta (s.str.) bouceki (P. Masner, 1960) Ingroup Typical of species group Not in FC Not dissectedNordlanderiana grunini (Belizin, 1968) Ingroup Type species of genus Not in FC Not exhaustively dissected

Fig. 1. Quasimodoana decipiens (Forster, 1869), male.



Etymology. The name is derived from Quasimodo inVictor Hugo’s Notre Dame de Paris, in reference to thehunchback appearance of these wasps. The gender isfeminine.

Description. Compactly built eucoiline wasps, withbrown or black body, length 1.8–2.5 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, evenlyrounded or more or less conical, with small protrudingocelli. Occiput with several hairs, without carina. Scatteredhairs on head mostly short.Mandible basally dark and hairy, apically light and

glabrous. Maxillary palpus four-segmented, terminal seg-ment enlarged towards apex, with an erect seta. Labialpalpus two-segmented.Female antenna with 13 articles, first flagellomere more

or less shortened, apical seven to nine flagellomeres forminga rather indistinct club. Male antenna with 15 articles,second flagellomere more or less modified (asymmetric,elongated, curved), with placoid sensillae on all flagello-meres.Mesosoma high, usually humped (anterior end of meso-

scutum abruptly rising). Entire mesosoma usually stronglysclerotised, black, with bulging sclerites and strong ridges.Pronotum with several hairs, without distinct ridges lateralto pronotal plate, but with strong wrinkles ventrally.Pronotal plate of varying shape, always with submediandepressions laterally open but narrow, the dorsal rims ofthese depressions weakly delineated (ventrolateral partsof dorsal area of pronotal plate sticking close to surface ofpronotum without a distinct edge) (Fig. 3). Mesoscutumwith four rows of hair punctures; two submedian onesrepresenting rudimentary notauli, and two lateral ones inposterior half.Scutellum large, posterior part in lateral view protruding

(over propodeum) in a subrectangular shape, in dorsal viewmore or less evenly rounded. Sculpture of scutellum punc-

tate-reticulate, asymmetrical and with punctures of differentsizes intermixed, but largely with finer punctures nearscutellar plate and increasingly rough towards the lateraledge, and longitudinally reticulate on lateral surfaces. Lateralbars rather short, indistinctly or rather weakly striate.Scutellar plate raised, oval-shaped, with a distinct marginand slightly concave disc with irregularly distributed hairs insmall pits, and a large posterior glandular release pit.Mesopleuron glabrous, with ridges along lower margin

and across lower third (precoxal carina and mesopleuralcarina), and a diagonal groove from middle of anteriormargin to middle of dorsal margin (by the forewinginsertion). Subalar groove well developed but withoutsubalar pit.Metapleuron with varying amount of hairs, with several

ridges extending forwards and upwards from posteriormargin, with depressed areas in between (Fig. 4). Antero-ventral cavity round. Posteroventral corner largely glabrous

Fig. 3. Quasimodoana gibba sp.n., pronotal plate.

Fig. 4. Quasimodoana decipiens, metapleuron, posterolateral view.Fig. 2. Quasimodoana decipiens, male, face and basal parts ofantennae.



and directed posterolaterally, with a concave surface, form-ing a well-developed metapleural triangle. Anterodorsallyfrom the metapleural triangle runs a more or less developedcarina (which is probably homologous with ‘ridge 3’ sensuNordlander (1980) in Leptopilina, but which here is moredorsally oriented than in other eucoilines), running almostparallel to the posterior edge, and separated from it by astrongly depressed area, dorsally adjoining the median meta-pleural carina or disappearing by the metapleural groove.Area anterior to this subposterior carina also depressed butless sharply delimited. Median metapleural carina andmetapleural groove varying. Subalar groove deep and wide,the calyptral carina delimiting it ventrally strong. Calyptradistinctly protruding. Propodeal complex, including smalleror larger parts of metapleura, with long hairs. Propodeal ridgesglabrous, lyre-shaped (with an angle between the divergentlower thirds and the slightly concavely rounded upper two-thirds), joined by nucha ventrally but separate dorsally, onlyweakly 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 andscattered hairs apically. Hind coxae with an additionaldistinct small dorsolateral tuft of hairs; middle coxae lackingsuch a tuft. Claws small and simple.Wings well developed, with short pubescence over entire

surface. Ciliation along fringe well developed (intermediatebetween the short pubescence of Trybliographa and the longof Leptopilina). Forewings large and rather long, apicallyrounded, with ordinary figitid nervature (lacking costal vein,with distinct R þ Sc and marginal cell forming a W-likeshape). Marginal cell closed, neither narrowed nor deepened(similar to Trybliographa, probably plesiomorphic), curva-ture of 2r and Rs varying, M þ Cu1, Cu1, Rs þ M and Mreduced, nebulous or indistinct, 1A and Cu1a entirelyindistinct. 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 to5 (females) or 3 to 4 (males)) varying, weak or rather densebut never felt-like, complete except for a very short inter-ruption dorsally, with very thin second row of long hairs.Female metasoma large, often strikingly long, usually withan outstretched, pointedly oval form. Hypopygium pro-truding, with a ventral row of setae. Ovipositor (terebra)long and strong, protruding. Male metasoma small, usuallymore or less quadrangular, suggesting diamond-shape,length approximately the same as mesosoma but height

distinctly less. (Note, however, that overall metasomalshapes in eucoilines may vary substantially between speci-mens, depending on the degree and pattern of telescoping ofthe posterior tergites.)

Distribution. Holarctic, with the single previouslydescribed species European, and one herein describedspecies (plus some undescribed species) Nearctic.

Biology. The single European species was collected inopen deciduous forest. The Nearctic specimens usually lacksuch information. Hosts are unknown.All specimens studied were collected between June and

October.

Quasimodoana decipiens (Forster, 1855) comb.n.

Originally described as Eucoila decipiens by Forster(1855: 255f)

Redescription. 2.0–2.5 mm long, dark brown with lighterbrown antennae and light brown legs and mandibles.Compound eye relatively small (in males approximately

1.5# wider than genae, with longest diameter distinctly lessthan twice the distance from upper rim to vertex, in femalesomewhat larger) and moderately protruding. Antennalsockets more or less protruding between eyes, face belowthem with faint medial vertical carina, and broad, distinctsubantennal striae (Fig. 2). Vertex somewhat conical. Scat-tered hairs on head short around ocelli, compound eyes andantennal 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 anddenser setae; third flagellomere (fifth antennomere) formingtransitory article, as long as club articles (longer than thepreceding one) but not or only slightly rounded, with a fewplacoid sensillae and varying density of setae. Flagellomeressubequal in length, except for first flagellomere (thirdantennomere) more or less distinctly shorter than others(Fig. 5). Male antenna longer than body, with a moderatelymodified second flagellomere (fourth antennomere) – inneredge weakly bicurved, outer edge distinctly but not stronglyoutward-curved, making the article asymmetrical and api-cally enlarged – not or very slightly longer than theneighbouring articles (Fig. 2).

Fig. 5. Quasimodoana decipiens, female antenna.



Pronotal plate wide, with narrow but laterally opensubmedian depressions, very weakly striate anterior flange,dorsal area with an irregular pattern of hair punctures, dorsalmargin bilobed, not projecting above pronotal margin.Lateral parts of pronotal plate usually distinctly elevatedabove surface of pronotum, with distinct but very short tracesof ridges discernible at junction. Pronotum uneven withscattered hairs, anterolateral parts with long, more or lessdense hairs, ventral parts usually strongly wrinkled.Mesoscutum markedly humped.Scutellum rather large, produced posteriorly, roundedly

rectangular in lateral view, with an irregular punctate-reticulate sculpture. Lateral bars basally more or less weaklywrinkled but not striate, with enlarged ventral lobes.Scutellar plate of intermediate size, ovoid shape, withdistinct margin and flat or slightly concave surface, poste-rior third with a large glandular release pit, other parts withan 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. Ventralthird of the metapleuron, anterior to the metapleural tri-angle, more or less entirely depressed. Median metapleuralcarina rather weak, not reaching anterior margin, angled inthe posterior part that forms the dorsal limit of thesubposterior depression. Posterior parts dorsal to thatcarina only weakly depressed. Metapleural groove usuallyindistinct. One small tuft of hairs in posteroventral corner(anterior to calyptral base) consisiting of only a few singlehairs, 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 MandM nebulousand rather short,M þ Cu1, Cu1 andCu1a not clearly visible.Female metasoma large, outstretched, pointed oval. Male

metasoma remarkably small. Surface mostly diaphanic,with very few and weak punctures, slightly more distincton posterior tergites.

Type specimen. A female, glued to a triangle, with leftantenna broken after sixth article, carrying a label ‘EucoiladecipiensM.’ apparently by Forster, a red ‘Typus’ label, anda lectotype label added by GN 1978 (with unique signatureEm 385). As no lectotype designation was published at thetime, the specimen lacked lectotype status. The specimen isthe only decipiens in the Forster collection, but that may beinsufficient reason to assume that there are no syntypes [inaccordance with Recommendation 73F of the InternationalCode of Zoological Nomenclature (ICZN, 1999)]. Thus, inorder to prevent any ambiguity in the interpretation of thespecies should any such specimens surface, we designatea lectotype in conformity with Recommendation 74G(ICZN, 2003).

Distribution. Europe. Specimens studied by the authors(listed below) from south and middle Sweden, from Ger-many (type locality) and from France. Also recorded fromthe Czech Republic (Mikula, 1989), but the identity of thesespecimens 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, Aa-chen, before 1855 (Forster) (ZMHB).Additional material: FRANCE: male, Mertzwiller,

9.ix.1980 (Huggert) (coll. GN); SWEDEN: female Smaland,Odensviholm, 23.vii.1947 (Jansson) (coll. GN); female, GotskaSandon 7.vii.1946 (Jansson) (coll. GN); one female, onemale, Gotska Sandon, Lilla lovskogen, 8.vii.1946 (Jansson)(coll. GN); female, Gotska Sandon, Stora lovskogen,9.vii.1946 (Jansson) (coll. GN); three males, same locality,12.vii.1946 (Jansson) (coll. GN); male, same locality, 17.vii.1946 (Jansson) (coll. GN); two males, same locality, 15.viii.1946 (Jansson) (coll. GN); female, Gotska Sandon withoutdate (Jansson) (NHRS); three males, one female, GotskaSandon 11.viii.1963 (Sundholm) (coll. GN); male, Ostergot-land, S:t Anna, Svensmaro, Sanningholmen, 7.viii.1976(Wangsjo) (NHRS); female, Narke, Oset, 14.viii.1939(Jansson) (coll. GN); female, Narke, probably Orebro,Sommarbo (’Or.Somm.’), 3.vii.1943 (Jansson) (coll. GN);male, Sodermanland Vasbyon (or similar), 22.vii.1949(Jansson) (LZM); male, Sodermanland, Tockenon, 20.viii.1952 (Jansson) (coll. GN); male, Sodermanland, Sandemardeciduous forest, 18.viii.1976 (Nordlander) (coll. GN); male,Sodermanland, Galo, Stegsholm, Olsholmen oak forest,12.vii.1975 (Nordlander) (coll. GN); male, Sodermanland,Galo, Stegsholm grazed mixed oak forest, 29.vii.1975(Nordlander) (coll. GN); male Sodermanland, Dalaromalmen,Sjotappan oak grove, 3.x.1976 (Nordlander) (coll. GN); male,Sodermanland, Orno, Sundby, Naset damp mixed forest,13.viii.1980 (Nordlander) (coll. GN); male, Sodermanland,Ava, Stavholmen mixed oak forest, 19.ix.1976 (Nordlander)(coll. GN); male, same locality, 2.x.1976 (Nordlander) (coll.GN); female, labelled ‘Ent. Anst. Mjoberg’ (collected by EricMjoberg early 20th century; indicating either a collecting sitenear Entomological Institute, Uppland or merely the fact thatthe specimen was integrated into the collections of theEntomological Institute); male, Vastmanland, Stromsholmmixed oak forest (Huggert) (coll. GN); male, Vastmanland,Karrbo, Solbacken (RN 1552/6600) Malaise trap, 15–20.vii.1991 (Nilsson) (LZM).

Quasimodoana gibba sp.n.

Etymology. Gibba is the feminine form of gibbus (Latin) ¼‘humped’.

Diagnostic characters. The first distinguishing charac-ter from Q. decipiens is that the fourth flagellomere in



the female antenna is the transitory joint to the club,not the third as in Q. decipiens. Furthermore, Q. gibba isdistinguished by its larger eye size (distance from ventralrim of compound eye to top of head less than one-halfthe longest diameter of eye), and the denser pubescenceon the metapleuron, often obscuring other metapleuralcharacters (see below). Often the scutellum is smallerwith distinctly striate lateral bars (at most wrinkled inQ. decipiens) and a comparatively larger scutellar plate(surface of scutellum protruding beneath scutellar plateis less than one-half the width of the scutellar plate tothe sides, and often less than one-half the length of thescutellar plate to the posterior), and the marginal cell ofthe forewing is often longer than in Q. decipiens. Theseadditional characters are all rather variable, and it canbe difficult safely to assign some male specimens tospecies without taking the geographical origin intoaccount.

Description. 2.0–2.5 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 thantwice 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, andbroad, distinct subantennal striae, like in Q. decipiens.Vertex very slightly conical. Scattered hairs on head like inQ. decipiens.Female antenna shorter than body, club formed by

(seven-)eight rounded flagellomeres with placoid sensillaeand denser setae; fourth flagellomere (sixth antennomere) isa clear transitory segment, rounded like the true club articlesbut longer, with placoid sensilla only in distal three-quartersor so. Flagellomeres subequal in length, first often some-what shorter, fourth and 11th (first and last club article)somewhat longer than others (Fig. 6). Male antenna longerthan body with flagellomeres subequal in length, secondflagellomere (fourth antennomere) moderately modified,distinctly curved especially along outer edge, longer thanother articles.Pronotal plate wide with narrow, laterally open subme-

dian depressions. Anterior flange with distinct striae, dorsalarea with an irregular pattern of hair punctures, dorsalmargin bilobed, not projecting above pronotal margin

(Fig. 3). Lateral parts of pronotal plate weakly elevatedabove surface of pronotum, any ridges very indistinct.Pronotum usually uneven with scattered hairs, anterolateralparts more or less hairy, ventral parts usually stronglywrinkled.Mesoscutum more or less strongly humped.Scutellumusually smaller and narrower than inQ. decipiens,

roundedly rectangular in lateral view, not protrudingvery far posteriorly, with an irregular punctate-reticulatesculpture. Lateral bars smooth dorsally, weakly striatelaterally. Scutellar plate large, covering most of scutellarsurface (dorsal area exposed beneath it in dorsal viewusually less than one-half the width of the plate on thesides and up to one-half the length of the plate to theposterior) (Fig. 8).Metapleuron with well-developed narrow metapleural

triangle. Subposterior carina less well developed than inQ. decipiens. The whole posterior end of metapleuronheavily depressed, making less contrast between ventraland dorsal parts than in Q. decipiens. Median metapleuralcarina indistinct, but metapleural groove very distinct. Hairsmore abundant than in Q. decipiens, covering much ofventral parts and scattered along posterior parts.

Fig. 6. Quasimodoana gibba sp.n., female antenna.

Fig. 7. Quasimodoana decipiens, scutellum.



Propodeal complex covered with long hairs and roughpuncture, propodeal carinae lyre-shaped, usually narrowerthan 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# longerthan 2r). 2r rather evenly curved, Rs almost straight,distinctly curved only in apical end. Rs þ M and Mnebulous and rather short, M þ Cu1, Cu1 and Cu1a notclearly visible.Female metasoma large, outstretched, pointedly oval.

Male metasoma remarkably small. Surface mostly diaph-anic, with very few and weak punctures, slightly moredistinct on posterior tergites.

Type specimens. Holotype, female, U.S.A., Maryland,Bowie 9.v.1945, DDT Expedition (USNM) pinpoint-gluedon left side, upward-bent head but otherwise in goodcondition (Fig. 9); Paratypes, U.S.A.: female identical dataas holotype (USNM); male (damaged, lacking antennae and

most of legs) identical data as holotype (USNM); male,North Carolina, Mt Pisgah, 5200–5500 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//3GOak-hickory//GMNSF 1500, NE/Sweeps’ (Whittaker)(USNM).

Distribution. North America (see above). AdditionalNearctic specimens of Quasimodoana have been studied bythe authors. These specimens probably represent a few newspecies, but the scarcity of specimens and the difficulties inassociating the sexes mean that we prefer to refrain fromdescribing 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 intwo 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 thegenera Quasimodoana, Trybliographa and Leptopilina. Theonly conflict in the tree is within Trichoplastini, but no treeretains a monophyletic Trichoplasta. Of these analyses,treating the characters as unordered gave a tree-length of236 steps, a consistency index (CI) of 0.42, and a retentionindex (RI) of 0.56. Ordering the characters a priori (logicalordering) resulted in a tree-length of 246 steps, a CI of 0.44and a RI of 0.62. Re-ordering the characters a posteriori(phylogenetic ordering) as expected retained the shortestpossible tree-length, 236 steps, but with higher consistency

Fig. 8. Quasimodoana gibba sp.n., scutellum.

Fig. 9. Quasimodoana gibba sp.n., habitus, female.

Fig. 10. Quasimodoana gibba sp.n., habitus, male.



and retention, namely CI ¼ 0.46, RI ¼ 0.63. Support val-ues differed somewhat between these trees, for most groupsidentical or slightly higher with phylogenetically orderedcharacters, for Leptopilina substantially higher (0.79 unor-dered, 0.95 reordered), whereas Trichoplastini decreasedsomewhat 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 func-tions tried (k ¼ 2–6) (Fig. 12), but with a minor differencefor k ¼ 1 (Eucoila crassinerva grouping with Trybliographalongicornis, rendering Trybliographa non-monophyletic).Compared with the topologies from analyses under equalweights, the ingroup is retained as monophyletic, and one ofthe two alternative topologies inside Trichoplastini is fav-oured. Except for the fact that Trybliographa is not mono-phyletic under k ¼ 1, the topology is identical betweendifferent concavity functions. These trees had CI ¼ 0.46and RI ¼ 0.63, and their parsimony score ranged from$50.93 (k ¼ 1) to $68.08 (k ¼ 6). Support values differedbetween trees, usually in an apparently random way, butfor two groups of interest (Leptopilina and Eucoila þ

Trybliographa þ Quasimodoana) support had smooth peaksat intermediate k values.When the dataset was split into two partitions, one with

the 58 two-state characters, and the other with the 25multistate characters, support values diminished for somegroups, and some differences in topology were produced,but almost all groups of interest were still present for bothdatasets. The multistate character dataset was analysed bothas logically ordered and as phylogenetically ordered, withthe latter producing a clearly shorter tree (96 steps vs. 107)with one remarkable difference, namely that Quasimodoanagrouped with Leptopilina rather than with Trybliographa þEucoila in the logical-order tree (Figs 14, 15). The tree withonly two-state characters did not retain a monophyletic in-group, 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 0.92–1.0). A non-monophyletic Quasimodoanarequired eight extra steps, and a Quasimodoana withinTrichoplastini required ten extra steps. Quasimodoana hasfour unique synapomorphies, namely the distinct subanten-nal striae, the shortened first flagellomere, and two con-cerning the metapleuron; and four uniting it with

Q decipiens

Q gibba

T rapae

T cubitalis

T longicornis

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 ofanalyses under equal weights. Thick branches have bootstrapsupport values over 0.95 in all analyses (characters unordered,logically ordered, phylogenetically ordered).

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

R heptoma

Chrestosema

Glauraspidia

Fig. 12. Single most parsimonous tree of analyses under impliedweights (characters phylogenetically ordered) for k ¼ 2–6. Thickbranches have bootstrap support values over 0.90 in all analyses andover 0.95 in some.



Trybliographa þ Eucoila; four other characters are sharedwith parts or the whole of Leptopilina or Trichoplastini andresult as homoplasies in this analysis (Table 3).Trybliographa is monophyletic in many trees but always

with low support (often collapsed into a polytomy withEucoila in bootstrap trees), but Eucoila þ Trybliographa isretained as monophyletic and well supported in all trees (1.0in all analyses of the entire dataset); and all analyses of theentire dataset have Quasimodoana as the sister of Eucoila þTrybliographa (mostly intermediate support around 0.85).The internal topology in this group varies, with differentattachments of the root in different analyses.Leptopilina is monophyletic in all trees, but with varying

support (bootstrap 0.77–0.95). Inside Leptopilina, L. heter-otoma and L. boulardi constantly group together with goodsupport.In all trees except the two-state-only tree, Eucoilini comes

out as monophyletic, but with very little support (usuallyaround 0.6, but even lower with unordered-only or logicallyordered characters), being dependent on a small number ofcharacters (only three morphological synapomorphies, allin the same body part: characters 52, 53 and 56). A treewith Trichoplastini within Eucoilini (as a sistergroup ofQuasimodoana 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 0.96–1.0), but its internal topology variesand is never well supported; however, Trichoplasta (Tricho-plasta s.str. þ Armigerina) is never retained as monophy-letic without Nordlanderiana.

Discussion

The support found for the monophyly of the new genusseems sufficient, as does that for its inclusion in theEucoilini. The support for the monophyly of Eucoiliniremains weak. Possibly Trichoplastini is nested withinEucoilini, as suggested by other analyses, for exampleBuffington et al. (2007). In that analysis, too, morphologyfavoured inclusion of Leptopilina in a monophyletic groupwith Eucoila and Trybliographa, but the 28S molecular datastrongly favoured Leptopilina as paraphyletic vis-a-vis theTrichoplastini genera, and made the combined data analysisfavour that topology. Members of the ingroup (Eucoilini þTrichoplastini) share characters, the most important ofwhich may be the modified second flagellomere in males

Q decipiens

Q gibba

T rapae

T cubitalis

T longicornis

E crassinerva

L longipes

L heterotoma

L boulardi

Chrestosema

N grunini

T(A) sp

T(T) bouceki

R heptoma

Glauraspidia

Fig. 13. Single most parsimonous tree of analysis with two-statecharacter partition, equal weights.

Q decipiens

Q gibba

L longipes

L heterotoma

L boulardi

T rapae

T longicornis

E crassinerva

T cubitalis

N grunini

T(A) sp

T(T) bouceki

R heptoma

Chrestosema

Glauraspidia

Fig. 14. Single most parsimonous tree of analysis with multistatecharacter partition, logically 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 maleflagellomere modified, not the second. However, there ismuch homoplasy between genera of higher eucoilines (as

documented in Fontal-Cazalla et al., 2002), so relationshipsat that level can be convincingly resolved probably only ina comprehensive, combined analysis of morphological andpreferably multigene molecular data, and including othertaxa.Optimizing biological traits on the tree makes it probable

that Quasimodoana are parasitoids of some calyptrateDiptera larvae living in decaying or fresh plant or mush-room matter (as in most species of Trybliographa, Eucoilaand Leptopilinawith known life histories). The usual patternin Eucoilinae is one wasp genus for one host fly family, andthus the host choices of the relatives of Quasimodoana(Drosophilidae in Lepotopilina, Anthomyiidae in Trybliog-rapha, but several families in Eucoila) provide no ground forspeculation. The feeding habits of these flies (herbivorous,mycophagous or saprophagous, including coprophagousand necrophagous) are diverse enough, and the host recordsfew enough, to provide little ground for speculation onwhere the host shift occurred, whether in mushrooms (manyTrybliographa, some Leptopilina), animal dung (Eucoila,some Trybliographa), carcasses (Eucoila), roots or vegeta-bles (some Trybliographa), fruit (some Leptopilina), leaves ofherbs and ferns (some Trybliographa), conifer cones (someTrybliographa) 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 EucoiliniThomson, 1862, despite the fact that our analysis is consis-tent with the hypothesis of monophyly. Morphologicaldata at this point argue unequivocally for the inclusion ofLeptopilina into Eucoilini, but further research may neces-sitate a reclassification of Eucoilini that either excludesLeptopilina 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 withoutEucoila s.str. (Nordlander, 1981)? Trybliographa is a mono-phyletic sistergroup to Eucoila in most parsimonous trees,but bootstrap trees collapsed Trybliographa þ Eucoila into

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 multistatecharacter partition, phylogenetically ordered, equal weights.

Table 4. Some synapomorphies of Quasimodoana.

Characterno. State Description Shared with

15 2 Subantennal striae distinct Unique20 2 First flagellomere of female antenna shorter than other flagellomeres Unique56 2 Lower metapleural carina pointing high, almost vertically Unique57 2 Surface of metapleuron anterior to posteroventral corner strongly depressed Unique7 1 Hair punctures on lateral part of vertex distinct Trybliographa, Eucoila, Nordlanderiana23 0 Erect seta medially on apical segment of maxillary palp present Trybliographa, Eucoila25 1 First segment of labial palp longer than apical segment Trybliographa, Eucoila67 1 Dorseroventral hairpatch of metacoxa present and small Trybliographa, Eucoila6 1 Compound eye large, twice as long as gena Leptopilina, Trichoplasta bouceki28 1 Submedian pronotal depressions intermedially deep, with ventral edge

distinct but dorsal edge indistinctLeptopilina boulardi (Trichoplastini haveentirely shallow depressions)

33 1 Mesoscutum humped Leptopilina44 2 Posterior part of scutellum in lateral view distinctly produced posteriorly Trichoplastini



a polytomy. Considering the amount of variation within thelarge genus Trybliographa, solving this problem clearlyrequires the sampling of more taxa.The other issue is how to circumscribe Trichoplasta to be

monophyletic (cf. Nordlander & Grijpma, 1991). Armigerinahas been treated as a subgenus here, as suggested by itseffective treatment as a synonym of Trichoplasta in Fontal-Cazalla et al. (2002), whereas the status of Nordlanderiana asan independent genus hitherto has been unquestioned. In thepresent analysis, Armigerina groups with Nordlanderianarather than with Trichoplasta s.str., and there is no supportfor these three taxa together (a possible Trichoplasta s.l.)being distinct from Rhoptromeris. Sampling of more taxaclearly is needed to resolve this issue.

Acknowledgements

Valuable comments, access to unpublished data and otherkinds of help were provided by Matt Buffington. Twoanonymous reviewers helped to improve the manuscript.For hosting the museum visits by GN in 1978 and 1982, andfor the loan of material, we are grateful to Arnold Menke(USNM, Washington), Henry Townes (†) (AEI, in AnnArbor at the time), Lubomir Masner (CNC, Ottawa) and E.Konigsmann (†) (ZMHB, Berlin). We also thank BertViklund (NHRS, Stockholm) and Roy Danielsson (LZM,Lund) for hosting MF more recently. Stefan Gunnarssonand Gary Wife helped with the SEM imaging. The morerecent work was funded by the Swedish Research Council(grant to FR) and the Swedish Taxonomy Initiative (grantto GN and FR).

References

Buffington, M.L., Nylander, J.A.A. & Heraty, J.M. (2007) Thephylogeny and evolution of the figitidae (Hymenoptera: Cyn-ipoidea). Cladistics, 23, 403–431.

Fontal-Cazalla, F.M., Buffington, M.L., Nordlander, G. et al.(2002) Phylogeny of the eucoilinae (Hymenoptera: Cynipoidea:Figitidae). Cladistics, 18, 154–199.

Forster, A. (1855) Die 2te Centurie neuer Hymenopteren. Verhand-lungen des naturhistorischen Vereines der preussischen Rheinlandeund Westphalens, 12, 226–258.

Goloboff, P.A. (1993) Estimating character weights during treesearch. Cladistics, 9, 83–91.

ICZN (International Commission on Zoological Nomenclature)(1999) International Code of Zoological Nomenclature, 4th edn.International Trust for Zoological Nomenclature, London.

ICZN (International Commission on Zoological Nomenclature)(2003) Declaration 44. Amendment of article 74.7.3. Bulletin ofZoological Nomenclature, 60, 263.

Kovalev, O.V. (1989) [A new tribe and a new genus of eucoilid wasps(Hymenoptera, Cynipoidea, Eucoilidae)]. Trudy ZoologitseskogoInstituta an SSSR, Leningrad, 188, 109–113 (in Russian).

Lin, K.S. (1988) The Eucoilidae from Taiwan, I. (Hymenoptera:Cynipoidea). Journal of Taiwan Museum, 41, 1–66.

Maddison, W.P. & Maddison, D.R. (1999) MacClade, version3.08a. Sinauer Associates, Sunderland, MA.

Mickevich, M.F. (1982) Transformation series analysis. SystematicZoology, 31, 461–478.

Mickevich, M.F. & Lipscomb, D. (1991) Parsimony and the choicebetween different transformations for the same character set.Cladistics, 7, 111–139.

Mikula, P. (1989) Cynipoidea. Enumeratio Insectorum Bohemoslo-vakiae. Checklist of czechoslovakian insects III (Hymenoptera)(ed. by L. Ledivy). Acta Faunistica Entomologica Musei Natio-nalis Pragae, 19, 129–132

Nordlander, G. (1980) Revision of the genus Leptopilina Forster,1869, with notes on the status of some other genera (Hymenop-tera, Cynipoidea: Eucoilidae). Entomologica Scandinavica, 11,428–453.

Nordlander, G. (1981) A review of the genus Trybliographa Forster,1869 (Hymenoptera, Cynipoidea: Eucoilidae), EntomologicaScandinavica, 12, 381–402.

Nordlander, G. (1982a) Identities and relationships of the pre-viously confused genera Odonteucoila, Coneucoila, and Tricho-plasta (Hymenoptera, Cynipoidea: Eucoilidae). EntomologicaScandinavica, 13, 269–292.

Nordlander, G. (1982b) Systematics and phylogeny of an interre-lated group of genera within the family Eucoilidae (Insecta:Hymenoptera, Cynipoidea). PhD Thesis, Stockholm University.

Nordlander, G. & Grijpma, P. (1991) Systematics and biology ofRhoptromeris strobigena sp.n., a parasitoid of chloropids inhabit-ing conifer cones (Hymenoptera: Cynipoidea: Eucoilidae).Entomologica Scandinavica, 22, 209–218.

Quinlan, J. (1984) Stentorceps, a remarkable new genus of eucoi-lid (Hymenoptera) from Africa. Systematic Entomology, 9,479–485.

Quinlan, J. (1986) A key to the Afrotropical genera of Eucoilidae(Hymenoptera), with a revision of certain genera. Bulletin ofthe British Museum Natural History Entomology Series, 52,243–366.

Quinlan, J. (1988) A revision of some Afrotropical genera ofEucoilidae (Hymenoptera). Bulletin of the British Museum Nat-ural History Entomology Series, 56, 171–229.

Ronquist, F. (1995) Phylogeny and early evolution of the Cynipoi-dea. Systematic Entomology, 20, 309–335.

Ronquist, F. (1999) Phylogeny, classification and evolution of theCynipoidea. Zoologica Scripta, 28, 1–2, 139–164.

Ronquist, F. & Nordlander, G. (1989) Skeletal morphology of anarchaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). En-tomologica Scandinavica Supplement, 33, 1–60.

Schick, K.N. (1999) Phylogenetic studies of Eucoilinae and Liopteridparasitoids (Cynipoidea: Hymenoptera). PhD Thesis, Universityof California, Davis.

Schulmeister, S. (2003) Review of morphological evidence on thephylogeny of basal Hymenoptera (Insecta), with a discussion ofthe ordering of characters. Biological Journal of the LinneanSociety, 79, 209–243.

Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsi-mony (*and other methods), version 4.0b10. Sinauer Associates,Sunderland, MA.

Vardal, H. (2004) From insect parasitoids to gall inducers andinquilines. PhD Thesis, Uppsala University.

Weld, L.H. (1952) Cynipoidea (Hym.) 1905–1950. Michigan (pri-vately printed), Ann Arbor.

Wilkinson, M. (1992) Ordered versus unordered characters. Cladis-tics, 8, 375–385.

Accepted 25 September 2007



Appendix 2. Characters in the phylogenetic analysis.

Characters are listed in accordance with the headings inFontal-Cazalla et al. (2002). Those characters adopted fromthere are indicated as such within parentheses after thecharacter number. For each character the consistency index(CI) and retention index (RI) scored in this analysis are given.For multistate characters the order consideration is given.

General body measurements

1. Shape of mesosoma: 0 ¼ longer than high, 1 ¼ lengthand height approximately equal. CI ¼ 0.5, RI ¼ 0.75.

2. Body size: 0 ¼ small (length well under 2 mm), 1 ¼intermediate (approximately 2 mm), 2 ¼ large (clearlyover 2 mm). A priori and a posteriori ordered. CI¼ 0.33,RI ¼ 0.33.

Head

3. (4 unchanged) Size of ocelli: 0 ¼ small, 1 ¼ large. CI ¼0.25, RI ¼ 0.

4. (5 changed) Relative position of anterior ocellus: 0 ¼contiguous with hind ocelli, 1 ¼ clearly anterior to hindocelli. CI ¼ 0.5, RI ¼ 0.8.

5. (7 changed) Shape of compound eye in dorsal view:0 ¼ protruding, 1 ¼ not protruding. CI ¼ 0.33, RI ¼ 0.

6. Size of compound eye: 0 ¼ subequal to gena, 1 ¼ longerthan gena, 2 ¼ twice as long as gena. A priori ordered,a posteriori unordered. CI ¼ 0.4, RI ¼ 0.57.

7. (9 changed) Hair punctures on lateral part ofvertex: 0 ¼ indistinct or absent, 1 ¼ distinct. CI ¼ 0.5,RI ¼ 0.83.

8. (11 unchanged) Relative position of antennal socket:0 ¼ close to ocelli, 1 ¼ far from ocelli. CI ¼ 0.5, RI ¼ 0.

Appendix 1. Character matrix in phylogenetic analysis.

1 1 1 2 1 3 1

Quasimodoana decipiens 1 1 0 0 1 2 1 1 0 1 0 0 0 0 2 1 1 0 1 2 0 0 0 1 1 0 0 1 0 1 1 0 1 0 0 0 1 1 0 1Quasimodoana gibba 1 1 0 0 1 2 1 1 0 ? ? ? 0 0 2 1 1 0 1 2 0 0 0 0 1 0 0 1 0 1 1 0 1 0 0 0 1 1 0 1Trybliographa rapae 1 2 0 0 1 1 1 1 0 0 0 1 0 1 1 1 1 1 1 1 0 1 0 0 1 1 0 0 0 0 2 1 0 1 1 0 0 1 0 1Trybliographa longicornis 1 1 0 0 1 1 1 1 0 0 0 1 0 0 1 1 0 0 1 1 0 1 0 0 1 0 0 0 1 1 0 1 0 0 0 0 2 2 0 2Trybliographa cubitalis 1 1 0 0 1 1 1 1 0 0 0 1 0 1 1 1 1 0 1 1 0 1 0 0 1 1 0 0 0 0 2 1 0 0 0 0 0 2 0 1Eucoila crassinerva 1 2 1 0 0 1 1 0 0 0 0 1 0 0 1 1 1 1 1 1 0 1 0 0 1 1 0 0 0 0 2 0 0 1 1 0 2 2 0 1Leptopilina longipes 1 0 1 0 0 2 0 1 1 1 0 0 0 0 0 1 1 0 1 1 1 0 1 0 0 0 0 0 1 0 1 0 1 0 0 0 2 1 1 1Leptopilina heterotoma 1 0 1 0 0 2 0 1 1 1 0 0 1 0 1 1 0 1 0 0 1 0 1 0 0 0 0 0 0 0 1 0 1 0 0 1 1 1 1 1Leptopilina boulardi 1 0 0 1 1 2 0 1 1 1 0 0 0 0 1 1 0 1 0 1 1 0 1 0 0 0 0 1 1 0 1 0 1 0 0 1 1 0 1 2Nordlanderiana grunini 0 2 0 1 1 1 1 1 0 1 2 0 0 0 1 1 1 0 1 1 0 0 1 0 0 0 1 2 1 0 0 0 0 1 1 1 1 1 0 0Trichoplasta (Armigerina) sp. 0 1 0 1 1 1 0 1 0 1 1 0 0 1 0 1 0 0 1 1 0 0 1 0 0 0 1 2 1 0 1 1 0 1 1 1 1 1 0 0Trichoplasta (s. str.) bouceki 0 0 0 1 1 2 0 1 0 ? ? ? 0 1 0 1 0 0 1 1 0 0 1 0 0 0 1 2 1 0 1 0 0 1 1 1 1 0 0 0Rhoptromeris heptoma 0 0 0 1 1 0 0 1 0 1 0 0 0 1 0 1 0 0 1 1 1 0 1 0 0 0 1 2 0 0 1 0 0 1 1 1 1 0 0 0Glauraspidia microptera 1 1 0 1 1 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 1 0 1 0 0 1 0 0 0 1 0 0 0 0 0 0 0 2 1 1Chrestosema erythropum 0 1 1 0 1 2 0 1 1 1 2 0 1 0 0 0 1 0 1 0 1 0 1 0 0 0 0 0 0 1 1 0 0 0 0 0 1 1 1 1

Appendix 1. Continued

4 1 5 1 6 1 7 1 8 1

0 1 0 2 0 0 2 0 1 1 1 1 1 1 0 2 2 2 1 1 1 0 0 2 0 0 1 0 1 0 0 0 0 0 2 0 0 0 1 1 1 0 00 1 0 2 1 0 2 0 1 1 1 1 1 1 0 2 2 2 1 1 1 0 0 2 0 0 1 0 1 0 0 0 0 1 2 0 0 1 1 1 2 0 01 1 0 1 0 0 2 0 0 1 1 2 1 0 1 1 1 1 0 1 1 1 0 2 0 1 1 1 1 0 0 0 1 1 2 0 0 1 1 2 2 1 00 2 0 0 0 1 2 1 0 1 1 2 2 1 1 1 1 1 0 0 1 1 1 1 0 1 1 1 1 0 1 0 0 1 2 0 0 1 0 2 2 1 10 1 0 1 0 0 2 0 0 1 1 2 2 0 1 1 1 2 0 0 1 1 0 2 0 1 1 1 1 0 0 0 1 1 2 0 0 1 1 1 1 1 00 1 0 1 2 0 2 1 0 0 1 2 2 1 1 1 1 1 1 0 0 0 0 2 0 1 0 0 0 0 0 0 0 1 2 0 0 1 0 2 2 1 01 1 0 1 1 0 2 0 1 1 1 1 1 1 0 1 1 1 1 1 0 0 0 0 0 0 0 0 1 0 0 1 0 1 2 1 1 1 0 1 0 1 12 2 1 0 1 1 2 0 1 1 1 1 1 1 0 0 1 1 1 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 1 1 1 0 0 1 0 0 11 2 1 0 1 1 0 0 1 1 1 1 1 1 0 0 1 1 1 0 1 1 0 1 1 1 0 0 0 0 0 1 0 1 2 1 1 0 0 0 0 0 10 0 0 2 0 0 0 0 1 1 1 0 0 0 0 0 0 1 1 1 0 1 1 0 1 0 0 0 0 1 0 0 0 0 2 0 0 0 1 1 1 0 00 0 0 2 0 0 1 0 1 0 1 0 0 0 0 0 0 1 1 1 1 0 1 1 1 0 0 0 0 1 0 0 0 0 1 0 0 0 1 2 1 1 01 0 0 2 0 0 1 0 1 0 1 0 0 0 0 0 0 0 0 1 1 0 1 0 1 0 0 0 1 1 0 0 0 0 1 0 0 0 0 2 1 ? ?2 0 0 2 2 1 0 0 1 0 1 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0 0 0 0 1 0 1 0 0 1 0 0 0 0 2 1 1 10 2 0 0 0 0 2 0 1 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 2 1 1 0 1 0 1 0 0 0 0 0 0 2 2 1 11 1 0 1 2 0 2 0 1 1 1 0 0 1 0 0 0 0 1 1 0 0 0 2 0 0 2 1 1 0 0 0 0 1 1 1 0 1 0 2 2 1 1



9. (23 changed) Lateral margin of occiput: 0 ¼ not definedby carina, 1 ¼ defined by carina. CI ¼ 0.5, RI ¼ 0.67.

10. (28 unchanged) Length of gula: 0 ¼ short, 1 ¼ long.CI ¼ 1, RI ¼ 1.

11. Sculpture of gula: 0 ¼ smooth, 1 ¼ with few striae,2 ¼ with several striae. A priori ordered, a posterioriunordered. CI ¼ 0.5, RI ¼ 0.33.

12. (30 unchanged) Shape of hypostomal carina medially:0 ¼ straight, 1 ¼ distinctly angled. CI ¼ 1, RI ¼ 1.

13. Angle of genal ventral margin: 0 ¼ straight angle tohead outline, 1 ¼ open angle to head outline. CI ¼ 0.33,RI ¼ 0.

14. Shape of ventral margin of clypeus: 0 ¼ more or lessevenly rounded 1 ¼ distinctly emarginate. CI ¼ 0.33,RI ¼ 0.5.

15. Subantennal striae: 0 ¼ absent, 1 ¼ present but weaklydelineated, 2 ¼ distinct. A priori and a posterioriordered. CI ¼ 0.5, RI ¼ 0.67.

Antennae

16. (36 unchanged) Shape of second flagellomere of maleantenna: 0 ¼ cylindrical, 1 ¼ slightly asymmetric. CI ¼0.5, RI ¼ 0.

17. Number of female antennal club joints: 0 ¼ 7 or fewer,1 ¼ 8 or more. CI ¼ 0.25, RI ¼ 0.5.

18. Shape of clavomeres of female antenna: 0 ¼ weaklyenlarged compared with flagellomeres, longer thanwide, 1 ¼ strongly enlarged, width equal or subequalto length. CI ¼ 0.33, RI ¼ 0.33.

19. Shape of apical clavomere of female antenna: 0 ¼length subequal to width, 1 ¼ distinctly longer thanwide. CI ¼ 1, RI ¼ 1.

20. Length of first flagellomere of female antenna: 0 ¼distinctly longer than others, 1 ¼ equal to others, 2 ¼shorter than others. A priori and a posteriori ordered.CI ¼ 0.5, RI ¼ 0.33.

Labio-maxillary complex

21. (47 changed) Angle of distal margin of subapicalsegment of maxillary palp and movement of apicalsegment: 0 ¼ margin slants inwards and apex bendsinwards, 1 ¼ margin straight or slants outwards, apexbends outwards. CI ¼ 0.5, RI ¼ 0.8.

22. (48 changed) Relative length of apical segment ofmaxillary palp: 0 ¼ longer than preceding segment,1 ¼ shorter. CI ¼ 1, RI ¼ 1.

23. (51 unchanged) Erect seta medially on apical segment ofmaxillary palp: 0 ¼ present, 1 ¼ absent. CI ¼ 1, RI ¼ 1.

24. Shape of apical segment of maxillary palp: 0 ¼ more orless parallel-sided, 1 ¼ distinctly enlarged towardsapex. CI ¼ 0.5, RI ¼ 0.75.

25. (53 unchanged) Length of first segment of labial palp:0 ¼ short, 1 ¼ longer than apical segment. CI ¼ 1,RI ¼ 1.

26. Shape of apical segment of labial palp: 0 ¼ cylindricalor barrel-shaped, widest proximal to middle, 1 ¼ enlarg-ened, widest point in apical half. CI ¼ 0.33, RI ¼ 0.33.

Pronotum

27. (56 unchanged) Shape of submedian pronotal depres-sion (lateral pit of pronotal plate) laterally: 0 ¼ open,1 ¼ closed. CI ¼ 1, RI ¼ 1.

28. (57 changed) Depth of submedian pronotal depression(lateral pit of pronotal plate): 0 ¼ deep all over, ventraland dorsal edges distinct, 1 ¼ intermediate, ventraledge distinct but dorsal edge indistinct, 2 ¼ shallow,both ventral and dorsal edges indistinct. A prioriordered, a posteriori unordered. CI ¼ 0.5, RI ¼ 0.78.

29. Shape of dorsal margin of pronotal plate: 0 ¼ rounded,1 ¼ straight, parallellotrapezic. CI ¼ 0.25, RI ¼ 0.4.

30. Height of pronotal plate: 0 ¼ reaching pronotal-meso-scutal suture, 1 ¼ surpassing pronotal-mesoscutalsuture. CI ¼ 0.25, RI ¼ 0.25.

31. (62 unchanged) Ridges extending posteriorly fromlateral margin of pronotal plate: 0 ¼ absent, 1 ¼ short,2 ¼ reaching dorsal margin. A priori ordered, a poste-riori unordered. CI ¼ 0.4, RI ¼ 0.25.

32. Distribution of hairs on pronotum: 0 ¼ not extendinginto posteroventral part, 1 ¼ extending all the way tomesopleural border. CI ¼ 0.5, RI ¼ 0.67.

Mesoscutum

33. Shape of mesoscutum: 0 ¼ smooth, slowly rising, 1 ¼humped, abruptly rising. CI ¼ 0.5, RI ¼ 0.75.

34. Length of mesoscutum: 0 ¼ far shorter than entiremesosoma height, 1 ¼ approaching entire mesosomaheight. CI ¼ 0.33, RI ¼ 0.6.

35. Width of mesoscutum: 0 ¼ wider than long, 1 ¼ widthnot surpassing length. CI ¼ 0.33, RI ¼ 0.6.

36. (71 changed) Parascutal carina sinuation: 0 ¼ dis-tinctly, 1 ¼ less sinuate. CI ¼ 0.5, RI ¼ 0.8.

Scutellar-Axillar Complex

37. (72 changed) Length of longitudal carina or septumseparating scutellar foveae: 0 ¼ short, 1 ¼ intermedi-ate, 2 ¼ long. A priori ordered, a posteriori unordered.CI ¼ 0.4, RI ¼ 0.25

38. (75 changed) Striate sculpture on lateral bar: 0 ¼absent, 1 ¼ weak, 2 ¼ strong. A priori and a posterioriordered. CI ¼ 0.33, RI ¼ 0.2.

39. Length of lateral bar: 0 ¼ long, 1 ¼ short. CI ¼ 0.5,RI ¼ 0.75.

40. (80 unchanged) Shape of rim of scutellar plate in lateralview: 0 ¼ flat, 1 ¼ slightly convex, 2 ¼ distinctly con-vex. A priori and a posteriori ordered. CI ¼ 0.67,RI ¼ 0.75.

41. Length and shape of scutellar plate: 0 ¼ short ovoid,1 ¼ slightly outstretched ellipse, 2 ¼ long ellipse, ap-proaching posterior end of scutellum. A priori anda posteriori ordered. CI ¼ 0.33, RI ¼ 0.43.

42. Height of scutellar plate: 0 ¼ directly onto dorsalsurface, 1 ¼ slightly elevated dorsally, 2 ¼ stronglyelevated dorsally. A priori and a posteriori ordered.CI ¼ 0.5, RI ¼ 0.67.



43. (81 changed) Shape of dorsal surface of scutellar plate:0 ¼ partly concave, 1 ¼ flat or slightly convex. CI ¼ 1,RI ¼ 1.

44. (86 changed) Shape of dorsoposterior part of scutellumin lateral view: 0 ¼ truncate, 1 ¼ rounded, slightlyproduced posteriorly, 2 ¼ distinctly produced posteri-orly. A priori and a posteriori ordered. CI ¼ 0.4,RI ¼ 0.62.

45. Shape of dorsolateral margin of scutellum: 0 ¼rounded, more or less indiscernible, 1 ¼ weakly angled,although distinct by different sculpture of dorsal andlateral parts, 2 ¼ sharply angled. A priori ordered,a posteriori unordered. CI ¼ 0.29, RI ¼ 0.17.

46. Sculpture of dorsal surface of scutellum: 0 ¼ roughlypunctured, 1 ¼ partly or wholly smooth. CI ¼ 0.33,RI ¼ 0.33.

47. Structure of axillula: 0 ¼ shallow, not sharply defined,1 ¼ deep, but defining margin incomplete, 2 ¼ verydeep, defining margin sharp. A priori ordered, a poste-riori unordered. CI ¼ 0.4, RI ¼ 0.5.

48. Shape of axillula in lateral view: 0 ¼ not longer thanhigh, 1 ¼ clearly longer than high. CI ¼ 0.5, RI ¼ 0.

Mesopectus (Mesopleuron and Mesosubpleuron)

49. (91 changed) Subalar pit: 0 ¼ present in subalar groove,1 ¼ absent. CI ¼ 1, RI ¼ 1.

50. Position of mesopleural carina: 0 ¼ one-quarter or lessof total height of mesopleuron below carina, 1 ¼ one-third or more of height of mesopleuron below carina.CI ¼ 0.33, RI ¼ 0.5.

Metapectal-Propodeal Complex

51. (part of 107 changed) Ridge of posteroventral corner ofmetapleuron: 0 ¼ low, 1 ¼ high. CI ¼ 1, RI ¼ 0.

52. (part of 107 changed) Orientation of area beyondposteroventral ridge: 0 ¼ posteriorly, 1 ¼ posterolater-ally but concave, 2 ¼ posterolaterally and partly con-cave, partly convex. A priori and a posteriori ordered.CI ¼ 1, RI ¼ 1.

53. (part of 107 changed) Hairs of posteroventral corner ofmetapleuron: 0 ¼ on whole surface, 1 ¼ only in dorsalpart, 2 ¼ absent. A priori and a posteriori ordered.CI ¼ 0.57, RI ¼ 0.86

54. Structure of posterolateral angle of metapleuron indorsal part: 0 ¼ weakly demarcated, open angle, 1 ¼sharply demarcated, sharp angle. CI ¼ 0.5, RI ¼ 0.83

55. Shape of anteroventral cavity of metapleuron: 0 ¼ovoid, fairly circular, 1 ¼ outstretched, far higher thanwide. CI ¼ 1, RI ¼ 1.

56. Orientation of lower metapleural carina (Nordlander’s‘ridge 3’): 0 ¼ straight, pointing low, towards dorsaledge of anteroventral cavity, 1 ¼ more or less curved,pointing higher, towards median pit of mesopleural/metapleural suture, 2 ¼ curved, pointing even higher,towards anterodorsal corner of metapleura. A prioriand a posteriori ordered. CI ¼ 0.67, RI ¼ 0.86.

57. Surface of metapleuron anterior to posteroventral cor-ner: 0 ¼ not depressed, 1 ¼ weakly depressed, 2 ¼

strongly depressed. A priori and a posteriori ordered.CI ¼ 1, RI ¼ 1

58. Surface of metapleuron dorsal to posteroventral corner:0 ¼ not depressed, 1 ¼ weakly depressed, 2 ¼ stronglydepressed. A priori and a posteriori ordered. CI ¼ 0.5,RI ¼ 0.6

59. Number of carinae on ventral part of metapleuron:0 ¼ basic pattern of carinae present or indicated, but noadditional carinae (with higher metapleural carina(‘ridge 1’), median (‘ridge 2’), lower (‘ridge 3’) andventral (metacoxal rim) carina), 1 ¼ with additionalcarinae. CI ¼ 0.33, RI ¼ 0.5.

60. Hairs of median part of metapleuron: 0 ¼ no hairsanterior to posterior carina ventral to median meta-pleural carina, 1 ¼ with a few hairs anterior to posteriorcarina in median part. CI ¼ 0.33, RI ¼ 0.5.

61. Hairs of dorsal part of metapleuron: 0 ¼ restricted toimmediate base of calyptra, 1 ¼ protruding anteriorly,covering at least one-third of length of metapleuron.CI ¼ 0.4, RI ¼ 0.

62. (111 unchanged) Shape of calyptra in posterior view:0 ¼ rounded, 1 ¼ elongate. CI ¼ 0.25, RI ¼ 0.4.

63. Shape of propodeum in lateral view: 0 ¼ short andsteep, 1 ¼ outstretched posteriorly. CI ¼ 0.5, RI ¼ 0.8.

64. (113 changed) Shape of lateral propodeal carina: 0 ¼straight, 1 ¼ curved, 2 ¼ angled. A priori and a poste-riori unordered. A priori ordered, a posteriori unor-dered. CI ¼ 0.25, RI ¼ 0.33.

65. (114 unchanged) Dorsal extent of lateral propodealcarina: 0 ¼ not reaching scutellum, 1 ¼ reaching scu-tellum. CI ¼ 0.5, RI ¼ 0.8.

66. Hairs of median part of posterior face of propodeum:0 ¼ present, 1 ¼ absent, at least ventrally. CI ¼ 0.5,RI ¼ 0.75.

Legs

67. Size of posterolateral hair patch of mesocoxae: 0 ¼absent, 1 ¼ present and small, 2 ¼ present and large.A priori ordered, a posteriori unordered. CI ¼ 0.4,RI ¼ 0.57.

68. Dorseroventral hairpatch of metacoxae: 0 ¼ weak, 1 ¼strong. CI ¼ 0.33, RI ¼ 0.5.

69. Shape of metacoxae: 0 ¼ strong, only slightly longerthan wide, length subequal to ‘metapleuron’, 1 ¼ lessstrong, almost twice as long as wide, longer than‘metapleuron’. CI ¼ 0.25, RI ¼ 0.4.

70. (122 unchanged) Shape of metatarsal claw: 0 ¼ basestrongly expanded and apex strongly bent, 1 ¼ baseweakly expanded and apex slightly bent. CI ¼ 1, RI¼ 1.

71. Size of arolium: 0 ¼ smaller than or subequal to claw,1 ¼ distinctly larger than claw. CI ¼ 0.67, RI ¼ 0.

Wings

72. (128 with states lumped) Hair fringe along apicalmargin of forewing: 0 ¼ short, absent or moderatelylong (hairs not longer than first abscissa of radius),



1 ¼ very long (hairs longer than first abscissa). CI ¼0.67, RI ¼ 0.67.

73. Closure of marginal cell (R1 vein reaching Rs or not);0 ¼ closed, 1 ¼ open. CI ¼ 0.5, RI ¼ 0.5.

74. Shape of marginal cell: 0 ¼ shallow, angle of secondabscissa open (or straight after distinct curve of radius),1 ¼ deep, angle of second abscissa straight. CI ¼ 0.33,RI ¼ 0.67.

75. Curvature of second abscissa of radius: 0 ¼ almoststraight, 1 ¼ distinctly curved in apical end, 2 ¼ curvedin entire length. A priori unordered, a posterioriordered. CI ¼ 0.5, RI ¼ 0.6.

Metasoma

76. (135 unchanged) Shape of posterior part of femalepetiole: 0 ¼ not abruptly widened, 1 ¼ abruptly wid-ened. CI ¼ 0.5, RI ¼ 0.67.

77. (136 changed) Ventral flange of annulus of femalepetiole: 0 ¼ absent or small and narrow, 1 ¼ largeand broad. CI ¼ 1, RI ¼ 1.

78. Shape of large tergite (syntergum) of female metasoma:0 ¼ truncate posteriorly, 1 ¼ rounded posteriorly,whole tergite more or less crescent-shaped. CI ¼ 0.33,RI ¼ 0.67.

79. Shape of hypopygium: 0 ¼ small, weakly protruding,1 ¼ large, strongly protruding. CI ¼ 0.33, RI ¼ 0.6.

80. Hairy ring of female metasoma; 0 ¼ weak, 1 ¼ strong,2 ¼ very strong, woolly. A priori and a posterioriordered. CI ¼ 0.4, RI ¼ 0.5

81. Hairy ring of male metasoma: 0 ¼ weak, 1 ¼ strong,2 ¼ very strong, woolly. A priori ordered, a posterioriunordered. CI ¼ 0.4, RI ¼ 0.57.

82. Shape of ventral edge of large tergite (syntergum) ofmale metasoma: 0 ¼ broadly biangular, 1 ¼ sharplyrounded. CI ¼ 0.33, RI ¼ 0.5.

83. Size of second tergite of male metasoma (postsynter-gum; tergite of sixth abdominal segment): 0 ¼ approx-imately one-third of height of metasoma, 1 ¼approximately one-half of height of metasoma. CI ¼0.33, RI ¼ 0.67.



Documents

NRM - Forshage et al systent 2008Correspondence: Mattias Forshage, Department of Systematic Zoology, Evolutionary Biology Centre, Norbyva¨gen 18D, SE-75236 Uppsala, Sweden. E-mail: