A basal chalcidoid (Insecta: Hymenoptera) from the earliestCretaceous or latest Jurassic of MongoliaALEXANDR P. RASNITSYN, HASAN H. BASIBUYUK and DONALD L. J. QUICKE
Rasnitsyn, A.P., Basibuyuk, H.H. & Quicke, D.L.J.: A basal chalcidoid (Insecta: Hymenop-tera) from the earliest Cretaceous or latest Jurassic of Mongolia. Insect Syst. Evol. XX;XX-XX. Copenhagen xxxx. ISSN 1399-560X
Abstract – Khutelchalcis gobiensis gen. et sp. n. (Khutelchalcididae fam. n.), based on a tinyimpression fossil collected from the lowermost Cretaceous or uppermost Jurassic deposits ofKhutel-Khara in East Mongolia, was studied using an environmental chamber scanning elec-tron microscope. Character analysis suggests that the fossil belongs to a putative basal groupof the Chalcidoidea. The greatly reduced wing venation, apparent presence of multiporousplate sensillae with long aperture on the flagellomeres, and a high antennal insertion all sug-gest that the new taxa is related to the Chalcidoidea. The angulation of fore wing vein Cu atthe M+Cu fork is similar to that of Jurassic Jurapriidae, Cretaceous Serphitidae as well as var-ious chalcidoids. This character might be a synapomorphy for Chalcidoidea + Serphitoidea(including Mymarommatidae) + Jurapriidae, which in turn are possibly a sister clade toPlatygastroidea. The small size of this species is discussed in relation to conflicting scenariosconcerning the plesiomorphic biology of the chalcidoid+proctotrupoid linage. Mymarommat-idae and Serphitidae are combined in a single superfamily, Serphitoidea Brues, 1937 (= My-marommatoidea Debauche, 1948, syn. nov.).
A. P. Rasnitsyn, Palaeontological Institute, Russian Academy of Sciences, 117647 Moscow,Russia. (e-mail: [email protected]) and Department of Palaeontology, The Natural HistoryMuseum, London SW7 5BD, UK; H. H. Basibuyuk, Department of Biology, Faculty ofScience and Literature, Cumhuriyet University, 58140-Sivas, Turkey. (e-mail: [email protected]); D.L.J. Quicke, Department of Biological Sciences, Imperial CollegeLondon, Silwood Park Campus, Ascot SL5 7PY, UK and Department of Entomology, TheNatural History Museum, London SW7 5BD, UK. ([email protected]).
Insect Syst.Evol.
Introduction
The Chalcidoidea is one of the largest parasiticwasp superfamilies, and comprises more than21,000 described species, though the actual num-ber is undoubtedly very considerably greater thanthis (Noyes 1978, 1998, LaSalle & Gauld 1991).They occur in almost all terrestrial and a fewaquatic habitats and display a huge array of biolo-gies. Yet despite this, their fossil record is verysparse and the oldest known fossils date from theUpper Cretaceous (Yoshimoto 1975; Rasnitsyn1980) whereas phylogenetic considerations sug-gest that they should be a good deal older (Boucek1988; Gibson et al. 1999; Broad et al. submitted).One probable reason for this is that they are typi-cally small to very small insects and so are noteasy to detect in sedimentary deposits. This argu-
ment, however, assumes that the earliest chalci-doids were indeed small insects as are the majori-ty of extant species. Until recently, experts in thegroup have generally suggested that the lessderived of the living species are in fact rather larg-er-bodied robust taxa (e.g. Cleonyminae) that aretypically idiobiont ectoparasitoids of wood-boringhosts, and these would not propose such preserva-tional problems. Thus, if this scenario was correct,it would be harder to explain the absence of chal-cidoids in the earlier fossil record.
Recently, phylogenetic analysis of the familiesof Hymenoptera (Ronquist et al. 1999), and of thefamilies of Chalcidoidea (Campbell et al. 2000)have suggested an alternative scenario, that is, thatthe ancestral chalcidoid may have been a small-bodied endoparasitoid such as an egg parasitoid.
2 Rasnitsyn, A. P. et al. INSECT SYST. EVOL. 35.2 (2004)
Fig. 1. Khutelchalcis gobiensis Rasnitsyn, Basibuyuk & Quicke sp. n. Line drawing based on the part (half-imprintwith body structures looking normal, not mirrored) and, below, counterpart. Symbols: a1 - scape, a2 - pedicel, ax -axilla, cx3 - hind coxa, f1 - fore femur, ta1-3 – fore, mid and hind tarsi, ti1-3 - fore, mid and hind tibiae, N1-3 - pro-,meso- and metanotum (the latter probably with metapostnotum delimited behind), o - eye, ov - ovipositor (possiblycombined V1, V2 and maybe V3), pp? - supposed prepectus, s - spiracular excision of pronotum, scl - mesoscutellum,vp - mesothoracic ventropleuron (ventral surface of mesothorax), Vr1 - supposed upper margin of first valvifer, Vr2- second valvifer, V3 - ovipositor sheath; vein symbols conventional. In wing, dotted line shows sharp concave fur-row, dashed line - rounded convex fold.
Further, Broad et al. (submitted) have shown thatin order to reconstruct the ancestral chalcidoid asan ectoparasitoid on existing phylogenies requiresextremly high weighting against transitions fromecto- to endoparasitoid life styles as opposed totransitions in the other direction.
The phylogenetic position and internal phyloge-ny of Chalcidoidea have been recently reviewedand updated (Gibson 1999, Gibson et al. 1999).These results are rather disparate concerning rela-tionships within the superfamily but might bemore promising in respect to rooting the group.This is because Gibson suggests selection betweentwo potential sister groups, Platygastroidea andMymarommatidae, which might be closely related(Rasnitsyn 1988). We do not examine here thealternate hypothesis of Gibson (1999) on platygas-troid monophyly including Pelecinidae, Procrot-rupidae and Vanhorniidae which is a topic requir-ing a separate study.
Here we describe a new family of fossil Hymen-optera from the Jurassic/Cretaceous boundary inMongolia based on an impression fossil, thatappears to be either a chalcidoid or at least a mem-ber of the chalcidoid + proctotrupoid s.l. clade. Itssmall size is consistent with the more recent viewsabout chalcidoid ancestral biology describedabove, rather than the more traditional ‘story’ thatwas probably largely based on the dogma that themost basal member of a given clade of parasiticwasps would have to have been an ectoparasitoidof a concealed, wood-borring host (Quicke et al.2000). It can also shed some light on the problemof chalcidoid sister group relationships as it isolder than any other known Chalcidoidea andsharing characters that are characteristic of eitherChalcidoidea or various Platygastroidea s.l. Phylo-genetic relationships of the new family and of theChalcidoidea discussed here is not supported byparsimony analysis because of incomplete preser-vation state which only permits a very limited setof characters to be used, and these are insufficientfor such a formal calculation.
Material and methods
The present publication is based on a well pre-served and almost complete part and counterpartfossil impressed in a small piece of white inter-basalt mudstone from the lower Tsagan TsabFormation in 75 km SE Sain Shand (East GobiAymag, Mongolia). These deposits have yielded
more than 3,000 fossil insects that reveal bothJurassic and Cretaceous affinities and therefore areconsidered to represent the earliest Cretaceous(Ponomarenko 1990, Rasnitsyn et al. 1998). Thecollection, including the fossil under description,is kept in the Palaeontological Institute, RussianAcademy of Science, Moscow (abbreviated belowas PIN).
The fossil was studied using binocular lightmicroscopes in PIN and at The Natural HistoryMuseum in London, and with an EnvironmentalChamber Mark ISI ABT 55 scanning electron mi-croscope in at The Natural History Museum. Lightmicroscopy revealed relatively little because of thelow colour contrast and confusing background tex-ture of the type specimens. However, examinationusing environmental chamber SEM under highvoltage and very short working distance showedconsiderably more detail and has enabled us todescribe the new taxon.
Family Khutelchalcididae Rasnitsyn,Basibuyuk & Quicke, fam. n.
Type genus. – Khutelchalcis Rasnitsyn, Basibuyuk& Quicke, gen. n. by present designation andmonotypy.
Diagnosis. – Size small (body length of the onlyknown specimen less than 2 mm; Figs. 1-3). An-tennal attachment high (at midheight of head andabove midheight of eye, Fig. 4), antenna geni-culate, scape moderately large, single anellus pos-sibly present (The antennal segments interpretedhere as pedicel and anellus can be easily taken asordinary scape and pedicel respectively. However,the two large pyriform structures proximal to thepedicel can hardly be anything other than scapes.These two segments are placed in correct positionof a scape, have similar form, and show the samecolour pattern under polarised illumination as therest of body. Therefore, these are unlikely to becracks of the matrix or other strange structures).Flagellomeres as seen from inside (Fig. 5) withlong slit-like structures of varying length up tomore than half of segment length, only compara-ble with apertures of longitutinal multiporousplate sensillae as figured by Basibuyuk & Quicke(1999: figs 2F, 4G,) for various Chalcidoidea andfor Cynipidae (but not for other Cynipoidea exam-ined: id., figs. 2G, 4H, 4I). Head orthognathous.Pronotum short medially, widely emarginate atposterodorsal angle to harbour mesothoracic spir-
INSECT SYST. EVOL. 35:2 (2004) A basal chalcidoid 3
acle (Figs. 1, 3, 6: sculptured mesopleural integu-ment is distinctly, smoothly bent inward, that is,toward the observer, at the margin of excision, thusindicating the true excision and not just a brokenoff part of the integument, nor any other structurelike tegula superimposing the body wall). Freeprepectus, unknown if external or not, apparentlydelimited as elongate structure extending down-ward from hind end of putative spiracular emar-gination (Fig. 6). Legs, as preserved, not distinc-tive. Fore wing venation (Figs. 1-3) similar to thatof Scelionidae (Fig. 7): costal space moderatelynarrow, margined anteriorly with distinct vein C,pterostigma not prominent (possibly absent), Ronly slightly incrassate and 2rs not at all incrassatecompared to other veins, first section of RS almostparallel to M (but disconnected unlike in Scelion-
idae), distal abscissa of RS short, subparallel andrather close to but not approaching to wing margin(marginal cell narrow), 2rrs short, subperpendicu-lar to RS; RS+M and M distad of 2rrs (possiblyexcept a short section very far from base of M) andall crossveins except 2rrs absent. Unlike in Scel-ionidae, vein M+Cu and Cu join at an angle. Hindwing unknown. Metasoma rather spindle-shaped,apically acuminate at least in side view, withoutevident petiole, no terga or sterna particularlylarge and none occupying more than half metaso-ma length, with metasomal apex probably open atrest. Ovipositor possibly internal, with at least itsmost part covered by long apical sternum, withvalvifer apparently not coiled cynipoid-like.
Composition. – Type genus only.
4 Rasnitsyn, A. P. et al. INSECT SYST. EVOL. 35.2 (2004)
Fig. 2. SEM photo of the part. [kchalcis2.*]
Comparison. – The Khutelchalcididae appears tobelong to the Proctotrupoidea s.l. + Chalcidoideagroup of families as indicated by its small size,reduced wing venation and ovipositor not extend-ing beyond apex of metasoma. The new family isfurther tentatively attributed to Chalcidoideabecause of its putative multiporous plate sensillae
interpreted after the long inner aperture, the raresynapomorphy otherwise known only in chalci-doids and, homoplastically, in the higher (phy-tophagous) Cynipoidea of the family Cynipidae. Ifour interpretation of the fossil is correct in respectof the presence of the spiracular excision, an indi-cator of the chalcidoid synapomorphy of a dorsal
INSECT SYST. EVOL. 35:2 (2004) A basal chalcidoid 5
Fig 3. Composed SEM photo of the counterpart. For symbols see Fig. 1.
spiracular position, and of the free prepectus, aplesiomorphy lost in all extant Proctotrupomorphaother than the Monomachidae, Chalcidoidea, andpossibly Mymarommatidae, monophyly of Khu-telchalcididae within Chalcidoidea is very likely.Within Chalcidoidea the new family differs fromall other families in its far more plesiomorphicwing venation which is more complete and hasveins of more uniform thickness than in any extanttaxa. It differs from the majority of chalcidoids in
the presence of single antennal anellus whichlooks like a delimited basal part of the first flagel-lomere rather than a real anellar segment (freeanelli are also absent in Mymaridae, Leucospid-idae, Eucharitidae: Eucharitinae; see Gibson, 1986,for details). No definitive synapomorphy has beenfound as characterising Khutelchalcididae, thoughthis could be because of the incomplete preserva-tion state of the only known fossil.
6 Rasnitsyn, A. P. et al. INSECT SYST. EVOL. 35.2 (2004)
Fig. 4. SEM photo and morphological interpretation of the head, antenna and partial thorax of the counterpart
Fig. 5. SEM photo of antenna of the counterpart and its morphological interpretation.
Genus Khutelchalcis Rasnitsyn, Basibuyuk &Quicke, gen. n.
Type species. – Khutelchalcis gobiensis Rasnitsyn,Basibuyuk & Quicke, sp. n. by present designa-tion and monotypy.
Diagnosis. – As in family.
Description. – Very small, stout insect with com-paratively long and wide fore wings. Femaleantenna inserted at the level of upper 0.3 of eyeheight, with scape subclavate, pedicel long, sub-conical, flagellomeres 1-3 long cylindrical, (onlyflagellomeres 1-3 and the base of the 4th are pre-served). Head capsule almost twice as high as longin side view, somewhat protruding above antennalbases, widest near upper margin, with large com-pound eye, with ommatidia convex and large (with10-12 across shortest eye diameter), temple mod-erately narrow. Nothing apparent of usual lines/sutures on mesonotum except for a straight trans-scutal suture delimiting a nearly rectangularscutellum with deep lateral foveae, and small, non-protruding axillae. Metanotum wide and ribbon-like. Propodeum short. Fore femur as long asheigth of head, slightly narrowing toward apex,with both upper and lower contours weakly con-vex. Fore tibia thin. Fore wing only slightly short-er than body, twice as long as wide, with fore mar-gin weakly concave for basal 0.7 and hind marginconvex. Vein R straight before RS base, shortbetween it and pre-pterostigmal break. Distal
abscissa of RS 0.5 times as long as R before break,receiving 2rrs near its one third, 2rrs almostapproaching pre-pterostigmal break. With twosubparallel, sharp, concave folds posterior to and,less distinctly, anterior to vein Cu. Posteromedialand posterobasal wing areas with several addition-al longitudinal veins of more or less questionableidentity: centrally there is possibly a short distalsection of M, basally, possibly a long part of veinA, and subbasally a vein or perhaps a short convexfold of obscure homology. Outer third of wingwith approximately 6 fan-like diverging convexfolds. Metasoma much longer than head andmesosoma combined, as preserved, much higherthan thorax (possibly distended due to postmortemdecomposition), highest subbasally, rounded bas-ally, acuminate apically. First descernable tergitewith a transverse row of pits near base dorsally.Ovipositor occupying all visible sternal part ofmetasoma but not extending beyond last sternum,with sheath probably short.
Etymology. – From Khutel-Khara-Ula Mts. and Chalcis F.
Gender. – Feminine.
Composition. – Type species only.
Khutelchalcis gobiensis Rasnitsyn, Basibuyuk& Quicke, sp. n.
Type material. – Holotype female, PIN no. 3965/ 429,Palaeontological Institute, Russian Academy ofSciences, Moscow, Russia.
INSECT SYST. EVOL. 35:2 (2004) A basal chalcidoid 7
Fig. 6. SEM photo of counterpart head and thorax and its morphological interpretation.]
Locality and horizon. – Khutel-Khara-Ula (knownalso as Khara-Khutul-Ula) Mts. 70 km SW of SainShand, East Gobi Aymag, Mongolia. LowermostCretaceous, low in Tsagan Tsab Formation.
Description of holotype. – Female. Head and tho-racic integument reticulate, somewhat scaly in lat-eral pronotum and anterior mesonotum, less regu-lar (more rugose) further backward in preservedthoracic surface (Figs. 3, 6). Metasoma smooth aspreserved (Figs. 2, 3). No colour pattern pre-served. Proportions of scape, pedicel, anellus andflagellomeres 1-3 as 1: 0.45: 0.1: 0.75: 0.65: 0.6(Figs. 4, 5). Scape ca. 0.4 times as long as heightof head, 3x as long as wide, pedicel narrowed to-wards base, almost twice as long as wide, ca. 0.7xscape width, flagellomeres 1-4 about half as wideas pedicel. Apparently first tergum comprisingabout 0.4x metasomal length and much more thanhalf its height, following two discernible terga ofsubequal length followed by one long or, rather,two (or more) shorter terga (Figs. 1-3). Three lastvisible sterna comprising some three quarter ofvisible sternal part of metasoma, apical sternumwith lower contour concave in posterior two third.Ovipositor sheath narrowly rounded apically(Figs. 1, 2).
Measurements (mm): body length, as preserved,1.3, head height 0.3, fore femur length 0.3, fore-wing length 1.1, width 0.55-0.6 (different wings),
metasoma (as preserved, up to end of apical ter-gum) 0.75.
Etymology. – From Gobi Desert, the type locality.
Discussion
The fossil species described here is much olderthan any other known Chalcidoidea, whose oldestrecord otherwise is from the Upper Cretaceous(Yoshimoto, 1975; Rasnitsyn, 1980). This mayexplain the outstanding symplesiomorphies of thefamily. Indeed, if it was not for flagellomeres mostprobably bearing multiporous plate sensillae withlong aperture, posterodorsal pronotal emarginationand supposedly free, large prepectus, Khutelchal-cis would be easily taken as one more extinctgenus of Scelionidae, a common Cretaceous fami-ly known from near the Jurassic/Cretaceousboundary (in Baissa, East Siberia, in deposits sup-posedly of Berriassian age (Rasnitsyn, Jarzem-bowski and Ross, 1998), and in Khotont in CentralMongolia, Fig. 7, whose age can be even the latestJurassic (Lukashevich, 1996)). Yet it is putativelysynapomorphic with Chalcidoidea in the characterstates described above that together uniquely char-acterise Chalcidoidea (see the section Compari-son). The alternative hypothesis, that multiporousplate sensillae with long aperture constitute asynapomorphy for Khutelchalcididae and Cynipo-idea and thus gained homoplastically with Chal-cidoidea, cannot be definitely ruled out but seemsfar less likely because the only gall wasps knownto have a long inner aperture to the multiporousplate sensilla are members of the most advancedphytophagous family, Cynipidae. This would infermonophyly of Khutelchalcis directly with Cynip-idae well within Cynipoidea, as the only alterna-tive to its monophyly with Chalcidoidea. And yet,Khutelchalcis shows no cynipoid synapomorphyother than the very presence of multiporous platesensillae, and no synapomorphies of Cynipidaeitself.
Within the Chalcidoidea the new family may bethe sister group of all other chalcidoids includingMymaridae because of its comparatively short (forchalcidoids) multiporous plate sensillae and itsmore plesiomorphic wing venation, particularly inits more complete and almost uniformly thick setof veins.
Discovery of the Khutelchalcididae also poten-tially sheds some light on the relationship of Chal-cidoidea to the other proctotrupomorph families,
8 Rasnitsyn, A. P. et al. INSECT SYST. EVOL. 35.2 (2004)
Fig. 7. Undescribed representative of Scelionidae, PIN4703/10, body length 1.9 mm, Knotont in Ara KhangaiAimag, Mongolia, lowermost Cretaceous. C – costalvein.
which is still an area of considerable debate(Whitfield 1992; Dowton & Austin 1994; Dowtonet al. 1997; Basibuyuk & Quicke 1997, 1999; Ras-nitsyn 1988, 2000; Gibson 1999, Gibson et al.1999; Ronquist et al. 1999; Early et al. 2001). Awealth of cladograms have been produced usingdifferent methods (intuitive vs. parsimonous) andmaterial (molecular, morphological and ethologi-cal characters, either separately or in combina-tion). These studies show the Chalcidoidea (usual-ly together with Mymarommatidae, when the lat-ter is considered separately or at all) variously asforming a monophyletic group with the Platy-gastroidea (Whitfield 1992; Dowton & Austin1994, 2001: figs 3-5, 7-8, 11, 12; Dowton et al.1997: fig. 2; Rasnitsyn 1988, 2000), or with Platy-gastroidea + Ceraphronoidea sensu stricto (Ron-
quist et al. 1999), or with Platygastroidea, Diapri-idae and Ceraphronoidea (Basibuyuk & Quicke1997), or with Cynipoidea and Ceraphronoidea(Basibuyuk & Quicke 1999), or with Cynipoidea(Dowton & Austin 2001: figs 6, 9, 10, 13), or withCynipoidea, Platygastroidea, Diapriidae, Maamin-gidae, and Ceraphronoidea (Basibuyuk et al.2000), or with Diapriidae + Monomachidae +Maamingidae (Dowton & Austin 2001: fig. 2)!Controversy also surrounds the relationships of thePlatygastroidea which appear sometimes as thesister group of the Chalcidoidea, or with Chalci-doidea+Ceraphronoidea and/or with Cynipoidea,Diapriidae and Maamingidae), with Cynipoidea(Dowton & Austin 2001: fig. 2), Ceraphronoidea(id., fig. 9), or Pelecinidae (id., fig. 10).
Before proceeding further we should considertwo extinct families omitted from the majority ofpublications cited above, viz. the Serphitidae andJurapriidae. The Serphitidae was established basedon the Upper Cretaceous Serphites paradoxusBrues (Fig. 8) which was originally comparedwith Helorus Latr., Acanthoserphus Dodd, Van-horniidae and Roproniidae (Brues in Carpenter etal. 1937). Subsequently, Kozlov and Rasnitsyn(1979) added more Lower Cretaceous taxa andlumped Serphitidae with Mymarommatidae, basedon their two-segmented petioles and the presenceof apparently intermediate fossils, and they con-sidered this enlarged Serphitidae to be related tothe Scelionidae. Rasnitsyn (1980) further pro-posed (erroneously so: cf. Gibson, 1986) Serphit-idae sensu lato and Scelionidae to be monophylet-ic with Mymaridae which was thus removed fromChalcidoidea, though later Rasnitsyn (1988)agreed with Gibson (1986) and replaced Mymar-idae in its former superfamily and considered theMymarommatidae as a family of obscure relation-ship. Mymarommatidae, as well as Scelionidae +Platygastridae, have subsequently been elevated tosuperfamily status (Gibson 1993, and Masner1993, respectively). However, the position ofSerphitidae sensu stricto relative to the new super-families has remained unclear, though morerecently Rasnitsyn (2000) considered Serphitidaeand Mymarommatidae to form a monophyleticgroup, and these collectively forming the sistergroup of the Chalcidoidea. In the mean time, theSerphitidae have been discovered from moreancient, Lower Cretaceous deposits of Spain(Martínez-Delclòs et al. 1998) which is consistentwith the above. Equally Serphitidae from the
INSECT SYST. EVOL. 35:2 (2004) A basal chalcidoid 9
Fig. 8. Serphites paradoxus Brues, female, UpperCretaceous (Campanian) of Medicine Hat (Alberta,Canada). Courtesy Lubomir Masner.
Burmese amber (Rasnitsyn & Ross 2000) is nowfound to come from the Lower and not UpperCretaceous (Cruickshank & Ko 2003).
The monotypic, Upper Jurassic family Jurapri-idae (Fig. 9) was described (Rasnitsyn 1985) asthe stem group of Diaprioidea (established by Ras-nitsyn 1980) but this arrangement was later aban-doned (Rasnitsyn 1988), and it has subsequentlybeen suggested to be a stem group for a lineageincluding only the Chalcidoidea, Serphitidae sensustricto, and what now constitute the superfamiliesMymarommatoidea and Platygastroidea (Rasnit-syn 1988). The main reasons for this proposalwere the putative synapomorphies of the genicu-late antenna, with moderately elongate scape, andthe hind wing with RS leaving R far basad of thehamuli.
Returning to the phylogenetic significance ofKhutelchalcis, the first observation to be men-tioned is that it shows no obvious similarity toCeraphronoidea given the available characters. In-stead, the two taxa demonstrate contrasting vena-tional synapomorphies. Characteristic of Khutel-
chalcis are the chalcidoid synapomorphy in highantennal attachment on the head and putative earlystages of changes that culminated in the wellknown chalcidoid synapomorphies, viz. elongateflagellar sensillae (probably multiporous) and forewing with pterostigma lost and vein C much weak-ened (while R is only incipiently incrassated,costal space still wide, discal veins only moderate-ly reduced, and 2r-rs and, partially, RS preservedand retain their angular connection). In contrast,Ceraphronoidea show a plesiomorphic antennalattachment position (adjacent to the clypeus) andan alternative direction in the changes in venation.In particular, the least derived members of Mega-spilidae and the extinct Stigmaphronidae have lostthe costal space, vein C being appressed to, orfused with, R which is hypertrophied and thick,the pterostigma is also hypertrophied, 2r-rs isalined with the distal portion of RS, and all veinsbehind R+RS are lost, at least as tubular veins.
Similarly, Khutelchalcis shows no specific puta-tive synapomorphies with other suggested chalci-doid sister groups, viz. Cynipoidea, Diapriidae,Monomachidae and Maamingidae, except for thecynipoid flagellomeres bearing multiporous platesensilla which differ in having short apertures intheir groundplan. We have failed to identify anyother specific similarity between Khutelchalcisand Cynipoidea, unlike those between Khutel-chalcis and Chalcidoidea. There are a few wellknown putative synapomorphies of Chalcidoideapresent in Khutelchalcis and not in Cynipoidea,e.g. enlarged scape and greater reduction of discalveins (incomplete basal vein, lost RS+M and atleast the greater distal part of vein M).
A new putative synapomorphy of Khutelchalcis,Jurapria, Serphitidae and Chalcidoidea is of par-ticular interest, viz the angulation (or conspicuousbend in the case of Jurapria) of vein Cu at its forkwith vein M. This contrasts with the straight veinCu which is characteristic of the majority of Proc-totrupomorpha (except some Ropronia Pro-vancher, Monomachus Klug, Austroserphus Dodd,which probably represent reversals). A straightvein Cu has been proposed as a synapomorphy forthe Proctotrupomorpha by Rasnitsyn (1980, 1988)who overlooked the other exceptions apart thanRopronia. An angular or conspicuously bent veinCu appears in some more or less advanced groups(Ropronia Provancher, Jurapria, Serphites, Khu-telchalcis and in various Chalcidoidea, see below)but not in the putative proctotrupomorph stem
10 Rasnitsyn, A. P. et al. INSECT SYST. EVOL. 35.2 (2004)
Fig. 9. Jurapria sibirica Rasnitsyn, holotype female,body length 2.4 mm, Upper Jurassic of Uda(Transbaikalia, Russia).
family, Mesoserphidae, which is diverse enoughthat we might expect it to display a wide range ofvariation (Rasnitsyn 1980, 1985, 1986, 1990,1994; Darling & Sharkey 1990; Rasnitsyn et al.1998; Rasnitsyn & Martínez-Delclòs 2000; Zhang& Zhang 2000). In Ropronia, Monomachus, andAustroserphus the angulation occurs only in casesin which crossvein cu-a is shifted distad of theM&Cu fork, as is confirmed by examples whencrossvein cu-a is interstitial rather than postfurcal(cf. Ropronia pedunculata Provancher vs. otherspecies of Ropronia; Monomachus figured byNaumann & Masner 1985, vs. other figures of thegenus, e.g. in Goulet and Huber 1993, fig. 205;and Austroserphus vs. Proctotrupes Latreille). Adistal position of cu-a is unknown in theMesoserphidae, therefore the above cases of angu-lar vein Cu can reasonably be ascribed to inde-pendent distad shifts of cu-a.
In contrast, Jurapria, Serphites, and Khutelchal-cis lack cu-a, and this could therefore indicate aseparate origin of their bent or angular vein Cu andthat it could constitute a synapomorphy for them,the more so since some Chalcidoidea have a spec-tral rudiment of vein Cu also with a distinct,though less conspicuous bend, which is compara-ble to the condition seen in one of the wings of K.gobiensis (the more caudad in Figs. 1-3). This wasillustrated by Ferrièrre & Kerrich (1958: figs. 63,64, 71, I-IV) for Cleonymidae, Chalcididae,Eucharidae and Perilampidae, by Askew (1968:figs. 28-31, 50-51) for Eulophidae, by Bouc̆ek &Rasplus (1991: figs. 3, 79, 253, 285 and many oth-ers) for Pteromalidae, and also was observed byAPR for Torymus spp., Podagrion pachymerusWalker, Eucharis adscendens F., Stilbula sp.,Encyrtus sp., Norbanus sp., Entedontinae gen.indet. in the collection of the Natural HistoryMuseum, London.
Gibson (1986) inferred Serphitidae to be sym-plesiomorphic with respect to the Chalcidoidea +Mymarommatidae, in contrast to the Scelionidae(and by inference also to the Platygastridae) inhaving digitiform cerci, a conspicuously devel-oped posterior metasomal spiracle, and in the api-cal metasomal sternum being rather freely mov-able relative to the tergum. Jurapriidae can beadded to Chalcidoidea + Mymarommatidae atleast in respect of the apical sternal character.
Gibson (in review of this manuscript) has indi-cated a synapomorphy which is incompatible withour taxonomic hypothesis, viz. the loss of vein C
INSECT SYST. EVOL. 35:2 (2004) A basal chalcidoid 11
Fig 10. Putative cladogram of the chalcidoid +mymarommatoid + platygastroid clade: Kh – Khutel-chalcididae, Ch – remaining Chalcidoidea, Se – Ser-phitidae, Mt – Mymarommatidae, Ja – Jurapriidae, Pa –Platygastroidea. Putative per node synapomorphies or(for nod 1 only) groundplan character states:
11. Antenna with 15 segments at least in female sex,strongly geniculate, subclavate in female, with scapemoderately long; forewing with RS+M and all cross-veins other than 2rrs and incomplete 1mcu lost astubular veins; hindwing with RS (possibly also Aand cua) lost as tubular veins, rm subvertical, meet-ing R far basad of distal hamuli; female metasomaloose apically; ovipositor internal; possibly(unknown for Jurapriidae): male genitalia tubular,with cuspides lost.
12. Forewing with Cu bent at fork with M.13. Antenna with 13- or 14 segments (depending on
whether 14-segmented antenna in a few advancedChalcidoidea is groundplan or reversal for the clade),with scape long; 1st metasomal segment modifiedinto petiole (it is unknown, however, which of thetwo states characteristic of the subclades is plesio-morphic: either tube-like petiole, as in clade 7, orring-like one which is probably groundplan state ofclade 4).
14. Antenna attached high above clypeus, with multipo-rous plate sensilla; prepectus possibly external (ten-tatively identified in Khutelchalcididae, internal inRotoitidae); possibly other non-venational chalci-doid synapomorphies listed by Rasnitsyn (1988:node 66) (unknown for Khutelchalcididae),.
15. No family level synapomorphies found.16. All tubular veins lost except R and, in fore wing
only, 2r-rs and basal vein, forewing R and 2r-rsincrassate.
17. Metasomal segments 1 and 2 thin tube-like.18. Antenna with 10 segments in female, 9 in male;
pterostigma very large, 2r-rs very short.19. Numerous synapomorphies as scrutinised by Gibson
(1986).10. No family level synapomorphies found.11. Antenna 14-segmented; pronotum short medially,
rigidly connected with mesonotum; mesofurcal-mesotrochanteral muscle modified into pleuro-trochanteral; hind wing narrow subbasally, withmaximum width beyond vannus; metasomal spira-cles lost; apical metasomal tergum and sternum co-adapted, only slightly movable one against another;ovipositor internal, modified as described by Austinand Field (1997).
in Chalcidoidea, Mymarommatoidea, and Platy-gastroidea, as opposed to Khutelchalcis and Ser-phitidae which retain it. However, this inferencedoes not hold true when applied to fossils, forsome ancient Scelionidae preserved this vein aswell (Fig. 7). This implies that vein C is not lost inthe ground plan of the Platygastroidea or in that ofthe above clade embracing it. Loss of vein C iswidely accepted to have occurred independently invarious hymenopterans, particularly those ofminute body size or those that have supposedlyoriginated from minute ancestors, like Cynipoidea(except Archaeocynipidae), many Stephanidae,Carminator Shaw in Megalyridae, Loboscelidi-inae in Chrysididae, and so on. The cases ofChalcidoidea other than Khutelchalcis, Mymar-ommatidae, Platygastridae and advanced Scelion-idae represent apparently a simple addition to thatlist.
Gibson (id.) draws attention to the similarity be-tween the structure of metasoma of Khutelchalcisand Scelionidae in that the terga and sterna beingsubequal height (joined midheight) with the apicalones, equally extending backward and not veryeasily movable in respect each other. The similari-ty is evident but most probably plesiomorphic,based on the similar metasomal structure seen inJurapriidae and Serphitidae.
Relying particularly on the characters availablefrom the fossils, we hypothesise the following re-lationships: Platygastroidea, (Jurapriidae, ((Ser-phitidae, Mymarommatidae), Chalcidoidea) (Fig.10). Of these, the sistergroup position of Platygast-roidea is the least well supported of our inferences.The phylogenetic hypothesis presented here sug-gests that the Serphitidae and Mymarommatidaeshould be combined in a single superfamily, thevalid name for which is Serphitoidea Brues, 1937(= Mymarommatoidea Debauche, 1948, syn.nov.). As regards the Jurapriidae, we prefer to treatit as a family incertae sedis associated temporari-ly with the Serphitoidea until this enigmatic fami-ly is better known.
Acknowledgements
This work was supported in part by a Leverhulme Trustresearch grant to DLJQ and Mike Fitton (London)which, among other things, sponsored visits by APR tothe UK and a British Council short visit/travel grant toHHB and DLJQ. David Orme assisted with montagingSEM images of the new taxon.
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Revised manuscript accepted January 2004
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