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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 The occurrence and phylogenetic implications of the ovipositor clip within the Figitidae (Insecta: Hymenoptera: Cynipoidea) Matthew L. Buffington a a Department of Entomology, University of California, Riverside, CA

Online Publication Date: 01 January 2007 To cite this Article: Buffington, Matthew L. (2007) 'The occurrence and phylogenetic implications of the ovipositor clip within the Figitidae (Insecta: Hymenoptera: Cynipoidea)', Journal of Natural History, 41:33, 2267 - 2282 To link to this article: DOI: 10.1080/00222930701579732 URL: http://dx.doi.org/10.1080/00222930701579732

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Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 ora fNtrlHistory Natural of Journal orsodne ate .Bfigo,Sseai noooyLb SA / mtsna MH 0h&Constitution 10.1080/00222930701579732 online & DOI: 1464-5262 10th print/ISSN NMNH, [email protected] 0022-2933 Smithsonian Email: ISSN c/o USA. 20013, USDA, DC Lab, Washington, Entomology NW, Systematic Ave Buffington, L. Matthew Correspondence: 180 Some basal 1999). a by Ronquist characterized 1989; within further Nordlander plane, are vertical and a Cynipidae (Ronquist along and ovipositor, metasoma Figitidae the flattened of base laterally the a around al. coiled are et valves 2003). Quicke ovipositor Quicke 1988; ovipositor and Fergusson on Heraty the ; 1995; focused understand 1981 Ronquist have better Browning 1999; Parasitic to are studies 1994, and order 1988). several that 1992, Austin In Hymenoptera, 1999). ovipositor 1961; Bolton al. of (Scudder the and et forms morphology (Quicke of (Gauld parasitic attacked adaptations of host order of morphological evolution type the of the of to variety linked success a factors as display evolutionary cited been Hymenoptera the have Hymenoptera to within ovipositor the leading of function and structure The Introduction prevent of Keywords: dipterous evolution semi-concealed and competing of The two oviposit colonization Eucoilinae. to successful quickly and the on Eucoilinae. Figitinae for to clip by responsible in ovipositor order larvae probably parallel the is in in clip of clip evolved ovipositor presence/absence the structure the the concealed this of and lack suggests fully Mapping Anacharitinae yet the phylogenies host. attack subfamilies, hosts the that two with that exposed Figitids here fighting clip. hypothesized fully attack is the that attack It clip. possess Eucoilinae Aspicerinae, ovipositor and to the Figitinae is lacked found All all structure were hosts oviposition. morphological hosts during This host Figitidae. dipterous parasitoid throughout semi-concealed surveyed the is restrain clip to ovipositor postulated the of presence The Abstract USA ( CA, Riverside, California, of University Entomology, of Department BUFFINGTON L. MATTHEW Cynipoidea) Hymenoptera: Figitidae (Insecta: the the within of clip implications ovipositor phylogenetic and occurrence The h eaoa(oqit19) h Figitidae The from exerted 1999). when position (Ronquist ventral a valves paired metasoma fused the the in and results the position Charipinae) dorsal and a Anacharitinae assuming figitid valves in (absent condition This valves. cetd1 uy2007) July 10 Accepted h yiodapsesauiu vpstrmrhlg nta h is n second and first the that in morphology ovipositor unique a possess Cynipoidea The onbot opooy parasitoid. morphology, Koinobiont, 07 13–6:2267–2282 41(33–36): 2007; , # 07Tyo Francis & Taylor 2007 sensu oqit(99 r further are (1999) Ronquist u ws fteovipositor the of twist Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 av.I olwu ae,vnLnee ta.(99 lbrtdo h tutr and structure the on elaborated host (1999) potential al. of from et escape learned by Lenteren possessed the females the clip van prevent ovipositor older preventing the paper, to that of was follow-up structure function and a this means host, use In physical would-be successfully larva. some the to that from 1998). how al. ovipositor hypothesized experience et colony the Lenteren eventually paper fly (van was of that herself ovipositor the removal cleaning the of through wasp ordeal, the larvae, authors this of from escaping The hours perish several the not by behind did followed wasp wasp removed the female If result the ( The medium. 1998). of growth larva al. host dragging et Lenteren the the van if 1974; was al. oviposition et This of (Samson-Bosthuizen parasitization 1999). stages escape early 1998, the eucoiline al. the in et of ovipositor behaviour Lenteren oviposition (van the on clip heterotoma studies ovipositor after described the was Figitidae, feature 1999). some (Ronquist by valvula oviposition third possessed during the ovipositor of base the the of at flexion weakness of ventral point allows a that possessing members all by characterized n rnmsineeto irsoy(E)t eelacnaiyo h eta ufc of surface of ventral ovipositor the the on on concavity valve a reveal fused to the (TEM) microscopy electron transmission and Kelner-Pillout). and Carton, Qik 97 a etrne l 98 ucee l 1999). al. et Quicke restraint 1998; term host al. and for the first used et and the ovipositor Lenteren 2) was hymenopteran Figure van this a 1998, known, of 1997; as al. structure al. (Quicke far physical et et So a Lenteren structure. of Lenteren the van description van explain only (cf. to (cf. clip coined teeth was this clip’’ of in ‘‘ovipositor pinched row was a cuticle with ovipositing host of interfaced Freezing end 1). free Figure 1998, distal whose lobe flexible 2268 h noooyRsac uem CRvrie(iesd,C) ld specimens slide CA); are in or (Riverside, specimens TX) Riverside voucher Station, UC (College All Collection Museum, 1. Aspicerinae). Insect Appendix Research University (e.g. in A&M Entomology is clades Texas more the examined the sampled speciose species were in Eucoilinae) all less deposited (e.g. per of rich species this than species list (one in particularly genera thoroughly A sampled are 40 were subfamilies. that of clades (2007) total seven Additionally, a al. in for et Parnipinae, classified Buffington and genus) by Pycnostigminae recovered for Figitidae except study of clades major All terminology (1998). sampling other al. Taxon et (1989); Lenteren van Nordlander and (2002) and al. Ronquist et Fontal-Cazalla follows follows terminology Ovipositor family. methods the across and of across Materials clip understanding preference host the better of a of and for clip presence allowing the ovipositor map the the Figitidae, to of the used examine evolution is across the broader (forthcoming) to clip both al. much necessary et the a Buffington of was in Therefore, presence/absence Figitidae fauna taxa. of phylogeny eucoiline figitid The and Figitidae. the figitine of several on survey structure notch-like a heterotoma h ou fti ae so nte nqemrhlgclfaueo h ovipositor the of feature morphological unique another on is paper this of focus The a etrne l 19)ue ih irsoy cnigeeto irsoy(SEM), microscopy electron scanning microscopy, light used (1998) al. et Lenteren Van ro oti ok h nykonfgtdt oss h vpstrci is clip ovipositor the possess to figitid known only the work, this to Prior .L Buffington L. M. lhuhFrusn(98 n oqit(95 ohmnintepeec of presence the mention both (1995) Ronquist and (1988) Fergusson although , Tosn eeldta eti av eaewsswr nbet eoetheir remove to unable were wasps female naive certain that revealed (Thomson) etpln heterotoma Leptopilina .heterotoma L. etpln heterotoma Leptopilina olwdb E,rvae htthe that revealed SEM, by followed , h ocvt a oee ya by covered was concavity The . Drosophila and .boulardi L. p)atmtdto attempted sp.) Leptopilina Leptopilina (Barbotin, Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 nw o uolneadFgtne nyrcrsi hc ulfe systematist expanded be qualified would list a are The included. record, records. which were the host records in erroneous co-authored secondary of hosts or if records possibility parasitoids, considerably semi-concealed the reared Only eliminate the of to of Figitinae. records shown identification and the host in Eucoilinae unpublished participated and for published known summarizes I Table records Host nos (voucher Riverside UC Museum, Research Entomology 56878–56906). the in deposited are examined apical Ser, slot, S, lobe; L, valvulae; second 2vlv, valvulae; first 1vlv, 10 (A). bar: in area Scale boxed serration. of of clip enlargement ovipositor (B) the of side; micrographs Electron 1. Figure m m. curneadpyoeei mlctoso h vpstrclip ovipositor the of implications phylogenetic and Occurrence Neralsia Fgtne.()Oeve foioio i,lateral tip, ovipositor of Overview (A) (Figitinae). 2269 Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 oto a civduigAt-otg .50(yotc t,U) mgswere Images UK). Ltd, (Synoptics 5.0 v. Auto-Montage using achieved was control 2270 atrdb V Y7 -C iia aeafte oteAiso- i 2/3 a via Axioskop-2 the to fitted 1360 is camera camera digital JVC the 3-CCD of resolution KY-70 The c-mount. JVC reduction a by Hoyer’s- of captured Coverslips index. Glyptol optical with mounting superior sealed latter its mounting were The to Hoyer’s was specimens 56907–56933). due or mounted ovipositor nos preferred TAMU) The ultimately voucher at ethanol. was UCR; prepared medium 95% at containing (specimens containing prepared glass euparal (specimens glass watch either medium watch a to second to transferred a transferred was then to ovipositor then entire ethanol, the 80% exposed, the from Once separated vices. was specimen pin each of ethanol metasoma or The water pin. mesosoma with insect hydrated their were from dissection removed for and candidates as identified specimens Dried bar: Scale teeth. microscopy T, preparation—light sensillae; Se, Specimen lip; lateral LL, lobe; L, valvulae; second 2vlv, (A). in 10 area of clip boxed ovipositor of the enlargment of micrographs Electron 2. Figure mgswr atrdtruhaZisAiso- sn I ihig mgswere Images lighting. DIC using Axioskop-2 Zeiss a through captured were Images m m. .L Buffington L. M. + ed n isce n8%ehnli ac ls sn netpn itdto fitted pins insect using glass watch a in ethanol 80% in dissected and head, Trybliographa H . Eciia) A vriw eta ie (B) side; ventral Overview, (A) (Eucoilinae). 6 04pxl.Iaecapture Image pixels. 1024 6 Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 Eucoilinae Borgmeier Eucoilinae iii ao otHs ih Reference niche Host Host Figitinae hosts. dipteran semi-concealed attack that taxon clips Figitid ovipositor possessing Figitidae of Summary I. Table Triplasta Trichoplasta heptoma Rhoptromeris strobigena Rhoptromeris dnoeaanastrephae Odontosema odadel semirufa Nordlandiella anastrephae Lopheucoila fimbriata Leptopilina heterotoma Leptopilina Kleidotoma liooajaponica Kleidotoma Hexacola Ganaspis tritoma Eutrias etei euxestae Dettmeria ‘‘ keilini Eucoila Dicerataspis Cothonaspis Aganaspis Xyalophora Figites iie anthomyiarum Figites easahyalinipennis Neralsia Eucoila (Hartig) Grijpma and Nordlander Borgmeier (Kieffer) (Rohwer) (Kieffer) (Thompson) Huzimatu (Bouche Ashmead p.Anthomyiidae, spp. ’sp nhmide Muscidae, Anthomyiidae, spp. ’’ p ucdeRtigvgtto Dı vegetation Rotting Muscidae sp. p.Msia,DoohldeRtigvgtto hvue(97,Dı (1977), Shivpuje vegetation Rotting Drosophilidae Muscidae, spp. p.Ehdia,ClrpdeAgebd erse 18) Simmonds (1989), Beardsley beds Algae Chloropidae Ephydridae, spp. ´) p.TprtdeFut(ngon)Watne l 18) io and Jiron (1981), al. et Wharton ground) (on Fruit Tephritidae spp. p.Atoyia,Sepsidae, Anthomyiidae, spp. p.Msia ugcrinBike(91,Tre tal. et Turner (1961), Blickle Dung/carrion Muscidae spp. p esdeDn/aro ige l 18) Nordlander (1982), al. et Figg Dung/carrion Sepsidae sp. p rspiia otn eeainWatne l (1998) al. et Wharton vegetation Rotting Drosophilidae sp. pMsia,Drosophilidae, Muscidae, sp ife nhmideRtigvgtto elnadBuePuie (1913) Pluvinel Baume and Keilin vegetation Rotting Anthomyiidae Kieffer hmo esde acpaia ugcrinBue(1986) Blume Dung/carrion Sarcophagidae Sepsidae, Thomson curneadpyoeei mlctoso h vpstrclip ovipositor the of implications phylogenetic and Occurrence hooia oiesNrlne n rjm (1991) Grijpma and Nordlander plant grasses, Cereal Conifers Chloropidae Chloropidae ehiia ri o rud hro ta.(91 1998) (1981, al. et Wharton ground) (on Fruit Tephritidae ocaia ri o rud hro ta.(1998) al. flower et Asteraceous Wharton ground) (on Fruit (1975) Boness vegetation Rotting references and (1980) Nordlander Agromyzidae vegetation Rotting Lonchaeidae Drosophilidae Drosophilidae pyrdeAgebd uias (1940) Huzimatsu beds Algae Ephydridae ttdeFut(ngon)Vlcne(1986) Valicente ground) (on Fruit Otitidae nhmideCrinJms(1928) James Carrion Anthomyiidae ucdeDn ob n olce 16), (1969) Hoelscher and Combs Dung Muscidae Lonchaeidae pyrde Canacidae Ephydridae, Sarcophagidae, Muscidae, Callophoridae Sepsidae, Sarcophagidae, Sarcophagidae Muscidae, Calliphoridae, heads otn vegetation, Rotting la es dung beds, Algae ugcrinTre ta.(98,Fg tal. et Figg (1968), al. et Turner Dung/carrion ugcrinTopo 15) Wharton (1955), Thompson Dung/carrion material fungi carrion ee 12) Nordlander (1923), Meyer odadr(1982) Nordlander ufntn(2004a) Buffington enr(98,Fg ta.(1982), al. et Figg (1968), Depner a n alro(1996) Gallardo and ´az 17),Jnele l (1999) al. et Jonsell , (1978) hri,vnApe ta.1991 al. et Alphen van therein, lm 18) erse (1993) Beardsley (1986), Blume (1952) alro(1996) Gallardo odadr(1982) Nordlander (1986), Blume (1982), (1982) ezn(99,Orsi(1995) Ovruski (1989), Mexzon (1979) Wharton (1973), Wylie (1968), (1979) lm (1986) Blume (1972), Morgan and Thomas a and ´az 2271 Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 h vpstrci ( clip ovipositor The clip ovipositor the of Presence the of either in Results topology tree that affect noted not be did should analysis it analyses. the and Bayesian two from 163), or alternative character the (character parsimony examining this character between for clip of morphological ovipositor here topology a removal the of included as tree presence/absence included The are 2007). in was both al. et but difference (Buffington scenarios minor is The evolutionary was (2007) D2 characters). al. hypotheses 28S et figitid total competing Buffington of of (1855 of analyses phylogeny based) partitions total-evidence The clip. Bayesian on mapping ovipositor for one used based the were of (2007) based, al. presence/absence et parsimony Buffington the from relationships (one sub-group and hypotheses subfamily phylogenetic Two UC Museum, mapping Research Character Entomology the at deposited were deposited images 56877). All are no. beam. (voucher Phillips electron stubs A kV Riverside 10 interval. SEM a each at MorphBank. using cloud imaging ion on for the s used 30 to was relative three SEM stubs for the XL-30 Reference gold–palladium of covered angle with stubs the sputter-coated SEM Heraty changing and to intervals, (HMDS; mounted adhesive then mounting hexamethyldisilazane were double-sided Samples to dehydration. by final transferred for second were 1998) the Hawks Following samples and above. (95%), described as bath dissected ethanol were Specimens SEM. using niche examined Host species, Two micrcoscopy electron preparation—scanning Specimen image. (www.morphbank.com). focused composite MorphBank a on produce to deposited ‘‘montaged’’ are and images planes focal Host All of series a as captured a taxon Figitid Continued. I. Table 2272 Fgr )wt h xeto of exception the with 4) (Figure diinlfaueo h ardvle( l) h pclsrain(e,Fgr B,was 1B), feature. Figure this (Ser, possess serration to apical observed also the were vlv), eucoilines (1 all not valve and but paired 4) Some (Figure the for observed. figitine clip of the the feature description the additional between matched clip and similar, the entirely heterotoma was of clades 5) morphology (Figure eucoiline The 7). (Figure Eucoilinae hsseisi rbbynot probably is species This Trybliographa rbigah trichopsila Trybliographa (Hartig) .L Buffington L. M. a vnLnee ta.19) ihnteFigitinae the Within 1998). al. et Lenteren (van p.Atoyia eeain ug odadr(1982) Nordlander fungi Vegetation, Anthomyiidae spp. Neralsia sensu p Fgtne and (Figitinae) sp. alpoia,Muscidae, Calliphoridae, .trichopsila T. Sarcophagidae a etrne l 98 a on oocri l Figitinae all in occur to found was 1998) al. et Lenteren van Lonchidia u pce of species a but rbigah rapae Trybliographa n n prioy rto(aein ldsof clades (Bayesian) two or (parsimony) one and , ugcrinScesaa(1974) Sychevskaya Dung/carrion + Eucoila 3 O,1S n opooia data morphological and 18S, COI, D3, M osae esnlcommunication). personal Forshage, (M. etod(uolne,were (Eucoilinae), Westwood + sieia ld,an clade, Aspicerinae L. Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 ( Anacharis iue3 xmlso iiia htlc h vpstrci.(–)Aahrtne (A) Anacharitinae: (A–C) clip. ovipositor the lack that Figitidae of Examples 3. Figure sieia Fgr D ) hrpne(iue3) mriia Fgr 3G), (Figure Emargininae 3F), (Figure the Eucoilinae, Charipinae the and E), Among 3H), (Figure 3D, Thrasorinae (Figure Aspicerinae oooy h ldswoetx oss h vpstrci r eaae ynumerous by separated (Swofford are optimization DELTRAN clip or ovipositor ACCTRAN Figitinae, the either For In clip. possess trees. the clip taxa evidence lack ovipositor total that whose the 7B) clades (Figure of clades Bayesian bars) the and (black 7A) topology, (Figure presence parsimony the best mapping the of on result the summarizes the 7 Figure of portion cavity the mapping of Character depth the C–D). near 4A, are (Figure the E) Figitinae 3A–E, Aspicerinae, Figure in 3D, valve, (Figure Within clip (fused serrations ovipositor Thrasorinae. vlv 2 these and lack of that of Aspicerinae, tip groups the depth Anacharatinae, at figitid the serrations among these of including feature presence recovered H), the common (2007) is A al. clip B). et ovipositor 7A, the Buffington (Figure clip. sister-groups ovipositor as the groups lacked clades 6D–H) (Figure Thoreauella ebr fFgtdeta aka vpstrci nld ncaiia Fgr 3A–C), (Figure Anacharitinae include clip ovipositor an lack that Figitidae of Members D )Apcrne (D) Aspicerinae: E) (D, ; ;()Trsrne( Thrasorinae (H) ); curneadpyoeei mlctoso h vpstrclip ovipositor the of implications phylogenetic and Occurrence Euceroptres Melanips Gronotoma Lonchidia .2vv ue vpstrvalve. ovipositor fused vlv, 2 ). (E) ; cretycasfe sFgtne o shown). not Figitinae; as classified (currently Callaspidea ru Fgr AC and 6A–C) (Figure group F hrpne( Charipinae (F) ; Alloxysta Aegilips Zaeucoila ;()Emargininae (G) ); (B) , Xyalaspis group 2273 (C) , Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 ru ote(oeEciia ( Eucoilinae (core the to group ufmle n eoeigteeciie( figitid all eucoiline to the sister-group as recovering Parnipinae the and recovering subfamilies including reasons, several for analysis and Eucoilinae core in gain (parallel rfrnefratcigsm-ocae a eie bv)dpeoslra.Evidence larvae. dipterous above) defined (as semi-concealed attacking foliovores. external for type of this preference predators of and Examples niche. foliovores its external within likely include refuge less niche seeking either by host-feeding is parasitization and An of escape predaceous) fruit. this to or rotting of able phytophagous and (either not Examples algae, externally or substrate. feeds roots, the that host dung, one into the carrion, is fleeing caveat, include host a by niche As parasitoid previously host-feeding parasitoid. the of the mines, from by type parasitoid escape detected the stem be readily if could and can but host olfaction), larva the leaf by substrate, (e.g. host galls, parasitoid the a enters include by space. detection niche niche obvious prevents the feeding substrate from A of away seeds. and moving type insects, freely parasitized from this host of the prevents Examples that encasement of a to form refers discussion following The parsimony, Discussion steps (two tree presented Bayesian phylogenies the competing over two preferred the again Bayesian). on is steps clip three tree ovipositor parsimony the the map here, to required steps of at gain in (single ACCTRAN loss both secondary in with steps two node showing Eucoilinae differently, tree Bayesian eucoiline Zamischus h vpstrci asa igegi ttebs f(oeEciia ( Eucoilinae (core of tree, base parsimony the at the gain or in single parsimony Eucoilinae a using as Within maps maps clip clade). clip the ovipositor Figitinae of the gain, presence (single the how analyses affect Bayesian not does 1987) Maddison and valve. ovipositor fused vlv, 2 clip. (A) the Figitinae. of within extent present clip ovipositor the (D) of Examples 4. Figure 2274 ufntne l 20)fvue h asmn-ae eut ftettlevidence total the of results parsimony-based the favoured (2007) al. et Buffington h curneo h vpstrci ihnFgtdei togycreae iha with correlated strongly is Figitidae within clip ovipositor the of occurrence The Xyalophora .L Buffington L. M. ru)cae(CTA rDLRN.Teoioio lpmp nt the to on maps clip ovipositor The DELTRAN). or (ACCTRAN clade group) h iwn nl n()i osladntltrl h seikadlnsaematt eiet the delineate to meant are lines and asterisk the lateral; not and dorsal is (B) in angle viewing The . Kleidotoma semi-concealed ocae host concealed Kleidotoma Gronotoma + Gronotoma Zamischus oti n hs edn eaiu ihna within behaviour feeding whose one is host + sahs hs edn ih swti a within is niche feeding whose host a as Zamischus + ru)cae ae ntenumber the on Based clade. group) + Zaeucoila Zaeucoila Neralsia groups). (B) ; rus rDELTRAN or groups) ru)caea sister- as clade group) Amphithectus Kleidotoma (C) ; group Trischiza exposed + ; Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 Dieucoila mundata ot a eie bv)uieslylc h vpstrci.Teefgtd include figitids These clip. ovipositor the on lack inducers concealed gall attacking universally cynipid Figitids attack above) I. Table which defined in Parnipinae, summarized (as is preferences hosts host these supporting (A) Eucoilinae. within clip ovipositor the of Examples 5. Figure vpstrvalve. ovipositor ivsAde 01 n haoia hc takhmnpea alidcr on inducers gall hymenopteran attack which Thrasorinae and 2001) Nieves-Aldrey (D) ; ortr (H) Foerster; Nordlandiella Triplasta E ‘‘ (E) ; curneadpyoeei mlctoso h vpstrclip ovipositor the of implications phylogenetic and Occurrence (I) ; Eucoila Glauraspidia ’’ impatiens (J) ; Trybliographa a;(F) Say; Papaver dnoeaanastrephae Odontosema (K) ; gnsi daci Aganaspis Kleidotoma Ppvrca)(oqitand (Ronquist (Papaveraceae) (L) ; Wl) (B) (Weld); Aporeucoela ife;(G) Kieffer; Leptopilina l,fused vlv, 2 . Quercus Ganaspis 2275 (C) ; Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 orahtehs.Smlrmdfctoswr on nsm hlioda(eayand (Heraty al. et Chalcidoidea Quicke some tissues. plant in in oviposit found that were 1988) (Fergusson modifications Cynipoidea substrate (1999) a and Similar through 2003) al. host. sawing Quicke and and the et drilling denticles with of reach associated form Quicke the be the to to in host. that postulated came host, parasitoids were modifications the that these of concealed serrations cases, tips most encasing ovipositor a In with hosts. substrate the associated attacking concealed attack traits require the when morphological not various penetrate Furthermore, do the reviewed first hosts restraint. concealed must host escape on readily ovipositor specializing for cannot figitids hosts clip that Concealed ovipositor follows 1998). al. it et and and Lenteren parasitism, (van 1979) 1987). envenomation Burks Miller to and prior 1955; Cave 1979; (Thompson (Burks larvae larvae Neuroptera exposed Syrphidae attack exposed that Anacharitinae attack that clip. Aspicerinae ovipositor the lack also Fontal- 2004b; 2002), 2004a, al. 2002, et (Buffington plants Cazalla various on Agromyzidae leaf-mining attack Rnus 99.Wti uolne the Eucoilinae, Within 1999). (Ronquist iue6 xmlso uolneta akteoioio lp (A) clip. ovipositor the lack that (H) Eucoilinae of Gronotoma Examples 6. Figure 2276 p.(Fagaceae), spp. h vpstrci sascae ihhs etan uigteerysae foviposition of the stages early include the during These restraint host clip. with associated ovipositor is clip the ovipositor The lack also hosts exposed attack that Figitids Dicerataspis (C) ; .L Buffington L. M. l,fsdoioio valve. ovipositor fused vlv, 2 . Ealata (D) ; Mimosa Tropideucoila p.(ioaee,and (Mimosaceae), spp. (E) ; Rhabdeucoela Gronotoma (F) ; ohuol anastrephae Lopheucoila and Nothofagus aapdu nigrimanus Ganaspidium Zaeucoila p.(Nothofagaceae) spp. eu rus which groups, genus owr (G) Rohwer; Kefr;(B) (Kieffer); Dettmeria ; Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 hliodai pi ume Casn14;MneadEehi 91 swl as well as 1991) Evenhuis and Menke koinobiont 1940; of (Clausen mummies lineages aphid several in hosts. in exposed Chalcidoidea and common concealed were both attacked notches that sub-apical Ichneumonoidea endoparasitic noted also (1999) attacked: is ‘‘ host for the clip. clades which indicate ovipositor in bars type the Black niche unknown. possess to names ?, refer members inducer/inquilline; terminal tips gall to all tree G, next exposed; at which E, boxes Numbers semi-concealed; in analysis. (from S, Letters Figitidae Bayesian concealed; sampled. of C, (B) taxa relationships of analysis; for cladograms number Parsimony possible the (A) two indicate to forthcoming). on al. mapped clip et ovipositor Buffington the of Presence 7. Figure piiain ‘‘ optimization; adsn1987). Maddison hrpneseilz nhpraaiimo piohgu rcndeand Braconidae aphidophagous of hyperparasitism on specialize Charipinae + E’ niae rsneo hrce ae nDLRNotmzto Sofr and (Swofford optimization DELTRAN on based character of presence indicates DEL’’ curneadpyoeei mlctoso h vpstrclip ovipositor the of implications phylogenetic and Occurrence + C’ niae rsneo hrce ae nACCTRAN on based character of presence indicates ACC’’ 2277 Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 takn nuietfe goyi on agromyzid unidentified an attacking vpstraogteetr egho h ieutlahs av a eetd h same The of detected. was species larva unidentified host a an until for mine her with the observed it of probing was began length female behaviour entire the located, the was along mine the ovipositor Once antennae. the with surface observation). oe that noted on ovipositing 2278 l fteehssaesm-ocae nta hyd o pcfclycneltesle in themselves conceal specifically not do they that in and semi-concealed (Ephydridae) (Sarcophagidae), are beds carrion hosts and algae these Sepsidae) the from of (Muscidae, vary dung all within to hosts (Anthomyiidae) condition these plants for of plesiomorphic roots microhabitats the the Though is I. Table (e.g. clip attack ovipositor also the clip of ovipositor been Gronotoma have the lack they possessing since though eucoilines constraint 6F–H), (Figure other clip that Aganaspis ovipositor hosts the from lack reared taxa these Interestingly, re orahtehost. the reach to order Watne l 98.A nteohrfgtd htatc ocae ot,teovipositors the the hosts, concealed attack of that figitids members other 2004b; the the of some in members 2004a, As of 1998). though 2002, al. these et 2000, 2006), (Wharton of members (Buffington Ronquist of ( majority Agromyzidae and The 7). leaf-mining (Figure Buffington analyses parasitize Bayesian and clades parsimony both in Zaeucoila ieyt ecneldDpea(..oioio lplcig ohs etan needed). restraint host no lacking; clip ovipositor are (i.e. hosts that Diptera suggest concealed Unfortunately position be ). phylogenetic to are and E likely hosts 3G) 3D, Figure Though well- 1988, (Figure Aspicerinae. (Fergusson vlv possess morphology 2 for of and lacking do species Figitinae) currently for aspicerines unknown are in the for studies (as oviposition), clip observational vlv ovipositor oviposition (quick-strike 1 the lack on do may serrations they feeders Though developed anacharitines detritous I). 1940), typically (Table reason and soil are (Clausen the Chamaemyiids Eucoilinae same vegetation in and niche. or on on Figitinae host’s insects scavengers of the and personal hosts phytophagous in cyclorraphan of Buffington, found the semi-concealed predators be whereas the L. typically can and are Eucoilinae, M. hosts, and syrphids exposed 1999; Figitinae these of between (Ronquist hosts of difference larvae main mode The anacharitine Syrphidae observation). the and for Chamaemyiidae responsible probably are traits precocious both both oviposition. are and Miller, neuropterans hemerobiid G. these 1987; aggressive, of and Miller larvae and and instar chrysopid Cave later 1985; of The Lambdin and communication). parasitoids (Miller personal larvae quick-strike instar third are and both second are Anacharitinae hosts clip. these Ronquist ovipositor since system 1991; this in Evenhuis needed and not concealed. probably and (Menke is immobile clip Psylloidea ovipositor immobile An 1999). on parasitic Chalcidoidea Dettmeria h rcs vpsto eaiu of behaviour oviposition precise The ot o iiia ossigteoioio lp(iue ,5 r umrzdin summarized are 5) 4, (Figures clip ovipositor the possessing Figitidae for Hosts ebr fEciia htlc h vpstrci eogt the to belong clip ovipositor the lack that Eucoilinae of Members (including Aspicerinae the lack hosts exposed attack that Figitidae why explain to reasons other are There .L Buffington L. M. ruso eea ufntne l 20)rcvrdteecae ssister-groups as clades these recovered (2007) al. et Buffington genera. of groups , p. al ;Watne l 98.Ti a eepandb phylogenetic by explained be may This 1998). al. et Wharton I; Table spp., + G Lopheucoila Zaeucoila . Liriomyza nigrimanus Gronotoma ru ld teetx eogt the to belong taxa (these clade group Lonchidia aapdu nigrimanus Ganaspidium , p Armzde M ono,proa omncto) twas It communication). personal Johnson, (M. (Agromyzidae) sp. Melanips is oae h efmn teftruh‘dumn’ h leaf the ‘‘drumming’’ through itself mine leaf the located first Dicerataspis and Fgtne,Eagnne n ynsimne ovipositor Pycnostigminae, and Emargininae, (Figitinae), Zaeucoila ssgetdi ufntne l,20)atc exposed attack 2007) al., et Buffington in suggested as iue6–)aekonfo Tephritoidea from known are 6F–H) Figure , Dettmeria Celtis rusms is eert h efsraein surface leaf the penetrate first must groups p Umca)(.L ufntn personal Buffington, L. (M. (Ulmaceae) sp. , Kefr ( (Kieffer) Lopheucoila Agrostocynips Gronotoma and , Zaeucoila Dicerataspis ru)wsobserved was group) ru fgenera). of group ( Zaeucoila Zaeucoila Gronotoma sunknown. is group) group and Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 n tp,wt usqetls fteci nthe in the clip possess clip, the to of of origin Figitidae loss would (single core Eucoilinae subsequent Eucoilinae and and with all Figitinae Figitinae step), of between the between one clip ancestor requiring relationship the scenario, sister-group the of third a homology require A require the steps). would Forcing (two clip. (two loss, loss Eucoilinae ovipositor subsequent in of gain with parallel degree way gain a same single as highest the maps a in but as step), maps or (one clip Figitinae steps) ovipositor for the tree parsimony 7B), Figure the tree, as (Bayesian scenario alternate the eayfrteueo h mgn qimn eddt iueti ae.BbWharton, paper; this Bob of paper. reviews this final figure provided to Doug Forshage needed John Mattias and thank equipment and to (TAMU) imaging Vilhelmsen, of like Lars the Riley also both Heraty, of would Ed Riverside; John I use work. dissection. UC was the for at continued Work for specimens Pinto dearly. Heraty obtain my me help John guiding helped their and (UCR) in appreciate Heraty Yanega instrumental I John and were of 2000, guidance them of the at fall Woolley under the James continued and in Wharton after University Robert of system A&M direction character Texas the this under started in originally interest was research my on sparking work in seminal instrumental his was Lenteren van Joop five on accepted is clip Acknowledgements the of loss the suggest unless however, lineages trees, these Bayesian between lineages. and separate homologous parsimony not best is The clip clip. ovipositor the maximum possess allow the to since unmodified a necessary is within not ovipositor is the concealed restraint cases, fully of efficiency. these are method In ovipositional parasitized hosts a escape. readily leaf-mine), If readily or cannot itself. more gall host prevent defending a be to host (e.g. possible semi- may as substrate the quickly specific by attack other hosts as attacks inflicted some exposed parasitoid that injury with the fully cases, possible parasitoids subdued these contrast, In quickly with restraint. of not In host stages without if correlated venom). early parasitization is the (e.g. escape during one means could oviposition restrained been that that physically of suggest not hosts is would concealed type has host here types the This described which host system oviposition. in The various exist 1998). into should al. category behaviour et Lenteren ovipositing (van for proposed scheme classification A steps. 39 Conclusion additional an requires this tree, parsimony the On step). (excluding clade Figitinae parsimony) 7A, the (Figure to tree on preferred maps the clip On feature. ovipositor gall 1994). the the a of (Godfray in evolution this the host of do a interpretations pat; to cow able the A. be into host deeper to (R. the escape unlikely successfully can If is oviposit it observation). wasp, leaf-mine to the or personal crevices of Buffington, presence and the holes L. to detects through M. larva able pat is communication; wasp cow personal ovipositing the of Wharton, the of species however, centre pat, some the cow Arguably a enter parasitization. within escaping themselves ‘‘conceal’’ restrict Muscidae could that stuctures specialized h cr uolne( Eucoilinae (core the otEciia n l iiia htatc eicnelddpeoshssuniversally hosts dipterous semi-concealed attack that Figitinae all and Eucoilinae Most different two in results 7) (Figure clip ovipositor the of presence the of Mapping Leptopilina Kleidotoma curneadpyoeei mlctoso h vpstrclip ovipositor the of implications phylogenetic and Occurrence ehscniudt upr yrsac vrsne This since. ever research my support to continued has he ; group + Zamischus Gronotoma ru) uoln ld oese) In step). (one clade eucoiline group)) Lonchidia and Zaeucoila n tp swl as well as step) one ; rus(one groups 2279 Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 uk D 99 yioda n rmenK udPJ,SihD uk D dtr.Ctlgeof Catalogue editors. BD, Burks D, Smith Jr, on P Notes 1987. Hurd G. K, Miller Krombein R, Cave In: Cynipoidea. 1979. BD. Burks ufntnM oqitF 06 aiuo9:FmlaFgtde n Ferna In: Figitidae. Familia 97: Capitulo 2006. F. Ronquist Cynipoidea). (Hymenoptera: M, Figitidae of Buffington evolution and phylogeny The JM, Heraty JAA, Nylander M, Buffington of description The 2004b. M. Buffington ufntnM 02 ecito of description A 2002. M. Buffington on notes of taxonomic species and new International A genera ovipositors. 1993. to JW. insect key Beardsley along Cynipoidea), eggs (Hymenoptera: Eucoilidae of Hawaiian movement 1989. for JW. mechanism Beardsley A 1981. TO. Browning AD, Austin References grant Woolley. NSF-PEET J. and and Pinto Wharton J. and R. by Heraty to provided J. awarded was to 9712543 Funding awarded DEB references. 9978150 host BSR figitid grant critical NSF-PEET many provided Forshage Mattias 2280 enrK.16.Hmnpeosprstso h onfly, horn Northeast the in of fly parasites horn Hymenopterous with 1968. associated KR. found Depner parasitoids pupal Company. Hymenopterous Book 1969. McGraw-Hill CE. York: Hoelscher New RL, insects. Combs Entomophagous 1940. CP. Clausen of description The 2004a. M. Buffington ufntnM 00 h hlgn n lsiiaino the of classification and phylogeny The 2000. M. Buffington eayJ,Qik L.20.Pyoeei mlctoso vpstrsrcuei uhrtdeadPerilampidae and Eucharitidae in structure ovipositor Entomological of insects. implications Phylogenetic drying 2003. for DLJ. Quicke alternative JM, Heraty chemical Hexamethyldisilizane—a 1998. Press. Press. DH. University University Hawks Oxford Princeton Oxford: JM, NJ: Hymenoptera. Princeton, Heraty The Parasitoids. 1988. 1994. HCJ. editors, Godfray B, Bolton I, Gauld Ros-Farre J, Central Liljeblad in G, parasites insects Nordlander associated M, and Buffington flies FM, pasture Fontal-Cazalla of Bucoprophilous genitalia 1982. female GD. of Thomas importance RD, phylogenetic Hall of DE, structures Figg the of study comparative A 1988. NDM. Fergusson Dı lm R 96 aaie fDpeaascae ihbvn rpig napsuei atcnrlTexas. central east in pasture a Flu on von droppings Hochwassergenist im bovine Arthropoden with 1975. M. associated Boness Diptera of Parasites 1986. RR. Blume fly, face the of Parasites 1961. RL. Blickle a B alroF.19.SbecnpiesdlBai,prstie edı de parasitoides Brasil, del cinipoideos Sobre 1996. FE. Gallardo NB, ´az yeotr nAeianrho eio ahntn mtsna nttto rs.p1045–1108. p Press. Institution Smithsonian Washington: Mexico. of north America in Hymenoptera 829–838. p Colombia. de Nacional Universidad Hmnpea cnuoia)parasitizing Ichneumonidae) (Hymenoptera: Introduccio 23:1–29. Cladistics of species 106:192–198. Washington of of status iiia:Eciia)[hss.CleeSain ea & nvriy 8 p. 182 University. A&M Texas Station: College [thesis]. Eucoilinae) Figitidae: 29:165–194. Society Entomological Hawaiian the of Proceedings species. non-endemic apparently 10:93–108. Embryology and Morphology Insect of Journal issip.Junlo cnmcEtmlg 62:1234–1235. Entomology Economic of Journal Mississippi. 98:211–216. News Gronotoma Hmnpea hlioda.Junlo aua itr 37:1751–1764. History Natural of Journal Chalcidoidea). (Hymenoptera: Cladistics Figitidae). 109:369–374. News Cynipoidea: (Hymenoptera: Eucoilinae the of Phylogeny 2002. 18:154–199. F. Ronquist 55:476. Society Entomological Kansas the of Journal Missouri. 13:13–30. Entomology Systematic (Hymenoptera). Cynipoidea the 55:127–129. Argentina de Entomologica Societa Revista Cynipoidea). of larvae on parasitic aainEtmlgs 100:1057–1060. Entomologist Canadian otwsenEtmlgs 11:215–222. Entomologist Southwestern 54:802. noooia nttt 27:1–6. Institute Entomological .L Buffington L. M. Ganaspidium Agrostocynips nalsHmnpead aRegio la de Hymenoptera los a ´n Fo rtr rceig fteEtmlgclSceyo ahntn104:589–601. Washington of Society Entomological the of Proceedings ¨rster. ayr quadrituberculata Parydra ed(yeotr:Fgtde uolne.Poednso h noooia Society Entomological the of Proceedings Eucoilinae). Figitidae: (Hymenoptera: Weld iz(yeotr:Fgtde uolne.Zoaa408:1–11. Zootaxa Eucoilinae). Figitidae: (Hymenoptera: Diaz ncai melanoneura Anacharis Aegeseucoela Kleidotoma uc autumnalis Musca Preseucoela odadel semirufa Nordlandiella nNorpcl ooa oiddClmin eEntomologı de Colombiana Sociedad Bogota: Neotropical. ´n rmNrhAeia(yeotr:Cnpie:Eucoilidae), Cynipoidea: (Hymenoptera: America North from irmsposticus Micromus ufntn e ae ihtxnmcntso h ttsof status the on notes taxonomic with name, new Buffington, ow(itr:Ehdia) otiuin fteAmerican the of Contributions Ephydridae). (Diptera: Loew ufntn e eu,wt oe ntesau fNearctic of status the on notes with genus, new Buffington, nNwHmsie ora fEooi Entomology Economic of Journal Hampshire. New in , se.Bne olgsh Beitra Zoologische Bonner ¨ssen. Hmnpea iiia)and Figitidae) (Hymenoptera: amtbairritans Haemotobia Gronotoma Kefr,nwcmiain ihntso the on notes with combination, new (Kieffer), Nuotr:Hmrbia) Entomological Hemerobiidae). (Neuroptera: ,Nee-lryJ,Pjd-ilrJ, Pujade-Villar JL, Nieves-Aldrey P, ´ ru ( group peo seclrs(Hymenoptera, estercoleros ´pteros Dpea ucde,i Alberta. in Muscidae), (Diptera: ne ,SakyM,editors. MJ, Sharkey F, ´ndez s.l )o eea(Hymenoptera: genera of .) hrtpsmellicornis Charitopes g 26:383–401. ¨ge ay ´a Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 vuk M 95 ua n avlpplprstis(yeotr)otie from obtained (Hymenoptera) parasitoids larval-pupal and Pupal 1995. SM. Ovruski hmsG,Mra E 92 aaie ftehr l nMsor.Junlo cnmcEntomology Economic of Journal Missouri. in fly horn the of Parasites 1972. parsimony. CE. Wagner Morgan under GD, states Thomas of character biology ancestral The 1974. Reconstructing VI. 1987. Sychevskaya WP. Maddison Royal DL, shootfly the sorghum Swofford of of parasites natural Transactions new of Record ovipositor. 1977. PR. insect Shivpuje frit-fly the the of of Parasites 1952. morphology FJ. comparative Simmonds The of 1961. parasitization of GGE. Success Scudder 1974. cynipoid, K. archaic Bakker JC, an Lenteren Zoological of van Cynipoidea). morphology M, (Hymenoptera, Samson-Bosthuizen Skeletal Figitidae 1989. of G. subfamily Nordlander new F, 28:139–164. A Ronquist Scripta 2001. Zoologica Cynipoidea. Entomology JL. the Systematic Nieves-Aldrey of evolution (Hymenoptera). F, and Cynipoidea classification Ronquist the Phylogeny, 1999. of F. evolution Ronquist early and Phylogeny 1995. F. Ronquist with Hymenoptera parasitic relationships the and in function structure of and structure Ovipositor Ovipositor distribution 1994. 1999. L. and PV. Vilhelmsen A, Gaitens structure LeRalec SN, DLJ, the Quicke Ingram of JR, Tunstead implications MG, Phylogenetic Fitton 1992. DLJ, SN. Quicke Ingram MG, Hall. Fitton and Chapman DLJ, London: wasps. Quicke Parasitic 1997. DLJ. Quicke odadrG rjm .19.Sseaisadbooyof biology and Systematics 1991. P. Grijpma G, Nordlander odadrG 92 dniisadrltosiso h rvosycnue genera confused previously the of relationships and Identities 1982. G. Nordlander odadrG 98 eiino h eu hprmrsFese,16 ihrfrnet north-western to reference with 1869 Foerster, genus Rhoptromeris the of genus Revision 1980. the G. Nordlander of Revision 1978. G. Nordlander ilrG,LmdnP.18.Osrain on Observations 1985. PL. Lambdin GL, Miller elnD am-lvnlC 93 omslvrie tbooi ’ncnpd noohg.Bulletin entomophage. cynipide d’un and checklists, biologie species nomenclature, genera, et to key Charipidae: lavaraires American North 1991. Formes HH. Evenhuis AS, 1913. fungi. Menke wood-decaying C. in breeding Baume-Pluvinel insects of D, patterns species Colonization Keilin the 1999. of M. distribution Jonsson geographical G, Rica: Nordlander Costa with M, of larvae, Jonsell dipterous hymenopterans of Parasitoid parasites 1989. cynipid RG. certain Mexzon of LF, status Jiron economic and life-histories the On 1928. HC. James fly, cynipid new a of life-history The 1940. K. Huzimatsu ee .12.DeprstshnHmnpee e rtlee( Fritfliege der Hymenopteren parasitischen Die 1923. R. Meyer hooisihbtn oie oe Hmnpea yioda uolde.Etmlgc Scandinavica Entomologica Eucoilidae). Cynipoidea: (Hymenoptera: cones 22:209–218. conifer inhabiting chloropids aveo yatrpcfiso h aiySrohgde(itr) noooia eiw53:36–44. Review Entomological (Diptera). 65:169–174. Sarcophagidae family the of flies synanthropic of larvae 87:199–229. Biosciences Mathematical 88:52. News Entomological Anthomyiidae). 43:503–542. Research Entomological 113:2–40. London 24:67–85. of Zoology Society of Entomological Journal Netherlands experience. of matter a Cynip.): (Hym. 33:1–60. Supplement Scandinavica Entomologica Ibaliidae). 133:483–494. Society Linnean the of Journal Rendiconti Entomologia, di Italiana 20:309–335. Nazionale dell’Accademia Atti hypotheses. new 47:197–239. 28:635–682. of History exploration Natural of an Journal Ichneumondoidea. to reference special with 26:587–608. Hymenoptera, History within Natural of Journal Hymenoptera. the in valvilli ovipositor 40:367–370. Entomophaga capitata Trichoplasta eeoisstigma Hemerobius Hmnpea yioda uolde.Etmlgc cniaia11:428–453. Scandinavica Entomologica Eucoilidae). Cynipoidea: (Hymenoptera, uoenseis tde nEciia Hm:Cnpie)I.Etmlgc cniaia9:47–62. Scandinavica Entomologica II. Cynipoidea) (Hym.: Eucoilidae on Studies species. European noooi 9:111–120. Entomologie e pce of species new 47:27–88. a Belgique la de et France la de Scientifique 3:145–161. Conservation Insect of Journal 34:53–60. Entomophaga Tephritidae). (Diptera: flies 15:287–316. fruit Biology with Applied associated of Annals forms. larval new some of descriptions 15:457–480. 4) (Series Reports Scientific ahntn93:136–158. Washington Dp. ehiia)ppecletdi orlclte fTcmnPoic,Argentina. Province, Tucuman of localities four in collected pupae Tephritidae) (Dipt.: Hmnpea yioda uolde.Etmlgc cniaia13:269–292. Scandinavica Entomologica Eucoilidae). Cynipoidea: (Hymenoptera: Dilyta Nuotr:Hmrbia) noooia es96:93–97. News Entomological Hemerobiidae). (Neuroptera: curneadpyoeei mlctoso h vpstrclip ovipositor the of implications phylogenetic and Occurrence Fo rtr(yeotr:Cnpie) rceig fteEtmlgclSceyof Society Entomological the of Proceedings Cynipoidea). (Hymenoptera: ¨rster uol trichopsila Eucoila Leptopilina siel frit Oscinella ncai melanoneura Anacharis Fo rtr 89 ihntso h ttso oeohrgenera other some of status the on notes with 1869, ¨rster, liooajaponica Kleidotoma atg(yeotr,Cnpie) aaieo the of parasite a Cynipoidea), (Hymenoptera, Hartig L)i atr ot mrc.Aeia ultnof Bulletin American America. North Eastern in (L.) siooafrit Oscinosoma hprmrsstrobigena Rhoptromeris teioavrasoccata varia Atherigona Hmnpea iiia) aaieof parasite Figitidae), (Hymenoptera: .s.Thk meilUniversity, Imperial Tohoku sp. n. , ..Zishitfu Zeitschrift L.). Odonteucoila blarufipes Ibalia Anastrepha sueciabochei Pseudeucoila p . aaiodof parasitoid a n., sp. , p.and spp. od (Diptera: Rond. Coneucoela (Hymenoptera, rAngewandte ¨r Ceratitis 2281 Weld and , Downloaded By: [National Agricultural Library] At: 20:41 25 October 2007 fEtmlg,TxsAMUiest;ohrie h oce shue tthe at housed is voucher the otherwise, University; Riverside. Department UC the A&M at Museum, located Research Texas is Entomology specimen voucher Entomology, the indicates name of taxon a after asterisk An examined Figitidae of List 1. Appendix larval-pupal Pan and egg-pupal California. Hymenopterous Central 1981. WG. from Hart MM, Diptera Fischel RH, dung-breeding Rhode FE, of Gilstrap parasitoids RA, Wharton and of predators ovipositor Some multipurpose 1979. The RA. 1999. Wharton F. Bin R, Romani N, parasitoid Isidoro the JC, in clip Lenteren ovipositor van the of anatomy Functional 1998. F. Bin N, Isidoro the JC, of Lenteren selection van host and finding habitat Host 1991. I. Eijs G, Nordlander JJM, Alphen van between comparison parasitoide a do Diptera: Ocurrencia 1986. breeding FH. dung Valicente of parasitism insect Natural of 1968. predators RR. and Gerhardt parasites RP, Burton the EC, of Turner catalogue A editor. WR, Thompson In: Cynipidae. 1955. WR. Thompson yi G 93 aaie ffc fly, face of Parasites 1973. HG. Wylie fruit-infesting with associated (Cynipoidea) Eucoilidae Neotropical 1998. FE. Gilstrap SM, Ovruski RA, Wharton 2282 Thrasorinae Anacharitinae Charipinae Emargininae Aspicerinae Figitinae Eucoilinae elvle nai.Cnda noooit105:257–261. Entomologist Canadian Ontario. Belleville, of parasitoids 55:181–186. Entomologist Pacific Rendiconti Entomologia, di Italiana Nazionale dell’Accademia 47:265–277. Atti function. and form between relationships 27:263–268. Embryology and Morphology Insect of heterotoma 87:324–329. Oecology species. australis Leptopilina 15:391–392. Brasil 61:1012–1015. do Entomology Entomologica Economic of eluta Journal fly. Euxesta face the host, introduced 210–216. an Control. and Biological hosts of native Institute Commonwealth Ottawa: 3. Part 2, Section pests. ehiia,wt e eod rmAgnia oii n ot ia ora fHmnpeaResearch Hymenoptera of Journal Rica. Costa and Bolivia Argentina, from 7:102–115. records new with Tephritidae, 26:285–290. sp.; sp.*; (Kieffer); aapdu nigrimanus Ganaspidium (Yoshimoto); Xyalophora sp.; .L Buffington L. M. rbigah rapae Trybliographa Leptopilina Dettmeria : Topo)(yeotr:Eciia) tutr ogi saiglra.ItrainlJournal International larvae. escaping grip to structure a Eucoilidae), (Hymenoptera: (Thompson) : Amphithectus : : eaiu capitata Ceratitus : ow 88(itr:Oiia) arga eSt aos G ni aSociedade da Anais MG. Lagoas, Sete de regiao na Otitidae), (Diptera: 1868 Loew, gnsi daci Aganaspis loyt brassicae Alloxysta dnoeaanastrephae Odontosema Aspicera : sp. Plectocynips : Thoreauella eiisclarimontis Aegilips Ealata Hmnpea uolde ihacmaio ftence fEuropean of niches the of comparison a with Eucoilidae) (Hymenoptera: sp.; sp.*; sp.; sp.*; sp.; ohuol anastrephae Lopheucoila sp.; Callaspidea sp. and Westwood. Dicerataspis (Kieffer); (Weld)*; ‘uol’ impatients ‘‘Eucoila’’ uc autumnalis Musca Figites Euceroptres Anastrepha Ashmead; etei euxestae Dettmeria Kieffer; sp.; sp.; Glauraspidia Agrostocynips Borgmeier*; p.(itr:Tprtde nCsaRc.Entomophaga Rica. Costa in Tephritidae) (Diptera: spp. sp. sp.*; heolpi ambrosiae Phaenoglyphis Melanips Lonchidia Anacharis Dpea ucde,adascae itr near Diptera associated and Muscidae), (Diptera: ogee,13 Hmnpea uolde em Eucoilidae) (Hymenoptera: 1935 Borgmeier, Dieucoila (Say); sp. Rohwer*; sp.; Rhabdeucoela sp.*; sp.; sp.; Gronotoma Hexacharis aapsmundata Ganaspis Neralsia Aporeucoela sp.; odadel semirufa Nordlandiella (Ashmead). ioym pacifica Disorygma sp.*; sp.; etpln heterotoma Leptopilina sp.; sp.*; Drosophila sp.; Trischiza Xyalaspis Tropideucoila Kleidotoma Caleucoela Fo Leptopilina Leptopilina parasitoid ¨rster*; sp.; sp. :