MORPHOLOCY H1SТOLOGY AND FINE SТRUCI1JRE

Morphology and Sensilla of the Orbicula а Sclerite Between the Tarsal Claws in the Hyrnenoptera

HASAN Н ВАSlВUУUК12з DONAID L J QUICKEY ALEXANDR Р RASNIТSYN4 AND

MIKE С FIТfON2

Unit of Parasitoid Sуstешаtics CAВI Bioscience ПК Centre (Ascot) Dераrtшепt of Biology Iшреriаl College at Silwood Park Ascot Berkshire 8L5 7РУ UК and Dерartшепt ofEnton101ogy the Natural Нistpry Мusеuш

London SW7 5BD UК

Апп Entomol Soc Ат 93(3) 625-636 (2000) АВSТВЛCf ТЬе огЫспlа (шanuЬrium) is а dorsal sclerite between the tarsal claws in the Нуshyшепорtега It~ detailed external шогрhоlоgy was surveyed throughout the order using scanning electron rnicroscopy Four types of ргеsuшеd rnechanosensory sensilla were found two sensilla trichodea and two sensilla сашрапifогшiа Variation in number Ьаре orientation and associated structures ате discussed in the light of current phylogenetic llypotheses Тllere are geneally two sensilla tichodea А оп еасЬ orbicula and this is а putative synapomorphy for all of Hyrnenoptera excluding the Xyelinae ТЬе presence of а single sensilla trichodea А with а marked socket is postulated аБ а sупарошогрhу fo the Chalcidoidea Myrnarornrnatidae Diapriidae ап undescribed New Zealand fашilу of Proctotrupoidea and possibly Ceraphronidae Arrangernent of the sel1sila trichodeaB in а sil1g1e row supports а putative clade СОШРlisiпg the Chalcidoidea Мутаroшшаtidае Ceraphrol1idae Cynipoidea Diapriidae Platygastridae and the undescribed New Zealal1d family Presence оЕ а БрШ at the Ьзsе of the sensilla trichodea В is а putative БупароrnогрЬу for tlle Chalcidoidea al1d the undescribed New Zealal1d family PossessiOI1 of sensilla сarnрапifогшiа type В suggests а clade cornprising Chalcidoidea Diapriidae and 8celionidae

КЕУ WORD8 Нушепорtега phylogeny morphology orbicula sensilla

ТНЕ НICНER LEvEL phylogeny of the Нушепорtега has received increasing attention in the la5t 30 yr (Rasnitsyn 1969 Konigsmann 1976 1977 1978а 1978Ь 1980 1988 Brothers 1975 Brothers and Carpenter 1993 Dowton and Austin 1994 1995 1997 Dowton et Ш 1997 Vilhеlшsеп 1997 Ronquist et al 1999) but there is still по опе reasonably robust phylogenetic hypothesis of the order ТЬе monophyly of several шаjог parasitoid groups and their interrelationships are still sensitive to апу new test (Ronqllist et аl 1999) Formal cladistic reanalysis ofRasnitsyns (1988) data has shown that his intuitively drawn and widely cited set of phylogenetic relationships has по convincing Sllpport for шапу шаjог apocritan divisions (Rопqпist et аl 1999) Nevertheless Rasnitsyns phylogenetic hypothesis of lower Нушепорtега (Symphyta) рагshyticularly the branching order of пшjог groupings is sllpported Ьу this reanalysis (Ronquist et Ш 1999) as well as Ьу Vilhelmsens (1997) independent stlldy The other шаjог segment оfhушепорtегап phylogeny

I Dераrtшепt of Biology Imperial College Silwood Park Ascot Berkshire uк SL5 7РУ

2 Department of Entofl101ogy Natuтa Нistory МllsеШfl London uк SV75BD

Department ofBiology Cllmhllriyet University 58140-Sivas Ттshykev

4Paeontologica Institte Russian Academy of Sciences 117647 MQSCOV Russia

that ЬаБ good Sllpport is а part of the Aculeata The hypothesis that the Aculeata comprises two шаjог lineages (Rasnitsyn 1988) the Chrysidoidea and the Vespoidea +Apoidea has Ьееп preserved in subseshyqllent fогшal analyses (Ronqllist et al 1999) Relashytionships among families ofthe Chrysidoidea found Ьу Ronquist et аl (1999) were largely congruent with those in the hypothesis produced Ьу Brothers and Carpenter (1993) Rasnitsyns hypothesized геlаНопshyships between the groups of parasitoid Нушепорtега аге fragile and Ьауе largely failed to раББ the test of fогшal cladistic analysis (Ronquist et аl 1999) In these reanalyses Rasnitsyn s infraorders ЕvапiошогshyрЬа and Ргосtоtгнрошогрhа did not appear as mопоshyphyletic nor was the proposed sister group relationshyship between Ichпеuшопоidеа and AC1l1eata еП Sllppolied There is therefore still а need for шuсh further work оп new character sуstешs to provide data for а sound phylogeny ТЬе Нушепорtега is опе of the шоst biologically

diverse and species-rich grollPS ofinsects (Ia-alle and Gallld 1993) and possesses diverse БеПБОгу sуstешs Most studies to date have concentrated оп antennal sensilla (Basibuyuk and Quicke 1999a) with the senshysoгу sуstешs of other parts of the body largely being ignored Those othcr sуstешs that Ьауе Ьееп investishygated include the ovipositor (eg Ganesalingam 1972 Greaoy et al 1977 LeRalec and Wajnberg 1990 Leshy

0013-8746000625-1)636$02000 copy 2000 Entomologica1 Society of America

626 ANNALS OF ТНЕ ENTOMOLOGICAL SOCJEТY OF АМЕШСА Vol 93 по 3

Ralec et al 1996) whole body parts (Sсhшidt and Sшith 1987) and legs (Navasero andElzen 1991 Меуshyhofer et al1997) but in еасЬ case tllese Ьауе included only опе or а few species ofрашsitiс wasps Studies of l1Оst searching and detection Ьу parasitic wasps Ьауе largely Ьееп related to receptors оп the antcnna and ovipositor and the possible role of tarsal sепsШа has only recently Ьееп suggested (МеуЬofег et al 1997)

In tMs article уе survey the detailed шогрhоlоgу of the orbicula and its associated sепsiПа throughout the Нушепорtега Тhe orbicula is а dorsal cuticular scleshyrite situated basal to the агоliuш in tlle claw-bearing tarsal sеgшепt Уе report phylogenetic implications of огЬiculю structurcs including sensilla and discuss thcir evolution in thc light of current phylogenetic hypotheses Comparisons bctween sexes shov that orbicular scnsilla ате not diffегепtiаПу developed in fешаlеs and therefore probably do not play а particshyular role in I1О8t location in parasitic wasps

Materials and Methods

Scanning Electron Microsoopy Ргерагаtiоп Most specimens were examined Ьу sсаrшiпg electron mishycroscopy (SEM) but some binocular light microscopy observations were also involved particularly of large aculeate specimens Fore legs were гешоvеd апd transferred to absolute ethanol for at least 18 Ь then air-dried апd mounted before sputter coating In шоst cases ultrasonic сlеапiпg vas поt appHed to avoid breaking sensilla trichodea Rare sресimепs were ехshyашiпеd without coating usiпg an Епviгопшепtal СhашЬег МЮk АВТ 55 microscope Stапdагd SEM tесhпiquеs Ьауе the роtепtial drawback of fаiliпg to геуеаl sоше Нпе details such as the ргеsепсе of sепshysШа carnpaniformia especially iп sшall specimens vithout special treatment Тherefore the data ргеshysented for these sепsillа ShOllld Ье treated with some саге althollghwhеп visibIe their lосаtiопs were clear

Telminology and Material Exarnined IdепtiRсаshytiопs of sепsШа types are based оп Mclver (1985) and Frazier (1985) Microsculpture tегшiпоlоgy largely follows Аllеп and Ваll (1980) Material exarnil1ed is il1cluded as Аррепdiх 1

Results

Orbicula ТЬе orbicula (mal1llbrium) is а dorsal cllticular sclerite at the base of the mешЬгаl10US аюshyНит iп the telotarsus (clav-bearil1g sеgшепt) (Figs 1 апd 2 (Sсhшidt al1d Smith 1987 СанЫ and Воltоп 1988 Meyhofer et al 1997) In the Нушепорtега the orbicula is well developed althollgh its sculpture al1d the types пumЬеrs апd аггапgешепts of its sensilla уагу These are suшmаrizеd iп ТаЫе 1 апd describerl below

Surface Textllre of Olbicula (character 1) Insect il1tegumel1ts аге tlsually furпishеd with microsculpshyture геsultiпg fгош ерidегшаl сеll secretions Sculpshyture is llSually шапifеst as а polygonal and геgulю шеsh sуstеш but а wide range of variation and сотshyplete reduction mау осспr (Аllеп апd BalI1980) We

recognize three states as follows (1) scale-like with projection(s) а scale tips (Fig 1 state О in ТаЫе 1) (2) simple scales (Figs 2 апd 3 state 1 in ТаЫе 1) (3) virtually entirely SП1Qоth (Fig 4 state two in ТаЫе 1) State О is largely confined to the superfarnilies Xyeloidea Pamphilioidea Cephoidea and Тепthгеdishynoidea although it also occurs in а few Apocrita (ТаshyЫе 1) Тhe scales are usually regular in distriЬutiоп (Fig 1) Ьи ате SОlllеtiшеs шarkеdlу reduced Тhey are most evident оп the basal half of the orbicula Тhe scales usually possess а single plojection apically Ьи there are several iп а few tenthredinoids (Arge Blasshytiootorna Selaпdria) and ichneumonids (Labena Мetорiщ Phytodeitus Rhyssa) (Fig 1 arrow)

State опе occurs in тапу Symphyta and in some Apocrita notably in the Trigonalidae Stephanidae and тапу Ichnellmonoidea (Table 1) Tlйs usually forms а regнlar mesll Ьн both slightly traI1sverse (Fig 3) апd markedly elongate scu]pture aso occurs (Fig 2) А fev proctotrupoid families notably the Helorishydae and Roproniidae Ьауе weakly deve]oped scaleshylike ll1icrosculpture Ьн the basa half of the orbicula is sшооth in Heloridae and ШS SOll1e deI1tiform scaes in Rоргопiidае (Fig 5 aIТOW) Most parasitic taxa and а [еу savflies laуе а сошрlеtеlу slllooth orbicllla (state 2 ТаЫе 1)

Orbicular Sепsillа We Ilave identiRed four types of sепsillа оп the orbicllla Two are setiforlll sensilla trishychodea А (STA) and sепsillа trichodea В (SТE) and the other two аге dome-like sепsillа сашрапifогmiа А (SCA) al1d sensilla саmрапifО]miа В (SCB) In most cases сотраПsoп of their shapes reative роsitiопs ll11mber and socket stПlсturе enabled llS to postulate homologies Ьеtwееп taxa uпсегtaiпtiеs are indicated in the text vhепеvег песеssагу

STA (character 2) Tllese structures arise from flexshyible sockets Ьауе the seta truпk gradually tapering toward the tip and ате щиаllу strопglу 10пgitlldil1аllу grooved (Fig 2) Т1lеу сап Ье distingнished from sепsiJlа trichodea В (SТE) Ьу роssеssiоп of а wеПshydefined socket Ьу llSually Ьеiпg шuсh lопgег and thicker (as тисЬ as three tiшеs) and Ьу Ьеiпg located шоге distally (Figs 2 апd 4 STA cf SТE) They ате alvays Ьеп toward the атоНит to sоше degree al1d llsually overlap it (Fig 4) Iп а few taxa notably the Siricidae where there шау Ье опlу а single sel1silla trichodea or only опе type whеп tlere ате several relative size could поt Ье used to help determine which type they were altlIough other morphological aspects suggest that they ате STA (Fig 6)

STA аге ргеsепt iп almost аН Hymenoptera (ТаЫе 1) Sexual dimогрhisш occurs most поtаblу in the Siricidae in which females usually Ьауе по or оссаshysiопаllу only опе STA although males usually Ьауе hvo or шоге However Sirex fell1ales сап sOllletimes ЬаУе six STA (Fig 6) Similarly in some of tlIe Асshyuleata that display extreme sexual dimorpllism (eg Tiphiidae) fешаlеs uSllally have reduced orbicula апd thlls Ьауе по sепsillа trichodea

Most Hymenoptera possess two STA (see ТаЫе 1 and аЬоуе for ехсерtiопs) bllt only а single SТA is present iп аН chalcidoids (including mymarids ыlt

627 Мау 2000 BASIВUYUK ЕТ ЛL ОНВICULЛ OF HYМENOPТERA

ТаЫе l Character states Сог the orblcula tшd associated веп- ТаЫе 1 Continued Ша u the Hymenoptera

Charactersl

Characters(1 Таха Таха 2 3 4 5 6 7 8 9

2 3 4 5 6 7 8 9 Sclerogibbidae 1 2 3 О о о з О О

Xyelidae Dryinidae 2 2 3 О О О О О ХуеНпае О О О О О ЕmЬоlешidае 2 2 3 О о о о о 1vfacroxyelinae 1 2 О 2 О 01 Betlylidae 2 2З 3 О О О О О

Xyelecia 1 2 2 О О О Chrysididae 112 2 03 О О О 02 О О Pamphiliidae Tiphiidaeb 02 023 О О О

Cephalciinae О 2 О 2 О О Mutillidae 2 2 3 О О О о О Раmрhiliiпае О 2 23 О О О О Sapygidae 2 3 О О О

Megalodonteidae О 2 1 О О О О Seoliidae 1 2 3 О О О О Megalodnntes О 2 О Bradynobaenidae 2 02 01] О О О О

kuznetzovi Sierolomorphidae 2 3 О О О Argidae 2О О О О RllOpalosomatidae О 2 01 О О О О Bla5toeotomidae О 2 О О О Formicidae 2 2 03 О О О 02 О О Cimbicidae О 2 011 О О О О Pompilidae 2 3 3 О О О О О Diрriопidае О 2 23 О О ОО Vepidae 1 2 О О О О Pergidae Specidae О 2 3 О О

Acordulecerinae 1 2 3 О О О 2 О О Apidae 1 2 123 О О О PIylacteltgtphaga 1 2 О 2 О О Perga 1 2 О 2 О 01

Chamcters and character 5tate (1) Orbicua texture О sltulpshylорlщrоtоmа 1 2 О ОР о tured primarily vith scale but а least ome scale usually а Ьмal асеаТепthrеdiпidае О 2 О О О vith cutic1l1ar projections 1 sculptured entirely vith 5cales 2 almost Ccphidae О 2 03 О О О О О entirely mooth (2) Conspicllously socketed sensila trichodea АAnaxyelidae 2 О О О (STA) оп orbicula о попе 1 опе2 two 3 tlgtreeormore (3) Sensilla Siricidae 0123 О О О trichodae В о попе 1 опе 2 hvo 3 three or more (4) Sensilla Xiplgtydriidae trichodea В (two or тоте) О = scattered 1 symmetrical or formsXiphydriinae 1 2 01 О О О О а single ~ow (5) Sensilla trichodea В О simple 1 Ьщallу enlarged (6) Dereeyrtinae 1 2 1 О О О О Senilla trichodea В О witlOllt а Ьаза cuticular projection 1 а basalOrussidae 1 О О О О

Cuiglia 1 2 О О cuticHlar projection (7) Sensilla campalliformia А (with ап eneircling scшuinslаndii ridge) О попе 1 опе 2 (уо 3 three or тоте (8) SепsШа сат-

Stephanidae 1 02 03 О О О 02 О О paniformia В (vithout ап encirclillg ridge) О попе 1 two (9) Trigonalidae 1 213 3 О О О 2 О О Position of sensila саmрапifоrmiа [о sensilla trichodea А О anterior Megalyridae 1 ОЗ 3 О О О 1 О О 1 level with or posterior Еуапнаае 1 2 3 О О О О О ь Sexual diшоrрllism (see Results)

О

Aulacidae 1 2 3 О О О О О GastеruрШdае 1 2 3 О О О О О absent in sоше agaonids) (Fig 7) шушагошmаtidsIbaliidac 2 2 3 О О О О О Figitidae 2 2 013 1 1 О О О diapriids the undescribed New Zealand family and Liopteridae 2 2 3 1 1 О О О possibly Ceraphronidae This single sепsiПuт is al-Cyuipidae 2 2 023 1 1 О О О ways situated оп the apical halfofthe orbicula and hasMegaspilidae 2 3 3 О О О О О Ceraphronidae 2 1 013 О О О а distinct high соllаг to its socket wmch is angled Diapriidae 2 1 2 1 1 О О 1 О distally (Fig 8) This соllаг шау serve to restrict back-Heloridae 1 2 3 О о о О о ward movements oftlle sensillum Three or more STA Vаrtlюrniidае 2 2 3 1 О О О юе found in some Siricidae Trigonalidae MegalyrishyРеlееiпiclае 1 2 3 О О О О О Proctotrupidae 2 02 23 О О О О dae Mcgaspilidae Ichneumonoidea and some ас-Roproniidae 011 2 3 О О О О О uleate taxa most notabIy the Pompilidae In this last Мопоmасhidaе 1 2 3 О О О О ОР family there are usual1y six or more STA which formNev Zealand ЕатНу 2 1 2 1 1 1 О 1

а transverse apical row (Fig 9) Sсеliопidае 2 2 013 О О О О 1 О Platygastridae 2 07 2 ] 1 О О Diffelentiating between STА and SТВ in those taxa Chalcididae 2 ] 3 О 1 О 1 with three or more STA becomes increasingly diffishyО Leucospidae 2 1 023 О 1 О спlt and ассшаtе assessment of the number of еасЬEurytomidae 2 1 3 1 1 1 О 1 1 Ormyridae 2 1 3 1 1 1 О 1 type is not always possibIc (particularly in Braconidae Pteromalidae 2 1 3 1 1 О 1 1 and Megaspilidae) Sucll states are indicated Ьу () iп Eupelmidae 2 1 3 О 1 1 ТаЫс 1 (see characters 2 З 4 and 5)Епсуrtidае 2 1 3 1 STB (cllaracter 3) SТВ пsиаllу сап Ье differenti-Agonidae 2 О О Aphelinidae 2 ] 3 1 1 ated from the STA Ьу being shorter al1d more slender Elasmidae 2 1 2 1 1 О 1 having а less pronounced socket (not obvious in manyEulophidae 2 1 23 1 1 1 О 1 sшаП specimens) and а more posterior and irregularSigniphoridae 2 1 3 1 1 О Trichogrammatidae 2 1 2 1 1 07 1 distribution ТЬеу are usually а siшрlе cylindrical Ьюr Torymidae 2 1 3 2 1 О 1 in а socket (Figs 2 5 9) bllt in some taxa they аге Mymaridae 2 1 2 1 1 о l widened and complessed basally (Fig 10) ТЬе posshyМуmагоmшаtidае 2 1 2 1 sibility that there тау Ье two distinct types of SТВBraconidae 112 23 3 О О О 2 О 1

Liobracon 2 3 О О 3 О 01 cannot Ье determined withollt llltгastruсtшal and Ichneumonidae 01 213 0111213 О О О 23 О 01 electrophysiological work and we rccognize only one Plumatiidae 2 2 3 О О О О О type SТВ аге present only in а few sушрhуtan taxaScolebythidae 1 2 17 О О О О but аге posscssed Ьу тапу Apocrita (ТаЫе 1)

628 ANNALS OF ТНЕ ENTOMOLOGICAL SOCIEТY OF АМЕтСА Vol 93 по 3

Figs l-3 (1) ОгЫсlllа in Sеflщiа sp (Tenthredinidae) STA sensilla tгiсhоdеа А arrow head sho ing scale-like SСlllрtше Nith cllticlllar projections Ваг = 30 т (2)Orbicula in Arotes arnoentL~ (Ichneumonidae) STA three sensilla tJiсlюdеа А black aJrow sel1silla tlichodea В white arrow sel1silla саmралifогтiа [уре А Ваг = 38 т (3) Close-up of orbicula tехtше il1 Регgо dorsafis (Pergidae) showing scale-like sculpture and two sensilla сатралifогтiа А (апоws) Ваг = б т

ТЬеге аге t1Sшllу three ог тоге SТE (Figs 5 7 9 ТаЫе 1) although only two аге found in а few sawflies some parasitic wasps апс some tiphiids and apids А single SТE occurs in а питЬе of isolated cases (Figs 2 апс 4 ТаЫе 1)

AITangement ofSTB (character 4) STB аге usually (Iistributed irregularly (Figs 5 and 9) but they fопп а single IOW in almost аll cynipoids (except IbaJiiclae) chalcidoids (except at least some Chalcidiclae ап(1 Leucospidae ) Diapriidae Vanhorniidae the undeshyscribed NeN Zealand family Platygastridae and some Ploctotrupidae (Fig 7)

Basal Modifjcation of STB (characteI 5) ТЬе SТE аге wi(lened апс laterally compressed in the Chalshycidoidea Diapriidae Platygastridae the undescribed New Zealand family апс some Cynipoidea (Figs 7 and 10)

Spur of STB Socket (chaJacter 6) ТЬе se nsilla socket bears а spur Nhich appears to restrict its backshyward mobi1ity in most chaJcidoid families including

mymarids together ith the uшlеSСliЬеd New Zeashyап(l famil y (Figs 78 10) А similar spur is present in the Cynipoidea but it is situated тоге distantly from the sensilla ТЬе 110mology of the spurs in (Ье Суshynipoidea witIl those in the other taxa is therefore unceгtain

Sensilla Сшпрапifогтiа (SC) (characters 7 and 8) Two types of sensilla campanifOlmia аге distinguishshyаЫе апс аге termed Ьеге SCA and SCB SCA is usually а roundecl dome encircled Ьу а cuticular ridge (Figs 2 апcl 3) whereas SCB has по external el1circling lidge аl1с арреаг as а simple small tubercle (Figs 4 and 8) SCA aJe Гоиl1с il1 the Trigonalidae апс Pergidae (ехshycept lлрlчrоtОm()) апс in тапу species of BJashyconidae апс Ichl1eumol1idae ТЬеу also are present il1 some replesel1tatives of several isolated taxa (Xyelishydae Pamphiliklae Stephal1idae Megalyridae Sclerogibbidae Chrysididae апс Formicidae) (ТаЫе 1) SCB аге possessed Ьу the Chalcidoidea Diapliidae and some species of Sceliol1idae al1d possibly Ьу the

629 Мау 2000 BASIВUYUК ЕТ AL ORВICLJLA 0[ HVMENOPТElA

Figs4-7 (4) ОЪi Сl1 lа11 Triteleio sp (SceliOl1idae) vith two sel1silla trichodeaA (ореl1 апоs) sil1gle sensilla tichoclea В (hite arro) аl1сl two sel1silla campalliformia type В (white shadovecl bIack arrows) Bar = 8 fLl11 (5) Orbicula il1 RopTonio gопnаni (Roproniiclae) Ilote clel1tifoCI1l sca les (arтo) Ваг = 30 fLl1l (6) Orbiclla ill Ггсmех sp (Sricidae) shoving presl1l11ed six sel1silla trichodeaA Ваг = 50 fLl1l (7) Orbicl1la ill Ptoth -ix sp (Pteromalidae) sho ingsil1 g1e sel1silla trichodea А (ореl1 апоw) and а tansverse JO W of sellsilla tcl)Qclelt1 В (smзl1 arrows) Ваг = 13 fLl11

undescribed New Zealand family АI1 (аха that have sensiJla campalliformia Iave only опе type with tlle possible exceptioll иоьгасоn (Braconidae) (Fig11) The great majority oftaxa witJl SC (either А ог В) Ьауе two of them but only а sillg]e SCA is foulld ill megashyIyrids апс three ог тоге occur in some pergids Ьгаshycollids sclerogibbids апс тапу ichneumonids

Positioll of SC ( сhагасtег 9) SCA аге usua]]y роshysitioned iпеgulагlу апс distal to STA Howeer they сап Ье both distal аllcl basltll (о STA ill LVlacroxyel (MacroxyeJinae) Perga (Pergide) апе] тапу IchshyneumOlliclae (ТаЫе 1) Positiolls of SCB аге quite

cOllsistellt within the Chakidoideltl аllс Diapriidae ill vhich they аге always 10саtеclllttегоvепtгаJlу Jelative to the STA оП eitheJ siele of JateraJly compressed orbicula level witJl 01 sliglltly basal to it (Fig 8) TI1e SCB of sceliollids however ltге locateel clistal (о the STA (Fig 4)

Discussioll

Functiolla Implicatiolls The orbiculal sellsillt аге lttrgllably all mechalloreceptors (McIver 1985) 111 th e eulophicl Sympiesi~ seiceicomi~ Nees they have Ьееп

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Figs8-11 (8) 5e nsilla ill Jurywma sp (Ешуtоm idае) 5ТА sensilla tri chodea А wi th а wellmiddot (leve loped socke t 5ТВ sefJS illa tic llOdea В white 3IТOW se lls illa саmралi fогmi а type В Bar = 3 Lm (9) CeГOfo les sp (Роm рШdае) SllO v ing pos ition зп(1 аrrалgеmепt of sensilla tricllOdeaA Bar = 38 LПJ (10) Sелsillа tichodea В of Castmcantl-s рсетjlIШI (Pterom alidae) Sl10ving widened base and basal spur (ореп апоws) Вш = 3Lnl (11) Orbicula in иоЬгас(т sp (Braconidae) sho-ing bot11 types of se nsilla саmрап ifогmiа vl)ite апоws ргеSLmеd sел sillа campan iformia В white shadove(1 bIack arrovs sensilla campan ifomia А Ва = 20 Lm

suggested as шviпg а role [п de tecting vibration s ргоmiddot ducecl Ьу th e ir leafminer host (Meyhbfer e t al 1997) Althougl this тау gtell Ье the case theiI prese nce [п taxa with diffe le nt blologies (incllding phytoph agollS taxa a llC раГltlsitоid s of deeply conceaed hosts) sugshygests that they have otl er гoles as well ТЬе ргеsеп се of apparen tly identi ca sensila iп both sexes of тапу taxa fшthеl weake ns tle host locatio) hypotllesis Because ST and SC аге prese nt in th e most basal ineages ofthe H ymenoptera (eg MacJQxyeinae and Cephalciinae) we suggest that the iJ огigiп а l and ргеshy

cominant fun ction is топitогiпg moveme nts and s tгеss resulting fгош the asps own 10comotion anс possibly also subst)a te viblation that тау indicate the ргеsепсе of pote) tia pledatoгs Athough they тау play а role [п host de tection the ack of sexua diтогshyphism i) рагаs itiс taxa suggests that this is not а majoJ need Indeed the sexua dinlOrphism in th e phytophshyagous Siricidae and iп th e рагаs itiс Tipl iidae both cases vit females having fewe r STA suggests а gleate J iшрогtапсе fOl th e maes Моге obselvations оп paras itoid host sею-сhiпg behavioJ includi ng еесshy

631 Мау 2000 BASmUYUK ЕТ ЛL ORВICULA OF НУМENОPrERЛ

trophysiological studies are needed to determine tle exact function of these sensilla

Рhуlоgепеtic Imрliсаtiопs То assess рhуlоgепеtiс implications ofthi5 character system we have mapped the data onto the most recent phylogenetic hypothesis of the епtiге order re5ulting from implied weighting (Ronquist et а11999) Only the uпаmЬiguоus changes were indicated and polytomies were treated as Iard (ie аssumiпg multiple speciation events) (Fig 12) We have included the undescribed New Zеаlапd famshyНу iп the putative clade comprising Chalcidoidea Cershyарhroпоidеа Diapriidae and 50 оп because it shares at least опе putative sупароmогрhу with mепtiопеd taxa (Basibuyuk апd Quicke 1997) We also have plotshyted the data оп to Rasnitsyns (1988) intuitive hypothshye5i5 and оп to а tree resulting from reanцJysis of а соmЫпаtiоп ofRasnitsyns data (Ronquist ct а1 1999) with VHhelmsen s (1997) sуmрhуtап апd Brother5 and Сагрепtегs (1993) aculeate hypothese5 However the fit of the data were marginally better оп the latest hypothesis of Rопquist et а1 (1999)

Orbicular Sculpture The orbiculas of тanу of the most Ьа5аlliпеаgеs (eg the Xyelinae Pamphiloidea Cephoidea апd Tenthredinoidea other than pergids) have scale-like microsculpture with cuticular projecshytion5 (character 1 5tate О) sugge5ting thi5 to Ье the plesiomorphic condition for the order Simpfe scaleshylike 5culpture without рrojесt10ПS (character 1 5tate 1) appears first 1П the siricoid grade апd i5 геtаiпеd in the Evaniomorpha sеП5U Rasпitsуп рщ of the Procshytotrupoidea (РеlеС1П1dае Heloridae Rоргопiidае Мопоmасhidае) апd 1П mапу Iсhпеumопidае It also 15 ргеsепt 1П the Масгохуеliпае and Pergidae which сап Ье best сопsidегеd as iпdерепdепt acquisitions Further reduction of the microsculpture to а сотshypletely 5mooth orbicula (character 1 state 2) has арshyparently occurred оп at least three separate оссasiопs iп рщ of the miсгоhуmепорtегап lineage iпсll1diпg the Супiроidеа Diapriidae the uпdеsсгiЬеd New Zеаlапd family Platygastroidea Сегарhгопоidеа Муshymarommatidae апd Chalcidoidea iп the pl1tative Proctotrl1pidae + Vanhorniidae liпеаgе and in the Aculeata ТЬе presence of some simple scale-like sсulрtше (charactcr 1 state 1) in the Sclerogibbidae Scolebythidae апdVe5pidae сап best Ье considered as reversals (Rasnitsyn 1988 Brothers and Сагрепtег 1993 Ronquist et al 1999)

Sепsillа Trichodea А STА аге ргеsепt iп almost аН taxa оfНуmепорtега except Хуеliпае and their presshyепсе is therefore а putative synapomorphy for the Нуmепорtега ехсludiпg the ХуеНпае АЬsепсе of STA in а few isolated taxa in which they are и511аНу polymorphic (5iricids orussids stephanids megalyrshyids proctotrupids platygastrids ЬгаdупоЬаепids апd tiphiids) сап be5t Ье regarded as iпdерепdепt losscs (Fig 12) Тhe Пl1mЬег ofSTA i5 overwhelmingly two and this

сопditiоп (character 2 state 2) is most likely the gгоuпd рlап for tlle Нушепорtега as а whole Reducshytion iп the number of STA to опе (character 2 state 1) with а markedly raised socket is а pl1tative synashyротогрЬу for а clade соmрrisiпg Chalcidoidea Муshy

mагоmшаtоidеа Diapriidae Ceraphronidae and the undescribed New Zеalапd family but with опе revershysal (iп Супiроidса) and at least опс tгапsitiоп to other states (in Megaspilidae) Three ог more STA occur in some Trigonalidae Megalyridae Megaspilidae Brashyсопidае Ichпеumопidае апd Pompilidae bl1t iп most of the5e cases the Iikely gгоuпd рlап i5 two STA Ьеshycause that state occurs iп тanу trigопаlids iсhпеushymопids as well as in some Ьгасопids (eg аgаthidiпе gепега Labagathis and L~opronotum) ТЬе ргеsепсе of thrce or more STA in the Mega5pilidae and Pompilishydae i5 best cOl1Sidered to герге5епt two indерепdепt асquisitiопs (Fig 12)

Sensilla Trichodea В апdRelated Structures Тhese are present in most taxa but their two 5eparate аЬshysепсеs and variаtiоп in their пumЬег (character 3) appear to iпdiсаtе sоше hошорlаsу iп the Apocrita for the givеп 5et of рhуlоgепеtiс hypotheses However possession ofthree ог тоге STA (character 3 state 3) provides support for the monophyly of Apocrita (Fig 12) Аrrапgеmепt of the SТВ iп а siпglе row (Fig 7 character 4 state 1) 511pports the clade соmргisiпg the Chalcidoidea Mymarommatoidea Ceraphronoidea Platygastroidea Cynipoidea Diapriidae and the uпshydescribed New Zealand fашilу (Fig 12) This state a1so occurs iп the Vanhorniidae and Pamphilioidea Тhe оссипепсе ofdi5persed SТВ (character 4 state О) iп the Chalcididae Lel1cospidac and Ibaliidae (which аге relatively larger iп 5ize) within thi5 clade тау sl1ggest sizе-dерепdепt сопvегgепсе for this characshyter but there i5 а gгоwiпg body ofеvidепсе supporting some of the relationships amопg the аЬоуе putative clade (Basibuyнk апd Quicke 1997 1999Ь GiЬsоп 1999 Rопquist 1999 Ronquist et а1 1999) It should Ье поtеd that SТВ a1so аге di5persed in some scelionids ТЬе SТВ in some of these taxa are modified basally (character 5 state 1) being laterally compressed and widепеd These modified SТВ аге ргеsепt in the Diashypriidae the uпdеsсгiЬеd New Zealand family Platyshygastridae and in most Chalcidoidea and Cynipoidea

Some гесепtlу iпvеstigаtеd iпdерепdепt character suite5 5uggest аltегпаtivе геlаtiопshiрs Ьеtwееп the аЬоуе taxa А row of erect setae opposing the distal lamui iп the hind wiпg was fоuпd in the Chalshycidoidea Сегарhгопоidеа Diapriidae апd the undeshyscribed New Zealand family (Basibl1yuk and Quicke 1997) апd simultaneo1lS апtеппal gгооmiпg оссшs iп the Megaspilidae Chalcidoidea Мушагidае Суshynipoidea Proctotrupoidea апd sоше Platygastroidea (behavior 4 Basibl1Yl1k and Ql1icke 1999Ь) Gibson (1999) has reviewed hypotheses сопсегпiпg the sister grol1p геlаtiОП5Ырs of chalcidoid wasps Не did not fiпd сопviпсiпg support for RasnitsYl1s (1988) platyshygastroid + (chalcidoids + mушагоmmаtоids) hypothshyesis bl1t instead for а clade compri5ing Platygastroishydea Proctotrupoidea Pelecinidae and Vanhorniidae Не a1so sнggested that the Chalcidoidea and Муshymагоmшаtоidеаmау Ье тоге closely related to either the Diapriidae or Сегарhroпidае rather than to t1le Platygastroidea

We fОl1пd а spur at the base of еас SТВ (character 6) which seems to restrict the mobility of the 1Iюг

ф

w IltJ

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+ + + ~ ~~v~+ь+ + ~I + + ~ ь I + + b~ middotamp~~vI+ + $+o~ ~I+ ~~~ + + ~Ifgtb+ ~~ +lt q + + +~f + bfgt 90 ~ьф()~ ZmiddotI+ I +

raquo~+О~~vt~(~lt-~()Ь~~~+~~tь~ь~~Ь~1ь~~O~~~~~t~~fmiddot~~о$~+ ~~$Ь~Ь~~~~~О~()Ь~~=~~~$ь~ьt~~~~Ь~~~О--~Ь~~~~~ЬО~6~~~q~1~~~~~~~~~~~lt)~O~~~~t~q~~lt~qОltgtO~О~~ltgt~~O~middotampQvQОО-gtltf~q~~amplt~~~~~~lti)-~О~~~ltlt6~~fI~o~~Q~Qo~~~iо~lt)ф1~~о~lti~1lti-gt~(~OJ

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sect n

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bull bull bull bull bull 11 JAI 1 1 1 1 1 1 1 I 1 1 +1 2

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2

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сgtFig12 Unambiguous character state changes оЕ orbicula and associated structures superimposed оп the most recent phylogenetic hypothesis ofRonquist et аl (1999) resulting

ijfrom impJied weight analysis Numbers оп the cladogram correspond to the character numbers (ТаЫе 1) Character state changes for each character are explained in the text

w

~- -А д j ~ л ~ r ~ _ ~ gt -- -1 iI- _- 1 1-1-1 1- 1 ~ olJc j J

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

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Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

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DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

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Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

Konigsmann Е 1976 Das phylogenetische Systcm der Нуshymenoptelmiddota Теil IEinftihrung Grundplanmerkmale Schwestergruppe und Fossilfunde Dtsch Епtопюl Z23 251-279

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Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

626 ANNALS OF ТНЕ ENTOMOLOGICAL SOCJEТY OF АМЕШСА Vol 93 по 3

Ralec et al 1996) whole body parts (Sсhшidt and Sшith 1987) and legs (Navasero andElzen 1991 Меуshyhofer et al1997) but in еасЬ case tllese Ьауе included only опе or а few species ofрашsitiс wasps Studies of l1Оst searching and detection Ьу parasitic wasps Ьауе largely Ьееп related to receptors оп the antcnna and ovipositor and the possible role of tarsal sепsШа has only recently Ьееп suggested (МеуЬofег et al 1997)

In tMs article уе survey the detailed шогрhоlоgу of the orbicula and its associated sепsiПа throughout the Нушепорtега Тhe orbicula is а dorsal cuticular scleshyrite situated basal to the агоliuш in tlle claw-bearing tarsal sеgшепt Уе report phylogenetic implications of огЬiculю structurcs including sensilla and discuss thcir evolution in thc light of current phylogenetic hypotheses Comparisons bctween sexes shov that orbicular scnsilla ате not diffегепtiаПу developed in fешаlеs and therefore probably do not play а particshyular role in I1О8t location in parasitic wasps

Materials and Methods

Scanning Electron Microsoopy Ргерагаtiоп Most specimens were examined Ьу sсаrшiпg electron mishycroscopy (SEM) but some binocular light microscopy observations were also involved particularly of large aculeate specimens Fore legs were гешоvеd апd transferred to absolute ethanol for at least 18 Ь then air-dried апd mounted before sputter coating In шоst cases ultrasonic сlеапiпg vas поt appHed to avoid breaking sensilla trichodea Rare sресimепs were ехshyашiпеd without coating usiпg an Епviгопшепtal СhашЬег МЮk АВТ 55 microscope Stапdагd SEM tесhпiquеs Ьауе the роtепtial drawback of fаiliпg to геуеаl sоше Нпе details such as the ргеsепсе of sепshysШа carnpaniformia especially iп sшall specimens vithout special treatment Тherefore the data ргеshysented for these sепsillа ShOllld Ье treated with some саге althollghwhеп visibIe their lосаtiопs were clear

Telminology and Material Exarnined IdепtiRсаshytiопs of sепsШа types are based оп Mclver (1985) and Frazier (1985) Microsculpture tегшiпоlоgy largely follows Аllеп and Ваll (1980) Material exarnil1ed is il1cluded as Аррепdiх 1

Results

Orbicula ТЬе orbicula (mal1llbrium) is а dorsal cllticular sclerite at the base of the mешЬгаl10US аюshyНит iп the telotarsus (clav-bearil1g sеgшепt) (Figs 1 апd 2 (Sсhшidt al1d Smith 1987 СанЫ and Воltоп 1988 Meyhofer et al 1997) In the Нушепорtега the orbicula is well developed althollgh its sculpture al1d the types пumЬеrs апd аггапgешепts of its sensilla уагу These are suшmаrizеd iп ТаЫе 1 апd describerl below

Surface Textllre of Olbicula (character 1) Insect il1tegumel1ts аге tlsually furпishеd with microsculpshyture геsultiпg fгош ерidегшаl сеll secretions Sculpshyture is llSually шапifеst as а polygonal and геgulю шеsh sуstеш but а wide range of variation and сотshyplete reduction mау осспr (Аllеп апd BalI1980) We

recognize three states as follows (1) scale-like with projection(s) а scale tips (Fig 1 state О in ТаЫе 1) (2) simple scales (Figs 2 апd 3 state 1 in ТаЫе 1) (3) virtually entirely SП1Qоth (Fig 4 state two in ТаЫе 1) State О is largely confined to the superfarnilies Xyeloidea Pamphilioidea Cephoidea and Тепthгеdishynoidea although it also occurs in а few Apocrita (ТаshyЫе 1) Тhe scales are usually regular in distriЬutiоп (Fig 1) Ьи ате SОlllеtiшеs шarkеdlу reduced Тhey are most evident оп the basal half of the orbicula Тhe scales usually possess а single plojection apically Ьи there are several iп а few tenthredinoids (Arge Blasshytiootorna Selaпdria) and ichneumonids (Labena Мetорiщ Phytodeitus Rhyssa) (Fig 1 arrow)

State опе occurs in тапу Symphyta and in some Apocrita notably in the Trigonalidae Stephanidae and тапу Ichnellmonoidea (Table 1) Tlйs usually forms а regнlar mesll Ьн both slightly traI1sverse (Fig 3) апd markedly elongate scu]pture aso occurs (Fig 2) А fev proctotrupoid families notably the Helorishydae and Roproniidae Ьауе weakly deve]oped scaleshylike ll1icrosculpture Ьн the basa half of the orbicula is sшооth in Heloridae and ШS SOll1e deI1tiform scaes in Rоргопiidае (Fig 5 aIТOW) Most parasitic taxa and а [еу savflies laуе а сошрlеtеlу slllooth orbicllla (state 2 ТаЫе 1)

Orbicular Sепsillа We Ilave identiRed four types of sепsillа оп the orbicllla Two are setiforlll sensilla trishychodea А (STA) and sепsillа trichodea В (SТE) and the other two аге dome-like sепsillа сашрапifогmiа А (SCA) al1d sensilla саmрапifО]miа В (SCB) In most cases сотраПsoп of their shapes reative роsitiопs ll11mber and socket stПlсturе enabled llS to postulate homologies Ьеtwееп taxa uпсегtaiпtiеs are indicated in the text vhепеvег песеssагу

STA (character 2) Tllese structures arise from flexshyible sockets Ьауе the seta truпk gradually tapering toward the tip and ате щиаllу strопglу 10пgitlldil1аllу grooved (Fig 2) Т1lеу сап Ье distingнished from sепsiJlа trichodea В (SТE) Ьу роssеssiоп of а wеПshydefined socket Ьу llSually Ьеiпg шuсh lопgег and thicker (as тисЬ as three tiшеs) and Ьу Ьеiпg located шоге distally (Figs 2 апd 4 STA cf SТE) They ате alvays Ьеп toward the атоНит to sоше degree al1d llsually overlap it (Fig 4) Iп а few taxa notably the Siricidae where there шау Ье опlу а single sel1silla trichodea or only опе type whеп tlere ате several relative size could поt Ье used to help determine which type they were altlIough other morphological aspects suggest that they ате STA (Fig 6)

STA аге ргеsепt iп almost аН Hymenoptera (ТаЫе 1) Sexual dimогрhisш occurs most поtаblу in the Siricidae in which females usually Ьауе по or оссаshysiопаllу only опе STA although males usually Ьауе hvo or шоге However Sirex fell1ales сап sOllletimes ЬаУе six STA (Fig 6) Similarly in some of tlIe Асshyuleata that display extreme sexual dimorpllism (eg Tiphiidae) fешаlеs uSllally have reduced orbicula апd thlls Ьауе по sепsillа trichodea

Most Hymenoptera possess two STA (see ТаЫе 1 and аЬоуе for ехсерtiопs) bllt only а single SТA is present iп аН chalcidoids (including mymarids ыlt

627 Мау 2000 BASIВUYUK ЕТ ЛL ОНВICULЛ OF HYМENOPТERA

ТаЫе l Character states Сог the orblcula tшd associated веп- ТаЫе 1 Continued Ша u the Hymenoptera

Charactersl

Characters(1 Таха Таха 2 3 4 5 6 7 8 9

2 3 4 5 6 7 8 9 Sclerogibbidae 1 2 3 О о о з О О

Xyelidae Dryinidae 2 2 3 О О О О О ХуеНпае О О О О О ЕmЬоlешidае 2 2 3 О о о о о 1vfacroxyelinae 1 2 О 2 О 01 Betlylidae 2 2З 3 О О О О О

Xyelecia 1 2 2 О О О Chrysididae 112 2 03 О О О 02 О О Pamphiliidae Tiphiidaeb 02 023 О О О

Cephalciinae О 2 О 2 О О Mutillidae 2 2 3 О О О о О Раmрhiliiпае О 2 23 О О О О Sapygidae 2 3 О О О

Megalodonteidae О 2 1 О О О О Seoliidae 1 2 3 О О О О Megalodnntes О 2 О Bradynobaenidae 2 02 01] О О О О

kuznetzovi Sierolomorphidae 2 3 О О О Argidae 2О О О О RllOpalosomatidae О 2 01 О О О О Bla5toeotomidae О 2 О О О Formicidae 2 2 03 О О О 02 О О Cimbicidae О 2 011 О О О О Pompilidae 2 3 3 О О О О О Diрriопidае О 2 23 О О ОО Vepidae 1 2 О О О О Pergidae Specidae О 2 3 О О

Acordulecerinae 1 2 3 О О О 2 О О Apidae 1 2 123 О О О PIylacteltgtphaga 1 2 О 2 О О Perga 1 2 О 2 О 01

Chamcters and character 5tate (1) Orbicua texture О sltulpshylорlщrоtоmа 1 2 О ОР о tured primarily vith scale but а least ome scale usually а Ьмal асеаТепthrеdiпidае О 2 О О О vith cutic1l1ar projections 1 sculptured entirely vith 5cales 2 almost Ccphidae О 2 03 О О О О О entirely mooth (2) Conspicllously socketed sensila trichodea АAnaxyelidae 2 О О О (STA) оп orbicula о попе 1 опе2 two 3 tlgtreeormore (3) Sensilla Siricidae 0123 О О О trichodae В о попе 1 опе 2 hvo 3 three or more (4) Sensilla Xiplgtydriidae trichodea В (two or тоте) О = scattered 1 symmetrical or formsXiphydriinae 1 2 01 О О О О а single ~ow (5) Sensilla trichodea В О simple 1 Ьщallу enlarged (6) Dereeyrtinae 1 2 1 О О О О Senilla trichodea В О witlOllt а Ьаза cuticular projection 1 а basalOrussidae 1 О О О О

Cuiglia 1 2 О О cuticHlar projection (7) Sensilla campalliformia А (with ап eneircling scшuinslаndii ridge) О попе 1 опе 2 (уо 3 three or тоте (8) SепsШа сат-

Stephanidae 1 02 03 О О О 02 О О paniformia В (vithout ап encirclillg ridge) О попе 1 two (9) Trigonalidae 1 213 3 О О О 2 О О Position of sensila саmрапifоrmiа [о sensilla trichodea А О anterior Megalyridae 1 ОЗ 3 О О О 1 О О 1 level with or posterior Еуапнаае 1 2 3 О О О О О ь Sexual diшоrрllism (see Results)

О

Aulacidae 1 2 3 О О О О О GastеruрШdае 1 2 3 О О О О О absent in sоше agaonids) (Fig 7) шушагошmаtidsIbaliidac 2 2 3 О О О О О Figitidae 2 2 013 1 1 О О О diapriids the undescribed New Zealand family and Liopteridae 2 2 3 1 1 О О О possibly Ceraphronidae This single sепsiПuт is al-Cyuipidae 2 2 023 1 1 О О О ways situated оп the apical halfofthe orbicula and hasMegaspilidae 2 3 3 О О О О О Ceraphronidae 2 1 013 О О О а distinct high соllаг to its socket wmch is angled Diapriidae 2 1 2 1 1 О О 1 О distally (Fig 8) This соllаг шау serve to restrict back-Heloridae 1 2 3 О о о О о ward movements oftlle sensillum Three or more STA Vаrtlюrniidае 2 2 3 1 О О О юе found in some Siricidae Trigonalidae MegalyrishyРеlееiпiclае 1 2 3 О О О О О Proctotrupidae 2 02 23 О О О О dae Mcgaspilidae Ichneumonoidea and some ас-Roproniidae 011 2 3 О О О О О uleate taxa most notabIy the Pompilidae In this last Мопоmасhidaе 1 2 3 О О О О ОР family there are usual1y six or more STA which formNev Zealand ЕатНу 2 1 2 1 1 1 О 1

а transverse apical row (Fig 9) Sсеliопidае 2 2 013 О О О О 1 О Platygastridae 2 07 2 ] 1 О О Diffelentiating between STА and SТВ in those taxa Chalcididae 2 ] 3 О 1 О 1 with three or more STA becomes increasingly diffishyО Leucospidae 2 1 023 О 1 О спlt and ассшаtе assessment of the number of еасЬEurytomidae 2 1 3 1 1 1 О 1 1 Ormyridae 2 1 3 1 1 1 О 1 type is not always possibIc (particularly in Braconidae Pteromalidae 2 1 3 1 1 О 1 1 and Megaspilidae) Sucll states are indicated Ьу () iп Eupelmidae 2 1 3 О 1 1 ТаЫс 1 (see characters 2 З 4 and 5)Епсуrtidае 2 1 3 1 STB (cllaracter 3) SТВ пsиаllу сап Ье differenti-Agonidae 2 О О Aphelinidae 2 ] 3 1 1 ated from the STA Ьу being shorter al1d more slender Elasmidae 2 1 2 1 1 О 1 having а less pronounced socket (not obvious in manyEulophidae 2 1 23 1 1 1 О 1 sшаП specimens) and а more posterior and irregularSigniphoridae 2 1 3 1 1 О Trichogrammatidae 2 1 2 1 1 07 1 distribution ТЬеу are usually а siшрlе cylindrical Ьюr Torymidae 2 1 3 2 1 О 1 in а socket (Figs 2 5 9) bllt in some taxa they аге Mymaridae 2 1 2 1 1 о l widened and complessed basally (Fig 10) ТЬе posshyМуmагоmшаtidае 2 1 2 1 sibility that there тау Ье two distinct types of SТВBraconidae 112 23 3 О О О 2 О 1

Liobracon 2 3 О О 3 О 01 cannot Ье determined withollt llltгastruсtшal and Ichneumonidae 01 213 0111213 О О О 23 О 01 electrophysiological work and we rccognize only one Plumatiidae 2 2 3 О О О О О type SТВ аге present only in а few sушрhуtan taxaScolebythidae 1 2 17 О О О О but аге posscssed Ьу тапу Apocrita (ТаЫе 1)

628 ANNALS OF ТНЕ ENTOMOLOGICAL SOCIEТY OF АМЕтСА Vol 93 по 3

Figs l-3 (1) ОгЫсlllа in Sеflщiа sp (Tenthredinidae) STA sensilla tгiсhоdеа А arrow head sho ing scale-like SСlllрtше Nith cllticlllar projections Ваг = 30 т (2)Orbicula in Arotes arnoentL~ (Ichneumonidae) STA three sensilla tJiсlюdеа А black aJrow sel1silla tlichodea В white arrow sel1silla саmралifогтiа [уре А Ваг = 38 т (3) Close-up of orbicula tехtше il1 Регgо dorsafis (Pergidae) showing scale-like sculpture and two sensilla сатралifогтiа А (апоws) Ваг = б т

ТЬеге аге t1Sшllу three ог тоге SТE (Figs 5 7 9 ТаЫе 1) although only two аге found in а few sawflies some parasitic wasps апс some tiphiids and apids А single SТE occurs in а питЬе of isolated cases (Figs 2 апс 4 ТаЫе 1)

AITangement ofSTB (character 4) STB аге usually (Iistributed irregularly (Figs 5 and 9) but they fопп а single IOW in almost аll cynipoids (except IbaJiiclae) chalcidoids (except at least some Chalcidiclae ап(1 Leucospidae ) Diapriidae Vanhorniidae the undeshyscribed NeN Zealand family Platygastridae and some Ploctotrupidae (Fig 7)

Basal Modifjcation of STB (characteI 5) ТЬе SТE аге wi(lened апс laterally compressed in the Chalshycidoidea Diapriidae Platygastridae the undescribed New Zealand family апс some Cynipoidea (Figs 7 and 10)

Spur of STB Socket (chaJacter 6) ТЬе se nsilla socket bears а spur Nhich appears to restrict its backshyward mobi1ity in most chaJcidoid families including

mymarids together ith the uшlеSСliЬеd New Zeashyап(l famil y (Figs 78 10) А similar spur is present in the Cynipoidea but it is situated тоге distantly from the sensilla ТЬе 110mology of the spurs in (Ье Суshynipoidea witIl those in the other taxa is therefore unceгtain

Sensilla Сшпрапifогтiа (SC) (characters 7 and 8) Two types of sensilla campanifOlmia аге distinguishshyаЫе апс аге termed Ьеге SCA and SCB SCA is usually а roundecl dome encircled Ьу а cuticular ridge (Figs 2 апcl 3) whereas SCB has по external el1circling lidge аl1с арреаг as а simple small tubercle (Figs 4 and 8) SCA aJe Гоиl1с il1 the Trigonalidae апс Pergidae (ехshycept lлрlчrоtОm()) апс in тапу species of BJashyconidae апс Ichl1eumol1idae ТЬеу also are present il1 some replesel1tatives of several isolated taxa (Xyelishydae Pamphiliklae Stephal1idae Megalyridae Sclerogibbidae Chrysididae апс Formicidae) (ТаЫе 1) SCB аге possessed Ьу the Chalcidoidea Diapliidae and some species of Sceliol1idae al1d possibly Ьу the

629 Мау 2000 BASIВUYUК ЕТ AL ORВICLJLA 0[ HVMENOPТElA

Figs4-7 (4) ОЪi Сl1 lа11 Triteleio sp (SceliOl1idae) vith two sel1silla trichodeaA (ореl1 апоs) sil1gle sensilla tichoclea В (hite arro) аl1сl two sel1silla campalliformia type В (white shadovecl bIack arrows) Bar = 8 fLl11 (5) Orbicula il1 RopTonio gопnаni (Roproniiclae) Ilote clel1tifoCI1l sca les (arтo) Ваг = 30 fLl1l (6) Orbiclla ill Ггсmех sp (Sricidae) shoving presl1l11ed six sel1silla trichodeaA Ваг = 50 fLl1l (7) Orbicl1la ill Ptoth -ix sp (Pteromalidae) sho ingsil1 g1e sel1silla trichodea А (ореl1 апоw) and а tansverse JO W of sellsilla tcl)Qclelt1 В (smзl1 arrows) Ваг = 13 fLl11

undescribed New Zealand family АI1 (аха that have sensiJla campalliformia Iave only опе type with tlle possible exceptioll иоьгасоn (Braconidae) (Fig11) The great majority oftaxa witJl SC (either А ог В) Ьауе two of them but only а sillg]e SCA is foulld ill megashyIyrids апс three ог тоге occur in some pergids Ьгаshycollids sclerogibbids апс тапу ichneumonids

Positioll of SC ( сhагасtег 9) SCA аге usua]]y роshysitioned iпеgulагlу апс distal to STA Howeer they сап Ье both distal аllcl basltll (о STA ill LVlacroxyel (MacroxyeJinae) Perga (Pergide) апе] тапу IchshyneumOlliclae (ТаЫе 1) Positiolls of SCB аге quite

cOllsistellt within the Chakidoideltl аllс Diapriidae ill vhich they аге always 10саtеclllttегоvепtгаJlу Jelative to the STA оП eitheJ siele of JateraJly compressed orbicula level witJl 01 sliglltly basal to it (Fig 8) TI1e SCB of sceliollids however ltге locateel clistal (о the STA (Fig 4)

Discussioll

Functiolla Implicatiolls The orbiculal sellsillt аге lttrgllably all mechalloreceptors (McIver 1985) 111 th e eulophicl Sympiesi~ seiceicomi~ Nees they have Ьееп

630 ANNALS OF ТНЕ ENTOMOLOC1CAL SOCIEТY OF AМERJCA Vol 93 п о 3

Figs8-11 (8) 5e nsilla ill Jurywma sp (Ешуtоm idае) 5ТА sensilla tri chodea А wi th а wellmiddot (leve loped socke t 5ТВ sefJS illa tic llOdea В white 3IТOW se lls illa саmралi fогmi а type В Bar = 3 Lm (9) CeГOfo les sp (Роm рШdае) SllO v ing pos ition зп(1 аrrалgеmепt of sensilla tricllOdeaA Bar = 38 LПJ (10) Sелsillа tichodea В of Castmcantl-s рсетjlIШI (Pterom alidae) Sl10ving widened base and basal spur (ореп апоws) Вш = 3Lnl (11) Orbicula in иоЬгас(т sp (Braconidae) sho-ing bot11 types of se nsilla саmрап ifогmiа vl)ite апоws ргеSLmеd sел sillа campan iformia В white shadove(1 bIack arrovs sensilla campan ifomia А Ва = 20 Lm

suggested as шviпg а role [п de tecting vibration s ргоmiddot ducecl Ьу th e ir leafminer host (Meyhbfer e t al 1997) Althougl this тау gtell Ье the case theiI prese nce [п taxa with diffe le nt blologies (incllding phytoph agollS taxa a llC раГltlsitоid s of deeply conceaed hosts) sugshygests that they have otl er гoles as well ТЬе ргеsеп се of apparen tly identi ca sensila iп both sexes of тапу taxa fшthеl weake ns tle host locatio) hypotllesis Because ST and SC аге prese nt in th e most basal ineages ofthe H ymenoptera (eg MacJQxyeinae and Cephalciinae) we suggest that the iJ огigiп а l and ргеshy

cominant fun ction is топitогiпg moveme nts and s tгеss resulting fгош the asps own 10comotion anс possibly also subst)a te viblation that тау indicate the ргеsепсе of pote) tia pledatoгs Athough they тау play а role [п host de tection the ack of sexua diтогshyphism i) рагаs itiс taxa suggests that this is not а majoJ need Indeed the sexua dinlOrphism in th e phytophshyagous Siricidae and iп th e рагаs itiс Tipl iidae both cases vit females having fewe r STA suggests а gleate J iшрогtапсе fOl th e maes Моге obselvations оп paras itoid host sею-сhiпg behavioJ includi ng еесshy

631 Мау 2000 BASmUYUK ЕТ ЛL ORВICULA OF НУМENОPrERЛ

trophysiological studies are needed to determine tle exact function of these sensilla

Рhуlоgепеtic Imрliсаtiопs То assess рhуlоgепеtiс implications ofthi5 character system we have mapped the data onto the most recent phylogenetic hypothesis of the епtiге order re5ulting from implied weighting (Ronquist et а11999) Only the uпаmЬiguоus changes were indicated and polytomies were treated as Iard (ie аssumiпg multiple speciation events) (Fig 12) We have included the undescribed New Zеаlапd famshyНу iп the putative clade comprising Chalcidoidea Cershyарhroпоidеа Diapriidae and 50 оп because it shares at least опе putative sупароmогрhу with mепtiопеd taxa (Basibuyuk апd Quicke 1997) We also have plotshyted the data оп to Rasnitsyns (1988) intuitive hypothshye5i5 and оп to а tree resulting from reanцJysis of а соmЫпаtiоп ofRasnitsyns data (Ronquist ct а1 1999) with VHhelmsen s (1997) sуmрhуtап апd Brother5 and Сагрепtегs (1993) aculeate hypothese5 However the fit of the data were marginally better оп the latest hypothesis of Rопquist et а1 (1999)

Orbicular Sculpture The orbiculas of тanу of the most Ьа5аlliпеаgеs (eg the Xyelinae Pamphiloidea Cephoidea апd Tenthredinoidea other than pergids) have scale-like microsculpture with cuticular projecshytion5 (character 1 5tate О) sugge5ting thi5 to Ье the plesiomorphic condition for the order Simpfe scaleshylike 5culpture without рrojесt10ПS (character 1 5tate 1) appears first 1П the siricoid grade апd i5 геtаiпеd in the Evaniomorpha sеП5U Rasпitsуп рщ of the Procshytotrupoidea (РеlеС1П1dае Heloridae Rоргопiidае Мопоmасhidае) апd 1П mапу Iсhпеumопidае It also 15 ргеsепt 1П the Масгохуеliпае and Pergidae which сап Ье best сопsidегеd as iпdерепdепt acquisitions Further reduction of the microsculpture to а сотshypletely 5mooth orbicula (character 1 state 2) has арshyparently occurred оп at least three separate оссasiопs iп рщ of the miсгоhуmепорtегап lineage iпсll1diпg the Супiроidеа Diapriidae the uпdеsсгiЬеd New Zеаlапd family Platygastroidea Сегарhгопоidеа Муshymarommatidae апd Chalcidoidea iп the pl1tative Proctotrl1pidae + Vanhorniidae liпеаgе and in the Aculeata ТЬе presence of some simple scale-like sсulрtше (charactcr 1 state 1) in the Sclerogibbidae Scolebythidae апdVe5pidae сап best Ье considered as reversals (Rasnitsyn 1988 Brothers and Сагрепtег 1993 Ronquist et al 1999)

Sепsillа Trichodea А STА аге ргеsепt iп almost аН taxa оfНуmепорtега except Хуеliпае and their presshyепсе is therefore а putative synapomorphy for the Нуmепорtега ехсludiпg the ХуеНпае АЬsепсе of STA in а few isolated taxa in which they are и511аНу polymorphic (5iricids orussids stephanids megalyrshyids proctotrupids platygastrids ЬгаdупоЬаепids апd tiphiids) сап be5t Ье regarded as iпdерепdепt losscs (Fig 12) Тhe Пl1mЬег ofSTA i5 overwhelmingly two and this

сопditiоп (character 2 state 2) is most likely the gгоuпd рlап for tlle Нушепорtега as а whole Reducshytion iп the number of STA to опе (character 2 state 1) with а markedly raised socket is а pl1tative synashyротогрЬу for а clade соmрrisiпg Chalcidoidea Муshy

mагоmшаtоidеа Diapriidae Ceraphronidae and the undescribed New Zеalапd family but with опе revershysal (iп Супiроidса) and at least опс tгапsitiоп to other states (in Megaspilidae) Three ог more STA occur in some Trigonalidae Megalyridae Megaspilidae Brashyсопidае Ichпеumопidае апd Pompilidae bl1t iп most of the5e cases the Iikely gгоuпd рlап i5 two STA Ьеshycause that state occurs iп тanу trigопаlids iсhпеushymопids as well as in some Ьгасопids (eg аgаthidiпе gепега Labagathis and L~opronotum) ТЬе ргеsепсе of thrce or more STA in the Mega5pilidae and Pompilishydae i5 best cOl1Sidered to герге5епt two indерепdепt асquisitiопs (Fig 12)

Sensilla Trichodea В апdRelated Structures Тhese are present in most taxa but their two 5eparate аЬshysепсеs and variаtiоп in their пumЬег (character 3) appear to iпdiсаtе sоше hошорlаsу iп the Apocrita for the givеп 5et of рhуlоgепеtiс hypotheses However possession ofthree ог тоге STA (character 3 state 3) provides support for the monophyly of Apocrita (Fig 12) Аrrапgеmепt of the SТВ iп а siпglе row (Fig 7 character 4 state 1) 511pports the clade соmргisiпg the Chalcidoidea Mymarommatoidea Ceraphronoidea Platygastroidea Cynipoidea Diapriidae and the uпshydescribed New Zealand fашilу (Fig 12) This state a1so occurs iп the Vanhorniidae and Pamphilioidea Тhe оссипепсе ofdi5persed SТВ (character 4 state О) iп the Chalcididae Lel1cospidac and Ibaliidae (which аге relatively larger iп 5ize) within thi5 clade тау sl1ggest sizе-dерепdепt сопvегgепсе for this characshyter but there i5 а gгоwiпg body ofеvidепсе supporting some of the relationships amопg the аЬоуе putative clade (Basibuyнk апd Quicke 1997 1999Ь GiЬsоп 1999 Rопquist 1999 Ronquist et а1 1999) It should Ье поtеd that SТВ a1so аге di5persed in some scelionids ТЬе SТВ in some of these taxa are modified basally (character 5 state 1) being laterally compressed and widепеd These modified SТВ аге ргеsепt in the Diashypriidae the uпdеsсгiЬеd New Zealand family Platyshygastridae and in most Chalcidoidea and Cynipoidea

Some гесепtlу iпvеstigаtеd iпdерепdепt character suite5 5uggest аltегпаtivе геlаtiопshiрs Ьеtwееп the аЬоуе taxa А row of erect setae opposing the distal lamui iп the hind wiпg was fоuпd in the Chalshycidoidea Сегарhгопоidеа Diapriidae апd the undeshyscribed New Zealand family (Basibl1yuk and Quicke 1997) апd simultaneo1lS апtеппal gгооmiпg оссшs iп the Megaspilidae Chalcidoidea Мушагidае Суshynipoidea Proctotrupoidea апd sоше Platygastroidea (behavior 4 Basibl1Yl1k and Ql1icke 1999Ь) Gibson (1999) has reviewed hypotheses сопсегпiпg the sister grol1p геlаtiОП5Ырs of chalcidoid wasps Не did not fiпd сопviпсiпg support for RasnitsYl1s (1988) platyshygastroid + (chalcidoids + mушагоmmаtоids) hypothshyesis bl1t instead for а clade compri5ing Platygastroishydea Proctotrupoidea Pelecinidae and Vanhorniidae Не a1so sнggested that the Chalcidoidea and Муshymагоmшаtоidеаmау Ье тоге closely related to either the Diapriidae or Сегарhroпidае rather than to t1le Platygastroidea

We fОl1пd а spur at the base of еас SТВ (character 6) which seems to restrict the mobility of the 1Iюг

ф

w IltJ

ья

+ + + ~ ~~v~+ь+ + ~I + + ~ ь I + + b~ middotamp~~vI+ + $+o~ ~I+ ~~~ + + ~Ifgtb+ ~~ +lt q + + +~f + bfgt 90 ~ьф()~ ZmiddotI+ I +

raquo~+О~~vt~(~lt-~()Ь~~~+~~tь~ь~~Ь~1ь~~O~~~~~t~~fmiddot~~о$~+ ~~$Ь~Ь~~~~~О~()Ь~~=~~~$ь~ьt~~~~Ь~~~О--~Ь~~~~~ЬО~6~~~q~1~~~~~~~~~~~lt)~O~~~~t~q~~lt~qОltgtO~О~~ltgt~~O~middotampQvQОО-gtltf~q~~amplt~~~~~~lti)-~О~~~ltlt6~~fI~o~~Q~Qo~~~iо~lt)ф1~~о~lti~1lti-gt~(~OJ

gtz z gtshyt о

~ tj z о о r о

~ ~ rл о

sect n

о

gt

bull bull bull bull bull 11 JAI 1 1 1 1 1 1 1 I 1 1 +1 2

7 I I 6

7Т I т

2

4

4Т I I I I I I I

7 I I I I I I 1 3 gJ (j gtshy

~ ф

сgtFig12 Unambiguous character state changes оЕ orbicula and associated structures superimposed оп the most recent phylogenetic hypothesis ofRonquist et аl (1999) resulting

ijfrom impJied weight analysis Numbers оп the cladogram correspond to the character numbers (ТаЫе 1) Character state changes for each character are explained in the text

w

~- -А д j ~ л ~ r ~ _ ~ gt -- -1 iI- _- 1 1-1-1 1- 1 ~ olJc j J

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

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Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

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Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

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Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

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Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

627 Мау 2000 BASIВUYUK ЕТ ЛL ОНВICULЛ OF HYМENOPТERA

ТаЫе l Character states Сог the orblcula tшd associated веп- ТаЫе 1 Continued Ша u the Hymenoptera

Charactersl

Characters(1 Таха Таха 2 3 4 5 6 7 8 9

2 3 4 5 6 7 8 9 Sclerogibbidae 1 2 3 О о о з О О

Xyelidae Dryinidae 2 2 3 О О О О О ХуеНпае О О О О О ЕmЬоlешidае 2 2 3 О о о о о 1vfacroxyelinae 1 2 О 2 О 01 Betlylidae 2 2З 3 О О О О О

Xyelecia 1 2 2 О О О Chrysididae 112 2 03 О О О 02 О О Pamphiliidae Tiphiidaeb 02 023 О О О

Cephalciinae О 2 О 2 О О Mutillidae 2 2 3 О О О о О Раmрhiliiпае О 2 23 О О О О Sapygidae 2 3 О О О

Megalodonteidae О 2 1 О О О О Seoliidae 1 2 3 О О О О Megalodnntes О 2 О Bradynobaenidae 2 02 01] О О О О

kuznetzovi Sierolomorphidae 2 3 О О О Argidae 2О О О О RllOpalosomatidae О 2 01 О О О О Bla5toeotomidae О 2 О О О Formicidae 2 2 03 О О О 02 О О Cimbicidae О 2 011 О О О О Pompilidae 2 3 3 О О О О О Diрriопidае О 2 23 О О ОО Vepidae 1 2 О О О О Pergidae Specidae О 2 3 О О

Acordulecerinae 1 2 3 О О О 2 О О Apidae 1 2 123 О О О PIylacteltgtphaga 1 2 О 2 О О Perga 1 2 О 2 О 01

Chamcters and character 5tate (1) Orbicua texture О sltulpshylорlщrоtоmа 1 2 О ОР о tured primarily vith scale but а least ome scale usually а Ьмal асеаТепthrеdiпidае О 2 О О О vith cutic1l1ar projections 1 sculptured entirely vith 5cales 2 almost Ccphidae О 2 03 О О О О О entirely mooth (2) Conspicllously socketed sensila trichodea АAnaxyelidae 2 О О О (STA) оп orbicula о попе 1 опе2 two 3 tlgtreeormore (3) Sensilla Siricidae 0123 О О О trichodae В о попе 1 опе 2 hvo 3 three or more (4) Sensilla Xiplgtydriidae trichodea В (two or тоте) О = scattered 1 symmetrical or formsXiphydriinae 1 2 01 О О О О а single ~ow (5) Sensilla trichodea В О simple 1 Ьщallу enlarged (6) Dereeyrtinae 1 2 1 О О О О Senilla trichodea В О witlOllt а Ьаза cuticular projection 1 а basalOrussidae 1 О О О О

Cuiglia 1 2 О О cuticHlar projection (7) Sensilla campalliformia А (with ап eneircling scшuinslаndii ridge) О попе 1 опе 2 (уо 3 three or тоте (8) SепsШа сат-

Stephanidae 1 02 03 О О О 02 О О paniformia В (vithout ап encirclillg ridge) О попе 1 two (9) Trigonalidae 1 213 3 О О О 2 О О Position of sensila саmрапifоrmiа [о sensilla trichodea А О anterior Megalyridae 1 ОЗ 3 О О О 1 О О 1 level with or posterior Еуапнаае 1 2 3 О О О О О ь Sexual diшоrрllism (see Results)

О

Aulacidae 1 2 3 О О О О О GastеruрШdае 1 2 3 О О О О О absent in sоше agaonids) (Fig 7) шушагошmаtidsIbaliidac 2 2 3 О О О О О Figitidae 2 2 013 1 1 О О О diapriids the undescribed New Zealand family and Liopteridae 2 2 3 1 1 О О О possibly Ceraphronidae This single sепsiПuт is al-Cyuipidae 2 2 023 1 1 О О О ways situated оп the apical halfofthe orbicula and hasMegaspilidae 2 3 3 О О О О О Ceraphronidae 2 1 013 О О О а distinct high соllаг to its socket wmch is angled Diapriidae 2 1 2 1 1 О О 1 О distally (Fig 8) This соllаг шау serve to restrict back-Heloridae 1 2 3 О о о О о ward movements oftlle sensillum Three or more STA Vаrtlюrniidае 2 2 3 1 О О О юе found in some Siricidae Trigonalidae MegalyrishyРеlееiпiclае 1 2 3 О О О О О Proctotrupidae 2 02 23 О О О О dae Mcgaspilidae Ichneumonoidea and some ас-Roproniidae 011 2 3 О О О О О uleate taxa most notabIy the Pompilidae In this last Мопоmасhidaе 1 2 3 О О О О ОР family there are usual1y six or more STA which formNev Zealand ЕатНу 2 1 2 1 1 1 О 1

а transverse apical row (Fig 9) Sсеliопidае 2 2 013 О О О О 1 О Platygastridae 2 07 2 ] 1 О О Diffelentiating between STА and SТВ in those taxa Chalcididae 2 ] 3 О 1 О 1 with three or more STA becomes increasingly diffishyО Leucospidae 2 1 023 О 1 О спlt and ассшаtе assessment of the number of еасЬEurytomidae 2 1 3 1 1 1 О 1 1 Ormyridae 2 1 3 1 1 1 О 1 type is not always possibIc (particularly in Braconidae Pteromalidae 2 1 3 1 1 О 1 1 and Megaspilidae) Sucll states are indicated Ьу () iп Eupelmidae 2 1 3 О 1 1 ТаЫс 1 (see characters 2 З 4 and 5)Епсуrtidае 2 1 3 1 STB (cllaracter 3) SТВ пsиаllу сап Ье differenti-Agonidae 2 О О Aphelinidae 2 ] 3 1 1 ated from the STA Ьу being shorter al1d more slender Elasmidae 2 1 2 1 1 О 1 having а less pronounced socket (not obvious in manyEulophidae 2 1 23 1 1 1 О 1 sшаП specimens) and а more posterior and irregularSigniphoridae 2 1 3 1 1 О Trichogrammatidae 2 1 2 1 1 07 1 distribution ТЬеу are usually а siшрlе cylindrical Ьюr Torymidae 2 1 3 2 1 О 1 in а socket (Figs 2 5 9) bllt in some taxa they аге Mymaridae 2 1 2 1 1 о l widened and complessed basally (Fig 10) ТЬе posshyМуmагоmшаtidае 2 1 2 1 sibility that there тау Ье two distinct types of SТВBraconidae 112 23 3 О О О 2 О 1

Liobracon 2 3 О О 3 О 01 cannot Ье determined withollt llltгastruсtшal and Ichneumonidae 01 213 0111213 О О О 23 О 01 electrophysiological work and we rccognize only one Plumatiidae 2 2 3 О О О О О type SТВ аге present only in а few sушрhуtan taxaScolebythidae 1 2 17 О О О О but аге posscssed Ьу тапу Apocrita (ТаЫе 1)

628 ANNALS OF ТНЕ ENTOMOLOGICAL SOCIEТY OF АМЕтСА Vol 93 по 3

Figs l-3 (1) ОгЫсlllа in Sеflщiа sp (Tenthredinidae) STA sensilla tгiсhоdеа А arrow head sho ing scale-like SСlllрtше Nith cllticlllar projections Ваг = 30 т (2)Orbicula in Arotes arnoentL~ (Ichneumonidae) STA three sensilla tJiсlюdеа А black aJrow sel1silla tlichodea В white arrow sel1silla саmралifогтiа [уре А Ваг = 38 т (3) Close-up of orbicula tехtше il1 Регgо dorsafis (Pergidae) showing scale-like sculpture and two sensilla сатралifогтiа А (апоws) Ваг = б т

ТЬеге аге t1Sшllу three ог тоге SТE (Figs 5 7 9 ТаЫе 1) although only two аге found in а few sawflies some parasitic wasps апс some tiphiids and apids А single SТE occurs in а питЬе of isolated cases (Figs 2 апс 4 ТаЫе 1)

AITangement ofSTB (character 4) STB аге usually (Iistributed irregularly (Figs 5 and 9) but they fопп а single IOW in almost аll cynipoids (except IbaJiiclae) chalcidoids (except at least some Chalcidiclae ап(1 Leucospidae ) Diapriidae Vanhorniidae the undeshyscribed NeN Zealand family Platygastridae and some Ploctotrupidae (Fig 7)

Basal Modifjcation of STB (characteI 5) ТЬе SТE аге wi(lened апс laterally compressed in the Chalshycidoidea Diapriidae Platygastridae the undescribed New Zealand family апс some Cynipoidea (Figs 7 and 10)

Spur of STB Socket (chaJacter 6) ТЬе se nsilla socket bears а spur Nhich appears to restrict its backshyward mobi1ity in most chaJcidoid families including

mymarids together ith the uшlеSСliЬеd New Zeashyап(l famil y (Figs 78 10) А similar spur is present in the Cynipoidea but it is situated тоге distantly from the sensilla ТЬе 110mology of the spurs in (Ье Суshynipoidea witIl those in the other taxa is therefore unceгtain

Sensilla Сшпрапifогтiа (SC) (characters 7 and 8) Two types of sensilla campanifOlmia аге distinguishshyаЫе апс аге termed Ьеге SCA and SCB SCA is usually а roundecl dome encircled Ьу а cuticular ridge (Figs 2 апcl 3) whereas SCB has по external el1circling lidge аl1с арреаг as а simple small tubercle (Figs 4 and 8) SCA aJe Гоиl1с il1 the Trigonalidae апс Pergidae (ехshycept lлрlчrоtОm()) апс in тапу species of BJashyconidae апс Ichl1eumol1idae ТЬеу also are present il1 some replesel1tatives of several isolated taxa (Xyelishydae Pamphiliklae Stephal1idae Megalyridae Sclerogibbidae Chrysididae апс Formicidae) (ТаЫе 1) SCB аге possessed Ьу the Chalcidoidea Diapliidae and some species of Sceliol1idae al1d possibly Ьу the

629 Мау 2000 BASIВUYUК ЕТ AL ORВICLJLA 0[ HVMENOPТElA

Figs4-7 (4) ОЪi Сl1 lа11 Triteleio sp (SceliOl1idae) vith two sel1silla trichodeaA (ореl1 апоs) sil1gle sensilla tichoclea В (hite arro) аl1сl two sel1silla campalliformia type В (white shadovecl bIack arrows) Bar = 8 fLl11 (5) Orbicula il1 RopTonio gопnаni (Roproniiclae) Ilote clel1tifoCI1l sca les (arтo) Ваг = 30 fLl1l (6) Orbiclla ill Ггсmех sp (Sricidae) shoving presl1l11ed six sel1silla trichodeaA Ваг = 50 fLl1l (7) Orbicl1la ill Ptoth -ix sp (Pteromalidae) sho ingsil1 g1e sel1silla trichodea А (ореl1 апоw) and а tansverse JO W of sellsilla tcl)Qclelt1 В (smзl1 arrows) Ваг = 13 fLl11

undescribed New Zealand family АI1 (аха that have sensiJla campalliformia Iave only опе type with tlle possible exceptioll иоьгасоn (Braconidae) (Fig11) The great majority oftaxa witJl SC (either А ог В) Ьауе two of them but only а sillg]e SCA is foulld ill megashyIyrids апс three ог тоге occur in some pergids Ьгаshycollids sclerogibbids апс тапу ichneumonids

Positioll of SC ( сhагасtег 9) SCA аге usua]]y роshysitioned iпеgulагlу апс distal to STA Howeer they сап Ье both distal аllcl basltll (о STA ill LVlacroxyel (MacroxyeJinae) Perga (Pergide) апе] тапу IchshyneumOlliclae (ТаЫе 1) Positiolls of SCB аге quite

cOllsistellt within the Chakidoideltl аllс Diapriidae ill vhich they аге always 10саtеclllttегоvепtгаJlу Jelative to the STA оП eitheJ siele of JateraJly compressed orbicula level witJl 01 sliglltly basal to it (Fig 8) TI1e SCB of sceliollids however ltге locateel clistal (о the STA (Fig 4)

Discussioll

Functiolla Implicatiolls The orbiculal sellsillt аге lttrgllably all mechalloreceptors (McIver 1985) 111 th e eulophicl Sympiesi~ seiceicomi~ Nees they have Ьееп

630 ANNALS OF ТНЕ ENTOMOLOC1CAL SOCIEТY OF AМERJCA Vol 93 п о 3

Figs8-11 (8) 5e nsilla ill Jurywma sp (Ешуtоm idае) 5ТА sensilla tri chodea А wi th а wellmiddot (leve loped socke t 5ТВ sefJS illa tic llOdea В white 3IТOW se lls illa саmралi fогmi а type В Bar = 3 Lm (9) CeГOfo les sp (Роm рШdае) SllO v ing pos ition зп(1 аrrалgеmепt of sensilla tricllOdeaA Bar = 38 LПJ (10) Sелsillа tichodea В of Castmcantl-s рсетjlIШI (Pterom alidae) Sl10ving widened base and basal spur (ореп апоws) Вш = 3Lnl (11) Orbicula in иоЬгас(т sp (Braconidae) sho-ing bot11 types of se nsilla саmрап ifогmiа vl)ite апоws ргеSLmеd sел sillа campan iformia В white shadove(1 bIack arrovs sensilla campan ifomia А Ва = 20 Lm

suggested as шviпg а role [п de tecting vibration s ргоmiddot ducecl Ьу th e ir leafminer host (Meyhbfer e t al 1997) Althougl this тау gtell Ье the case theiI prese nce [п taxa with diffe le nt blologies (incllding phytoph agollS taxa a llC раГltlsitоid s of deeply conceaed hosts) sugshygests that they have otl er гoles as well ТЬе ргеsеп се of apparen tly identi ca sensila iп both sexes of тапу taxa fшthеl weake ns tle host locatio) hypotllesis Because ST and SC аге prese nt in th e most basal ineages ofthe H ymenoptera (eg MacJQxyeinae and Cephalciinae) we suggest that the iJ огigiп а l and ргеshy

cominant fun ction is топitогiпg moveme nts and s tгеss resulting fгош the asps own 10comotion anс possibly also subst)a te viblation that тау indicate the ргеsепсе of pote) tia pledatoгs Athough they тау play а role [п host de tection the ack of sexua diтогshyphism i) рагаs itiс taxa suggests that this is not а majoJ need Indeed the sexua dinlOrphism in th e phytophshyagous Siricidae and iп th e рагаs itiс Tipl iidae both cases vit females having fewe r STA suggests а gleate J iшрогtапсе fOl th e maes Моге obselvations оп paras itoid host sею-сhiпg behavioJ includi ng еесshy

631 Мау 2000 BASmUYUK ЕТ ЛL ORВICULA OF НУМENОPrERЛ

trophysiological studies are needed to determine tle exact function of these sensilla

Рhуlоgепеtic Imрliсаtiопs То assess рhуlоgепеtiс implications ofthi5 character system we have mapped the data onto the most recent phylogenetic hypothesis of the епtiге order re5ulting from implied weighting (Ronquist et а11999) Only the uпаmЬiguоus changes were indicated and polytomies were treated as Iard (ie аssumiпg multiple speciation events) (Fig 12) We have included the undescribed New Zеаlапd famshyНу iп the putative clade comprising Chalcidoidea Cershyарhroпоidеа Diapriidae and 50 оп because it shares at least опе putative sупароmогрhу with mепtiопеd taxa (Basibuyuk апd Quicke 1997) We also have plotshyted the data оп to Rasnitsyns (1988) intuitive hypothshye5i5 and оп to а tree resulting from reanцJysis of а соmЫпаtiоп ofRasnitsyns data (Ronquist ct а1 1999) with VHhelmsen s (1997) sуmрhуtап апd Brother5 and Сагрепtегs (1993) aculeate hypothese5 However the fit of the data were marginally better оп the latest hypothesis of Rопquist et а1 (1999)

Orbicular Sculpture The orbiculas of тanу of the most Ьа5аlliпеаgеs (eg the Xyelinae Pamphiloidea Cephoidea апd Tenthredinoidea other than pergids) have scale-like microsculpture with cuticular projecshytion5 (character 1 5tate О) sugge5ting thi5 to Ье the plesiomorphic condition for the order Simpfe scaleshylike 5culpture without рrojесt10ПS (character 1 5tate 1) appears first 1П the siricoid grade апd i5 геtаiпеd in the Evaniomorpha sеП5U Rasпitsуп рщ of the Procshytotrupoidea (РеlеС1П1dае Heloridae Rоргопiidае Мопоmасhidае) апd 1П mапу Iсhпеumопidае It also 15 ргеsепt 1П the Масгохуеliпае and Pergidae which сап Ье best сопsidегеd as iпdерепdепt acquisitions Further reduction of the microsculpture to а сотshypletely 5mooth orbicula (character 1 state 2) has арshyparently occurred оп at least three separate оссasiопs iп рщ of the miсгоhуmепорtегап lineage iпсll1diпg the Супiроidеа Diapriidae the uпdеsсгiЬеd New Zеаlапd family Platygastroidea Сегарhгопоidеа Муshymarommatidae апd Chalcidoidea iп the pl1tative Proctotrl1pidae + Vanhorniidae liпеаgе and in the Aculeata ТЬе presence of some simple scale-like sсulрtше (charactcr 1 state 1) in the Sclerogibbidae Scolebythidae апdVe5pidae сап best Ье considered as reversals (Rasnitsyn 1988 Brothers and Сагрепtег 1993 Ronquist et al 1999)

Sепsillа Trichodea А STА аге ргеsепt iп almost аН taxa оfНуmепорtега except Хуеliпае and their presshyепсе is therefore а putative synapomorphy for the Нуmепорtега ехсludiпg the ХуеНпае АЬsепсе of STA in а few isolated taxa in which they are и511аНу polymorphic (5iricids orussids stephanids megalyrshyids proctotrupids platygastrids ЬгаdупоЬаепids апd tiphiids) сап be5t Ье regarded as iпdерепdепt losscs (Fig 12) Тhe Пl1mЬег ofSTA i5 overwhelmingly two and this

сопditiоп (character 2 state 2) is most likely the gгоuпd рlап for tlle Нушепорtега as а whole Reducshytion iп the number of STA to опе (character 2 state 1) with а markedly raised socket is а pl1tative synashyротогрЬу for а clade соmрrisiпg Chalcidoidea Муshy

mагоmшаtоidеа Diapriidae Ceraphronidae and the undescribed New Zеalапd family but with опе revershysal (iп Супiроidса) and at least опс tгапsitiоп to other states (in Megaspilidae) Three ог more STA occur in some Trigonalidae Megalyridae Megaspilidae Brashyсопidае Ichпеumопidае апd Pompilidae bl1t iп most of the5e cases the Iikely gгоuпd рlап i5 two STA Ьеshycause that state occurs iп тanу trigопаlids iсhпеushymопids as well as in some Ьгасопids (eg аgаthidiпе gепега Labagathis and L~opronotum) ТЬе ргеsепсе of thrce or more STA in the Mega5pilidae and Pompilishydae i5 best cOl1Sidered to герге5епt two indерепdепt асquisitiопs (Fig 12)

Sensilla Trichodea В апdRelated Structures Тhese are present in most taxa but their two 5eparate аЬshysепсеs and variаtiоп in their пumЬег (character 3) appear to iпdiсаtе sоше hошорlаsу iп the Apocrita for the givеп 5et of рhуlоgепеtiс hypotheses However possession ofthree ог тоге STA (character 3 state 3) provides support for the monophyly of Apocrita (Fig 12) Аrrапgеmепt of the SТВ iп а siпglе row (Fig 7 character 4 state 1) 511pports the clade соmргisiпg the Chalcidoidea Mymarommatoidea Ceraphronoidea Platygastroidea Cynipoidea Diapriidae and the uпshydescribed New Zealand fашilу (Fig 12) This state a1so occurs iп the Vanhorniidae and Pamphilioidea Тhe оссипепсе ofdi5persed SТВ (character 4 state О) iп the Chalcididae Lel1cospidac and Ibaliidae (which аге relatively larger iп 5ize) within thi5 clade тау sl1ggest sizе-dерепdепt сопvегgепсе for this characshyter but there i5 а gгоwiпg body ofеvidепсе supporting some of the relationships amопg the аЬоуе putative clade (Basibuyнk апd Quicke 1997 1999Ь GiЬsоп 1999 Rопquist 1999 Ronquist et а1 1999) It should Ье поtеd that SТВ a1so аге di5persed in some scelionids ТЬе SТВ in some of these taxa are modified basally (character 5 state 1) being laterally compressed and widепеd These modified SТВ аге ргеsепt in the Diashypriidae the uпdеsсгiЬеd New Zealand family Platyshygastridae and in most Chalcidoidea and Cynipoidea

Some гесепtlу iпvеstigаtеd iпdерепdепt character suite5 5uggest аltегпаtivе геlаtiопshiрs Ьеtwееп the аЬоуе taxa А row of erect setae opposing the distal lamui iп the hind wiпg was fоuпd in the Chalshycidoidea Сегарhгопоidеа Diapriidae апd the undeshyscribed New Zealand family (Basibl1yuk and Quicke 1997) апd simultaneo1lS апtеппal gгооmiпg оссшs iп the Megaspilidae Chalcidoidea Мушагidае Суshynipoidea Proctotrupoidea апd sоше Platygastroidea (behavior 4 Basibl1Yl1k and Ql1icke 1999Ь) Gibson (1999) has reviewed hypotheses сопсегпiпg the sister grol1p геlаtiОП5Ырs of chalcidoid wasps Не did not fiпd сопviпсiпg support for RasnitsYl1s (1988) platyshygastroid + (chalcidoids + mушагоmmаtоids) hypothshyesis bl1t instead for а clade compri5ing Platygastroishydea Proctotrupoidea Pelecinidae and Vanhorniidae Не a1so sнggested that the Chalcidoidea and Муshymагоmшаtоidеаmау Ье тоге closely related to either the Diapriidae or Сегарhroпidае rather than to t1le Platygastroidea

We fОl1пd а spur at the base of еас SТВ (character 6) which seems to restrict the mobility of the 1Iюг

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raquo~+О~~vt~(~lt-~()Ь~~~+~~tь~ь~~Ь~1ь~~O~~~~~t~~fmiddot~~о$~+ ~~$Ь~Ь~~~~~О~()Ь~~=~~~$ь~ьt~~~~Ь~~~О--~Ь~~~~~ЬО~6~~~q~1~~~~~~~~~~~lt)~O~~~~t~q~~lt~qОltgtO~О~~ltgt~~O~middotampQvQОО-gtltf~q~~amplt~~~~~~lti)-~О~~~ltlt6~~fI~o~~Q~Qo~~~iо~lt)ф1~~о~lti~1lti-gt~(~OJ

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bull bull bull bull bull 11 JAI 1 1 1 1 1 1 1 I 1 1 +1 2

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сgtFig12 Unambiguous character state changes оЕ orbicula and associated structures superimposed оп the most recent phylogenetic hypothesis ofRonquist et аl (1999) resulting

ijfrom impJied weight analysis Numbers оп the cladogram correspond to the character numbers (ТаЫе 1) Character state changes for each character are explained in the text

w

~- -А д j ~ л ~ r ~ _ ~ gt -- -1 iI- _- 1 1-1-1 1- 1 ~ olJc j J

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

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Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

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DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

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Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

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Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

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Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

628 ANNALS OF ТНЕ ENTOMOLOGICAL SOCIEТY OF АМЕтСА Vol 93 по 3

Figs l-3 (1) ОгЫсlllа in Sеflщiа sp (Tenthredinidae) STA sensilla tгiсhоdеа А arrow head sho ing scale-like SСlllрtше Nith cllticlllar projections Ваг = 30 т (2)Orbicula in Arotes arnoentL~ (Ichneumonidae) STA three sensilla tJiсlюdеа А black aJrow sel1silla tlichodea В white arrow sel1silla саmралifогтiа [уре А Ваг = 38 т (3) Close-up of orbicula tехtше il1 Регgо dorsafis (Pergidae) showing scale-like sculpture and two sensilla сатралifогтiа А (апоws) Ваг = б т

ТЬеге аге t1Sшllу three ог тоге SТE (Figs 5 7 9 ТаЫе 1) although only two аге found in а few sawflies some parasitic wasps апс some tiphiids and apids А single SТE occurs in а питЬе of isolated cases (Figs 2 апс 4 ТаЫе 1)

AITangement ofSTB (character 4) STB аге usually (Iistributed irregularly (Figs 5 and 9) but they fопп а single IOW in almost аll cynipoids (except IbaJiiclae) chalcidoids (except at least some Chalcidiclae ап(1 Leucospidae ) Diapriidae Vanhorniidae the undeshyscribed NeN Zealand family Platygastridae and some Ploctotrupidae (Fig 7)

Basal Modifjcation of STB (characteI 5) ТЬе SТE аге wi(lened апс laterally compressed in the Chalshycidoidea Diapriidae Platygastridae the undescribed New Zealand family апс some Cynipoidea (Figs 7 and 10)

Spur of STB Socket (chaJacter 6) ТЬе se nsilla socket bears а spur Nhich appears to restrict its backshyward mobi1ity in most chaJcidoid families including

mymarids together ith the uшlеSСliЬеd New Zeashyап(l famil y (Figs 78 10) А similar spur is present in the Cynipoidea but it is situated тоге distantly from the sensilla ТЬе 110mology of the spurs in (Ье Суshynipoidea witIl those in the other taxa is therefore unceгtain

Sensilla Сшпрапifогтiа (SC) (characters 7 and 8) Two types of sensilla campanifOlmia аге distinguishshyаЫе апс аге termed Ьеге SCA and SCB SCA is usually а roundecl dome encircled Ьу а cuticular ridge (Figs 2 апcl 3) whereas SCB has по external el1circling lidge аl1с арреаг as а simple small tubercle (Figs 4 and 8) SCA aJe Гоиl1с il1 the Trigonalidae апс Pergidae (ехshycept lлрlчrоtОm()) апс in тапу species of BJashyconidae апс Ichl1eumol1idae ТЬеу also are present il1 some replesel1tatives of several isolated taxa (Xyelishydae Pamphiliklae Stephal1idae Megalyridae Sclerogibbidae Chrysididae апс Formicidae) (ТаЫе 1) SCB аге possessed Ьу the Chalcidoidea Diapliidae and some species of Sceliol1idae al1d possibly Ьу the

629 Мау 2000 BASIВUYUК ЕТ AL ORВICLJLA 0[ HVMENOPТElA

Figs4-7 (4) ОЪi Сl1 lа11 Triteleio sp (SceliOl1idae) vith two sel1silla trichodeaA (ореl1 апоs) sil1gle sensilla tichoclea В (hite arro) аl1сl two sel1silla campalliformia type В (white shadovecl bIack arrows) Bar = 8 fLl11 (5) Orbicula il1 RopTonio gопnаni (Roproniiclae) Ilote clel1tifoCI1l sca les (arтo) Ваг = 30 fLl1l (6) Orbiclla ill Ггсmех sp (Sricidae) shoving presl1l11ed six sel1silla trichodeaA Ваг = 50 fLl1l (7) Orbicl1la ill Ptoth -ix sp (Pteromalidae) sho ingsil1 g1e sel1silla trichodea А (ореl1 апоw) and а tansverse JO W of sellsilla tcl)Qclelt1 В (smзl1 arrows) Ваг = 13 fLl11

undescribed New Zealand family АI1 (аха that have sensiJla campalliformia Iave only опе type with tlle possible exceptioll иоьгасоn (Braconidae) (Fig11) The great majority oftaxa witJl SC (either А ог В) Ьауе two of them but only а sillg]e SCA is foulld ill megashyIyrids апс three ог тоге occur in some pergids Ьгаshycollids sclerogibbids апс тапу ichneumonids

Positioll of SC ( сhагасtег 9) SCA аге usua]]y роshysitioned iпеgulагlу апс distal to STA Howeer they сап Ье both distal аllcl basltll (о STA ill LVlacroxyel (MacroxyeJinae) Perga (Pergide) апе] тапу IchshyneumOlliclae (ТаЫе 1) Positiolls of SCB аге quite

cOllsistellt within the Chakidoideltl аllс Diapriidae ill vhich they аге always 10саtеclllttегоvепtгаJlу Jelative to the STA оП eitheJ siele of JateraJly compressed orbicula level witJl 01 sliglltly basal to it (Fig 8) TI1e SCB of sceliollids however ltге locateel clistal (о the STA (Fig 4)

Discussioll

Functiolla Implicatiolls The orbiculal sellsillt аге lttrgllably all mechalloreceptors (McIver 1985) 111 th e eulophicl Sympiesi~ seiceicomi~ Nees they have Ьееп

630 ANNALS OF ТНЕ ENTOMOLOC1CAL SOCIEТY OF AМERJCA Vol 93 п о 3

Figs8-11 (8) 5e nsilla ill Jurywma sp (Ешуtоm idае) 5ТА sensilla tri chodea А wi th а wellmiddot (leve loped socke t 5ТВ sefJS illa tic llOdea В white 3IТOW se lls illa саmралi fогmi а type В Bar = 3 Lm (9) CeГOfo les sp (Роm рШdае) SllO v ing pos ition зп(1 аrrалgеmепt of sensilla tricllOdeaA Bar = 38 LПJ (10) Sелsillа tichodea В of Castmcantl-s рсетjlIШI (Pterom alidae) Sl10ving widened base and basal spur (ореп апоws) Вш = 3Lnl (11) Orbicula in иоЬгас(т sp (Braconidae) sho-ing bot11 types of se nsilla саmрап ifогmiа vl)ite апоws ргеSLmеd sел sillа campan iformia В white shadove(1 bIack arrovs sensilla campan ifomia А Ва = 20 Lm

suggested as шviпg а role [п de tecting vibration s ргоmiddot ducecl Ьу th e ir leafminer host (Meyhbfer e t al 1997) Althougl this тау gtell Ье the case theiI prese nce [п taxa with diffe le nt blologies (incllding phytoph agollS taxa a llC раГltlsitоid s of deeply conceaed hosts) sugshygests that they have otl er гoles as well ТЬе ргеsеп се of apparen tly identi ca sensila iп both sexes of тапу taxa fшthеl weake ns tle host locatio) hypotllesis Because ST and SC аге prese nt in th e most basal ineages ofthe H ymenoptera (eg MacJQxyeinae and Cephalciinae) we suggest that the iJ огigiп а l and ргеshy

cominant fun ction is топitогiпg moveme nts and s tгеss resulting fгош the asps own 10comotion anс possibly also subst)a te viblation that тау indicate the ргеsепсе of pote) tia pledatoгs Athough they тау play а role [п host de tection the ack of sexua diтогshyphism i) рагаs itiс taxa suggests that this is not а majoJ need Indeed the sexua dinlOrphism in th e phytophshyagous Siricidae and iп th e рагаs itiс Tipl iidae both cases vit females having fewe r STA suggests а gleate J iшрогtапсе fOl th e maes Моге obselvations оп paras itoid host sею-сhiпg behavioJ includi ng еесshy

631 Мау 2000 BASmUYUK ЕТ ЛL ORВICULA OF НУМENОPrERЛ

trophysiological studies are needed to determine tle exact function of these sensilla

Рhуlоgепеtic Imрliсаtiопs То assess рhуlоgепеtiс implications ofthi5 character system we have mapped the data onto the most recent phylogenetic hypothesis of the епtiге order re5ulting from implied weighting (Ronquist et а11999) Only the uпаmЬiguоus changes were indicated and polytomies were treated as Iard (ie аssumiпg multiple speciation events) (Fig 12) We have included the undescribed New Zеаlапd famshyНу iп the putative clade comprising Chalcidoidea Cershyарhroпоidеа Diapriidae and 50 оп because it shares at least опе putative sупароmогрhу with mепtiопеd taxa (Basibuyuk апd Quicke 1997) We also have plotshyted the data оп to Rasnitsyns (1988) intuitive hypothshye5i5 and оп to а tree resulting from reanцJysis of а соmЫпаtiоп ofRasnitsyns data (Ronquist ct а1 1999) with VHhelmsen s (1997) sуmрhуtап апd Brother5 and Сагрепtегs (1993) aculeate hypothese5 However the fit of the data were marginally better оп the latest hypothesis of Rопquist et а1 (1999)

Orbicular Sculpture The orbiculas of тanу of the most Ьа5аlliпеаgеs (eg the Xyelinae Pamphiloidea Cephoidea апd Tenthredinoidea other than pergids) have scale-like microsculpture with cuticular projecshytion5 (character 1 5tate О) sugge5ting thi5 to Ье the plesiomorphic condition for the order Simpfe scaleshylike 5culpture without рrojесt10ПS (character 1 5tate 1) appears first 1П the siricoid grade апd i5 геtаiпеd in the Evaniomorpha sеП5U Rasпitsуп рщ of the Procshytotrupoidea (РеlеС1П1dае Heloridae Rоргопiidае Мопоmасhidае) апd 1П mапу Iсhпеumопidае It also 15 ргеsепt 1П the Масгохуеliпае and Pergidae which сап Ье best сопsidегеd as iпdерепdепt acquisitions Further reduction of the microsculpture to а сотshypletely 5mooth orbicula (character 1 state 2) has арshyparently occurred оп at least three separate оссasiопs iп рщ of the miсгоhуmепорtегап lineage iпсll1diпg the Супiроidеа Diapriidae the uпdеsсгiЬеd New Zеаlапd family Platygastroidea Сегарhгопоidеа Муshymarommatidae апd Chalcidoidea iп the pl1tative Proctotrl1pidae + Vanhorniidae liпеаgе and in the Aculeata ТЬе presence of some simple scale-like sсulрtше (charactcr 1 state 1) in the Sclerogibbidae Scolebythidae апdVe5pidae сап best Ье considered as reversals (Rasnitsyn 1988 Brothers and Сагрепtег 1993 Ronquist et al 1999)

Sепsillа Trichodea А STА аге ргеsепt iп almost аН taxa оfНуmепорtега except Хуеliпае and their presshyепсе is therefore а putative synapomorphy for the Нуmепорtега ехсludiпg the ХуеНпае АЬsепсе of STA in а few isolated taxa in which they are и511аНу polymorphic (5iricids orussids stephanids megalyrshyids proctotrupids platygastrids ЬгаdупоЬаепids апd tiphiids) сап be5t Ье regarded as iпdерепdепt losscs (Fig 12) Тhe Пl1mЬег ofSTA i5 overwhelmingly two and this

сопditiоп (character 2 state 2) is most likely the gгоuпd рlап for tlle Нушепорtега as а whole Reducshytion iп the number of STA to опе (character 2 state 1) with а markedly raised socket is а pl1tative synashyротогрЬу for а clade соmрrisiпg Chalcidoidea Муshy

mагоmшаtоidеа Diapriidae Ceraphronidae and the undescribed New Zеalапd family but with опе revershysal (iп Супiроidса) and at least опс tгапsitiоп to other states (in Megaspilidae) Three ог more STA occur in some Trigonalidae Megalyridae Megaspilidae Brashyсопidае Ichпеumопidае апd Pompilidae bl1t iп most of the5e cases the Iikely gгоuпd рlап i5 two STA Ьеshycause that state occurs iп тanу trigопаlids iсhпеushymопids as well as in some Ьгасопids (eg аgаthidiпе gепега Labagathis and L~opronotum) ТЬе ргеsепсе of thrce or more STA in the Mega5pilidae and Pompilishydae i5 best cOl1Sidered to герге5епt two indерепdепt асquisitiопs (Fig 12)

Sensilla Trichodea В апdRelated Structures Тhese are present in most taxa but their two 5eparate аЬshysепсеs and variаtiоп in their пumЬег (character 3) appear to iпdiсаtе sоше hошорlаsу iп the Apocrita for the givеп 5et of рhуlоgепеtiс hypotheses However possession ofthree ог тоге STA (character 3 state 3) provides support for the monophyly of Apocrita (Fig 12) Аrrапgеmепt of the SТВ iп а siпglе row (Fig 7 character 4 state 1) 511pports the clade соmргisiпg the Chalcidoidea Mymarommatoidea Ceraphronoidea Platygastroidea Cynipoidea Diapriidae and the uпshydescribed New Zealand fашilу (Fig 12) This state a1so occurs iп the Vanhorniidae and Pamphilioidea Тhe оссипепсе ofdi5persed SТВ (character 4 state О) iп the Chalcididae Lel1cospidac and Ibaliidae (which аге relatively larger iп 5ize) within thi5 clade тау sl1ggest sizе-dерепdепt сопvегgепсе for this characshyter but there i5 а gгоwiпg body ofеvidепсе supporting some of the relationships amопg the аЬоуе putative clade (Basibuyнk апd Quicke 1997 1999Ь GiЬsоп 1999 Rопquist 1999 Ronquist et а1 1999) It should Ье поtеd that SТВ a1so аге di5persed in some scelionids ТЬе SТВ in some of these taxa are modified basally (character 5 state 1) being laterally compressed and widепеd These modified SТВ аге ргеsепt in the Diashypriidae the uпdеsсгiЬеd New Zealand family Platyshygastridae and in most Chalcidoidea and Cynipoidea

Some гесепtlу iпvеstigаtеd iпdерепdепt character suite5 5uggest аltегпаtivе геlаtiопshiрs Ьеtwееп the аЬоуе taxa А row of erect setae opposing the distal lamui iп the hind wiпg was fоuпd in the Chalshycidoidea Сегарhгопоidеа Diapriidae апd the undeshyscribed New Zealand family (Basibl1yuk and Quicke 1997) апd simultaneo1lS апtеппal gгооmiпg оссшs iп the Megaspilidae Chalcidoidea Мушагidае Суshynipoidea Proctotrupoidea апd sоше Platygastroidea (behavior 4 Basibl1Yl1k and Ql1icke 1999Ь) Gibson (1999) has reviewed hypotheses сопсегпiпg the sister grol1p геlаtiОП5Ырs of chalcidoid wasps Не did not fiпd сопviпсiпg support for RasnitsYl1s (1988) platyshygastroid + (chalcidoids + mушагоmmаtоids) hypothshyesis bl1t instead for а clade compri5ing Platygastroishydea Proctotrupoidea Pelecinidae and Vanhorniidae Не a1so sнggested that the Chalcidoidea and Муshymагоmшаtоidеаmау Ье тоге closely related to either the Diapriidae or Сегарhroпidае rather than to t1le Platygastroidea

We fОl1пd а spur at the base of еас SТВ (character 6) which seems to restrict the mobility of the 1Iюг

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bull bull bull bull bull 11 JAI 1 1 1 1 1 1 1 I 1 1 +1 2

7 I I 6

7Т I т

2

4

4Т I I I I I I I

7 I I I I I I 1 3 gJ (j gtshy

~ ф

сgtFig12 Unambiguous character state changes оЕ orbicula and associated structures superimposed оп the most recent phylogenetic hypothesis ofRonquist et аl (1999) resulting

ijfrom impJied weight analysis Numbers оп the cladogram correspond to the character numbers (ТаЫе 1) Character state changes for each character are explained in the text

w

~- -А д j ~ л ~ r ~ _ ~ gt -- -1 iI- _- 1 1-1-1 1- 1 ~ olJc j J

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

References Cited

Аllеп R Т and G Е Ball 1980 Synopsis of Mexican taxa of the Loxandrus series (Coleoptera Carabidae Pteroshystichini) Trans Ат Entomol Soc 105481-575

Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

DowtOIl М and А D Austin 1994 Molecular phylogeny ofthe insect order Нушегюрtега apocritan relationships Proc Natl Acad Sci USA 91 9911-9915

DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

FrazierJ L 1985 Ncrvous system sensory system рр 287shy356 lп М S Bluт [ed] Fundamentals of insect physishyology Wiley Nev York

Ganesalingam У К 1972 Апаtошу апd histology of tlle БеПБе огgапs of the ovipositor of tlle ichncunlOnid wasp Devorgilla canescens J Inscct Physiol 18 1857-1868

Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

Konigsmann Е 1976 Das phylogenetische Systcm der Нуshymenoptelmiddota Теil IEinftihrung Grundplanmerkmale Schwestergruppe und Fossilfunde Dtsch Епtопюl Z23 251-279

Konig~mann Е 1977 Das phylogenetische System der Нуshyшепорtега Теil 2 Sympllyta Dtsch Entomol Z 24 1-40

Копigsmanп Е 1978а Das phylogenetische Sуstеш dcr Нуmепорtега Teil 3 TerebraIltes (Unterordnung Ароshycrita) Dtsch Entomol Z 25 1-55

Konigsmann Е 1978Ь Das рhуlоgепеtisсhе Sуstеш der HYlllenoptera Теil 4 Aculeata (Uпtегоrdпuпg Ароshycrita) Dtsch Entomol Z 25 365-435

LaSalle J апd 1 D Gauld 1993 Нуmспорtега апd biodishyversity сАВ Iпtегпаtiопаl Wallingford

LeRalec А апd Е Wajnberg 1990 Sensory receptors of the ovipositor of Trichogram11Ul trшidis (Нут Trishyсhоgгаmшаtidас) Епtоmорhаgа 35 293-299

LeRalec А J М Rabasse апd Е Vajnberg 1996 Сотshyparativc morphology of the oipositor of Боте parasitic

634 ANNAI5 Ш ТНЕ ENTOMOLOGICAI~ SOCIEТY OF AМERICA Vol 93 по 3

Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

Mclver S В 1985 Mechanoreception рр 71-132 lп G А Kerkut and L I Gilbert [eds] Comprehensive insect physiology biochemistry and pharmacology vol 6 Pershygamon Oxford

МеуЬOfес В J Сшш and S Dorn 1997 МесЬапо- and chemoreceptors and their possible role in host-location behavior of Sympiesis seпceklJrnis (Нушепорtеrа Euloshyphidae) Апп Entomol Soc Аm 90 208-219

Navasero В С and G W Elzen 1991 Sensilla оп the antennae foretarsi and palpi of Microplitis croceipes (Cresson) (Hymenoptera Braconidae) Proc Entomol Soc Wash 93 373-347

Rasnitsyn А Р 1969 Oтigin and evolution of the lower Hymel1optera Trans Paleol1tol Iпst Acad Sci USSR 123 1-196

Rasnitsyn А Р 1980 Origin and evolution of Нуmепорshytera Тсanо Paleol1tol Inst Acad Sci USSR 174 1-192

Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

629 Мау 2000 BASIВUYUК ЕТ AL ORВICLJLA 0[ HVMENOPТElA

Figs4-7 (4) ОЪi Сl1 lа11 Triteleio sp (SceliOl1idae) vith two sel1silla trichodeaA (ореl1 апоs) sil1gle sensilla tichoclea В (hite arro) аl1сl two sel1silla campalliformia type В (white shadovecl bIack arrows) Bar = 8 fLl11 (5) Orbicula il1 RopTonio gопnаni (Roproniiclae) Ilote clel1tifoCI1l sca les (arтo) Ваг = 30 fLl1l (6) Orbiclla ill Ггсmех sp (Sricidae) shoving presl1l11ed six sel1silla trichodeaA Ваг = 50 fLl1l (7) Orbicl1la ill Ptoth -ix sp (Pteromalidae) sho ingsil1 g1e sel1silla trichodea А (ореl1 апоw) and а tansverse JO W of sellsilla tcl)Qclelt1 В (smзl1 arrows) Ваг = 13 fLl11

undescribed New Zealand family АI1 (аха that have sensiJla campalliformia Iave only опе type with tlle possible exceptioll иоьгасоn (Braconidae) (Fig11) The great majority oftaxa witJl SC (either А ог В) Ьауе two of them but only а sillg]e SCA is foulld ill megashyIyrids апс three ог тоге occur in some pergids Ьгаshycollids sclerogibbids апс тапу ichneumonids

Positioll of SC ( сhагасtег 9) SCA аге usua]]y роshysitioned iпеgulагlу апс distal to STA Howeer they сап Ье both distal аllcl basltll (о STA ill LVlacroxyel (MacroxyeJinae) Perga (Pergide) апе] тапу IchshyneumOlliclae (ТаЫе 1) Positiolls of SCB аге quite

cOllsistellt within the Chakidoideltl аllс Diapriidae ill vhich they аге always 10саtеclllttегоvепtгаJlу Jelative to the STA оП eitheJ siele of JateraJly compressed orbicula level witJl 01 sliglltly basal to it (Fig 8) TI1e SCB of sceliollids however ltге locateel clistal (о the STA (Fig 4)

Discussioll

Functiolla Implicatiolls The orbiculal sellsillt аге lttrgllably all mechalloreceptors (McIver 1985) 111 th e eulophicl Sympiesi~ seiceicomi~ Nees they have Ьееп

630 ANNALS OF ТНЕ ENTOMOLOC1CAL SOCIEТY OF AМERJCA Vol 93 п о 3

Figs8-11 (8) 5e nsilla ill Jurywma sp (Ешуtоm idае) 5ТА sensilla tri chodea А wi th а wellmiddot (leve loped socke t 5ТВ sefJS illa tic llOdea В white 3IТOW se lls illa саmралi fогmi а type В Bar = 3 Lm (9) CeГOfo les sp (Роm рШdае) SllO v ing pos ition зп(1 аrrалgеmепt of sensilla tricllOdeaA Bar = 38 LПJ (10) Sелsillа tichodea В of Castmcantl-s рсетjlIШI (Pterom alidae) Sl10ving widened base and basal spur (ореп апоws) Вш = 3Lnl (11) Orbicula in иоЬгас(т sp (Braconidae) sho-ing bot11 types of se nsilla саmрап ifогmiа vl)ite апоws ргеSLmеd sел sillа campan iformia В white shadove(1 bIack arrovs sensilla campan ifomia А Ва = 20 Lm

suggested as шviпg а role [п de tecting vibration s ргоmiddot ducecl Ьу th e ir leafminer host (Meyhbfer e t al 1997) Althougl this тау gtell Ье the case theiI prese nce [п taxa with diffe le nt blologies (incllding phytoph agollS taxa a llC раГltlsitоid s of deeply conceaed hosts) sugshygests that they have otl er гoles as well ТЬе ргеsеп се of apparen tly identi ca sensila iп both sexes of тапу taxa fшthеl weake ns tle host locatio) hypotllesis Because ST and SC аге prese nt in th e most basal ineages ofthe H ymenoptera (eg MacJQxyeinae and Cephalciinae) we suggest that the iJ огigiп а l and ргеshy

cominant fun ction is топitогiпg moveme nts and s tгеss resulting fгош the asps own 10comotion anс possibly also subst)a te viblation that тау indicate the ргеsепсе of pote) tia pledatoгs Athough they тау play а role [п host de tection the ack of sexua diтогshyphism i) рагаs itiс taxa suggests that this is not а majoJ need Indeed the sexua dinlOrphism in th e phytophshyagous Siricidae and iп th e рагаs itiс Tipl iidae both cases vit females having fewe r STA suggests а gleate J iшрогtапсе fOl th e maes Моге obselvations оп paras itoid host sею-сhiпg behavioJ includi ng еесshy

631 Мау 2000 BASmUYUK ЕТ ЛL ORВICULA OF НУМENОPrERЛ

trophysiological studies are needed to determine tle exact function of these sensilla

Рhуlоgепеtic Imрliсаtiопs То assess рhуlоgепеtiс implications ofthi5 character system we have mapped the data onto the most recent phylogenetic hypothesis of the епtiге order re5ulting from implied weighting (Ronquist et а11999) Only the uпаmЬiguоus changes were indicated and polytomies were treated as Iard (ie аssumiпg multiple speciation events) (Fig 12) We have included the undescribed New Zеаlапd famshyНу iп the putative clade comprising Chalcidoidea Cershyарhroпоidеа Diapriidae and 50 оп because it shares at least опе putative sупароmогрhу with mепtiопеd taxa (Basibuyuk апd Quicke 1997) We also have plotshyted the data оп to Rasnitsyns (1988) intuitive hypothshye5i5 and оп to а tree resulting from reanцJysis of а соmЫпаtiоп ofRasnitsyns data (Ronquist ct а1 1999) with VHhelmsen s (1997) sуmрhуtап апd Brother5 and Сагрепtегs (1993) aculeate hypothese5 However the fit of the data were marginally better оп the latest hypothesis of Rопquist et а1 (1999)

Orbicular Sculpture The orbiculas of тanу of the most Ьа5аlliпеаgеs (eg the Xyelinae Pamphiloidea Cephoidea апd Tenthredinoidea other than pergids) have scale-like microsculpture with cuticular projecshytion5 (character 1 5tate О) sugge5ting thi5 to Ье the plesiomorphic condition for the order Simpfe scaleshylike 5culpture without рrojесt10ПS (character 1 5tate 1) appears first 1П the siricoid grade апd i5 геtаiпеd in the Evaniomorpha sеП5U Rasпitsуп рщ of the Procshytotrupoidea (РеlеС1П1dае Heloridae Rоргопiidае Мопоmасhidае) апd 1П mапу Iсhпеumопidае It also 15 ргеsепt 1П the Масгохуеliпае and Pergidae which сап Ье best сопsidегеd as iпdерепdепt acquisitions Further reduction of the microsculpture to а сотshypletely 5mooth orbicula (character 1 state 2) has арshyparently occurred оп at least three separate оссasiопs iп рщ of the miсгоhуmепорtегап lineage iпсll1diпg the Супiроidеа Diapriidae the uпdеsсгiЬеd New Zеаlапd family Platygastroidea Сегарhгопоidеа Муshymarommatidae апd Chalcidoidea iп the pl1tative Proctotrl1pidae + Vanhorniidae liпеаgе and in the Aculeata ТЬе presence of some simple scale-like sсulрtше (charactcr 1 state 1) in the Sclerogibbidae Scolebythidae апdVe5pidae сап best Ье considered as reversals (Rasnitsyn 1988 Brothers and Сагрепtег 1993 Ronquist et al 1999)

Sепsillа Trichodea А STА аге ргеsепt iп almost аН taxa оfНуmепорtега except Хуеliпае and their presshyепсе is therefore а putative synapomorphy for the Нуmепорtега ехсludiпg the ХуеНпае АЬsепсе of STA in а few isolated taxa in which they are и511аНу polymorphic (5iricids orussids stephanids megalyrshyids proctotrupids platygastrids ЬгаdупоЬаепids апd tiphiids) сап be5t Ье regarded as iпdерепdепt losscs (Fig 12) Тhe Пl1mЬег ofSTA i5 overwhelmingly two and this

сопditiоп (character 2 state 2) is most likely the gгоuпd рlап for tlle Нушепорtега as а whole Reducshytion iп the number of STA to опе (character 2 state 1) with а markedly raised socket is а pl1tative synashyротогрЬу for а clade соmрrisiпg Chalcidoidea Муshy

mагоmшаtоidеа Diapriidae Ceraphronidae and the undescribed New Zеalапd family but with опе revershysal (iп Супiроidса) and at least опс tгапsitiоп to other states (in Megaspilidae) Three ог more STA occur in some Trigonalidae Megalyridae Megaspilidae Brashyсопidае Ichпеumопidае апd Pompilidae bl1t iп most of the5e cases the Iikely gгоuпd рlап i5 two STA Ьеshycause that state occurs iп тanу trigопаlids iсhпеushymопids as well as in some Ьгасопids (eg аgаthidiпе gепега Labagathis and L~opronotum) ТЬе ргеsепсе of thrce or more STA in the Mega5pilidae and Pompilishydae i5 best cOl1Sidered to герге5епt two indерепdепt асquisitiопs (Fig 12)

Sensilla Trichodea В апdRelated Structures Тhese are present in most taxa but their two 5eparate аЬshysепсеs and variаtiоп in their пumЬег (character 3) appear to iпdiсаtе sоше hошорlаsу iп the Apocrita for the givеп 5et of рhуlоgепеtiс hypotheses However possession ofthree ог тоге STA (character 3 state 3) provides support for the monophyly of Apocrita (Fig 12) Аrrапgеmепt of the SТВ iп а siпglе row (Fig 7 character 4 state 1) 511pports the clade соmргisiпg the Chalcidoidea Mymarommatoidea Ceraphronoidea Platygastroidea Cynipoidea Diapriidae and the uпshydescribed New Zealand fашilу (Fig 12) This state a1so occurs iп the Vanhorniidae and Pamphilioidea Тhe оссипепсе ofdi5persed SТВ (character 4 state О) iп the Chalcididae Lel1cospidac and Ibaliidae (which аге relatively larger iп 5ize) within thi5 clade тау sl1ggest sizе-dерепdепt сопvегgепсе for this characshyter but there i5 а gгоwiпg body ofеvidепсе supporting some of the relationships amопg the аЬоуе putative clade (Basibuyнk апd Quicke 1997 1999Ь GiЬsоп 1999 Rопquist 1999 Ronquist et а1 1999) It should Ье поtеd that SТВ a1so аге di5persed in some scelionids ТЬе SТВ in some of these taxa are modified basally (character 5 state 1) being laterally compressed and widепеd These modified SТВ аге ргеsепt in the Diashypriidae the uпdеsсгiЬеd New Zealand family Platyshygastridae and in most Chalcidoidea and Cynipoidea

Some гесепtlу iпvеstigаtеd iпdерепdепt character suite5 5uggest аltегпаtivе геlаtiопshiрs Ьеtwееп the аЬоуе taxa А row of erect setae opposing the distal lamui iп the hind wiпg was fоuпd in the Chalshycidoidea Сегарhгопоidеа Diapriidae апd the undeshyscribed New Zealand family (Basibl1yuk and Quicke 1997) апd simultaneo1lS апtеппal gгооmiпg оссшs iп the Megaspilidae Chalcidoidea Мушагidае Суshynipoidea Proctotrupoidea апd sоше Platygastroidea (behavior 4 Basibl1Yl1k and Ql1icke 1999Ь) Gibson (1999) has reviewed hypotheses сопсегпiпg the sister grol1p геlаtiОП5Ырs of chalcidoid wasps Не did not fiпd сопviпсiпg support for RasnitsYl1s (1988) platyshygastroid + (chalcidoids + mушагоmmаtоids) hypothshyesis bl1t instead for а clade compri5ing Platygastroishydea Proctotrupoidea Pelecinidae and Vanhorniidae Не a1so sнggested that the Chalcidoidea and Муshymагоmшаtоidеаmау Ье тоге closely related to either the Diapriidae or Сегарhroпidае rather than to t1le Platygastroidea

We fОl1пd а spur at the base of еас SТВ (character 6) which seems to restrict the mobility of the 1Iюг

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w IltJ

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+ + + ~ ~~v~+ь+ + ~I + + ~ ь I + + b~ middotamp~~vI+ + $+o~ ~I+ ~~~ + + ~Ifgtb+ ~~ +lt q + + +~f + bfgt 90 ~ьф()~ ZmiddotI+ I +

raquo~+О~~vt~(~lt-~()Ь~~~+~~tь~ь~~Ь~1ь~~O~~~~~t~~fmiddot~~о$~+ ~~$Ь~Ь~~~~~О~()Ь~~=~~~$ь~ьt~~~~Ь~~~О--~Ь~~~~~ЬО~6~~~q~1~~~~~~~~~~~lt)~O~~~~t~q~~lt~qОltgtO~О~~ltgt~~O~middotampQvQОО-gtltf~q~~amplt~~~~~~lti)-~О~~~ltlt6~~fI~o~~Q~Qo~~~iо~lt)ф1~~о~lti~1lti-gt~(~OJ

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sect n

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bull bull bull bull bull 11 JAI 1 1 1 1 1 1 1 I 1 1 +1 2

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2

4

4Т I I I I I I I

7 I I I I I I 1 3 gJ (j gtshy

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сgtFig12 Unambiguous character state changes оЕ orbicula and associated structures superimposed оп the most recent phylogenetic hypothesis ofRonquist et аl (1999) resulting

ijfrom impJied weight analysis Numbers оп the cladogram correspond to the character numbers (ТаЫе 1) Character state changes for each character are explained in the text

w

~- -А д j ~ л ~ r ~ _ ~ gt -- -1 iI- _- 1 1-1-1 1- 1 ~ olJc j J

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

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Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

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DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

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Ganesalingam У К 1972 Апаtошу апd histology of tlle БеПБе огgапs of the ovipositor of tlle ichncunlOnid wasp Devorgilla canescens J Inscct Physiol 18 1857-1868

Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

Konigsmann Е 1976 Das phylogenetische Systcm der Нуshymenoptelmiddota Теil IEinftihrung Grundplanmerkmale Schwestergruppe und Fossilfunde Dtsch Епtопюl Z23 251-279

Konig~mann Е 1977 Das phylogenetische System der Нуshyшепорtега Теil 2 Sympllyta Dtsch Entomol Z 24 1-40

Копigsmanп Е 1978а Das phylogenetische Sуstеш dcr Нуmепорtега Teil 3 TerebraIltes (Unterordnung Ароshycrita) Dtsch Entomol Z 25 1-55

Konigsmann Е 1978Ь Das рhуlоgепеtisсhе Sуstеш der HYlllenoptera Теil 4 Aculeata (Uпtегоrdпuпg Ароshycrita) Dtsch Entomol Z 25 365-435

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Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

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Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

630 ANNALS OF ТНЕ ENTOMOLOC1CAL SOCIEТY OF AМERJCA Vol 93 п о 3

Figs8-11 (8) 5e nsilla ill Jurywma sp (Ешуtоm idае) 5ТА sensilla tri chodea А wi th а wellmiddot (leve loped socke t 5ТВ sefJS illa tic llOdea В white 3IТOW se lls illa саmралi fогmi а type В Bar = 3 Lm (9) CeГOfo les sp (Роm рШdае) SllO v ing pos ition зп(1 аrrалgеmепt of sensilla tricllOdeaA Bar = 38 LПJ (10) Sелsillа tichodea В of Castmcantl-s рсетjlIШI (Pterom alidae) Sl10ving widened base and basal spur (ореп апоws) Вш = 3Lnl (11) Orbicula in иоЬгас(т sp (Braconidae) sho-ing bot11 types of se nsilla саmрап ifогmiа vl)ite апоws ргеSLmеd sел sillа campan iformia В white shadove(1 bIack arrovs sensilla campan ifomia А Ва = 20 Lm

suggested as шviпg а role [п de tecting vibration s ргоmiddot ducecl Ьу th e ir leafminer host (Meyhbfer e t al 1997) Althougl this тау gtell Ье the case theiI prese nce [п taxa with diffe le nt blologies (incllding phytoph agollS taxa a llC раГltlsitоid s of deeply conceaed hosts) sugshygests that they have otl er гoles as well ТЬе ргеsеп се of apparen tly identi ca sensila iп both sexes of тапу taxa fшthеl weake ns tle host locatio) hypotllesis Because ST and SC аге prese nt in th e most basal ineages ofthe H ymenoptera (eg MacJQxyeinae and Cephalciinae) we suggest that the iJ огigiп а l and ргеshy

cominant fun ction is топitогiпg moveme nts and s tгеss resulting fгош the asps own 10comotion anс possibly also subst)a te viblation that тау indicate the ргеsепсе of pote) tia pledatoгs Athough they тау play а role [п host de tection the ack of sexua diтогshyphism i) рагаs itiс taxa suggests that this is not а majoJ need Indeed the sexua dinlOrphism in th e phytophshyagous Siricidae and iп th e рагаs itiс Tipl iidae both cases vit females having fewe r STA suggests а gleate J iшрогtапсе fOl th e maes Моге obselvations оп paras itoid host sею-сhiпg behavioJ includi ng еесshy

631 Мау 2000 BASmUYUK ЕТ ЛL ORВICULA OF НУМENОPrERЛ

trophysiological studies are needed to determine tle exact function of these sensilla

Рhуlоgепеtic Imрliсаtiопs То assess рhуlоgепеtiс implications ofthi5 character system we have mapped the data onto the most recent phylogenetic hypothesis of the епtiге order re5ulting from implied weighting (Ronquist et а11999) Only the uпаmЬiguоus changes were indicated and polytomies were treated as Iard (ie аssumiпg multiple speciation events) (Fig 12) We have included the undescribed New Zеаlапd famshyНу iп the putative clade comprising Chalcidoidea Cershyарhroпоidеа Diapriidae and 50 оп because it shares at least опе putative sупароmогрhу with mепtiопеd taxa (Basibuyuk апd Quicke 1997) We also have plotshyted the data оп to Rasnitsyns (1988) intuitive hypothshye5i5 and оп to а tree resulting from reanцJysis of а соmЫпаtiоп ofRasnitsyns data (Ronquist ct а1 1999) with VHhelmsen s (1997) sуmрhуtап апd Brother5 and Сагрепtегs (1993) aculeate hypothese5 However the fit of the data were marginally better оп the latest hypothesis of Rопquist et а1 (1999)

Orbicular Sculpture The orbiculas of тanу of the most Ьа5аlliпеаgеs (eg the Xyelinae Pamphiloidea Cephoidea апd Tenthredinoidea other than pergids) have scale-like microsculpture with cuticular projecshytion5 (character 1 5tate О) sugge5ting thi5 to Ье the plesiomorphic condition for the order Simpfe scaleshylike 5culpture without рrojесt10ПS (character 1 5tate 1) appears first 1П the siricoid grade апd i5 геtаiпеd in the Evaniomorpha sеП5U Rasпitsуп рщ of the Procshytotrupoidea (РеlеС1П1dае Heloridae Rоргопiidае Мопоmасhidае) апd 1П mапу Iсhпеumопidае It also 15 ргеsепt 1П the Масгохуеliпае and Pergidae which сап Ье best сопsidегеd as iпdерепdепt acquisitions Further reduction of the microsculpture to а сотshypletely 5mooth orbicula (character 1 state 2) has арshyparently occurred оп at least three separate оссasiопs iп рщ of the miсгоhуmепорtегап lineage iпсll1diпg the Супiроidеа Diapriidae the uпdеsсгiЬеd New Zеаlапd family Platygastroidea Сегарhгопоidеа Муshymarommatidae апd Chalcidoidea iп the pl1tative Proctotrl1pidae + Vanhorniidae liпеаgе and in the Aculeata ТЬе presence of some simple scale-like sсulрtше (charactcr 1 state 1) in the Sclerogibbidae Scolebythidae апdVe5pidae сап best Ье considered as reversals (Rasnitsyn 1988 Brothers and Сагрепtег 1993 Ronquist et al 1999)

Sепsillа Trichodea А STА аге ргеsепt iп almost аН taxa оfНуmепорtега except Хуеliпае and their presshyепсе is therefore а putative synapomorphy for the Нуmепорtега ехсludiпg the ХуеНпае АЬsепсе of STA in а few isolated taxa in which they are и511аНу polymorphic (5iricids orussids stephanids megalyrshyids proctotrupids platygastrids ЬгаdупоЬаепids апd tiphiids) сап be5t Ье regarded as iпdерепdепt losscs (Fig 12) Тhe Пl1mЬег ofSTA i5 overwhelmingly two and this

сопditiоп (character 2 state 2) is most likely the gгоuпd рlап for tlle Нушепорtега as а whole Reducshytion iп the number of STA to опе (character 2 state 1) with а markedly raised socket is а pl1tative synashyротогрЬу for а clade соmрrisiпg Chalcidoidea Муshy

mагоmшаtоidеа Diapriidae Ceraphronidae and the undescribed New Zеalапd family but with опе revershysal (iп Супiроidса) and at least опс tгапsitiоп to other states (in Megaspilidae) Three ог more STA occur in some Trigonalidae Megalyridae Megaspilidae Brashyсопidае Ichпеumопidае апd Pompilidae bl1t iп most of the5e cases the Iikely gгоuпd рlап i5 two STA Ьеshycause that state occurs iп тanу trigопаlids iсhпеushymопids as well as in some Ьгасопids (eg аgаthidiпе gепега Labagathis and L~opronotum) ТЬе ргеsепсе of thrce or more STA in the Mega5pilidae and Pompilishydae i5 best cOl1Sidered to герге5епt two indерепdепt асquisitiопs (Fig 12)

Sensilla Trichodea В апdRelated Structures Тhese are present in most taxa but their two 5eparate аЬshysепсеs and variаtiоп in their пumЬег (character 3) appear to iпdiсаtе sоше hошорlаsу iп the Apocrita for the givеп 5et of рhуlоgепеtiс hypotheses However possession ofthree ог тоге STA (character 3 state 3) provides support for the monophyly of Apocrita (Fig 12) Аrrапgеmепt of the SТВ iп а siпglе row (Fig 7 character 4 state 1) 511pports the clade соmргisiпg the Chalcidoidea Mymarommatoidea Ceraphronoidea Platygastroidea Cynipoidea Diapriidae and the uпshydescribed New Zealand fашilу (Fig 12) This state a1so occurs iп the Vanhorniidae and Pamphilioidea Тhe оссипепсе ofdi5persed SТВ (character 4 state О) iп the Chalcididae Lel1cospidac and Ibaliidae (which аге relatively larger iп 5ize) within thi5 clade тау sl1ggest sizе-dерепdепt сопvегgепсе for this characshyter but there i5 а gгоwiпg body ofеvidепсе supporting some of the relationships amопg the аЬоуе putative clade (Basibuyнk апd Quicke 1997 1999Ь GiЬsоп 1999 Rопquist 1999 Ronquist et а1 1999) It should Ье поtеd that SТВ a1so аге di5persed in some scelionids ТЬе SТВ in some of these taxa are modified basally (character 5 state 1) being laterally compressed and widепеd These modified SТВ аге ргеsепt in the Diashypriidae the uпdеsсгiЬеd New Zealand family Platyshygastridae and in most Chalcidoidea and Cynipoidea

Some гесепtlу iпvеstigаtеd iпdерепdепt character suite5 5uggest аltегпаtivе геlаtiопshiрs Ьеtwееп the аЬоуе taxa А row of erect setae opposing the distal lamui iп the hind wiпg was fоuпd in the Chalshycidoidea Сегарhгопоidеа Diapriidae апd the undeshyscribed New Zealand family (Basibl1yuk and Quicke 1997) апd simultaneo1lS апtеппal gгооmiпg оссшs iп the Megaspilidae Chalcidoidea Мушагidае Суshynipoidea Proctotrupoidea апd sоше Platygastroidea (behavior 4 Basibl1Yl1k and Ql1icke 1999Ь) Gibson (1999) has reviewed hypotheses сопсегпiпg the sister grol1p геlаtiОП5Ырs of chalcidoid wasps Не did not fiпd сопviпсiпg support for RasnitsYl1s (1988) platyshygastroid + (chalcidoids + mушагоmmаtоids) hypothshyesis bl1t instead for а clade compri5ing Platygastroishydea Proctotrupoidea Pelecinidae and Vanhorniidae Не a1so sнggested that the Chalcidoidea and Муshymагоmшаtоidеаmау Ье тоге closely related to either the Diapriidae or Сегарhroпidае rather than to t1le Platygastroidea

We fОl1пd а spur at the base of еас SТВ (character 6) which seems to restrict the mobility of the 1Iюг

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raquo~+О~~vt~(~lt-~()Ь~~~+~~tь~ь~~Ь~1ь~~O~~~~~t~~fmiddot~~о$~+ ~~$Ь~Ь~~~~~О~()Ь~~=~~~$ь~ьt~~~~Ь~~~О--~Ь~~~~~ЬО~6~~~q~1~~~~~~~~~~~lt)~O~~~~t~q~~lt~qОltgtO~О~~ltgt~~O~middotampQvQОО-gtltf~q~~amplt~~~~~~lti)-~О~~~ltlt6~~fI~o~~Q~Qo~~~iо~lt)ф1~~о~lti~1lti-gt~(~OJ

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сgtFig12 Unambiguous character state changes оЕ orbicula and associated structures superimposed оп the most recent phylogenetic hypothesis ofRonquist et аl (1999) resulting

ijfrom impJied weight analysis Numbers оп the cladogram correspond to the character numbers (ТаЫе 1) Character state changes for each character are explained in the text

w

~- -А д j ~ л ~ r ~ _ ~ gt -- -1 iI- _- 1 1-1-1 1- 1 ~ olJc j J

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

References Cited

Аllеп R Т and G Е Ball 1980 Synopsis of Mexican taxa of the Loxandrus series (Coleoptera Carabidae Pteroshystichini) Trans Ат Entomol Soc 105481-575

Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

DowtOIl М and А D Austin 1994 Molecular phylogeny ofthe insect order Нушегюрtега apocritan relationships Proc Natl Acad Sci USA 91 9911-9915

DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

FrazierJ L 1985 Ncrvous system sensory system рр 287shy356 lп М S Bluт [ed] Fundamentals of insect physishyology Wiley Nev York

Ganesalingam У К 1972 Апаtошу апd histology of tlle БеПБе огgапs of the ovipositor of tlle ichncunlOnid wasp Devorgilla canescens J Inscct Physiol 18 1857-1868

Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

Konigsmann Е 1976 Das phylogenetische Systcm der Нуshymenoptelmiddota Теil IEinftihrung Grundplanmerkmale Schwestergruppe und Fossilfunde Dtsch Епtопюl Z23 251-279

Konig~mann Е 1977 Das phylogenetische System der Нуshyшепорtега Теil 2 Sympllyta Dtsch Entomol Z 24 1-40

Копigsmanп Е 1978а Das phylogenetische Sуstеш dcr Нуmепорtега Teil 3 TerebraIltes (Unterordnung Ароshycrita) Dtsch Entomol Z 25 1-55

Konigsmann Е 1978Ь Das рhуlоgепеtisсhе Sуstеш der HYlllenoptera Теil 4 Aculeata (Uпtегоrdпuпg Ароshycrita) Dtsch Entomol Z 25 365-435

LaSalle J апd 1 D Gauld 1993 Нуmспорtега апd biodishyversity сАВ Iпtегпаtiопаl Wallingford

LeRalec А апd Е Wajnberg 1990 Sensory receptors of the ovipositor of Trichogram11Ul trшidis (Нут Trishyсhоgгаmшаtidас) Епtоmорhаgа 35 293-299

LeRalec А J М Rabasse апd Е Vajnberg 1996 Сотshyparativc morphology of the oipositor of Боте parasitic

634 ANNAI5 Ш ТНЕ ENTOMOLOGICAI~ SOCIEТY OF AМERICA Vol 93 по 3

Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

Mclver S В 1985 Mechanoreception рр 71-132 lп G А Kerkut and L I Gilbert [eds] Comprehensive insect physiology biochemistry and pharmacology vol 6 Pershygamon Oxford

МеуЬOfес В J Сшш and S Dorn 1997 МесЬапо- and chemoreceptors and their possible role in host-location behavior of Sympiesis seпceklJrnis (Нушепорtеrа Euloshyphidae) Апп Entomol Soc Аm 90 208-219

Navasero В С and G W Elzen 1991 Sensilla оп the antennae foretarsi and palpi of Microplitis croceipes (Cresson) (Hymenoptera Braconidae) Proc Entomol Soc Wash 93 373-347

Rasnitsyn А Р 1969 Oтigin and evolution of the lower Hymel1optera Trans Paleol1tol Iпst Acad Sci USSR 123 1-196

Rasnitsyn А Р 1980 Origin and evolution of Нуmепорshytera Тсanо Paleol1tol Inst Acad Sci USSR 174 1-192

Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

631 Мау 2000 BASmUYUK ЕТ ЛL ORВICULA OF НУМENОPrERЛ

trophysiological studies are needed to determine tle exact function of these sensilla

Рhуlоgепеtic Imрliсаtiопs То assess рhуlоgепеtiс implications ofthi5 character system we have mapped the data onto the most recent phylogenetic hypothesis of the епtiге order re5ulting from implied weighting (Ronquist et а11999) Only the uпаmЬiguоus changes were indicated and polytomies were treated as Iard (ie аssumiпg multiple speciation events) (Fig 12) We have included the undescribed New Zеаlапd famshyНу iп the putative clade comprising Chalcidoidea Cershyарhroпоidеа Diapriidae and 50 оп because it shares at least опе putative sупароmогрhу with mепtiопеd taxa (Basibuyuk апd Quicke 1997) We also have plotshyted the data оп to Rasnitsyns (1988) intuitive hypothshye5i5 and оп to а tree resulting from reanцJysis of а соmЫпаtiоп ofRasnitsyns data (Ronquist ct а1 1999) with VHhelmsen s (1997) sуmрhуtап апd Brother5 and Сагрепtегs (1993) aculeate hypothese5 However the fit of the data were marginally better оп the latest hypothesis of Rопquist et а1 (1999)

Orbicular Sculpture The orbiculas of тanу of the most Ьа5аlliпеаgеs (eg the Xyelinae Pamphiloidea Cephoidea апd Tenthredinoidea other than pergids) have scale-like microsculpture with cuticular projecshytion5 (character 1 5tate О) sugge5ting thi5 to Ье the plesiomorphic condition for the order Simpfe scaleshylike 5culpture without рrojесt10ПS (character 1 5tate 1) appears first 1П the siricoid grade апd i5 геtаiпеd in the Evaniomorpha sеП5U Rasпitsуп рщ of the Procshytotrupoidea (РеlеС1П1dае Heloridae Rоргопiidае Мопоmасhidае) апd 1П mапу Iсhпеumопidае It also 15 ргеsепt 1П the Масгохуеliпае and Pergidae which сап Ье best сопsidегеd as iпdерепdепt acquisitions Further reduction of the microsculpture to а сотshypletely 5mooth orbicula (character 1 state 2) has арshyparently occurred оп at least three separate оссasiопs iп рщ of the miсгоhуmепорtегап lineage iпсll1diпg the Супiроidеа Diapriidae the uпdеsсгiЬеd New Zеаlапd family Platygastroidea Сегарhгопоidеа Муshymarommatidae апd Chalcidoidea iп the pl1tative Proctotrl1pidae + Vanhorniidae liпеаgе and in the Aculeata ТЬе presence of some simple scale-like sсulрtше (charactcr 1 state 1) in the Sclerogibbidae Scolebythidae апdVe5pidae сап best Ье considered as reversals (Rasnitsyn 1988 Brothers and Сагрепtег 1993 Ronquist et al 1999)

Sепsillа Trichodea А STА аге ргеsепt iп almost аН taxa оfНуmепорtега except Хуеliпае and their presshyепсе is therefore а putative synapomorphy for the Нуmепорtега ехсludiпg the ХуеНпае АЬsепсе of STA in а few isolated taxa in which they are и511аНу polymorphic (5iricids orussids stephanids megalyrshyids proctotrupids platygastrids ЬгаdупоЬаепids апd tiphiids) сап be5t Ье regarded as iпdерепdепt losscs (Fig 12) Тhe Пl1mЬег ofSTA i5 overwhelmingly two and this

сопditiоп (character 2 state 2) is most likely the gгоuпd рlап for tlle Нушепорtега as а whole Reducshytion iп the number of STA to опе (character 2 state 1) with а markedly raised socket is а pl1tative synashyротогрЬу for а clade соmрrisiпg Chalcidoidea Муshy

mагоmшаtоidеа Diapriidae Ceraphronidae and the undescribed New Zеalапd family but with опе revershysal (iп Супiроidса) and at least опс tгапsitiоп to other states (in Megaspilidae) Three ог more STA occur in some Trigonalidae Megalyridae Megaspilidae Brashyсопidае Ichпеumопidае апd Pompilidae bl1t iп most of the5e cases the Iikely gгоuпd рlап i5 two STA Ьеshycause that state occurs iп тanу trigопаlids iсhпеushymопids as well as in some Ьгасопids (eg аgаthidiпе gепега Labagathis and L~opronotum) ТЬе ргеsепсе of thrce or more STA in the Mega5pilidae and Pompilishydae i5 best cOl1Sidered to герге5епt two indерепdепt асquisitiопs (Fig 12)

Sensilla Trichodea В апdRelated Structures Тhese are present in most taxa but their two 5eparate аЬshysепсеs and variаtiоп in their пumЬег (character 3) appear to iпdiсаtе sоше hошорlаsу iп the Apocrita for the givеп 5et of рhуlоgепеtiс hypotheses However possession ofthree ог тоге STA (character 3 state 3) provides support for the monophyly of Apocrita (Fig 12) Аrrапgеmепt of the SТВ iп а siпglе row (Fig 7 character 4 state 1) 511pports the clade соmргisiпg the Chalcidoidea Mymarommatoidea Ceraphronoidea Platygastroidea Cynipoidea Diapriidae and the uпshydescribed New Zealand fашilу (Fig 12) This state a1so occurs iп the Vanhorniidae and Pamphilioidea Тhe оссипепсе ofdi5persed SТВ (character 4 state О) iп the Chalcididae Lel1cospidac and Ibaliidae (which аге relatively larger iп 5ize) within thi5 clade тау sl1ggest sizе-dерепdепt сопvегgепсе for this characshyter but there i5 а gгоwiпg body ofеvidепсе supporting some of the relationships amопg the аЬоуе putative clade (Basibuyнk апd Quicke 1997 1999Ь GiЬsоп 1999 Rопquist 1999 Ronquist et а1 1999) It should Ье поtеd that SТВ a1so аге di5persed in some scelionids ТЬе SТВ in some of these taxa are modified basally (character 5 state 1) being laterally compressed and widепеd These modified SТВ аге ргеsепt in the Diashypriidae the uпdеsсгiЬеd New Zealand family Platyshygastridae and in most Chalcidoidea and Cynipoidea

Some гесепtlу iпvеstigаtеd iпdерепdепt character suite5 5uggest аltегпаtivе геlаtiопshiрs Ьеtwееп the аЬоуе taxa А row of erect setae opposing the distal lamui iп the hind wiпg was fоuпd in the Chalshycidoidea Сегарhгопоidеа Diapriidae апd the undeshyscribed New Zealand family (Basibl1yuk and Quicke 1997) апd simultaneo1lS апtеппal gгооmiпg оссшs iп the Megaspilidae Chalcidoidea Мушагidае Суshynipoidea Proctotrupoidea апd sоше Platygastroidea (behavior 4 Basibl1Yl1k and Ql1icke 1999Ь) Gibson (1999) has reviewed hypotheses сопсегпiпg the sister grol1p геlаtiОП5Ырs of chalcidoid wasps Не did not fiпd сопviпсiпg support for RasnitsYl1s (1988) platyshygastroid + (chalcidoids + mушагоmmаtоids) hypothshyesis bl1t instead for а clade compri5ing Platygastroishydea Proctotrupoidea Pelecinidae and Vanhorniidae Не a1so sнggested that the Chalcidoidea and Муshymагоmшаtоidеаmау Ье тоге closely related to either the Diapriidae or Сегарhroпidае rather than to t1le Platygastroidea

We fОl1пd а spur at the base of еас SТВ (character 6) which seems to restrict the mobility of the 1Iюг

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+ + + ~ ~~v~+ь+ + ~I + + ~ ь I + + b~ middotamp~~vI+ + $+o~ ~I+ ~~~ + + ~Ifgtb+ ~~ +lt q + + +~f + bfgt 90 ~ьф()~ ZmiddotI+ I +

raquo~+О~~vt~(~lt-~()Ь~~~+~~tь~ь~~Ь~1ь~~O~~~~~t~~fmiddot~~о$~+ ~~$Ь~Ь~~~~~О~()Ь~~=~~~$ь~ьt~~~~Ь~~~О--~Ь~~~~~ЬО~6~~~q~1~~~~~~~~~~~lt)~O~~~~t~q~~lt~qОltgtO~О~~ltgt~~O~middotampQvQОО-gtltf~q~~amplt~~~~~~lti)-~О~~~ltlt6~~fI~o~~Q~Qo~~~iо~lt)ф1~~о~lti~1lti-gt~(~OJ

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7 I I 6

7Т I т

2

4

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7 I I I I I I 1 3 gJ (j gtshy

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сgtFig12 Unambiguous character state changes оЕ orbicula and associated structures superimposed оп the most recent phylogenetic hypothesis ofRonquist et аl (1999) resulting

ijfrom impJied weight analysis Numbers оп the cladogram correspond to the character numbers (ТаЫе 1) Character state changes for each character are explained in the text

w

~- -А д j ~ л ~ r ~ _ ~ gt -- -1 iI- _- 1 1-1-1 1- 1 ~ olJc j J

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

References Cited

Аllеп R Т and G Е Ball 1980 Synopsis of Mexican taxa of the Loxandrus series (Coleoptera Carabidae Pteroshystichini) Trans Ат Entomol Soc 105481-575

Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

DowtOIl М and А D Austin 1994 Molecular phylogeny ofthe insect order Нушегюрtега apocritan relationships Proc Natl Acad Sci USA 91 9911-9915

DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

FrazierJ L 1985 Ncrvous system sensory system рр 287shy356 lп М S Bluт [ed] Fundamentals of insect physishyology Wiley Nev York

Ganesalingam У К 1972 Апаtошу апd histology of tlle БеПБе огgапs of the ovipositor of tlle ichncunlOnid wasp Devorgilla canescens J Inscct Physiol 18 1857-1868

Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

Konigsmann Е 1976 Das phylogenetische Systcm der Нуshymenoptelmiddota Теil IEinftihrung Grundplanmerkmale Schwestergruppe und Fossilfunde Dtsch Епtопюl Z23 251-279

Konig~mann Е 1977 Das phylogenetische System der Нуshyшепорtега Теil 2 Sympllyta Dtsch Entomol Z 24 1-40

Копigsmanп Е 1978а Das phylogenetische Sуstеш dcr Нуmепорtега Teil 3 TerebraIltes (Unterordnung Ароshycrita) Dtsch Entomol Z 25 1-55

Konigsmann Е 1978Ь Das рhуlоgепеtisсhе Sуstеш der HYlllenoptera Теil 4 Aculeata (Uпtегоrdпuпg Ароshycrita) Dtsch Entomol Z 25 365-435

LaSalle J апd 1 D Gauld 1993 Нуmспорtега апd biodishyversity сАВ Iпtегпаtiопаl Wallingford

LeRalec А апd Е Wajnberg 1990 Sensory receptors of the ovipositor of Trichogram11Ul trшidis (Нут Trishyсhоgгаmшаtidас) Епtоmорhаgа 35 293-299

LeRalec А J М Rabasse апd Е Vajnberg 1996 Сотshyparativc morphology of the oipositor of Боте parasitic

634 ANNAI5 Ш ТНЕ ENTOMOLOGICAI~ SOCIEТY OF AМERICA Vol 93 по 3

Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

Mclver S В 1985 Mechanoreception рр 71-132 lп G А Kerkut and L I Gilbert [eds] Comprehensive insect physiology biochemistry and pharmacology vol 6 Pershygamon Oxford

МеуЬOfес В J Сшш and S Dorn 1997 МесЬапо- and chemoreceptors and their possible role in host-location behavior of Sympiesis seпceklJrnis (Нушепорtеrа Euloshyphidae) Апп Entomol Soc Аm 90 208-219

Navasero В С and G W Elzen 1991 Sensilla оп the antennae foretarsi and palpi of Microplitis croceipes (Cresson) (Hymenoptera Braconidae) Proc Entomol Soc Wash 93 373-347

Rasnitsyn А Р 1969 Oтigin and evolution of the lower Hymel1optera Trans Paleol1tol Iпst Acad Sci USSR 123 1-196

Rasnitsyn А Р 1980 Origin and evolution of Нуmепорshytera Тсanо Paleol1tol Inst Acad Sci USSR 174 1-192

Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

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633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

References Cited

Аllеп R Т and G Е Ball 1980 Synopsis of Mexican taxa of the Loxandrus series (Coleoptera Carabidae Pteroshystichini) Trans Ат Entomol Soc 105481-575

Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

DowtOIl М and А D Austin 1994 Molecular phylogeny ofthe insect order Нушегюрtега apocritan relationships Proc Natl Acad Sci USA 91 9911-9915

DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

FrazierJ L 1985 Ncrvous system sensory system рр 287shy356 lп М S Bluт [ed] Fundamentals of insect physishyology Wiley Nev York

Ganesalingam У К 1972 Апаtошу апd histology of tlle БеПБе огgапs of the ovipositor of tlle ichncunlOnid wasp Devorgilla canescens J Inscct Physiol 18 1857-1868

Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

Konigsmann Е 1976 Das phylogenetische Systcm der Нуshymenoptelmiddota Теil IEinftihrung Grundplanmerkmale Schwestergruppe und Fossilfunde Dtsch Епtопюl Z23 251-279

Konig~mann Е 1977 Das phylogenetische System der Нуshyшепорtега Теil 2 Sympllyta Dtsch Entomol Z 24 1-40

Копigsmanп Е 1978а Das phylogenetische Sуstеш dcr Нуmепорtега Teil 3 TerebraIltes (Unterordnung Ароshycrita) Dtsch Entomol Z 25 1-55

Konigsmann Е 1978Ь Das рhуlоgепеtisсhе Sуstеш der HYlllenoptera Теil 4 Aculeata (Uпtегоrdпuпg Ароshycrita) Dtsch Entomol Z 25 365-435

LaSalle J апd 1 D Gauld 1993 Нуmспорtега апd biodishyversity сАВ Iпtегпаtiопаl Wallingford

LeRalec А апd Е Wajnberg 1990 Sensory receptors of the ovipositor of Trichogram11Ul trшidis (Нут Trishyсhоgгаmшаtidас) Епtоmорhаgа 35 293-299

LeRalec А J М Rabasse апd Е Vajnberg 1996 Сотshyparativc morphology of the oipositor of Боте parasitic

634 ANNAI5 Ш ТНЕ ENTOMOLOGICAI~ SOCIEТY OF AМERICA Vol 93 по 3

Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

Mclver S В 1985 Mechanoreception рр 71-132 lп G А Kerkut and L I Gilbert [eds] Comprehensive insect physiology biochemistry and pharmacology vol 6 Pershygamon Oxford

МеуЬOfес В J Сшш and S Dorn 1997 МесЬапо- and chemoreceptors and their possible role in host-location behavior of Sympiesis seпceklJrnis (Нушепорtеrа Euloshyphidae) Апп Entomol Soc Аm 90 208-219

Navasero В С and G W Elzen 1991 Sensilla оп the antennae foretarsi and palpi of Microplitis croceipes (Cresson) (Hymenoptera Braconidae) Proc Entomol Soc Wash 93 373-347

Rasnitsyn А Р 1969 Oтigin and evolution of the lower Hymel1optera Trans Paleol1tol Iпst Acad Sci USSR 123 1-196

Rasnitsyn А Р 1980 Origin and evolution of Нуmепорshytera Тсanо Paleol1tol Inst Acad Sci USSR 174 1-192

Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

633 Мау 2000 BASllilTYUK ЕТ ЛL ORВICULA Ш НУМЕКОРТЕRЛ

bull

truпk to keep the seta роiпtiпg distally (Figs 8 апd 10) These БРШБ are found in the Chalcidoidea (with ехshyсерtiопs of Боте Chalcididae апd Leucospidae) апd iп the uпdеsсriЬеd New Zеаhшd [атНу (Fig 12) А siшilаr structure but with а different оriепtаtiоп a1so is present in some membels of the Супiроidеа

Sепsillа Campaniformia ТЬе presence апd пumЬеr of sепsillа with ап епсirсliпg ridge (Figs 2 апd 3 character 7) iп а few sуmрhуtап апd acu1eate апd sеvеrа1попасulеаtе аросritап fаmШеs do поt provide support for апу major clade iп a1tell1ative hypotheses (Rаsпitsуп 1988 Brothers апd Carpel1ter 1993 УНshyhе1msеп 1997 Rопquist et а1 1999) except that of mопорhу1у of the Ichl1еumопоidеа Its ОССllпеl1се iп some [ашiliеБ оfЕvапiоmоrphа ие the Megalyridae Тrigопаlidае апd iп Боше members of Stерhапidае) шау iпdicаtе rеlаtiопshiрs amol1g these fаmШеs Howshyever iп the givеп hypothesis (Rопqпist et al 1999) it reqllires tVO extra steps assumiпg that this character is fixed онсе iH the Apocrita c1ade al1d sпЬsеqllенt1у lost iп the Aculeata clade апd iл the liпеagе iпсludiпg Еvапiоidса + rеmаiпiпg taxa (Fig 12) ТЬе ртеБСl1се of sепsillа саmрапifоrmiа without ап

епсirсliпg ridge (Figs 4 апd 8 charactcr 8) has Ьееп rcportcd prcviollsly iп ап e1l10phid wasp (Meyhofer et

al 1997) апd appears to Ье рrеsепt iп almost аН Chalshycidoidea апd Diapriidae iп mапу Sceliol1idae апd possibly also iп thc ппdеsсriЬеd Ncw Zеalапd fапlilу Prеsепсс iп ti1is last family bas yet to Ьс сопfirmсd Ьу stпdуiпg тоте matcrial рartiспlarlу sресimепs freshly kil1ed апd preserved iп еthапоl Howevcr it сап ЬС proposed as а рпtаtivс sупаРОПlОrphу for tl1c СЬаlshycidoidea Diapriidae апd Sсеliопidае

Тl1e роsitiоп (character 9) of thc sCl1silla camparlishyformia does поt provide Sllpport for апу major grollpshyiпg iп the givеп рhуlоgепу (Rопquist ct аl 1999) Nevcrthcless iл the Chalcidoidca the пumЬеr апd роsitiоп of SCB arc сопsistеllt alld ате probably апshytapomorphic for tllis sllpcrfamily (ТаЫс 1) ТО assess thc validity of these plltative sупароmоrshy

phies а рarsimопу апаlуsis is needed that iпсludеs аН data that ЬаУС rесепtlу Ьесоше avai1able Such ап апаlуsis is Сllrrепtlу Ьсiпg carried out al1d the [иП data set is ауаНаЫе (httрwwwbiоiсаснkrеsеarсhl dlqhymphylpau)

Асkпоwlеdgmепts

We thank the staff~ of electron microscopy and photoshygraphic units in NlLf (London) for their assistance Kees уап Achterberg Nando Bin Barry Bolton John LaSalle John Noyes ~ssirno Оlшi Andy Polaszek Frank уап Уееп АпshyпеНе Walker апd Dougas Yu donated Боте of the material studied John LaSalle and Апdу Polaszek are thапkеd for sоше idепtifiсаtiопs This work was supported iп part Ьу а Lеvегlшlmе Trust research grant to DJ~JQ and MGF and Ьу the NERC Iпitiаtivе iп Тахопоту

References Cited

Аllеп R Т and G Е Ball 1980 Synopsis of Mexican taxa of the Loxandrus series (Coleoptera Carabidae Pteroshystichini) Trans Ат Entomol Soc 105481-575

Basibuyuk Н Н and DLJ Quicke 1997 Hamuli iп the Нушепорtега (fusecta) and their phylogenetic iшрliсаshytion5 J Nat Hist 31 1563-1585

Basibuyuk Н Н and DLJ Quicke 1999а Gross morpholshyogy of пшltiрогоus plate sensilla in the Hymenoptera (Insecta) Zool Scr 28 51-67

Basibuyuk Н Н and DLJ Quicke 1999Ь Gгоошiпg ЬеshyIшviоuгs i11 the Hyme110ptera (Iпsесtа) potential рЬуshylogenetic significance Zool J ипп Soc 125 349-382

Brotllers D J 1975 Ply]ogeny aIld cla5sification of tlle aculeate Hymenoptera with special reference to Mutilshylidae КаПБ Sci Bull 50 483-648

Brotllers D J and J М Carpenter ]993 Рhуlоgепу of Aculeata Chrysidoidea апd Vespoidea (Hymenoptera) J Нуrп Res 2 227-304

DowtOIl М and А D Austin 1994 Molecular phylogeny ofthe insect order Нушегюрtега apocritan relationships Proc Natl Acad Sci USA 91 9911-9915

DОVltоп М апd А D Austin 1995 Increased genetic dishyversity ill mitochondrial genes is correlated with the еуоshylution ofparasitism in the Hymenoptela J Mol Evol 41 958-965

Dowton М andA D Austin 1997 Evidcnce for AT-transshyversion bias ill wasp (Hymenoptera Symphyta) mitoshychondrial gcnes and its iшрliсаtiОIlS [or tlle origin оЕ parasitism J Mol Evol 44 398-405

Dowton М А D Austin N DilIon alld Е Bartowsky 1997 Molecular phylogeny of tllC apocritan wasps the Procshytotrupomorpha and ЕvапiОПlOrphа Syst Епtотоl 22 245-255

FrazierJ L 1985 Ncrvous system sensory system рр 287shy356 lп М S Bluт [ed] Fundamentals of insect physishyology Wiley Nev York

Ganesalingam У К 1972 Апаtошу апd histology of tlle БеПБе огgапs of the ovipositor of tlle ichncunlOnid wasp Devorgilla canescens J Inscct Physiol 18 1857-1868

Gauld 1 D апd В Bolton 1988 Тhe Hymcnoptera British Мusеuш Natural History-Oxford University Press Lonshydon

Gibson GAP 1999 Sister-group relationships ofthe Platyshygastroidea and Chalcidoidea (Hymenoptera)-an altershynate llypothesis to Rasnitsyn (1988) Zoologica Scr 28

Greany Р D S D Нawke Т С Carlysle and D W Anthony 1977 Sense organs in tlш ovipositor of Bioshysteres (Opius) lDngicaudatus а parasite of the Caribbean fruit fly Аrwstrерlю sшрensа Апп Епtошоl Soc Ат 70 319-321

Konigsmann Е 1976 Das phylogenetische Systcm der Нуshymenoptelmiddota Теil IEinftihrung Grundplanmerkmale Schwestergruppe und Fossilfunde Dtsch Епtопюl Z23 251-279

Konig~mann Е 1977 Das phylogenetische System der Нуshyшепорtега Теil 2 Sympllyta Dtsch Entomol Z 24 1-40

Копigsmanп Е 1978а Das phylogenetische Sуstеш dcr Нуmепорtега Teil 3 TerebraIltes (Unterordnung Ароshycrita) Dtsch Entomol Z 25 1-55

Konigsmann Е 1978Ь Das рhуlоgепеtisсhе Sуstеш der HYlllenoptera Теil 4 Aculeata (Uпtегоrdпuпg Ароshycrita) Dtsch Entomol Z 25 365-435

LaSalle J апd 1 D Gauld 1993 Нуmспорtега апd biodishyversity сАВ Iпtегпаtiопаl Wallingford

LeRalec А апd Е Wajnberg 1990 Sensory receptors of the ovipositor of Trichogram11Ul trшidis (Нут Trishyсhоgгаmшаtidас) Епtоmорhаgа 35 293-299

LeRalec А J М Rabasse апd Е Vajnberg 1996 Сотshyparativc morphology of the oipositor of Боте parasitic

634 ANNAI5 Ш ТНЕ ENTOMOLOGICAI~ SOCIEТY OF AМERICA Vol 93 по 3

Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

Mclver S В 1985 Mechanoreception рр 71-132 lп G А Kerkut and L I Gilbert [eds] Comprehensive insect physiology biochemistry and pharmacology vol 6 Pershygamon Oxford

МеуЬOfес В J Сшш and S Dorn 1997 МесЬапо- and chemoreceptors and their possible role in host-location behavior of Sympiesis seпceklJrnis (Нушепорtеrа Euloshyphidae) Апп Entomol Soc Аm 90 208-219

Navasero В С and G W Elzen 1991 Sensilla оп the antennae foretarsi and palpi of Microplitis croceipes (Cresson) (Hymenoptera Braconidae) Proc Entomol Soc Wash 93 373-347

Rasnitsyn А Р 1969 Oтigin and evolution of the lower Hymel1optera Trans Paleol1tol Iпst Acad Sci USSR 123 1-196

Rasnitsyn А Р 1980 Origin and evolution of Нуmепорshytera Тсanо Paleol1tol Inst Acad Sci USSR 174 1-192

Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

634 ANNAI5 Ш ТНЕ ENTOMOLOGICAI~ SOCIEТY OF AМERICA Vol 93 по 3

Hymenoptera in relation to characteristics of their host Сап Entomol 128 413-433

Mclver S В 1985 Mechanoreception рр 71-132 lп G А Kerkut and L I Gilbert [eds] Comprehensive insect physiology biochemistry and pharmacology vol 6 Pershygamon Oxford

МеуЬOfес В J Сшш and S Dorn 1997 МесЬапо- and chemoreceptors and their possible role in host-location behavior of Sympiesis seпceklJrnis (Нушепорtеrа Euloshyphidae) Апп Entomol Soc Аm 90 208-219

Navasero В С and G W Elzen 1991 Sensilla оп the antennae foretarsi and palpi of Microplitis croceipes (Cresson) (Hymenoptera Braconidae) Proc Entomol Soc Wash 93 373-347

Rasnitsyn А Р 1969 Oтigin and evolution of the lower Hymel1optera Trans Paleol1tol Iпst Acad Sci USSR 123 1-196

Rasnitsyn А Р 1980 Origin and evolution of Нуmепорshytera Тсanо Paleol1tol Inst Acad Sci USSR 174 1-192

Appendix 1 List of Material Examined

Xyeloidea Xyelidae Macroxyela ор ~ USA М аепеа (Norton) lt N earctic М ferruginea (Say) lt N Атепса Megaxyela bicowripes (Rohwer) lt USA М gigantea Mocsary lt Косеа Pleoneura соshyniferaum Hartig lt Еигоре Р numidica Benson ~ N Мпса Xyela sp о USA Х julii (Brebisson) lt пк Xyelecia пеагспса Ross lt W Nеюmiddotсtiс

Pamphilioidea Меgаlоdопtеsidае Megawdontes esshycalerai Копоw lt Israel М kuznetzovi Dоvпаг-Zароlshyskij lt о Turkmenistan М merceti Konow lt Spain М olivieri (Brul1e) lt Israel М рhаeniсiш (LepeleshyНег) о lt Тшkеу М turcicus (Mocsary) lt Turkey Pamphiliidae Acantholyda erythrocephala (L) lt Сапаdа А jlavomarginata Маа lt Сhiпа Celiikptera nwculipennis (Stеiп) lt Turkey Neurotoma sаltlшm L lt Turkey Cephalcia aruensis Panzer lt Italy Раmрhiliш fumipennis Curtis ~ UK Р stramineipes (Hartig) lt Czechosovakia Раmрhiliш sylvaticus (L) lt UK

Tenthredinoidea Argidae Arge суапосгосеа (Forshyster) ~ France А ustulata (L) ~ UX Scobina poecila Кlug lt МеЫсо Blastocotomidae Blasticoshytoma filiceti расфса MaIaise lt Japan В nipponica Takauchi lt Japan Cimbicidae Abia candens Konow ~ UX АЬш sericea (L) lt Еигоре Cimbex sp ~ UK Trialwsoma sp ~ Norway Diprionidae Diprion pini (L) о UK Macrodiprion петorаиоУ Enslin lt Сеттanу Мопосtenш juniperi L lt SwitshyzerJand Neodiprion sertifer (Geoffroy lt UX Pergishydae Lophyrotoma zonalis (Rohwer) lt Australia Perga dorsalis Leach lt Australia Р1lуlaсteорюgа sp ~ AustraIia gen sp (Acordlllecerinae) lt Brasil Tenthredinidae Aneugmenus ор о UK Аllапtш sp lt пк Сгоеsш sp lt UK Dolerus ор lt UX Еиига sp lt Sweden Мопорlюdnoides sp lt UK Nеmatш sp lt UX Pri~tiphora pallidiventris FalJen ~Netherlands Priophoms sp lt UX Selandria sp

Rasnitsyn А Р 1988 АI1 ol1tline of evolutiol1 of the l1opterols insects (Order Vespida) Oтient Insects 22 115-145

Ronqlist F ]999 Phylogeny of the Hymenoptera (Insecshyta) the state of the art Zool Scr 28

Ronquist F А Р Rasnitsyn А Воу К Eriksson and М Lindgren 1999 Phylogeny of the Hymenoptera а саshydistic reanaysis ofRasnitsyns (1988) data Zool Scr 28

Sсhrпidt J М and J В Srпith 1987 The external БеПБОТУ morphology of the legs and hairplate system of fешае Trichogramma minutus Вilеу (Hymenoptera Tтishyсhоgrашmаtidае) Proc В Soc Lond Ser В Тахоп 232 323-366

Vilhеlrпsеп L 1997 The phylogeny oflower Hymenoptera (Insecta) with а оиmmату of the eary еvопtiопary ЫБshytory of the order J Zool Syst Evol ВеБ 35 49-70

Received for publication 27 August 1999 accepted 8 DeceuromshyЬеУ 1999

lt UK Tenthredo sp ~ UX Trichwcampus sp ~ пк СерЬоЫеа Cephidae Calameuta filiformis Evershy

smann lt UX С filum Gllssokovskii lt Rllssia С hemorrlOidalis (F) lt Turkey С idolon Rossi lt Тшkеу СерIШ pygmaeus (L) lt пк Hartigia sp lt USA Н linearis Schrank ~ UX Jапшfеmоratш Curtis ~ UX Trachelu~ tаЬidш (F) ~ пк

Siricoidea sl Siricidae Eriotremex fоrmоsапш (Matsumura) lt Oriental Sirex edwardsii Brulle ~

Uruguay S noаШо F lt о UX Тшkеу Тгетех sp lt о USA Т apicalis Маtsumша lt о Japan 1 columba (L) 2 о USA Т jUscicomis F lt Czechoslovakia Urocerus аntenulаtш Marlatt lt Jashyрап И alblcornis (F) lt USA и californicus Noton lt о USA и cressoni Norton lt о USA И gigL~ (L) ~ о UX и taxodii (Ashmead) 2 USA Xeris ор lt UX x11Wrrisoni (Cresson) ~ о USA Х spectrum (L) ~ о Turkey Х tarsalis (Cresson) lt о USA Xiphydriidae Вгасhухiрhш grandis Philshyippi lt СЬНе Derecyta jakowlewi Tschitscherine lt N eotropics Xiplydria prowngata (Geoffroy) 2 UK gensp (Xiphydriinae) lt USA Апахуеlidае Synshytexis libacedrii Rohwer lt о USA

Orussoidea Orнssidae Сhаliпш haugi BIlYsson lt W АfПса С schulthessi Guiglia ~ о S Africa Guiglia schauinslandi (Ashmead) lt SA Mocsarya mеtаЛiса (Mocsary) lt Sri Lапkа М syriaca Benson lt о Greece Тшkеу Stirocorsia maculipennis (Smith) lt New Guinea S trifasciata (Cameron) lt М New Guinea ОrшsшаЬiеtiпш (Scopoli) lt Palaeshyarctic О occidentalis (Cresson) lt о USA

Trigonaloidea Тгigопаlidае Lycogaster pullata Shllckard lt N America Orthogonalys pltlchella (Cresson) lt USA Poeciwgonaws costalis (Cresshy50П) lt USA Trigonalis hahnii (Schulz) ~ Nethshyerlands

Evanioidea Evaniidae Evania appendigaster (L) 2 USA Evaniella sp ~ Zimbawe Semaeomiya sp lt Peru Aulacidae Aularus striаtш Jurine lt UK

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

Мау 2000 BASIВUi1IK ЕТ AL ОRВIСШl OF HYMENOPТERA 635

Pristaulacus ambiguus Schletterer Brasil Pristaulacus j sp ~ по data Gasteruptriidae Gasteruption assector

(L) о uK Hyptiogaster arenicolo Тшпег о Апsshytralia Н usymmetrica Turner ~ AustraIia

Megalyroidea Megalyridae Carminator ater Shaw ~ F Thailand Dinapsis ochulalzirta Hedqvist lt Madagascaг Megalyra fusciipennis Vestwood ~Ausshytralia

Stephanoidea Stephanidae Diastephanus sp lt Thai1and Parastephanellus spp 2 ~ lt Brasi1 Peru Stephanus froggatti Сашегоп ~ Solomon Islands

Ceraphronoidea Ceraphronidae Aphanogmus sp ~ USA Ceraphron spp ~NepaI С aguinaldi Desshysart ~Malaysia Megaspilidac gen sp 2 lt lt USA Dendrocerus aphidium (Rondani) О UK D сагрепshyteri (Curtis) lt Ux Trichostereisglabra (Boheman) ~ Сгеесе

Cynipoidea Cynipidae gen sp nr Aulacidea ~ UXAndricus kollari (Hartig) ~ UX NеurotепlS sp lt UK Figitidae Allltgtxysta ramulifera (Thomson) ~ UX А - (sp close to ramulifera) ~ UK A1Jictrix Westwood~ UКАпасfюгisеuсhаrioidеs (Dalman) ~ UX Callagtpidia defonscoloтbei Dahlbom ~ UX Figites anthomyiarum ВоисЬе ~ UX Ююрtrоmeris heptltgtma (Hartig) ~ по data Ibaliidae lbalia leucoshyspoides (Носkепwаrt) ~ UX Hopteridae Liapteron sp ~ Malaysia

Chalcidoidea Аgаопidае Ceratosolen sp ~ Кепуа Арhеliпidае Aphytis sp ~ UK Encarsia inaron (Walker) ~ UK Chalci(lidae Brachymeria sp ~ Кепуа Сопит sp lt Argentina Elasmidae Elasm11S sp ~Italy Encyrtidae Сюгеiа inepta (Dalman) ~ UK EuJophidae Aprostocetus microcosтu~ (Girault) ~ Апstгаliа Euderus sp lt UX Eupelectrus sp lt Costa Бiса Pediobius jUfI)Ш (СаЬап) lt Sudan Tetshyгаsticfzш schoenobii Ferriere ~ MaIaysia Eupelmishydae Еuреlтш sp ~ India Metapelma spectabile Westwood ~ USA Phlebopenes sp lt Argentina Eurytomidae Bephratelloides paraguayensis (Crawshyford) ~ Trinidad ЕUПjtаmа spp ~India UX Leushycospidae Leucospis slossonae Weld ~ USA Polisshyt01narpha surinamensis (Vestwood) lt Colombia Р сопuга Boucek ~ Brasil Mymaridae gen sp laquo Costa Бiса Мутаг sp ~ ПК Gonatocerus sp ~ UK Ormyridae Ormyrus sp ~ Sri Lапkа О nшshydttlus (F) ~ UK Ptcromalidae Gastracanthus pulshycherrimus Westwood ~ UK Plutotftrix sp ~ UK Pteromalus sp lt Malasia Pteramalus luzonen~is Саshyhan ~ Malaysia Spalangia sp о Кепуа Trichomashylopsis sp ~ Italy Signiphoridae Signiphora sp ~по data Trichogrammatidae gen sp ~ Costa Бiса Trishychogramma ]юumiеri Pintureau amp Babault ~Coтoгo 15 Torymidae Megustigml1S sp ~ Singapore Мymarоmшаtоidеа Myrnaгommatidae gen sp

uпkпоwп Platygastroidea Platygastridae gen sp ~ UK

Platygustersp 2 ~ ~UX Р phragmitis (Schrank) о Bohemia Leptacis sp ~ Peru Sсеliопidае gen sp ~ о Nepal лсапthosсеliо sp ~ Реш Gryon sp ~ Pcru С misellum Haliday ~ UX Telenamus sp о UK Т zygaenae Кieffer F Dаgеstап TrissoZcussp ~ UX Trimorus sp ~ UK Scelio sp ~ UX

Рrосtоtrпроidеа Diapriidae gen sp (Рsiliпi) о Кепуа Basalys sp lt uK Coptera occidentalis Muesebeck ~ Italy Sрilomiсп1S sp ~ UK S апshynulicomis Кieffer ~ UK Мinюрriа barbata Borgshyшеiеr ~ Brasil Platymischus dilatatus Westwood О UK Propsi[ltgtmma coluтbianum (Ashmead) ~ USA Heloridae НеlопlS spp ~ О UX Sweden Pelecinidae Pelecinus polyturator (Drury) о Сапshyada Proctotrupidae СоdпlS ligаtш Nees ~ UK Сryрtоsегрhш sp ~ UK Proctotrupes gladiator НаshyJiday lt UX Roproniidae Ropronia garmani Ashshymead ~ USA Vanhorniidae Vanhomia eucnemidashyrum Crawford ~ Сапаdа

Iсhпеumопоidеа Вгаеопidае Agathis сlошtflаliаshynиs (Ratzeburg) ~ по data Braconfulvipes Nees ~ Netherlands Charmon eruentat1S (Haliday) ~ Коshyсеа Coccygiclium sp laquo Rhodesia Coeloide-s sp ~ UK Disophrys caesa Кlug F lt Sраiп Doryctesfаrtш (Provancher) ~ Canada Glyptapanteles obliquae (Wilk) ~ Тпdiа lsopronotum foveola (Вшllе) ~ Sierra Lеопе Honwlobus sp О Malaysia LabagatJlis rufoatra Enderlein ~ Peru Liobracon sp ~ Brasil Macrocentrus иnеаПs Nees ~ Hungary Meteorus vershysicolor (Wesmael) lt Zaire Руgоstоlш sticticus (F) ~ NеthегJапds Rogas sp ~ NepaJ Stantonia sp lt Zimbawe TrachypetllS clavatus Gucrin lt Australia

Ichпеuшопidае АсrоtарftШ tiblalis (Cameron) lt Costa Бiса Arotes аmоеnш Сгеssоп ~Canada Ватуshy[ура wlcata (Provancher) ~ USA Brachycyrtus sp ~ Нuпgаrу Diacritus асiсulаtш (УоllепЬоуеп) lt UK Ganode-s matai Gauld ~ Costa Бiса Labena sp ~ Costa Бiса Labium longiceps (Cameron) о Tasshymania Меtорiш sp laquo Сатесоип Opfzion sp ~ UK Рltуtodietш fumiferanae Rohwer ~ UK Rhyssa ретshysuasoria (L) lt France T1Uronia sp ~ Brasil Тпshyclistus podagricus (Gravenhorst) lt UX ТПjрlюn signator Gravenhorst О UK

Chrysidoidea Bethylidae Bethylus fuscicomis Оиshyппе) о UK Pristocera laticomis Кieffer о NyasshysaIand PristolJethylus ceresensis Turner laquo S АЫса Trachepyris prepinquus Тигпес lt S Africa Chrysishydidae Adelphe anisornarphae Krombein ~ USA ТriсhПjsis суапеа (L) ~ UX Dryiпidае Alphadryishynus lamelati Olmi lt Brasil Сhеlogyпш cameroni (Кieffer) ~ о UK Меgаdryiпш magniicus Richshyards ~Агgепtiпа Mesodryinus dayi Olmi ~ Greece Prenanteon sp о UK Embolemidae лтрuliсоmоrshypha tаiшапensis Оlшi lt Taiwan ЕmЬоleтnш sp lt Canada Е rudii Westwood ~ UK Plumariidae gen sp ~ Nambia Plttmarius sp о Аrgепtiпе Plushymoroides andalgalensis Brothers о Argentine Sclerogibbidae gen sp (Sclerogibbini) ~ Nepal Sclerogibba citipes КromЬеiп ~ Тпта S magretti Кieffer ~ о Уеmеп Seolebythidae Clystopsenella sp ~ Costa Rica Scolebytu~ medecusS11s Evans о Madagascar Ycaploca evansi Nagy ~ S АЫса

Vespoidea Bradynobaenidae ВrаclупоЬаепш сапш Vestwood о Chile В gayi Spinola о Chile Chyshyphote~ sp о UsA Doryleika sp О Оmап Typhoctes sp о Costa Бiса Т pecuiaris Cresson ~ UsA Formicidae Aenictus sp lt Саmегоuп DоlichоdепlS cl1spidatus (Smith) ~Воrпео Dоrуlшfulvus (Westshy

shy

636 AclNAI~S OF 11ш ENTOMOLOGICAL SOCIEТY OF AMERICA Vol 93 по 3 ~ t

wood) lt Ghana Gnamptogenys annulata (Мауг) lt S Атепса Labidu~ praedator (Smith) lt Бгasil Messhysor galla (Мауг) lt Ghana Polyrhachis militaris (F) lt Ghana Tetrapanera rufonigra (Jerdon) lt India Mutillidae Oclontoplwtopsis aulus Бlаkе lt3 СЬНе Phatopsi~ imperialis Бlakе USA Pseudmnetlwca sanbornii acetis (Fох) lt USA Pseuclophatopsis bingshylютi Бisсhоff lt Saudi АгаЫа Р continua F lt lt3 Nigeria Timulla bagracla Cameron lt3 Боrnео Т ershyemita Mickel lt lt3 Philippines Pompilidae Anoplius nigerrimus (Scopoli) lt UX Ceropales sp lt UK Rhopalosomatidae Rhoplosoma sp lt3 Costa Юса Н nearcticum Бrиеs lt lt3 Costa Rica Н роеу Cres50n lt3 Сауmап I51 Paniscomimma erlangeriana Endershylein lt Angola Liosphex varius Townes lt USA Sapygidae Fedtschenkia antflracina (Ashmead) lt lt3 USA Polochrumrepanclum Spinola lt lt3 Poland Turkey Sapyga morawitzi Turner lt3 Turkey Scolishyidae Aurirneris africana Saussure lt lt3 Nyasaland Campsorneriella соlапБ (F) lt lt3 India Mega~colia

~ al~cto Smith lt Malaya М scutellaris СПЬ lt ~ Philippines Scolia glabula Раllа5 lt Turkey Sieroloshy morphidae Sierоlmnоrplю canaclensis Provancher lt

~USA S similis Ashmead lt Costa Юса Tiphiidae Aelurus gayi Spinola lt3 СЬНе Chrysothynnus inca

bull Тшпег lt3 Боliviа Myrmosa atra atra Panzer lt UK

Spilоtllуrщus bituberculata Тшпег lt lt3 Argentina S laetus Кlug lt3 СЬНе Tipllia femorata F lt3 UK ~

Vespidae Belonogastermeneliki Gribodo lt Ethioshy~

рЫа В saussurei Кiгby lt lt3 Yemen Bracllymenes j

clyschera Saussure lt Perи Mini1Ji bra~iliantlsm сотshyрааит (Fox) lt Агgепtiпа Stenogaster fugipennis ~ Gшsectгiп lt New Guiпеа

АроЫеа Apidae Apis dorsata F lt Java А melshylifera (L) lt France Trincostoma (Diagonozu) letshy r tow Vorbccki lt Congo Andrena sigmuncli Cockclcll ~

lt Canada Colletes daviesanus Smith lt lt3 UK Sphcshy i cidac Bembix sp lt Кспуа Ectemnius continuus (F) ~ lt UX

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