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A character analysis of the North American potter wasps (: ; Eumeninae)

Article in Journal of Natural History · October 1985 DOI: 10.1080/00222938500770551

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James Michael Carpenter Jeffrey Cumming American Museum of Natural History Agriculture and Agri-Food Canada

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The user has requested enhancement of the downloaded file. Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 A character analysis character A (Hymenoptera: Vespidae; Eumeninae) Vespidae; (Hymenoptera: JOURNAL Introduction important potential the wasps, some various invest!gate termed now characters well Progress here. genera which not uhr) by authors) some phylogenetic underway study 99 1981, 1979, The This remainder limited as seen However, of treats knowledge a they have new chaotic Eumeninae them aspects comprehensive paper OF in biological representatives for the evolutionary JAMES plan (Accepted ertc eea accompanies genera nearctic Museum on cladogram A cladistic analysis A cladistic the , Cambridge, here University are in to characters. NATURAL Smith Eumeninae been in relationships h senior the detail 26 the the of an the is (Parker states the several of genera are genera subfamily the of abundant similarly the 'stem-group' nearctic their phylogenetic is discussion M. and of 31 only Pterocheilus, evolutionary control for the family. of of first Comparative MA July HISTORY, CARPENTER unnatural generic any questions A behaviour 1966). let, Edmonton, Alberta, 43 Alcock of primary is the of author, primary of apparently analysed. fauna. application character idrd by hindered 02138 U.S.A. and Department and U.S.A. 02138 as 1984) significant nearly the and nearctic of They agents the a of the reclassification whole, of , importance North character very 1980). 1985, lineage Vespidae (Cowan (nonmonophyletic) and relationships The nearctic genera nearctic all the are are aim Zoology, (Bohart genera. ytm ae discussed, are systems portion diverse nonmonophyletic: Therefore, the of as and 19:877 results Eumeninae common of the But of is the American potter wasps potter American of their text. well interest states to 1978). cladistic the JEFFREY (Carpenter results of world However, Alberta of they group, Leptochilus avr University, Harvard sals the establish et as presented here presented characters higher can the Vespidae. 916 in the various no as al. predators of These to subfamily are be genera order appropriate. Eumeninae is presented. is Eumeninae (Zethinae nomenclatural with of analysis 1982) T6G evolutionary built genera classification, Zethus, relatively it M. a 1981). areas and draws greater Commonly used to study 2E3 of of and or CUMMING when ground-plan in is less Microdynerus. be Entomology, Eumeninae, presented. to are of , Also, as and are many by upon the Canada inclusive pursued Thus, investigation the species on unstudied the subjects previous indicated. a is comprehensive first biologists Raphiglossinae changes together framework a ecosystems, Eumeninae. rest which the known study the At richness states groups. This successfully. of higher-level and A least eight data , The addressing authors, the key are has currently as as we (Cowan require refer ground base potter of world either to made upon been than have and We the the as of to of is Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 the review as many generic did Vol. described categories" has the been 1980). morphological in splitting upgraded criticized Blfithgen criticized picture'. did the Taxonomic genera. adopted gist'), authors (1939b:98), rtczd it criticized names species, Van been proceed 878 of 'Eum6niens' the these Dalla into infrasubgeneric dividing ally innovation . Odynerus Saussure their defining the This The traditional The 1ihe (1938 B1/ithgen major not each indeed Eumeninae. 1: der four last specifically of into stated: Bequaert xxiii~xvii) which Torre of same level. Nearctic In Volume In value approach but historical difficulties Vecht his echoed even and form Bequaert to similarly the the (1875: resulted: directions the subgenera, characters 'divisions'. (Parker features). receive interpretations produced history was it for (including for Thus referring European applied 'Such he taxa basis of further, (1904) with fails subfamilies (1963) complexity (1918) classification . has 'overemphasizing different Saussure 148) discussed by at Saussure development this gave he some over 'first (1939: with such several as for 3 a,b) as in a 1966), added first in Parker The treated of continued used to other to indicated provided thus procedure, expecially a late generic He many regarded problem the to it. Gayella), a the from J.M. his guide attention. of species began the discourse 58), strongly characters trend Studies using others.., and (1875). similarly his B1/ithgen's as evolution of evolution of by partly tend (1852-1858, past in 'l~tudes' have as (1965), the neotropical the mr ntrl and natural 'more was 1953, although interpretations Carpenter h history the Saussure faunae, to classification many subgenera has a that this in this of to 45 analytically the names of the with multiplicity in group, prevailed the currently trend following otherwise Nevertheless, Willink however, extreme confuse species years. resisted Odynerus had the on the but it existing generic existing recurrent he recognized and nlig (1975) Snelling the study Odynerus monophyletic results always the treatment introduced Palearctic, eventually various some not but were for o. 1 Vol. of difficulty and their Prior species of in Oriental characters (1967) to rather than rather group splitting underway splitting of are by naming generally not Wesmael those nor 1939, are also from of eumenine positive ill are into the J. veins on natural role the also to by te workers other characters of defined M. divisions Bltithgen's only caused split unacceptable did a present workable of differences, currently 1938, and many far Ethiopian of adopted transitions 'l~tudes') names regional in that Region. of Cumming due classification and has Bliithgen groups he and Microdynerus classifying he results. Hypodynerus the by ignored generic leaves the (1836), relationships.' the further simplify taxonomy. in few or attempt used in therefore van Saussure genera (of. prdn of upgrading largest. forewing multitude of for in large Bohart of variable. part The and genera basis. approach groups' placed have recognized Saussure out der some The and Eumeninae, in many classification and genera have which and by ('a (as is (as his the in measure neotropical fauna his extended the Vecht by and to the He subsequent extreme Australian as German hs been thus Bohart proceeded (1982,1984). further the the complex were did and the and Saussure of and Soika and classification, of relate in h diversity the artificial Kurzenko almost and species divided the from many Thus subgenera. with the these details was group. and others and multiplicity cladogram Odynerus subgenera, not, recognized in to 18 Leptochilus, the however (1948, 1951) (1948, subdivision been taxa at taxa other partitioned the the mandibles both entomolo- Kurzenko has ultimately annectant insensibly genera eumenine divisions. placed ('l~tudes' formally Regions. workers. but because of without trend (1978 history Bohart 'larger (1980) added rarely form'. Soika latter these large This and and and will this has the for he by b) of of of in Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 method (apomorphic) Methods exploratory the usefulness pinned combine characters male literature recognized. nature in Information Drawings (van Kurzenko examined Alberta. all preserved of and Eumeninae, examined is of trenchant date. the Stereoscan (1978 a) opinion of permit these the common', plesiomorphic. computer States Eumeninae, much residues a this the tissues Cladistic Outgroups, More Due significance stem-group, species-group der phylogenetic Stenogastrinae genitalia. A serious present of group. specimens, study, the worse. of between comprehensive to Vecht and The the both than where or as programs. These programs. (1978) in character o the for of (synapomorphies) underlying were establishment 250 left the Kluge However, and but when rehydrated classification analysis analysis glycerin. male on a obstacles Guichard including Some and Study of Further, descriptive 148 in and Fischer and taxon large in electron unless of for these for catalogues available. lumpers as species cover made many the except the it arrangement aeoy (Parker category genitalia genera genitalia. and more well was made. analysis, number (Carpenter 200 can characters can old otherwise of Only the this only to the the dry by generic species (1980) microscope 84~ Farris as not for the and polarity new genus-group and evolutionary of be has 'larger and eea (plesiomorphic) general provide vertex, foveae splitting is included 1972) the the mounted camera employing were representatives a critically may Morphological examined, the splitters, of far of rendered 1100 truly explanatory Whereas characters. omit European progressive internal of key does key are Nearctic the taxa noted, 1969; previous phylogenetic dissected, and Nearctic and categories'. outweighed 1981). consisted was be gold the the species has much rational, at nearctic lucida the recognized 1966) Paravespa specimens)just and inferred the decided; stem-group the basis it this cf. resulted names understanding were coating structure criticism Wagner The n on and nearly incomprehensible Eumeninae U.S.S.R. not paraphyletic authors characters, more Department of may Farris of on slightly y distinguishing By classification power resolution of the for terminology pce. All species. Eumeninae Bequaert's and sectioning majority by exist approach with are (Krombein nearctic only in a and (Nelson and ultimately Euparagiinae, substantive a 78 behaviour, of the of many so have The Wild (Farris ('groups more currently 1982a) particular cleared even Blfithgen's program the character anterior those examination accessible, Cephalochilus, negative analyses of taxa provided by other by provided situation female of used of and cases the is and Platnick and (1939) for M5 rational were exist the characters Entomology, the 1979, is defined 1979). states in were resolve and otherwise having otherwise head stem-group Europe. to was problems. Eumeninae, to considered written as ofparaphyly reference dissection warm best cephalic examined only approach states, effects. were between Gayellini separate statement the lack 1982 b). taxonomy. for in Both capsule usually extracted to lsiiain As classification. are with means by foveae, Carpenter to other for lactophenol, of all implemented and were by Those synapomorphy 1981). group-defining unique The a are any foveae to the but Not a University of University J. genera, in for difference less (of valid genera (Masarinae) microscope. Tobias major they of the Cambridge is S. now out studied characters. in considered removal of semblance the specialists Palearctic piecemeal not alcoholic assessing from Thus today Farris of only the states general and ite in little do (1981). course to in Soika using areas or form were and and 879 not the the are the the the via on in as of of of in a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 for the nothing to is treated (1936, measure higher terminal 'Odynerus Raphiglossoides. the group' even of (1969). Raphiglossinae recognized recognized the homoplasy. swapping the highest character allowed performed, senior Eumeninae. of showed included Raphiglossinae within Swofford Characters that Eumeninae; approximation 'subpedunculate' by The this abandoned most 880 The primitive the the In 1. Zethinae, State Zethinae by Zethinae reason, is, wings character when important Second entire forewing 1974), the author vein refers the genera the Carpenter consistency consistency that study taxa in of University of and consistency s. l.' s. discussion, Ctenochilus indicate genus subfamily, in r-m this by using different using Alastor, Saussure The to Hundreds the it does the but two complex, to submarginal ('terms' having but of on refer that searching occurs rte (1903) Br6thes were state were have fuses and Illinois of characters the individual he This term relative as a as of Bohart request. this (1981) r-m that have considered to not the Zethinae acute for the for Saussure figured as of with two ('l~tudes' Vol. ('l~tudes' synonymized zethine. informal of has characters Eumenes as components and (Farris is cannot in of the New three, Natural occur they J.M. Nelson and second that included. a been for recognized importance mid-tibial groups examined cladograms and been Rs propodeal cell characters that for basis particular codings multiple as are York before Rs + combination It Zethinae in Carpenter split dividing petiolate. be designation 1969). 1875), Stange it similarly closest relatives closest and and Raphiglossinae it is 1979) submarginal History sister for one the to occurs that firmly more contains in The at Stony at from 3: under be (branches) Odynerus the spurs, Bequaert and outgroups. the by and Copies 325), that common parsimonious orifice relative of groups. were more tews have otherwise input (1965), Raphiglossinae the 'Eum6niens' and latter likely of In Saussure used established in traditional these Survey. and combinations characters the Eumeninae. are his and who indicates Hypalastoroides the of examined; a Eumeninae, closely Brook, ally of reaches cell (1Rs) is cell data character the as three . M. J. in to farthest variety The but two characters and the r defined are relatively considered by Richards of authors, However, data a with the old all The it sister-group paleontological (1875). form Zethinae it taxa, primary other matrix. with related Ruiz Cumming both and trees, vespid at in 2r. matrices s closely is PAUP apart of the may Ctenochilus the ite in little present. because may sense, of the of This were raphiglossines. respectively. the characters, plesiomorph genera (1942). uncertainty A petiolate with the it which one the characters. ahgosne by Raphiglossinae to by including be sections and in subfamily utpe iterations Multiple transitional presence may not (1962, has PAUP vary program is Astalor. unnatural. second of that related fig. venation the names the are presented apomorphic common reasonably of related other the evolved Raphiglossoides (a generally within category by 1. 'Eumenes its 'fixit6 its is, fig. was available anteriorly Raphiglossinae: condition of package and be submarginal as in to Ctenochilus The of genera Richards permits 34), elements genera This groupings excluded the placement monophyletic to blanket to a Zethinae so state Zethinae convergently (for considerable a genus. a here subgroup of term provide but provided resulted Zethus, constante', good within approach from included formerly by example ofvol. s. l.' s. (fig. of Hennig there has branch termed (1962) 'stem- of Soika terms D. close from were unit and and and and was and For cell 13); the the the the the as L. in of is it 1. a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 Hypalastoroides/ Microdynerus~ Leptochilus ~ Leptochiloides Maricopodynerus~ Eumenes Pseudodynerus~ -- Euodynerus~ Dolichodynerus Odynerus~ Gastrodynerus ~ Montezumia ,-~_~ Ancistrocerus, S m e r m S e Leucodynerus~ Cephalodynerus- Par Pseudepipona Stenodynerus~ Zethus Paranortonia~ n the in aaicoytrs Mcemns Paralastor). Micreumenes, Paramischocyttarus, the many H ypalastoroides. any some ) short autapomorphy Masarini Masarini Omicron), Leptochilus ancis t r o c Besides 2. truncate in weight Basal Polistinae differences g o d y d o n g at Vespidae. have and this and ~ angle the er condition erus-.-.~ as ~'- some u s point. the it stem-group, Fro. associated this a ~,,~,,~ Many of in very curvature second synapomorphy. 1. Polistinae. the angle (and Rs narrowly is Cladogram Eumeninae height probably may close is submarginal with this obtuse of Nearctic Rs. In relatives be of state the truncate, of the But M, plesiomorphic almost have or truncate or lengthened In the occurs in occurs stem-group the taking cell. ertc genera nearctic h Nearctic, the such Eumeninae it and curvature The acute perpendicular as this the Consequently, a (figs. angle Labus) second number (fig. in distribution Stenogastrinae of the 12, formed of the 14), of and it submarginal 13). Eumeninae. this Eumeninae. Eumeninae, of is s does as in Hypalastoroides. The Eumenes by vein an it the of Rs cannot Gayellini states autopomorphy also (often atr group, latter Euparagia, and cell. s. Vespinae into have M be I. angular (especially is There and accorded account, variable There M many some very 881 and are an in of is Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 number stem-group (cf most variation character others). Rs is length the angle border genera genera, possessing Head than distinction Montezumia have Montezumia and in and to 882 Hypalastoroides. towards Cowan hook rolled). hooked Montezumia World segment 21-23) loose and exclusive may (Odynerus generic of stem-group. have the an the reversal. is figs. 4. Nevertheless, ground-plan 3. Hypalastoroides other strongly the be 'zethines' Vespinae, a a Length arbitrary Male coiling Eumeninae; has is apical of genera (van which of simple long It in and modified synapomorphies, 1978). (Saussure length of of unique the the in form: states, the in may Soika appeared genera). between the dilectus, Zethus dilectus, a eumenine has antennal der is genera From and pterostigma; segments prestigma and arizonensis, very of posterior involving curved pterostigma curls (e.g. When the apex with apex male be have of M prestigma. imposition Vecht and 1979, in state contrary Pseudodynerus The in Monobia. the M a most more it elongate a 1852-1858, Synagris, Rhynchium) this, The over the ground-plan 'about such is is shape it (in several is in the og prestigma long apex. posterior beyond genera is bent back generally Bohart straight, 1963, condition (many often border). 'raphiglossines' modifications other modifications Vespidae J.M. which in parsimonious posteriorly the terminal evidently as 13 Pseudodynerus only to aatga of parastigma has The (some equal' the In upon prestigma. Microdynerus, This segments Parker slossonae), times. slightly nene of antennae Bohart apical such species Carpenter and 'basal the it occasionally Parker nearctic length border some trends, However, short is (most towards state also and species is continuous majority (some segment not Stange as Also, oriented 1966). A supported convergently segments angle oblique (1939 a) (1939 Zethus of have 'no in of generally still very Microdynerus of species 1965) varies interpretation of may as (and fauna, Pterocheilus therefore the the may the male and of quadrisectus, Eumeninae, the more European the considering realized 1965). a truncate'. retained been different is the of been Pterocheilus, prestigma outgroups most long (straighter), but pterostigma, long prestigma (fig. 13), (fig. and preceding be female large and within form J. eumenine of variation may state Polistinae than but by appear myriad. M. the Thus the used a Deuterodiscoelius prestigma Leptochilus + Kurzenko the other by acquired the a occur (Symmorphus). so Cumming synapomorphy from latter half' the half' authors) of It differences during natural and Pachodynerus erynnis, Pachodynerus and and for since it Saussure as (abscissa the hook do appears and state has state Monobia a segments, to may in genera, but They truncate a this characters the figs. Pachodynerus; this with fig. Leptochiloides. have have 'transitional' species be key some stem-group it copulation occurs in (about group oblique (1980). length not and group. within 14) sui relative is state include 19, Calligaster the character a probable formed relatives), ('l~tudes' the widespread in the been described as been appears short generis, and be of 20; forming angle of the The size hook is also have also equal in with possible of these antennae terminal for the for The A of Hypancistrocerus state) Pterocheilus approached the Paranortonia reduction to some the prestigma). coiling are number by (half apical (Cooper nature in Paranortonia, figs. these still the that an acute an that in with antenna to Vol. to and Most it pterostigma genera a R Eumeninae. Leptochilus may minor. separating be hook is within latter Polistinae extending or or to and indicated posterior it 19, antennal oblique. whereas segment no taxa, apically are 1: derived less greater curved specify of of be due of in 26). xxvi). 1955, clear even (figs. (and by may The The Old two and this size not not the the the all all A a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 more S This flagellomere primitive Pterocheilus flagellomere first elongation ment of ment be Eumeninae, variable been cannot group. the to many toothed the been 5-toothed Eumenes although reduction was Little mandible crossed incision reduction. character perpendicular Montezumia-Microdynerus Ancistrocerus World several the modification Pterocheilus. basal the In Montezumia, Euodynerus, ymmorphus). North be a 5. 6. number Striking sporadic extreme ground-plan flagellomere also lutae b Saussure by illustrated further a Mandible. teeth. attention reduced. egh of Length widespread derived. species traditional The species be occasions. a at mandible. or in applies it s. condition, American very mandible or to used rest, this in notch is is mandibles l., This of dimorphism the it is ('much nature in synapomorphy a not one Its with a were Pachodynerus, and up a and teeth short arista is Monobia, smaller character, He to his the as has At slight fourth The presence to may (almost not first always between to three state of few the Another a character n oil wasps social in However, 5 in characterized is key presence basal most, been of listed character to (e.g. Odynerus, longer teeth first the and length very related and found occur flagellomere. its other modification have number be (basal of to species grooves clear Parker flagellomere paid as presence, is so teeth of teeth but primitive quarters have is aaotna Pseudodynerus, Paranortonia, the Pseudepipona genera. impressive, the a by found type than broad') to broad') than long of in other perhaps often in social in of of teeth. component, genera 5-toothed it defining the ('l~tudes', just or to second but distinguishing the the Kurzenko Eumeninae; it only is tyloides eumine (reversal) that of as 1966, Nearctic penultimate) more of modifications state or Zethus the in mandibles correlated features what female, This interlock or this the otherwise ('generalized'), Bequaert Parker state such (Vespinae), the ridges of Pterocheilus, synapomorphic. more upper and occurs (much wasps. the in van but parsimonious scape; character taxonomy. the is on is Vol. form mandible Odynerus (fig. must have must as as only a but (1980), incorrect: third Dolichodynerus-Microdynerus showing Eumeninae is der the of the tooth found on found diagnose it general Paragymnomerus mandible (1966) is 'highly supported less and whether in with interest. the 1: fig. If of (1948) is is 18). more tooth more it in Vecht length associated O. teeth much Plate a the mandible. the but 22) the than has rest in has and and a 5-toothed dilectus, Some and may As used some semicircular condition some widespread: and basal modified' occurred while may Odynerus condition female, used to probably males in 1977). occur of of the Some more XX). been engage half shown Considering reduction Pterocheilus include regard the the the by be the other groups be exterior segments. or with Leptochiloides a the homoplasy the most but other male of scape. Parker It Eumeninae exaggerated attention has overlooked. a in while turned not mandible state few by and several the both When length also om of form males when a in arisen synapomorphy is this in a species genera 4-toothed inflection they in Carpenter mandible where clypeal quite characters. Pachodynerus, Vespidae, edge few Gymnomerus. Eumenines the mandibular the of the (1966) Ancistrocerus interlock. occurs the is Leptochiloides inward The mentioning the have of independently on are times. n elongate an an male stem-group eumenines common number of in is this the possessing range of the from extreme in mandibles emargination. structures does decussate component. the It apomorphic, considered been the mandible of an Zethus of number in the has scape) (1981), at tooth Moreover, where occurs one Develop- opposing the The Odynerus found are quite elongate males not. an male. form surface. of for 4 paid Due in of stem- teeth, of its angle teeth most deep may (e.g. first as The and and Old 883 this and (the this has are has the the the are the in to as In 4- to in of of a a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 teeth and by between the between evolution is tuberculocephalus-group, homoplasy apomorphy (bulging no other no margin and trait autapomorphy states are in similarly. times. 884 may slightly antennal apply Plagiolabra in Pterocheilus the separated visible light component, not spaced also Parker swollen of Odynerus, Microdynerus ocelli the 34, 41; tissue source quite the the eumenine taxonomy. 9. Finally, 7. 8. Presence Bohart Leptochilus-Microdynerus fovea are fovea a present of occurs be Cephalodynerus (platycerus Cephalodynerus of other occurs Cephalic similar state Antennocular Interantennal foveae latter (also Therefore in the to surrounds externally. the during copulation pits (1966) another different frons aibe in variable (fig. or good not Therefore the socket in and Vespidae. palearctic with is not), seen the they in two (1940). nearctic antennae in and literature for in several 35). of characters. present of very position as associated (raphiglossines'; characters a foveae. component, many situated basal these of synapomorphy, other foveae dense this groups form) are few species few characteristic Odynerus in the although diameter) In two does different this a this As several space. profusely character different carina. Pterocheilus A in first Pseudodynerus three projection other few species ('interantennal this pilosity, B1/ithgen vespids, and elongate genus. blunt of not the is J.M. and he condition close These are These Indeed, where with to palearctic (Cooper the one six is depression some groups form A genera, from occur reversal genera, and ewe the between groups stated, support despite is In homologous of ridge wide lineages. Carpenter of haired, genera to antennal is glandular the component. of Leptochilus set of Leptochilus, Leptochiloides Leptochilus, many and it (1983 as of of one diverging of this Pterocheilus in the midway a and and is the in parsimonious could convergence space at usually different the a little few 1955, (Zethus, Microdynerus, Leptochilus). Microdynerus, (Zethus, (some often it their female show another, this a apparent Symmorphus). and best covering a tubercle' has b) eumenines, primary listed deformiceps). slight male is genera. outgroups, few species the use of sockets first also (which group, and Cumming antennal tissue, weak). is unique occurs often been several Euodynerus, between absence visible related great channels et That supported character, third from at mandible depression above used be J. and al.). the reversal and much It of synapomorphies (Pterocheilus--although M. only can which considered A and interpretation in the that variation tooth is Parker generic exteriorly a the in these other genera slight but of socket many Pachodynerus carina more Cumming Euodynerus the in and and This be referred of which condition overall Odynerus. foveae the consequent of in of by the is the may most characterized the is near posterior is Leggett Euodynerus bidens. Pterocheilus bending Leptochiloides a neotropical Ancistrocerus similar pronounced level. 1966). of inflected could are Eumeninae, raised has and possible a vertex Leptochilus-Microdynerus and in the the function are as of (unanalysed the Ancistrocerus). raised to very evidently form. the a Similar the for the found product Secondary as nearctic This Zethus. occipital part bidens be internal in area pair of of behind height ocelli has the vertex in similar synapomorphy prep). eye area one viewed the is In is and as as Hypalastoroides of groups of fovea). in in only other comments often and a was was (fig. evolved they or of more Montezumia- genera, the a and small, females. the Symmorphus, Symmorphus, the organization on ground-plan or of margin of Most Usually Odynerus. pheromone in convergent more loss the tooth one Internally primitive) as There described noted the genera h frons the posterior so 1). cuticular They having carinate, occipital are such than is closely female a of and many is If fovea basal frons clear (figs. This syn- also well is this the are are for so, an by an as of in It it a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 (1977) due close both genera and (Gymnomerus It these invaginations Montezumia paraphyletic this have not 'rudimentary' but common has segmented homoplasy could ground-plan are Pseudepipona). as (1940) mandibles while Pterocheilus with Hawaiian flattened, the turgiceps). several chilus, While in remarkable an state taxonomy Raphiglossa, a the is emargination A 12. 13. necessarily 11. 10. in female. occurred to deep, Pachodynerus reduction the sexes. hairs to of reduced invaginations different also excavating hypothesized, here general Microdynerus) extreme Vertex and males, Occipital several Clypeus. Palpi. have a the different maxillary reduction broadly and semicircular While genus beinterpreted (fig. They maxillary since Carpenter compound and in for In treated + little and state The several tubercles. maxillary Synagris). genera. segments apparently forming and fringed the modification Monobia. genera specific correlated has development Pterocheilus having 15). The genal carina. are they Pseudopterocheilus the its truncate the of number form in of occasions Kurzenko are are has been Males palpi most type for close the common. various times, as and may in time with nest may (1981). the In characters eye a emargination margin. short, best been have of not In Forking some palpi). pointed Eumeninae. synapomorphy, The as exaggerated of has often (e.g. 4-segmented reversal characterize can the a of (fig. foveae relatives. of lower long of (GrisseU have only clypeal and an considered is wide the often of to palpal parallel these shapes deep (e.g. species, (1980) Following development Saussure Pterocheilus number 41). apomorphy Nearctic, We present tubercles be palpal In hairs is labial Nearctic are of clypeus. a clypeal range (fig. and been the loss Some reduced invaginations). similarly the Leptochiloides, agree emargination latter Ancistrocerus segments the These 1975). speculated the in and Since has (fig. it formula 36). basis in not occipital between most (e.g. palpi described. of labial the autapomorphic. may is psammophores (1852-1858). margin (Cephalodynerus species palpi the with generic do Often genera a evolved generally species, 16), Eumeninae although In has indicators forking male, are of It of in for to formula species not shaped are latter vary Symmorphus, part has palpi, producing is a some Bequaert of reduced are independent definition the as the psammophore carina have (as that diverge correlated must also 5 has 3-segmented, separation; tubercles been Reduction in of The waldenii, : 4-segmented few is posterior the within emargination quite author, of 3, occurs reduction Zethus). emargination the it received quite this the many From of as of a have have was labial a states 'zethine' genus different as from may (1918) in generic stem-group. a major head variable. 5:3 deformiceps, is of basket in Montezumia discussed variable genera are Pterocheilus 3 4-segmented it with evolved groups. important modifications, the several ocelli, in The Euodynerus palpi, most one found it segments (e.g. much is represent quite is in present from that the enlarged character has in interpreted and relationship. primitive Ctenochilus similar number presence another is synapomorphic used Leucodynerus), Certain two structure, of (e.g. but (e.g. while in more the distorted attention. genera. in the independently clearly Rhynchalastor by similarly Even the the Dolichodynerus on may in Pseudodynerus pairs m importance female and van and labial (Ischnocoelia, Zethus), palpi, and hairs stem-group), some convergence carrying pronounced in the (figs. of 'zethines', exoglyphus, copulation. states, of state this also as der eumenine produced The Monobia, ofpalpi here Odynerus segments the vertex a in and broadly palpi From fringed Bohart Zethus 37, on entails forked Lepto- unlike Vecht shape occur other most of such they as and 38). soil 885 the the for on in of of 6- is is it a a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 it angle component. character, convergent carina punctation, and this in Mesosoma morphy where in Antepipona recognized. (some Hypancistrocerus). Stenodynerus significance 886 preceding less problematic. foveae. e.g. some lateral Stenodynerus chiloides. species Microdynerus, carina Stenodynerus and Microdynerus component. it that 1939 punctation, cerus secondarily this Eustenancistrocerus, lissosomus this occurs the such seems The 14. Paraleptomenes, pattern few some surface Bohart pronounced group, a: (in are outgoings species Pronotal in punctures neotropical (see they fig. of It taxa type for species Microdynerus, Plagiolabra Less in these thus groups species best is has 13). loss loss below), of and to Most For these smooth some of The to Smeringodynerus may (1939a, was additionally + A as and of and component, in dense deep a these Beginning Leptochilus punctation part is either genera, largely interpreted punctation. (such pattern Eustenancistrocerus and of Eumenes, example, punctation interpretation that these accepted. of which considered eumenines genera degree Parancistrocerus vary species Parancistrocerus They species which central the the lateral Pseudonortonia) dissimilar of b). pronotal which stem-group. occur may as mesal smooth a first latter there J.M. have intraspecifically. These Leptochilus fork Microdynerus of vary transformation and with in supported of also where may the be Odynerus, foveae It punctation and appear in show group The uniformly as have punctation the is is of the Hypalastoroides, of Carpenter a some genera a faces. in is the and impressions and probably types pronotal establishes a is be primitive Dolichodynerus number the synapomorphy vertical, punctation great by the size, narrow, subgenus The punctation a clear the or latter of readily rather to-have species (fig. Gastrodynerus the The etc.). synapomorphy pronotal are genera pits smooth that are by are carina do Symmorphus, development approximation occurs taxonomic dense at lissosomus) extralimital 43), state, face). the a anterior of and another generally (figs. or mesal show not series, the feature It smooth labile least convergent distinguished of the extralimital of Onchopterocheilus. do less rounded, foveae. and one is structure mesal Ancistrocerus J. and in punctation There presence differ impressions 42, face more not Stenodynerus-Microdynerus both for Maricopodynerus difficult foveae M. states. dense. (D. Leucodynerus from Cephalodynerus of face fovea), if 43) matter; mesal linear have linear. the importance show space are Cumming it is of Leucodynerus turgiceps are Genera species appreciably dense distantly the and of is of but character and also a various pattern The of The of (fig. then thus groups the a the smooth by space, median to the but reversal. is Stenodynerus-Microdynerus of the various Stenodynerus ground-plan occurs they punctation of placement pattern and very two of density pronotum punctation faint (and attach 40) 'such transitions. quite interpreted the propodeal Eumenes-Microdynerus and (fig. types related significance, in which If Euodynerus. are (e.g. deep by in narrow or impressions face impressions genera from the species in it russipes. homology and Pterocheilus, It as 41), vandykei). and diverse is Blfithgen from any a absence is of Subancistrocerus, is many also central Dolichodynerus- Antepipona to also to strong and is genera some a sculpturing established female (Microdynerus + species character Pseudepipona, when quite Nevertheless, phylogenetic dense ground-plan of some have the submarginal Parancistro- equivocal as form occurs the In this species but However, group. species foveae of scattered In showing is thus synapo- (Bohart (1938b) general, present, smooth such a median degree Lepto- lateral where of sharp dense easily clade, occur more some and to and the are for on In as in of of of a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 pronotal useful most may Gayellini derived groups 'intermediate' majority where features posteNad of 'raphiglossines' posterior have level. has dynerus in exaggerated character Although ponent, dynerus, 'omaulus' case nearctic genera. species and encompassing annulatus-group cluniculus. some However, as shape quite Absence and and Eumeninae. the groups well 20. 15. 19. 17. 16. 18. happened some characterizes loss be for a Ancistrocerus variable species carina its varies indicator Parategula. Notaulices Pretegular Epicnemial Pronotal small, such notched) of Humeral These vandykei). Odynerus at of some genera as Cephalodynerus of characterizes presence face of and lobe social on of Zethus, secondary an that Coupled on Pachodynerus. this eumenines scattered the sculpturing as None Bohart is greatly, the incomplete (fig. flattened, of many a in in seem the of of the some common, do have blunt is carina wasps level: of regard is Euodynerus regard humeri carina. the and the + Eumenes, canna. carina. generally 40, Leptochiloides, defines and present This is cladogram not This generic carina. pronotal and with Pterocheilus to times, and a Allorhynchium genera present and absence 43) carina tegula Pachodynerus. other in more have (Polistinae). have parapsidal Eumenes, primitively is to abrupt to character scattered is can other A towards + Stange is and canna the A the the apomorphic, An A an usually the in this The Smeringodynerus, relationships. part canna Dolichodynerus raised of been elongate one, lead may split extends carina species. the related in lobe autapomorphy typical primitive complete outgroups helps pronotal extension characters, limited character, of hook carina the Nearctic (not of and great to 1965, including Paranortonia, independent be from species the Maricopodynerus, as fig. furrows. should area with (figs. It the have the of outgroups to and present the subgenus parategula to caudad some quite is which mesepisternum These majority phylogenetic this of 1 Bohart define ground-plan appearance a Eumenes loss but on state face. annuIatus, a punctation Chlorodynerus 3, it (or ground-plan a in of the and Eumeninae therefore this translucent the 5,40), the reaching sort species (e.g. Loss dorsally and has other has projects in the The are is of any mesepisternum the past several it indicates posterolateral events, (Soika 1982; stem Polistinae, (some difficult of the also Ancistrocerus, occurred Hypasastoroides seems Gastrodynerus. occurred which of by pronotal Dolichodynerus clearly dense is groups. the other it, of of and these Eumeninae be becomes significance. group. Soika of a fig. the Hypalastoroides incomplete occurs little and (fig. groups, Euodynerus a social 1982), stem-group. and trait, to lamella ground-plan treated the is carina, is close punctation to a 5) mesepisternum) for straight, grooves be independent best in 5) many complete It but occasionally from (1978 so determine. is Eumeninae. canna Reduction present in and Odynerus, useful wasps is thus found denser. relationship these corner ('prepectal' absence it a viewed apparently one with some along Leucodynerus, (Carpenter It the strong is some b). times This carina mexicanus, does Microdynerus on and has also a Mesal running only of have of in characters). mesoscutum. in caution. one very of character Many Hypalastoroides, the the the and a as the species may of Most Pseudodynerus. Pseudodynerus, the is occurs not Leptochiloides, independently. lamella the developments. covers partly It at is convergence. one), apomorphic, the is mesoscutum interruption is of outstanding inconsistent stem-group, between crest is secondarily Eumeninae some the seem pronotum, present genera not 1981). present present Bequaert; obliquely probably 'zethines' carina Dolicho- in in vertical but Gastro- specific it. (which of be of other other to some Also, com- The The The 887 the are the the the the be in in in in Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 cerus expanded narrow. externally This 'generalized" completely extreme developments mostly of groups. has the parategula posterolaterally related the some 'zethines' that Leptochilus. eoe small becomes parategula primitive this similar. However, has interpretation towards usually as transitional annulatus-group. (cluniculus specific Kurzenko Ancistrocerus. the two with parategula quite have 888 Masarini, only surface noted these With 21. greatest midpoint character, an been has other interpretation a a other different have Tegula. evenly short, at to weak lobe weak For and genera. A genera. In importance. have The the variation lost this genera have above. the in state and many Stenogastrinae Leptochilus considered 1980), characters, rounded some and and occurs oval varies, extralimital n evenly an example, states width scutum, exceeds or The convergent the partly eie assessment, revised level a from of perhaps convex In The externally an (cf. of overall plesiomorph, overall the the but pronounced genera evidently (fig. 31). (fig. palearctic and the Pachodynerus, The the emargination, and flattened Aside elongate secondary in it in shifted in lateral eumenines. figs. projects. form emargination of the cannot annulatus-group), retains tegula vertical small, O. some the with has length. the shape in shape smaller groups. This groups. smaller tegula crey reaches scarcely this tegula, similar tegula, primitive. with the because in plesiomorphic convex cinnabarinus, 27 J.M. genera from of posterior tegula shapes never rounded expansion and The a posteriorly. may species the posteriorly relatives. parategula posterior 33). a Leptochilus, e upheld. be is posterior this This parategula slight is However, aue of nature posterior Carpenter this, Polistinae not tegula condition other Although indicate These emarginate tegula, ofStenodynerus + usually the state. been Leptochiloides are This many is the is of more lobe the tegula the interior parategula and almost is Stenodynerus also is erythrocephalus lobe It genera relatively angle to condition. oval And There clear. more oval of states are states again same; shape The tegula The is of although considered cases relationship of there closely lobe, for is but the that perhaps + Vespinae, + h parategula, the different and great found approached partly the are which the tegula usually tegula angle. some where is example, adjoining even, universal. the rounding are When is indicating of states the of not . M. J. because length as are tegula is importance also (fig. few related Stenodynerus size n some in the also extends Insufficient in almost due difficult nearctic do This so tegula form of Parancistrocerus very generally of most it and species is part primitive in strongly in Cumming 29). adjoins the n attempt an parategula phylogenetically that Odynerus between and but many ml in small and the of caudally. other to different there Parancistrocerus in is the parategula, the number. It similar. in also occurs in to by fused the the also the in is considering caudally details tegula species is some margaretellus, to Stenodynerus + LeptochiIus with as Odynerus Odynerus, Pterocheilus in analogy each extralimital shorter Masarini. retained transitional have area attention longer tegula assess. A assess. the n abruptly and eumenine various is a is with in in and the groups Most For species similar seems scutum. is Most of group case length has of of 'raphiglossines' synapomorphy. been past Euparagiinae, Pterocheilus lengthened the the is a greatest with in in example, than Leptochilus is species when several species several in other been useful, these has are mesoscutum, genera have to flattened, In shape synapomorphy relation relation the and it many taxonomy, shown of states (fig. genera another but where The other nature of nature is be Parancistro- superficially been Leptochilus. Eumeninae, Euodynerus parategula, rounded made eumenines the this width the 31). genera obviously expanded mainly groups Gayellini although are Omicron different with and in vespids. it (fig. paid closely by to to tegula tegula Along genus is is of blunt some often both is The (e.g. and and has not but the the the the the 27) an an to so as at in of of of Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 eumenines Montezumia carina runs nearly slitlike (fig. become species acarinarium stenogastrines concave. with bordering (Soika 44, 45) the strongly orientation Masarinae, of subfamily. lobe have most with from within (fig. parategula morphies broadly allies. A in and eea are genera posterior one weakened, have ecdnrs Stenodynerus Leucodynerus characterizes genera. and posteriorly the Pseudepipona, convergent. the detail. the the 22. In The other description fossa 30). of some in the a the a Hypalastoroides sharp inner addition, has" 1978 narrow Vespidae, the these, general It Axillary concave as Leptochiloides narrow rounded Species shorter broader, vertically shape maximum A hand, lobe the s less is neotropical with the may recently 45). From this well). groups. (fig. 29), are displaced, fairly the transcutellar so a). of rbby independent probably The component (mites the tegula Pachodynerus. scutellar (fig. 44). This the transcutellar the This but tegula and that Eumenes shares some not less tegula with posterior be in discussed of state fossa. scutellum The than annulatus-group, deep the laterally and posterior longitudinally sloping of Euodynerus is been this character is a due as width social are very resembling homoplasy the apparently other the in such it scutellar details the in in groups. deeper Monobia (cf fossa has synapomorphy The crest is genus Pseudepipona occasionally used s. to Odynerus + Odynerus + general is fossa more lobe different. state more posteriorly Soika wasps I. as above expanded, lost states (fig. 44). an highly angle. concavity size carina have and is component, as has occurs in flat). are in might (Bohart elongate, Since present fall characterizing has evenly crest the some and a carina, Nearctic can delimited and not 1982). of some otherwise appear character shape usually a difficult This into can The derived become is the emargination, distinctive However, developments, the a few a Pterocheilus. Pterocheilus. be tegula This often be closely almost generally present species is and and for of in Gastrodynerus. extralimital broader tegula outgroups several 1982) primitive Those thus and also inwardly fossae identified is in but the Eumeninae Montezumia have these the gradually is to anteriorly flat the within been shallow located, that although shape to separate the similar adjoins related probably of be stem-group it in Stenodynerus convergence interpret, is species axilla the nearctic groups tegula, closer and taxa ('scutellar Dolichodynerus, the tegula was shape shortened overlooked. the of in treated directed A However, varies are and the in condition the as eumenines axilla trace in to so and a the swollen (fig. swollen to the compared smaller and to by tegula the with without variable. on is However, Stenodynerus, some have quadrilateral. Eumeninae. that they of Pseudepipona fauna, genus, the the these groups. these the scutellum, scutellum other n most and similar in the of the the and as posterior on of point pockets' and scutellum, s present. is and tegula and a Hypalastoroides, the ground-plan a the wing L. arizonae L. groups are transcutellar Variation shape. fossae short, basis a very scutellar but indicating in extralimital most the scutellum tegula where emargination its In with has some to synapomorphy 33). the dissimilar the base do does wide relatives Euparagia Leptochilus elongate so of es concave. less states of The lobe inwardly greatest the other of as Leucodynerus, Eumenes, states but not Euodynerus and Pseudodynerus- tegula This Cephalodynerus that placement is it than Other Duncan crest the in some posterior not emargination and slightly development also is transcutellar clearly relate serves state Allodynerus the carina moderately eumenines, n nearctic in genera the the in carina (fig. in state and reach posterior shape. utahensis (whether the differs width shape all nearctic directed synapo- remains and the fossa state Minixi and of 28). as 1939) more these state (figs. of vary lobe (e.g. The also and rest and has 889 the the the an on its In In of in in is is a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 Pachodynerus posteroventrally ment (metanotal (transmetanotal Leptochilus, present happened each bisecting (as Stenodynerus, independently of specific Symmorphus, XII) and is Euodynerus and vespids Anterhynchium, Synagris). Zethus This Euodynerus The times. slit enclosing is character similar Labus. as Other with most margined but Saussure 890 sloping have similar quite variable quite evidently the the in 23. The 24. Such 25. condition has euparagiines a strong shape is of the the of of some dorsum set metanotum, Propodeal seen Frequently in Here and Metanotum. aspects these metanotum Propodeal propodeum, occurred apomorphic characters transverse in have the margins crest of comments used some the the and many it Pseudodynerus, of in terms rugae 'raphiglossines' neotropical depressions is not (Syneuodynerus, however, the genera groups. metanotum. cannot the many propodeal Smeringodynerus where is of a erynnis, derived. not of these bisected Euodynerus of carinae the found the is one metanotum. rounded carina within times, many have the for metanotum shelf. thus in probably a carinae the but in concavity. the neotropical is metanotum characters also has ground-plan the that condition natural A the depressions the the declivity Hypalastoroides be other a found Euodynerus is uneven. The An the time rounded in metanotal and J.M. by is pointed Pseudodynerus). some of Duncan). propodeum dorsal various been concavity of a apply to apply possess considered located Euodynerus Presence extensive on The carinae, posterior concavity posterior + Dolichodynerus even synapomorphy shelflike varies independently, genera. Duncan the genus Pachodynerus, in Knemodynerus), independently Carpenter the genera and almost The of surface Pseudodynerus quadrisectus extensively, fossa and groups, plesiomorphic posterior several carinae. (figs. 4, have metanotum a dorsum of of them. anterior the within cryptieus) carina the strongly (see when and posterior These are dorsal of This state either Pachodynerus may (e.g. differ, is crenate the 1939), very has the has nearly bidens of presence a 6) discussion including eae groups, related Laterally very the The Development In and groove the angle dorsal the applies is is Parancistroeerus genera. be acts surface to and Euodynerus for and linking Pseudepipona angled important. a h nearctic the lateral angles lateral approached acquired the so punctation propodeum usually and J. with as narrow vertical propodeum highly {e.g. and posterior plesiomorphic the face and condition they still (the as M. developed Hypalastoroides. metanotum well or surface these may Pachodynerus. related to declivity, Euodynerus, there of other the a section). metanotum, Cumming constitutes absence borders of Gayellini it (all vary derived the the (variable continuation slope odrd by bordered by in be to of propodeum the dorsal margins or due The of bordered propodeum are is some many groups in lateral in in the fauna, primitive, genera, such is and has may ('l~tudes' Ancistrocerus. coarser + different both propodeum Eumeninae. some has raised of which the state. has of carinae to (found Stenodynerus). deeply approach cristate some the a in xrlmtl genera extralimital other Ancistrocerus). shelves groups. a further be the variability the the and metanotum teeth they also nearly more Various form previously longitudinal but the Cephalodynerus The laterally In almost propodeum is toothed in of Ancistrocerus, margins sloping but in a Vol. upright is in the same the Nearctic, form). excavated occurred fairly species outgroups). not are Dolichodynerus, short, and as the outgroups), metanotum clearly or aea carinae lateral form This vertical this) evidence are The loss is other well, less 3: to slitlike. constant Although presence by gradual. frequently or 184, gradually They character placed Develop- vertically but the medially. evidently Masarini differs rounded. then lamellae (such carinate derived, carinae, Certain groove spined. several as genera is of slope. plane plane other areas Plate very that The was and this it has but are as as of in in in in is a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 ('apical but project Montezumia in iated partly appears in membranous reduced. and (arizonensis). state. are realized the state that neotropical so Leptochilus, nerus some this nerus, does propodeum Leptochilus and Leptochiloides pronotal clade Stenodynerus-Leptochilus Zethus), It a euminine taxonomists, where separated carina separated Pseudepipona valvula but group). nerus. chodynerus, convergent is several Microdynerus that 26. Despite A outgroups, does is not not development slightly this is not into it more An enlarged, seems other fg. 3, (figs. It free where fusion fused it much is Propodeal is the scales" in A generally by also well but Both is not seem state fused species a a face in a differentiated raised Parker important lametla. 'zethines'; homoplasy, low component, evident: groups. only the few (fig. the and is occurs to (Willink angular, the correlate and projection developed 8). from development genera significant in is of are present but to and the states nearctic valvula ridge 3). is Although other derived convergently. derived of this and an and above Saussure) Hypalastoroides, loss of an informative differentiated it, (1966). problematic, but rather the is valvula. The upper other in Leucodynerus and The many aiu characters various character, variable have in cf 1982), character surrounding Pachodynerus therefore of rounded, well submarginal genera. in but elongate, the valvula is the these above whereas and they the Stange in parsimonious lamella extremely nearctic variable; This Stenodynerus + in fused component, lamella. Euodynerus the it the with valvuta, in are complex. are all parsimonious interpretation parsimonious itself, but The upper many has form genera in valvula Euodynerus + is A the valvula character shape Monobia, in 'zethines' where the to is fuses as contrary it valvula 1979), is Nearctic one other not the quadrate, differentiated the characters occurs the but has genera. it is variable a they most palearctic lamella has the but is small carina continuous and both and stem-group been of latter are not with type apomorphic. may the the apically characters Leptochilus with lack the propodeal interpretation do occurs in it do rather species in to Parancistrocerus, really the submarginal linking In A carina interpreted and Eumeninae valvula, (fig. of the main is clade not (Carpenter almost vary in not a further Bohart quadrate Smeringodynerus+ Pachodynerus. the submarginal associated species. developed fused are both group upper 9). somewhat are have occurred. similar. lamella is show projection. plesiomorphic of features somewhat, primitive may is and The somewhat Gastrodynerus, among its orifice, Euodynerus rectangular and so also approached modifications It genera. 1982). to the lamella In other a is projection shares as valvula it be subgroups. valvula considering 1981; carina This separated upper separated Smeringodynerus the not based the may evidence carina submarginal distinguishing with produced is Thus those truncate, present carina. Leptochiloides with state, Leptochilus Development The raised eumenines. as other Convergence otro margins posterior also melanized. particularly figs. 2, figs. in with be is raised on a is the (notably lamella loss above Dolichodynerus a (and genera Cephalodynerus, present pointed in synapomorphy elongate has that occurs in the and In the species of variable also in above propodeal Leucodynerus as in the other into 7). some may close the long above some valvula is characters to it carina the Increase in is possessing This and not Monobia plesiomorphic in Stenodynerus- is the The a lamella, Microdynerus a be lateral the + the valvula Marieopody- has apomorphic, characters of been in relationship. (it Montezumia valvula, more lesser in a Zethus simply genera, length. Cephalody- the annulatus- inferred number the iia to similar valvula Gastrody- palearctic is condition character the different- does a in used vatvula valvula weakly in (fig. carina, angles or of extent of of but latter some Doli- from (and very size, The and and and this less 891 not the the the by 4), in in of is is it Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 most than (Eumenes) various by the 892 h two the of the Pseudodynerus, present Metasoma (Carpenter eumenines propodeal miscooaster, characterize Secondary Eumeninae; dnrsPeohiu, yaatrie, mrnoyeu, Maricopodynerus Smerinoodynerus, Hypalastoroides, Odynerus+Pteroeheilus, including reduction none. This none. derived, This well important therefore states lamella aurata, tent state incorrect. used This questionably algeriensis transformation an and 1981). Leptochi'loides. the autapomorphy orifice 30. A 29. 31. 27. 28. Montezumia-Microdynerus as a is Odynerus is as Cephalodynerus, twice different found in character found Sexual This Hind-coxal a First generally Mid-tibial valvulae. Propodeal WiUink medially projections as a for few genera, Australozethus, Discoelius, Symmorphus Discoelius, Australozethus, of feature (where the Reduction to teeth has at the a and orifice. loss character 1981; fig. as might the segment dubious it other is montezuma), development in'raphiglossines' all in dimorphism stem-group, Raphiglossoides long is occurred of found form zeppelini. apomorphy must Leptochiloides series 1982). which otzma (arizonensis Montezumia to a nearly related A or stem-group it Odynerus separating eumenines spurs. This and lower be carina. Presence carina. The is orifice. of more be have as near 7). of primitively shape. then worth. to has Leucodynerus. as with in said spines, wide of the In is is J.M. universal. state one several well level The the Thus, evolved most groups much the created or been more the be in occurring projection The but the to are is An and is base less the genus the the as level (e.g. inferred in inferred primitive nearctic Carpenter apomorphic most character. of etc.). the have the general the and tooth 'zethines' Montezumia one (it the particularly some orifice elongate importance. times. parsimonious flat) component, legs. continuous a half in Alastor). by of such In of development is diagnosis most often of slight However, of the the it and a the many In under the species is actually as a eumenines the projection Aside is narrowly from quite number male. dorsal a as carina Flattening the of and Eumeninae. enlargement petiole wide affecting apomorphic and rounded variety projection major condition Thus eumenine does those times. Parazumia. this notable has Pseudodynerus-Montezumia of Monobia. variable, for the J. from border s it as Modifications it Raphiglossoides, as carina than acute) (fig. Euodynerus, Dolichodynerus M. could of it interpretation. formed of not a 'zethines' of ones rounded segment have from the may high mid-tibial three is (some above Cumming genera, a 3). and treating for Raphiglossinae it is differ condition, Kurzenko continuous is present femur. of must as state, be This to in is exhibit This one the present part hollowing lamella the the male the of Unfortunately, is narrowly species treated eumenine be in species with its II; segment differentiation submarginal above. from state lateral spur. be a is of The range production the presence spurs transitional, in fig. dentiform some an the an considered It as in which (1980) and of is as most subgenera plesiomorphic Monobia; mid-leg lamella exaggeration 2) seems elongate have projections acute various ground-plan of well The from probably Zethus I is synapomorphy classification, homoplasy, in has Elisella linae (segment the two in considered Willink Soika of as it distinction above hair carina too projection it mid-femur always some of component, in projections. around the acute) (Carpenter eumenines valvula, Psiliolossa but (guerreroi, and part Spinicoxa does an nor form inconsis- multiply patterns (1974) formed genera. I genera (fig. above (1982) upper of males of many but more been state does of and and and not but has the the the the 8). as of is is is is is a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 distinctions found it importance, most weighted sternum characterizes nineteenth Leptochilus-Microdynerus work. dynerus. treated segment in a segment ground-plan subfamily. anterior Labus greatest recognized gayellines, Symmorphus subdivision transverse this a Montezumia, odyneroides) with and that among Macrocalymma, probably more Euparagiinae occasions more Saussure similar segment with swells resemble less group does size, The 32. None than a a abruptly are sharp gradually more gradually more and in petiole, in The but very not Carina as these states shape width with part 'Zethinae' the I at least apically. Saussure the petiolate it probably of (l~tudes' are as within having rmtv for primitive if its carina Perhaps even stenogastrines seen as century small, this in particular elongate anterior so but wide of of only of Euparagiinae, a Paranot'tonia groups. from Pachodynerus allies, other in and well of near a his sharply it tergum of Odynerus. this indicates whereas somewhat narrower more towards to is unrelated derived Pachycoelius, that the the as A 'sessile' at studies differ 'zethines'. before Masarini, tergum little Vol. considered cup-shaped (inclusive other some be the Raphiglossa eumenines conical II the clade. (Wesmael the declivity. Eumeninae. first petiole. The it throughout heavily, (a form very sloping I. middle modified is 1: majority from state declivity the extreme Something feature the metasomas. departure what the Odynerus segment somewhat of state xxv). It component and Odynerus I. and groups of first useful but and eumenines, Usually apex, the Eumeninae. formed of In termed apex. the Pachycoelius anterior than lumping the It the primitive state that cf several first found polistines, 1836, The Ctenochilus). Nearctic and of segment, Euodynerus, state from the is in developments Transitions periole of of its at and figs. (cf has shape course also the from the However, have s. like Elisella, segment the nodular. and eumenine petiole than the the history length 'sessile') the Thus the of in Omicron may Saussure it declivity. l., also other remaining all in variable Eumeninae. narrows towards this and first a petiole could generic Macrocalymma some Eumeninae of the fully presumed and basis a This the the the narrower Soika and be Saussure en used been 'subsessile' even and the is of and genera. these tergum but of may presumed However, between It characteristic second, carinate apomorphic, sides Nortonia probably there as therefore represent and species of Zethus may for characterizes is of This it eumenine first in Macrocalymma 1852-1858). level. in Pseumenes broad might also petiolate 1969), usually its possible the have has the comparison pleisiomorphic separation of Labus). unite segment The are A (1852-1858, superficially fg 39) (fig. relatives segment but nearctic Eumenes different within the Odynerus group, metasoma. and can long evolved Psiliglossa be. primitive as actually influenced from Psiliglossa, many is details the be the various tergum the genera, that taxonomy, Although similar associated be In its have treated of petiole apex. has I form the Bequaert various have so second Eumenes, to Monobia Pseumenes (termed said allies shapes Eumenes group of this many to in on s. plesiomorphic in this a resembles 1875) group been condition. a be groups but Ancistrocerus are I. those Most form Some is to those to is state petiole much synapomorphy is having a as to into several is Raphiglossoides, more the of so usually may very a begins at with again primitive, fused species were 'zethines' presence also critical have made 'subsessile') derived variable other differ considerable (1918, traditionally Eumenes and Eumenes the n h mid- the in condition a species species subsequent of the that not composite short similar reduction the a 'zethines' noted found low there declivity with different the swelling Pseudo- a the has various shorter than greatly petiole in be to swells shape in state, 1925) level as of that that first first (e.g. (e.g. and s. 893 the the the the the s. are by to in in in of of l., is a I. Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 discussed lamella complete The generalizations margin various polarity, however, and If that in and (more variable ly anteriorly, development carina). sternum. Eoyeu, Ancistroeerus). (Euodynerus, a thickened, transverse usually Another tergum component, energetically certainly is that anterior is Masarini ground-plan below) been (1938a, them or Euparagiinae state, Ancistrocerus. group, 894 they angular plain rsn apically present broad other poorly 35. 34. In 33. Eumenes its give found details treated to one to and the are Apical Apical Eumenes First relatives), (fig. may of suoyeu quadrisectus Pseudodynerus other b) present, projecting that edge, but genera. appeared the An type are in the in nearctic loss developments in longitudinal may declivity developed, with different with developed, the indentation in but also so of 25). another. the hr there where appearance apically unfused apically its as eumenine found have of set occurs characteristic, the eumenines, furrow sternum. margin as of first and the occurs in have that He an a on this Vespidae, lamella role This but do other s. about considered A the primitive the Montezumia-Paranortonia Aneistrocerus). character but species, fusion elongate many tergum I. on caudally form a on Masarini. somewhat (1925) in in the in seems a in only of of and The also transverse of in many tergum it carina. may tergum tergum splitting the splitting generic the is The J.M. taxonomy furrow tergum from is do not do times, of Euodynerus, with is differ the carina members probably Ancistrocerus occurs where a genera related genera is less eonzd hs as this recognized a be petiole, n independent an of same no Those for other below partial character small a thickened into thickened Montezumia one tergum I, It Carpenter All present a use different I border I. even very similar thickened obviously diagnosis in and I. transverse carina (fig. in a seem may Some has Some type depression the in of thickness such as In in the groove groups. genera is a and plesiomorphic. of it. has some it apical these carina earinate incisions many delimiting sides been great be 39), and the may this (Eumenes). warranted. other where has This tndnrs + Parancistrocerus Stenodynerus a is to of one and variable and is strongly to is multiply Pachodynerus. degree are border genus. sternum cordlike. with of is an have also A lost depression 'zethines" Paranortonia many present (some the sclerite have of and is still Leptochilus Symmorphus group, subfamilies. occurs in occurs a modification, instead, the partial also species a J. phylogenetically mesal or so cordlike autapomorphy apical several occurred genera, first transparent an plications M. ones. in weighted tergum Some In within angles of aibe it variable and eumenines. neotropical been angular present A derived in is the may Cumming artificial Symmorphus Apical remains, longitudinal development and fusion thickened metasomal into found various modifications anterior While anterior the lamella the the Hypalastoroides same times but not but formation, groups, independently and indicate medially. instead Thus some component. Obliteration frequently. have genera. formed declivity too (Carpenter apically Montezumia-Paranortonia lamellae as between only a to scattered is Zethus plane division, This species is but carina a both heavily. this to Eumenes apex unrelated to of difficult translucent necessarily either carina overlap this correlated in furrow generic sternum this by Ever similar traces is where Thus the includes with state, and Euparagiinae on as of may a of and strong is is slightly the have (e.g. It clear 1981). representatives of the developed median the apomorphic, of the and various have since, genus. slightly raised have evolved, s. or is a to carinae the Eumenes be relationship. and may occurrences. the tergum similar to the I. Leptochilus translucent tergum probably a disc depression previously in determine tergum and punctures present the apical border the 'zethines' complete This Blfithgen with a sternum. a reflexed. sternum perhaps this relating apex strong- remain similar and Other of types, sharp stem- form have also into and and and (see s. has the the in of l., is is it a a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 just not concavity (Krombein association deutonymph chamber. adult times apex short dynerus contrary apex similar to tergum. reflexed blunt, extent Dolichodynerus-Microdynerus cerus occurs (unique characters, Gastrodynerus, length extreme similar. emarginate. rugae a from others used. other autapomorphy. length 10), occurs some determining Eumenes, 'zethines' + Monobia, strong an Another The The (Ancistrocerus 36. posterior that of Eumenes is within various In longitudinal component wasp characters lateral taxa apically in but (cf is the to to of eod tergum. Second not apex in type sculpture (fig. and A the in some transverse In variable where to fig. the one is may Soika a sternum slightly first in (e.g. Maricopodynerus, lengthening Paramischocyttarus, 1967). are between raised. Montezumia-Microdynerus major the A degree 3) similar to often Ancistroeerus. generic of seen), Bohart 2). to of some median the with tergum. also Ancistrocerus. this, the raised highly Eumenes). primitive the tndnrs aacsrcrs Dlcoyeu, etciu, Euo- Leptochilus, Dolichodynerus, Parancistrocerus, Stenodynerus+ the The This 1978 b). indicate Acaroidea. on (cf. of antilope). The sternum character within reflexed second but ridges packed A first a in anterior in the II), be groups genera, ridge to the relationships, mites the length strong and van strong condition longitudinal some area. the modified axillary an In h one the is carapacelike segment first Many first This relationship state, at highly derived groups evidently Parancistrocerus der tergum acarinarium, Stange occurs tergum and runs lamella is The It with including it depression margin the of of Leptochilus (cf. Leptochilus This sternum lamella tergum Other, may These is Vecht is an component, t the characters fossae in eumenines he genital base the of of no in is where Nearctic raised is winterschmidtiid genera (e.g. is in is again autapomorphy second 16) A (1965). elongate, be tergum but the various ridge approached also length multiply of not seem in Stenodynerus +Parancistrocerus, more 1981). running of present of tergum among presents were of the Zethus), more it Zethus first just chamber it either into the component, use as of state this without may may may tergum sternum. is Eumeninae to are interest. itself of long is there terminal previously anterior. Symmorphus groups tergum within tergum. the longer Parker than a genus be derived ata lamella partial others the unknown. (e.g. in laterally (fig. also be specially be associated and other sharp and second in autapomorphies, is Paranortonia, is of n the in petiole a noted that noted is most plesiomorphic specific relation a a be petiolate associated Eumenes + the 11) one Typically, this There to presence and than (figs. 9, 39) (figs. deutonymphs. Smeringodynerus, smooth is character; 1966). there characters This Finally, terga (a present, carina, the group, mentioned, not along of or may Symmorphus). short modified oriental accommodate Several in in and may both Stenodynerus + character with Maricopodynerus is value. to some, well It and have Stenogastrinae. second area, usually of an in and the the petiole h etr apex entire the is may however and Minixi be Anicistrocerus, with and the sexes under the the which correlated not sterna different a a second genus while (e.g. edges o the for at ridge acarin~rium as medially strong translucent and long elongate group of group second be some The segment, least has a of they the may is The (fig. the Cephalodynerus, transverse Acarozumia) and have present it is it much also the the petiole. of evolved may Mitrodynerus sternum, basis medially varies very carapacelike is also Montezumia Parancistro- longitudinal are overlapping 2), at of parts condition Eumeninae, the with tergum. propodeal to reduced dispersing it present been (twice Since taxa posterior being there not be value low serve as may may but a lamella tergum or of in This within many lesser other ridge used mite little very split and and (fig. the the are 895 the the the the or an be be In in in in of is Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 posteriorly genera the other strong In Stenodynerus. of Pseudonortonia. in evolved the one. Hypalastoroides-Microdynerus (traces This those Microdlvnerus dimorphism species), duran~joensis and 31), longitudinal not traces them Eumeninae, Effacement 896 Pterocheilus, metasomal and treated (VII condition. the ('nigricornis-group' Eumeninae. the it Pseudodynerus-Montezumia there the except ary Hypalastoroides-Microdynerus taxa several this is the 39. acarinarium of 38. 37. homoplasy, Pseudodynerus-Montezumia acarinarium, occur outgroups. loss which occurs thickness Other and cus in occurs is a cases supported with apomorphic el developed. well of context Transverse in Basomedian could depression o the for as Sexual acarinarium independently, in ridges transverse a few a species), l respectively). Vl atoyeu, Smeringodynerus Dolichodynerus, Gastrodynerus, , neotropical in synapomorphy Occasionally leading parts several sternum where the as of those In ridges externally Maricopodynerus be As component, presence of a the The tuberculocephalus~roup, the this Pachodynerus, dimorphism are The furrow sulcus unnatural. may such it the by in the groups Iutonidus), furrow characters, for characters, the to and groups. These groups. cladogram, may similar ridges of length present furrow II character longitudinal some form ridges the the base is groups be it authors) (fig. second may as or J.M. hollowing Symmorphus, Thus, on immediately (e.g. the not acarinarium the fortuitous with of of hrces linking characters species varies, absence the 24) is of to In varies, some at the terminal in can be second the even clade shares clade then the some placed only those Anterhynchium), Carpenter of Minixi metasomal is the is structural a various present and loss the Smeringodynerus, and metasoma component, component apomorphic of petiolate Montezumia-Microdynerus be component. Leptochilus second the from ucs A sulcus. base of include be example and of great further of to Stenodynerus + at of character attachment of ridges metasomal Smeringodynerus. both in visible raises homologous the mexicanum ('median some metasoma. a lesser the the specific the it of little in species Ancistrocerus importance sternum may details acarinarium and the the tergum. second genera. but base most sulcus tergum, the apomorphic in the a are longitudinal Pterocheilus sterna modification is albeit outgroups within second (present blunt, The interpretation J. extent be Eumeninae traces importance them slitlike distinguishing entrance' closely of question of as sternum. points. probably Ancistrocerus are M. very segment is the is Considering the the Gastrodynerus, Besides In in are a well sometimes with sometimes Parancistrocerus treated Cumming strongly the strongly quite in sternum. in n several in weak to all second female in stem-group, conflicts faint. in general, axillary In different related Microdynerus. These the Eumeninae, (Euparagiinae), the of paraphyly (present of in Parancistrocerus. median many as genera In addition, the beyond diverse. ('zethines', Parancistrocerus, beyond an Bohart (e.g. one. phylogenetic as scattered the It old Euodynerus component, truncate flattened tergum. has Typically, associations a is (lost to Parancistrocerus autapomorphy the with fossa species), in the the majority Ancistrocerus). there sense, other synapomorphy most of sulcus in with an Leptochilus. many a This also This diversity and 1952). the in smooth eesl is reversal many the being Hypalastoroides emarginate a with Eumenes (present The male in in In groups and nearctic parsimoniously is pce such species parategula central PsiligIossa sulcus. a usual relative distribution but at Paehodynerus Stenodynerus. profile (fig. profile Aneistrocerus this other The system of with little bidens. furrow species) absent extralimital the Thus, the apparently and eumenines not in it space applies has presence aculeate is in such s. Despite second- form African clear groups, base groove species ridges. of female length sexual found l), of of short apex, from most from both does to The The (fig. and has but the the the 3). to of of as as in of a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 in dissected relationships Leptochilus synapomorphy the should for the for figs. (dilectus base component. they appear examined associated membrane. association well received each characters. Stange primitive type in Vespidae the wall apodeme Eumeninae, and states widely volsella attention studies probably while common as male. of be switched). be This pce level species unfortunately +Polistinae+Vespinae part Parker terga well There cuspis + cuspis More 40. Masarinae, 34, supported, but supported, of of other This occurs the (Carpenter rounded be 'Eumeninae' 44, of the to Male the in 1965, as and inadequate are have male is (Carpenter of until nearctic important the (Snodgrass further is the species 1966) with 56). paramere, the in Bohart scattered lamina The dissected lamina completely little of If also te information other mar in this genus (Soika genus this in of sclerotization genitalia. labelled species papers Soika the this of has In development Bradley for these these the ventral but apically the qaretellus these and are importance specific Eumeninae, in taxonomy investigated error ventral a proves pce ic 1939. since species + cuspis. + and by 1981). lamina longitudinal Pterocheilus study. it Leptochilus-Microdynerus becomes of apparently is 1969; this 1969; 1981), species and structures of may of and the the is structures 1941) authors. was fused. Stange processes, (1922). the two The R. Zethus truncate possible have characters. projection to digitus ramus. Maricopodynerus Most These dissected) the be (cf. M. repeated by of the structure The of genera. became be present. structure in at the More more the lamina useful regarding dense Snodgrass Use Bohart, who Bohart, (1965) the cuspis is a is cuspis the the (using in authors and h New the Nearctic occurs-in carina, s a is generally anterior beyond and This stem-group that cuspis both are and 1978 the case, of generic level. Differences generic data Some rounded from cuspis well Cephalochilus, patches and character cuspis It illustrated synapomorphy of These is the is the extremely a the the form of Parker and b). is of apparently broad, have rather it are the established the observing Pterocheilus the apodeme form 1941: Eumeninae Stenodynerus not reduction World is the relatively specific the term cuspis This 'raphiglossines'). incorrectly the maricoporum relatively has required the naturalness of of cuspis structures of lamina also ignored in possible Euparagiinae male (1966: component. hair narrow, pincerlike of form plates in the basivolsella first consistently these the type and shows aibe in variable (e.g. the further or and that widespread at has and cuspis in by small longer Eumeninae. of genitalia consistent fig. before these, species-group lamina of aedeagus there the of structures 16-18 at otzma azteca), Montezumia the well in sclerotization the some and evidently with a are this differentiation 32). lamina the present their apex have digitus broad, lobe sternum study middle and than even Stenodynerus. is all any condition appears more instead In and sclerotized in In used the other is Eumene~Microdynerus no did those form fig. of the punctation and reduced the in synapomorphy attached to and lamina. interpretation all as in volsellaris. and anterior van blunt of are various The sclerotization to been Leptochilus these of 2 (the labels 2 (the or Old not ground-plan a other digitus is other the generalize the digitus the Parancistrocerus of characters. source this to the throughout important and their der less Stenogastrinae (cf. aedeagus World lamina). ot a lost, be receive is and characters phylogenetic In ground-plan genera. and century, sterna Vecht or the Bohart Also, rare. retained eumenines fused relation united of the Odynerus therefore This volsellar but form (cf. specific further genera, closely should can on apices found at mesal of much those in More at of They 1972: This with with figs. it and due has 897 the the the the the the the by be to in in of is Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 nest primitive cells nearctic spilogaster Ancistrocerus in Nearctic, evidently burrowing psammophore, a there the burrowing pattern'. is (Maindron the building burrowers genera placed in are burrowers are plan apparently and quite crop However, behaviour Nevertheless, present that: atrs of patterns and insufficiently Irregularly or Iwata Behaviour is well is concordance of not or Gayella. 898 + turret generally Parancistrocerus, erroneous builders, Montezumia However, Eumenines Similar 41. construction renter.Ancistrocerus only summaries several (Krombein architecture trait different is before digging (Grissell 1975), as 1975), (Grissell digging before 'it 1976, Nest known. support may knowledge. The may Iwata derived genera, only ground-nesting due apomorphic. in nest Kurzenko may in Euparagia and (probably shaped 1882, and assumed groups build groups problems architecture. evolution Zucchi Vespidae. be Stenogastrinae observations. detailed be phylogenetic from Pseudepipona variation this to tuberculocephalus in loam. As (1976) an for may the building behaviour such rent, renting 1979). (Evans (e.g. some Iwata behavioural is even less clear. less even is of both Evans outstanding this hypothesis masses those ground-plan Euodynerus et a make as be to Iwata (richardsi; believes Leptochiloides (1980) to turret al. Considering The within occur interpretation. years It It Iwata 1976): occurs be J.M. burrowing within classified 1973). as of provide nor The great tuberculocephalus and mud apparently 1976, nests and of complete primitive However, some eumenines. investigation. site derived + (one 1976, or primitive cells, before disputes Polistinae in (1976) Matthews burrowers, Carpenter building the plasticity, in structure nests tl leaves still autapomorphy. and The Gess species in and R. chimney state: species), the is the information Parker group'. into sandy Longair, whether plasticity the in (cf both mode Ancistrocerus. and mud and plasticity in enough plastered does majority s well). as and treatment whibh In this occurs in In renting variation this, to his the limits three + Vespinae all Vespinae + is that the (1974) Kurzenko Leucodynerus but and the soil renters at 1966). cells Gess renting have burrowing It not of Many fig. general group, generalizations us to of pers. comm.) pers. Vespidae, the is treating the occurs Eumeninae, nest of information also moisten J. and broad of the Further, mud, within is on to 45). burrow with difficult of evolved eumenines 1980). remarked: Cooper M. burrow majority very certain. other in of and various construction of many snl species single a use as the cu in occur does Both The the in (1980) Cumming in resin the Euparagia considerable the categories in builders. pre-existing and renting Not the Eumeninae build the genera. which burrowing unstable group and the to the from vdne is evidence other not building (1979) aspects various entrance and reports the or usrts as substrates, of soil Sphecidae'. has disagree uncritically in 'one renting all may expression Pterocheilus habits free nests construct species leaf stem-group the nest The so dnrs Stenodynerus Odynerus, neither the can wasps, with Masarini as Most Burrowing accumulated considers are and finds little be presumably generic of of burrowing material and in types primitive pattern site with cavities be homoplasy. to and made water of correlated with correlated (Cooper nesting far nesting may with most Masarini are of the with burrowing made be and of Pterocheilus This repeat construction a from of the Masarini, are renters. build also mud of literature classification. hypothesis intermediate. are in turret. Evans carried be the for an have or architecture into renting of species of behaviour, exclusively the re nests. free behaviour of type, n both and a all clear, immense: build to occurs 1979); details ability 'atavistic or the doubtful course a ground- the nesting light renters clearly In clarify paper, turret, (1977) In Other occur in close with that and and free soil but the are the the are the (cf A. of of of in of of is Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 cation provisioners (Odynerus, The Ancistrocerus provisioning commences. indifferently used genus chocyttarus, as as used. either and prey Pseumenes different 'intermediate' free shaped synapomorphy evolved oviposition Vespidae. been supersedure and resembled Spurway (1870) the competitive the stated species doubtless Euparagia the Euparagia 1976, 'irregular' a Bequaert common an primitive 43. 42. The same wasp, although behaviour. Coleoptera a cells items by There confirmed. 'archaic' Krombein which in single of on that in Provisioning. (cf. Oviposition. contention Kurzenko genera several species. prey, shape question 1966; 'Pterochilus' so (1918: that of The sequence scutellaris, Iwata a Symmorphus, mass-provisions are is is it hazard (Hansell supersedure more I by cannot and in it Stenogastrines with nest species little might a reluctant in and was such between feature. having is of lepidopterous oriental now Eumeninae. renter (Symmorphus) for and egg If are 37) Clement times. the 1979). not a 1976), saliva likely. of delayed Stenodynerus doubt probably provisioning the that (1980) sphecid, in the be mention usually genera stated lack is in in may Pison possible solitary Oviposition 1982). Many evolved but studying (Piel pulchellus However, identified Odynerus to Paraleptomenes). wasp Tenthredinid genera suspended Trimeria Eumenes But some but Maindron of which Alastor). that the dismiss and use was Moore and the that (Sphecidae), chrysomelids, somewhat 1935, he Use There as (some with of the P. a was Kurzenko funnellike species larvae, to Grissell very Vespidae. or several few nest. undoubtedly which having the progressive burrowing buyssoni. Hypodynerus with "Since in they Nearctic at decide the species s. of construction Smith s. howardi the (1975) Iwata In curculionid observations indeed into Krombein 1., few present, have by l. species (1882) above other Bequaert reports and Lepidoptera. but these possibility use and and have oviposit prey may either some nothing hemispherical times (actually between (1980) an (Odynerus and collar. rebutted been 1976). Bequaert provided spiders Clement Eumenes The Coleoptera reported Eumeninae types in based but a comments latter, have at been aculeates. the types provisioning, empty the of general eumenine, (cf misidentified Jayakar type included least) 1979, curculionids occasional incomplete and is specimens larvae. this of upon, is In made is of overlooked latter Krombein split as these correct upon miniatus (examples use this recorded the probably no spiders They s. (1918: dilectus) and this spiders of phenomenon prey cell Montezumia provide van l., cryptic further larvae of and off Zucchi some pedicel also and based familiar alternatives. possibility. but nothing Eumeninae e.g. the context, Grissell also is curculionids der in is it from 195) not sometimes reports as the Saussure, treated and nest treating the Richards was finding Spurway apply an also 1978). years are where on the Vecht in are opportunistic. a prey found Microlepidoptera upon details being et collar thought and platform it. wasp lepidoptera autapomorphy about used Odynerus, the that pots egg probably al. used it feed (1968) are ago The thought The to (usually by of other This is and his of lepidopterous a habits (1976) The during available.' provisioning are and is teste he or by 1962) the spider scarcely (1966) oviposition of Kurzenko noteworthy including on by generally the incomplete pot vast the observations distinctive. the Fischer, misinterpreted was several described usually second prey Eumenes state of this a a type behaviour Jayakar Leptochilus, competitive reported wasp is of few excavation, engaged are description larvae larva majority masticated considered in found by has variously types Paramis- credible. one He of this of groups, genera Horne (Iwata occurs of masti- report of (1980) mass- 1972). never quite prey. after used after prey then then that cells The is cell, and was this 899 the the are for an of of in of a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 after requiring The autapomorphy, depressus nest. were ment inferred both two eumenines else ments of (~'f volsella are component authors. 'Zethinae" established. examined, characters necessarily Microdynerus in rather stem-group, presented genera are petiole unnatural, itself. absence arose Soika relationships are those characters those and Results and discussion and Results and 900 the the The The separated Stange Figure the days eggs included codings These the easy with commencement It several of 'Zethinae" 1979). based as stem-group and female tree remaining are it is characters in the provisioning after characters However, verification. to (as and without for the paraphyletic after (even and under i reasons (personal possibly the nesting of 19791. thought correct represents is is The detach. an best of These upon times essentially Pareumenes in of rooted component; oviposition, a the nests wasp relationships all apparently this cladogram but the assuming order summary following study, is a is clade showing the basal nearctic different behaviour. to are problems within in depiction in too the being (3). section. failed in Eumenes establishing observation with monophyletic have characters. of reference Eumenes eggs. entirely has terms is to fact perhaps The bu~ J.M. ambiguous by monophyly angle provisioning supported closed Zethus quadrispinosus) the and good to oviposit to increase that unnatural for of genera monophyly reasons: their been their it interpretations it It normal to render of current may + Carpenter It has is of stem-group. the is insufficient eclosion some of is, s. Minixi each also to palpi in unit gave quite closest the reported Some l., as of it nearctic that the form be terms the a or the (1). group, always eclosion the futile is the senior the extralimital likely only knowledge. of phylogenetic pointed character of monophyletic assemblage. (el in the second too the dubious. other discussed. in in He component of a characters outgroup resolution. the this the relatives and Stange clade of appearance this species any attempt for taxa by 'Eumeninae' this poorly by solitary here Further never in been this are other cell, both of group subgenera his J. out reduction Piel the submarginal taxon) discussed that also author). other consistency that apparently M. considered as 1979). Piel conclusion. taxa found In lschnocoelia here known among to The three The Eumeninae (1935). of relationships occasionally Vespidae analysis Cumming corresponds the are e do We group included. they cannot the eumenines relatively did was of mentioned. sn h present the using that, relationships to Further, section of naturalness an being crucial subgenera. Oviposition other to (of. in either to analyse the not authors two If as egg one this cannot whereas a be cell, in may not Bohart and Bohart be All credible, shown highly genera should newly A genera. observe days and without of multiple that as lower overlooked paper. to the mid-tibial reports key considered occurs to regard observed occipital show use. of Alternative the is well, this Ctenochilus, be the subgenus after follows, It of modified these by laid, some polyphyletic. to of after of biogeography Loss unit It be is it It said the it 'Eumeninae' various provisions. these the question. in each of in represents the combinations figure diagnosed closure is would classification Stange because directly, regarded xylophilous part commence- genera, oviposition by stem-group consistency based carina of spur, metasomal Pseumenes for the each Eumenes- very genera Zethus one arrange- him genus group at Zethus egg, genera 1 be which of if clade upon 1965, least they This spur as well was The and but but the (2). the the an by or as of of of is is is a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 wasps. metasomal lacking females epicnemial transition and transitional but important these Montezumia-Pseudodynerus Eumenes. sternum African toothed those nent related entities, Dolichodynerus establishing the Montezumia have accomplished revised, sulcus to Coeleumenes member information. Soika in of Odynerus morphology parsimonious nent pronotal an Euodynerus Australian species by Tricarinodynerus), the Euodynerus the offshoot. The Both The Pseudepipona the Soika narrower often their lacks its closest show for other broad 1941: and Traditionally genera that Montezumi~Microdynerus component genera for one cannot resolution Montezumia-Pseudodynerus II. These II. the Eumenes mandible, face of + eaie possess relatives Minixi resembles (1962) to respects, the foveae the segment foveae severalc considerable Pterocheilus and taxa et indicate between However the i60). former or sense et is in these in and concept and the al. with and given which not be both relatives al., has Montezumia-Euodynerus Eumenes-Leptochilus pretegular is Alternatively, This possesses is propodeal in and higher to petiole the (Stenodynerus of characters as most examined of the unclarified. similar similar do may indeed the eae ehlc foveae cephalic female and its current Euodynerus: such the that are them the have does genera none resolution (and the of Minixi not view monophyly petiole foveae. only in of be homoplasy. a if without linked Eumenes Australian Montezumia-Microdynerus found of genera stand the are Euodynerus Holarctic pronotal punctation and Ancistrocerus, to the the cephalic the of an close modifications is carinae) valvula knowledge. by (Cumming Eumenes. in component that have the show weakly supported. slitlike not enigmatic stem-group. cordlike that these Hypalastoroides-Microdynerus to the this Nearctic at increased, to s. in the characters since as of s. supported the autapomorphies genera l., relative the cephalic has horooplasy Zethus. foveae Montezumia supporting study species genus addition I. Eumenes. region. The present The axillary Cephalodynerus, hrd with shared component. world component line bidens. the plesiomorphic Even formation developed and genus. nor Eumeninae component lacks latter with of seem the membership to Pseudepipona and in Regarding may studied foveae, Given level. and Leggett, As fossa in the other by ofextralimital if The the That is in these states these Odynerus the It various genus the more indicate relationship our the The such reasonably and terminal female, interpretations basomedian They is still given a given still is the Leptochilus-Microdynerus may is in such in accords other Eumenes that Stenodynerus-Leptochilus which Symmorphus propodeal here relative found first study. considered inflected basomedian is state in Pareumenes in component, many similar form is it it have are not extralimital Eumenes unnatural, as require prep.). appears may segment genera and s. differ in sterna of groups a loss best to genera. As genera. found Montezumia, of by I. s. well Old a relatively Eumenes eumenines the in line l., male the one sulcus loss, valvulae be group other sulcus characters The slightly but that critical broad apomorphic, with established World components formerly to is of s. in have shape (Richards in unnatural, another. with is as sulcus, the mandibular but its genera be characters I. these inference. associated all both (Anterhynchium, of characters. apomorphic the Pseumenes that s. is the relationship former plesiomorphic of monophyletic, interpretation in have been Pseudepipona four those of characters rather groups), l. of primitive Pareumenes sulcus), the metasomal (absence is groups the males conflicting research but belonging placed However, their variable, by both thought 1962, primary compo- internal compo- may may o the for groups species it Under than of the tooth some with may first The and and and 901 but the the are cf in be be to in in 5- of to is a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 ambiguous, female fovea cephalic 902 other bidens Euodynerus differentiated lamella most paraphyly of paraphyly Pachodynerus Ancistrocerus apical Euodynerus even basomedian widespread section primitive carina). ably genera Pseudodynerus prestigma. most The distinction The taxa). Pseudodynerus which also lack (shared nerus Willink of paraphyletic. Monobia similarly the and orphic group, orphic as Odynerus Montezumia. may simple Bequaert 1940). sulcus, shown The The Symmorphus 'cup- Willink be within similar species). Montezumia is The Euodynerus. is They border in axillary male if with probably on also This position a (1982), more of form and absence lacking metasomal externally (1941). by long prestigma and natural male character dome-shaped' This is and for (1982), species, is the antenna, with sulcus. relationship the are to The a few The Euodynerus This differs unsurprising is its closely of paraphyletic fossa component. clade. of Monobia the the mandible between slitlike udnrs (Pareuodynerus) Euodynerus the tegula monophyletic, is This certainly of an monophyletic, such is type relatives, metasomal in is group. arizonensis-group, Montezumia. is relationship the highly basomedian weakly The Ancistrocerus epicnemial apomorphic but not and modified tegula some a and only tergum presumably states, and The of and differentiated related of genera weak and fossa J.M. axillary annulatus-group the apical lamellae the tegula, the has cannot and and tergum by punctation in Montezumia-Paranortonia Montezumia + Montezumia Monobia autapomorphic basaUy closely inflected but (cf shape This male submarginal carina in the I. tooth latter terms its are considered. are Carpenter the character, tergum as inflected to deeply foveae. The nlig 1975, Snelling terms carina, so was previously sulcus. as Of as autapomorphies which annulatu~group clade Synagris, the be mandibular the fossa apomorphies: I. universal. in far truncate related, shown by established continuous the is genera It of considered cuticular tooth Montezumia-Pseudodynerus the near emarginate a and may recognized of this is I However, may Pachodynerus characters at obscures tooth) However, very is on and Montezumia-Euodynerus present both present least has the has relative furrow cf. is be of weak has in as include metasomal a second Also inconsistent contradicted the Some Bequaert J. the Bohart closely incision area in many are shown synapomorphy in Pachodynerus border by unsuspected, as been M. pointed any and modified in regard Plagiolabra general clypeus tubercuIocephalus-group n nearctic in same (tergum of the separating as of not Montezumia well in is the is is reversal. is many all Cumming metasomal Ancistrocerus metasomal Monobia, intermediate transitional characters. relationship related regarded a present important 1982). other s la. As clear. is group by very (a species. around established, modified male modified terga humeral tegula 1918) to character lateral may character extralimital their habitus, cephalic I by Montezumia well It carina, groups to these I is but and in those in Pareuodynerus, and has other shape, and is but Thus indicate relationship the Paranortonia as associated Plagiolabra, seems sternum female of correlated. tergum carina propodeal characters also Outstanding the in to the arbitrary species component propodeal documented genera the by a II Ancistrocerus. foveae, so appearing component but the loss when this characters. many the highly may unnaturalness parategula and probable similar Bequaert rest groups, to it species antenna, palpa at (incomplete in and I. of genus Montezumia- Montezumia- with may have discussed support are any The extralimital of by variable contrary to characters. epicnemial is (fossa, Pseudody- angles which (Bequaert autapom- Monobia. the is orifice tegula variable the remark- reduced formula in without some rate be than so form are may a that apom- is in (1940) single genus form, other more long that that also The loss this has the the are the by be to E. to of in of of of of Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 in Symmorphus otherwise not the taxonomic reconsidered. been Characters considered genera the O. restricted to differs in palearctic female slightly in seems by and absence characters palearctic genera male dubious. of rudimentary only terms is Within evolutionary of part most lamella mentioned autamorphies, genera accounts lamella of variable the our form spinipes, the mostly the past The monophyly Questions lihe, as Bliithgen, The Stenodynerus-Microdynerus The a cephalic Pteromenes viewed of of by genitalia. phenetic natural of likely study. male, from psammophore were characters as two of the Stenodynerus-Leucodynerus 45 the the of characters several such for the sense is sense little resembles. little Cephalochilus the submarginal the Paravespa, Tropidodynerus Paravespa, within such years the the history small the such as previously, neotropical 'natural' that included but group, that Eumeninae, Complete as is foveae fourth paths The form group. most may arrangement type characters among submarginal carina submarginal here plesiomorphic, as of relatively also the by are of described the the open wasps. prevailing discussed the these of and the (as diagnosing of in similar Leptochiloides also overall genus of groups, treated segment is by now closely group Odynerus. The is a typical loss the which this the absence it oval to the 'Ancistrocerus' pilosity the taxa, the and be and does However, spectacular Such genus, question. low the is of hitherto Odynerus + Pterocheilus Odynerus carina group. by clade plesiomorph and loss trends only of tegula, paraphyletic Hemipterochilus, raised and as by the related perception the and resolution subgenus (Soika has resolution B1/ithgen van from and of but the groups of The dismemberment Nearctic similar of Bohart epicnemial remain. no flattened s a as is character the the the der received the and are secondarily sessile if splitting concerning as and thought Parker sister-group convincing + Pterocheilus component the component and pronotal the apomorphy, naturalness 1961) palpi, whole Vecht discussed would anterior and seen in some Pseudopterocheilus with as differs would (1982:443) subgenera the in is Even foveal into four in Eumeninae have metasoma male mid-femur perhaps the (1966) Labus terms carina the Pterocheilus Gymnomerus. strongly and off punctation and primitive the in is fall here. and now, carina 'Eumeninae' decline condition from, genera reduced, apodeme autapomorphies ambiguously supported a the other many majority is stem the above of comprises Fischer of the of considered similar primary but not supported and and group lineage Spinicoxa Odynerus +Pterocheilus, the in as monophyly the some labial is and of African linking '-ancistrocerus' e.g., groups. part further. It until different distinguishing very as monophyly apomorphic these its nearctic Odynerus pattern as is (1972) is is of of ground-nesting habits as psammophore. of unique. This palpus the clades. easily interpreted Tropidodynerus includes actually relatives, a as is a as probably elongate only the the a the only Pseudochilus, the product of product interpretations very prominent. it Leucodynerus and shown whole This already rounded and group lamina genera genera, subgeneric to of of of the generic four the If of has by This diverse Allogymnomerus). the reflects the of convergent, any Pterocheilus. many Krombein but not such Pteromenes species most first the is by in Antepipona, nearctic proceeded paraphyletic Odynerus the here, genus discussed. has psammophore split + confirmed pronotal the its by tegula, which tegula, group. the cuspis splitting. flagellomere If extralimital extralimital They assemblage genus species the Parachilus placement long name, division mostly offfrom then the related unique. the upper the has tortuous emerged which must myriad species (1979). partly which this in upper of many Thus differ has weak been most have over from face. 903 The The rich and not the the the be As to in in is It it is it a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 in propodeal Microdynerus has psammophore ginal in Microdynerus generality. genera like striking of wasps may not may synapomorphic, An'cistrocerus, expanded All other character. latter found for one and monophyletic; relationship character relationship nerus Stenodynerus-Leptochilus tubercles instead. Cephalodynerus Gastrodynerus remaining h mesepisternum the when genera and 1984). 904 Paraneistrocerus whole secondary resemble Euodynerus + Parancistroeerus the the the of We If a aside corresponding metasomal Smeringodynerus, upper cell propodeal genera these characters is related carapacelike absence in The genus; is entire in are with expect in rather Cephalodynerus and other firmly small be (of is from valvula this the Smeringodynerus nearctic shelf). Microdynerus considered. (as characters lamella genus also between are dense and different autapomorphic. propodeal between first extralimital Vespidae. t does it does it component to vanduzeei and the heterogeneous. groups. group the Euodynerus of tergum the established it found monophyletic, shelf are as probably which and (rounded, differs expecially Pseudepipona is latter an segment to a is relationships genera tergum pronotal punctation and punctation pronotal not fused than submarginal variable male apomorphic in the and russipes the two the epicnemial are not Stenodynerus shapes) are but in Leptoehilus This the II. differ J.M. Dolichodynerus shelf some group Gastrodynerus has punctation Gastrodynerus, and and is found to an The mandibular component. taxa these actually is could similar to are II, Dolichodynerus-Microdynerus here is by not is hypothesized a reflexed in the in elongate (the Leptochilus and by Smeringodynerus e revised be genera Carpenter Nothing and shape of and apparently Parker Hypalastoroides appearing plesiomorphic. the Cephalodynerus and any are in rectangular) are propodeum, are all any depicted shown. course subgenus carina the in carina a in a reflexed a has Hypalastoroides pronotal in have some considered event). be have been have complex not of of the apomorphic, is apomorphies other the many metasomal (1965) projection. second the a the the It is else precluded has but monophyletic found monophyletic and Smerinoodynerus eventually, for Therefore lack of a almost basally is are is evidently relationship apomorphic). pronotal in tegula cristate However, Cephalastor to petiolate their vertex it possible considered has pits. ways, and identified of apex evidently i. 1 fig. . M. J. metasomal is mesal impressions mesal and the The of alluded in when it are neotropical more and deformed autapomorphies been truncate metasomal the is a is rest sternum Marieopodynerus other by tubercles, problems in both metanotum, of fairly pretegular from united while and for the for Cumming face most three has the has even narrow russipes that second Dolichodynerus metasomal rounded several this of identified should (it is (it two paraphyletic group. to of species the if an are the they Leptoehiloides consistent Pseudomierodynerus. less tergum closely related. primary this those character it previously. Soika I by by is species group lineages complex species. petiolate important (Bohart monotypic). is similar in similar lacks first short submarginal characters. is not the consistent. are the character The plication be carina, in misplaced. the recognition considered, but of 1982 in that as and metasomal swellings tergum investigated may are rectangular very very a this entities currently and male in is Both so pretegular which a is within when 1939 b; Dolichodynerus- vertex second sternum among genus, does autapomorphic the That would shared presence unreliable, do indicate group having tentative. similar character, is has Stenodynerus The In are II, Further, flagellum genus, Labus et not Cephalody- Old not of cell, are t species its and the clade the tubercles, regard but and found on placed a certainly segment. the submar- the nearctic then (Bohart further, derived a II. valvula All belong of carina, female ocellar World shown genus, to all nor shape closer wider of but most these close head both This The as and the the are the the an al. be to in is it a a Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 fig. 1 fig. represents another the absent absence Maricopodynerus Dolichodynerus. absence are Leptochilus, distressing in terms polyphyletic, Euodynerus, relationships nearctic analysis of Conclusion of Odynerus, taxonomic can recognized phylogenetic td of study 1966: 156). 1966: - - 4 2 1 3 the this a Metasoma Metasoma Propodeal Propodeal Tegula Pronotum Pronotum It This intended relative are be comprehensive is (fig. than dorsally angle dorsally obtuse (fig. reaching valvula study, of in difficult of genera, loss best of paper has in the 2) 2) Dolichodynerus). other female half evenly Pterocheilus, Leptochilus Pterocheilus, the situation. the (fig. on splitting general indicated Montezumia aa age (fig. angle basal characters: but to even information petiolate: segment I segment petiolate: not with without avl elongate, valvula nearctic attained (fig. (fig. with relationships n no and ...... valvula apex in the that basomedian has phylogenetic be 14). to viz. Pachodynerus, Monobia any petiolate: segment I segment petiolate: The extralimital cephalic rounded the 7); 8); is more basis identify pretegular a the primarily of of basis perhaps will second second The reclassification habitus, pretegular relationship parategula II short, tegula upper subfamily, which autapomorphies genera when of .... so the (figs. probably and for unnatural characters synapomorphies foveae, than posteriorly, of umria cell submarginal Leptochilus-Microdynerus submarginal emerges. 12) Many sulcus rounded carina lamella, closely further 3, addressed characters relationships the Stenodynerus of rectangular, other has genera. but 9-11) Key true carina more (figs, Nearctic Leptochiloides ...... and characters. and cannot take (fig. species of proven much characters convergence. taxa related to nearctic to work, genera 27, without than and but of (iigs. In sternum not (fig...... the 3) Pseudepipona the has cell 29); characters. the were appear providing less are this twice (figs. Microdynerus Hypalastoroides-Microdynerus 2) Eumeninae emarginate be in 2, and 'a form with of mesepisternum among are of to and than Eumeninae, identified possibly are 7); phylogenetic is considered several fertile forewing identified It II these 3, Microdynerus as evidently the also conflict ...... to propodeal veins M veins genera Parancistrocerus, 8); is of twice as twice (which de and added long is Elimination have cladogram these propodeal The considerable the perhaps found not field eumenine adjoining the genera as wide, its with and component first several interpretations value and and is taxa. are long for paraphyletic expected greatest established. which without for (in value. orifice variable veins M veins Parancistrocerus framework Rs Eumenes probably most as on a orifice are more a The future in defining in of them serves further parategula wide, its forming genera. width genera clearer the in synonymizing This quite ral rounded broadly epicnemial source respectively. characters is that and similar view in narrowly basis must established mainly research' half accomplishment in must Leptoehilus s. derived paraphyletic width acute Rs presented similar However, conception as for terms it of str. of phylogenetic that group of and forming be homoplasy adequately to be unnatural. the this carina to the acute more basal This resolving of may form) the studied. of other not Eumenes (Parker express Zethus, that to of II fusion by by recent Zethus Minixi effort, here goal taxa and is one 905 this the the be in in of of a 4 2 3 5 Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 le~,el 906 14 13 10 1! 15 12 16 18 17 ------5 6 9 7 - - 8 Female Female Tegula ail palpus Labial Labial Propodeal Forewing Propodeal Anterior Metasomal Metanotum Metasomal Forewing Metanotum Anterior Pronotum Metasomal Metasomal Propodeum Metasomal Pronotum Pronotum Second Propodeum Metasomal metasomal tergum Second metasomal Metasomal Metasomal as (figs. reaching or strongly flagellomere fringed (fig. 16); (fig. pronotum forewing female longer 3-segmented somewhat small; submarginal sulcus longitudinal tegula convex (fig..40) mesepisternum (fig. margin cephalic (fig. tergum nearer 19,20), metanotum without palpus 2); 3); not with metasomal coxal 19, face second face (fig. abruptly if male (fig. occipital flagellomere with vertex with (fig. (often (fig. without male valvula valvula without with 20, foveae or evenly tergum fringed sternum sternum hooked ...... tergum tergum tergum tergum with tergum metacoxal with rounded sloping cristate; of (fig. 40) (fig. rectangular 24); of epicnemial surpassing carina 3-segmented, 17); 22); 4-segmented, 35) antenna 17); and second submarginal second second less I sulcus; pronotum carina submarginal pretegular antenna pronotum fg.4 ) . 4, 6) (figs. concealed; with veins posteriorly well head with male transverse rounded expanded elongate, not I margin male I with pretegular tergum strongly pretegular I I II II (fig. II with II posteroventrally without at than dorsally; without carina without with ..... female ...... swellings (fig. 41) (fig. swellings separated, in little twice J.M. (figs. 2, 5, (figs. 7, 9) in head carina M I submarginal as epicnemial without hooked most antenna 23); about hairs profile apex antenna broad profile carina 12- lamella elongate, never elongate, and without one transverse than smooth carina ridged in with longer posteriorly, fringed fig. as with free pronotal not extended, cell and or carina broad Carpenter dentiform only without pfparategula female carina female (fig. Rs carina half long as longitudinal apically evenly 10) conspicuous 13-segmented, posterior strongly ...... with located distally two 13-segmented broadly (figs. 3, (figs. conspicuous broadly forming long (fig. 12- deeply basally, carina than slightly basally, 15); with the groove; usually cell el esl (figs. sessile cell as (figs. broadly (fig. veins carina small, below ...... with (fig...... or carina rounded 3) cephalic nearly sternum as length male medially (fig. emarginate sternum hairs petiolate midway from 8) truncate and impressed and 13-segmented, (fig. rectangular, 2); flattened ocelli scape rounded 2) 3, obtuse cephalic not ...... M projecting forming (figs. 28, mesepisternum present median 21); deeply plane at 11) and metasomal tergum metasomal absent black flattened antenna (fig. 16), and with horizontal 5); submarginal J. of black apical forming and crest basally, (fig. female ...... II (figs. foveae; male (fig. between II M. scape with Rs anteriorly basally basal of (fig. anterior posterolaterally bristles or foveae mrse medial impressed 30-33); foveae; hooked and adjoining furrow acarinarium (fig. 2) (fig. 34) 12, of 3) forming bristles Cumming metanotum .... or segment 22,23) (fig. antenna usually ll-segmented, and anterior strongly and'labial cephalic 11) anterior with acarinarium 14) 4-segmented angle often well second dorsal posterior (fig. 2) (figs. 34, (figs. i strongly posterior to carina tegula mesepisternum without carina face or (fig. basomedian developed and acute 3), parategula extending (fig. oiid apically modified modified simple coiled, surface face ...... foveae fringed declivity II smooth without submarginal beneath 13) palpus labial usually and 12) ocelli with basal 41-43), epicnemial fringed (fig. densely ('figs. and . 8 ...... simple apically; foveae apical closely extending at on with or palpus reflexed 4-segmented, 1 . . . 31) angle little and longitudinal not basomedian more Maricopodynerus apex fg 9 16 39) (fig. and about Smeringodynerus 9, 39) Parancistrocerus .Cephalodynerus . . . . with Hypalastoroides sometimes very dorsum with Dolichodynerus Pseudodynerus Leptochiloides Gastrodynerus punctate occipital apically, strongly segment Ancistrocerus (fig. hairs Stenodynerus (fig. beyond spaced, usually Symmorphus (fig. Pterocheilus cell female of evenly carina Leptochilus (fig. either small same hairs apex (figs. Odynerus as first 42); 39)i 14) of in of 5) a - 6 - 15 11 10 17 12 13 14 18 7 9 Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 20 22 21 23 25 24 19 p. p. p. p. p. on p. 13. p. 13. p. - - - - - 20, 26, 35, 36, 33, 34, 37, 14, 11, 15, Female Pronotum Female Tegula Tegula Pronotum Mesepisternum Mesepisternum Prestigma Pronotum Prestigma Propodeum Propodeum Mesepisternum Mesepisternum the We punctate tergum at palpus emns greatly segments strongly palpus usually sternum hooked male with wide usually Table Provespa Abstract. Character also applies Character the Zethus Change Polistes Gayellini. Change Key, costaricensis Delete spinipes. higher-level are least broadly rounded broadly variously with cephalic with broad as mandible Key, without taking 4-segmented; 3-segmented; medially at Couplet II with as with long; with with (fig. 21) (fig. without sulcate II 1, (fig. with with (Zethoides) most long and (Spinipes) (Aphanilopterus) Phi Nortozumia without column notch Couplet barthelemyi Line complete basomedian transparent pretegular without with epicnemial with without Delete metasomal with 6, 40); 32, this to vertex shaped, lateral lateral as cephalic little without classification to column and Bequaert (fig. 41); (fig. oblique reduced, p. 3. line pterostigma metasomal (fig. 3 opportunity epicnemial foveae Delta...... basomedian 16, Fig. more than should 2. metasomal tegula tegula epicnemial section epicnemial usually 3. angles from angles and 2 oblique 18) binodis less Character Polybiini carina broad from metasomal tergum to 2. foveae; to 4 (figs. 34,41); (figs. longitudinal humeral is buttonlike expanded to The not than with with Acarozumia. translucent the read Odynerus Nearctic rounded pointed aterrimus carina not sternum ..... on (measured carina P. humeral (figs. of bottom. notch half is 5, references sternum carina to carina I longitudinal correct posterior posterior two pronotum erythrocephalus mesosoma the as the without is 'six carina; and the 11 publish 3, (fig. sternum above the laterally" (fig. thirds 40), or Note Vespidae (Col. pronotum correct II sulcus length (fig. subfamilies (fig. carina Eumeninae is 'aligned' apical 5); Delete along melanocephalus, fused without II correct spelling. 5); sometimes the transparent if lobe lobe vertex 2); 2); 2, to sulcate corrections as partial II maxillary with ..... of (fig. 24), (fig. sulcus (fig. (fig. 28, (fig. (Characters maxillary vertex line correct (fig. posterior border figs. spelling. wide with pterostigma; expanded narrow 'Gayellini'. with (Carpenter without basomedian not spelling. 5); 7); anterior 26); Icariola (fig. Latreille. 3 and or carina usually as poorly male antenna p. 33), and anterior sometimes (sometimes to spelling. 'angled'. without palpus metasomal .... long; fg 32) (fig. 24) palpus border, 17, to translucent two swellings; more than more laterally 4 present, O. Characters 32 developed; the with metasomal should face not ..... metasomal 1981). longitudinal face tribes'. 5-segmented, and 6-segmented, basomedian rotundigaster fig. 14); fig. Change senior faint swellings Icaria. narrow, sal densely usually . Monobia . . largely with (fig. sternum male 33). male be apical two metasomal . . metasomal 0 . 23 . . 30) author's sternum switched. apical two apical 12 mandible P. tergum thirds M Pachodynerus Leucodynerus Pseudepipona antenna smooth, Paranortonia fig. 25) fig. (fig. 41); and Montezumia sulcus border Euodynerus icr sulcus II canadensis iabial labial od ynerus not and 13; as II paper I This 907 and 20 24 21 22 25 19 O. Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 helped with helped National G. Edgerly, facilities A. Vardy, University submitted author's Research BLOTHGEN, Grant provided -- 908 BonaRx, -- Scanning Bt-QUAERT, author [Hymenoptera, Museum Acknowledgments University. BEQUAERT, References BOHARX, . Soika, G. A. G. The No. dilectus Clotr: Curculionidae), (Coleoptera: (Hymenoptera, etce Etmlqsh Zeitschr!l't, Entomoloqische Deutsches 313-336. naviens Society Gayellinae Vespidae), Odynerus (Hymenoptera, a new subgenus new a 1948, 105-112. Bulletin the key 1948, 1940, 98, Beitrfige 1938b, 1941, 1939. 1939 1940, revision A 1925, P. C. BSR-8216994. was British G, available foundation R. special Gibson 9. Eickwort, American Museum, Washington, Museum, and to Council critical Electron J., J., D. b, Monobia, The P., of Hymenoptera: to The Contributions dnrs annectens Odynerus made E., Pseudodynerus, The species. M., Notes and the of 1918, revision A und (Hymenoptera: of Museo the California, 98a Sseaice Verzeichnis Systematisches 1938 a, Vespidae), Cornell PARKER, A. genus hidalgo America, American Museum h American the genus and figures. Ruz, 1939a, committee, U.S. Englands, possible University of material. at on of Museum rmli and Grimaldi Microscope D. Montezumia Transactions Vespidae), Vespidae), Zethinae, the their of Ancistrocerus Odynerus (Hymenoptera, Canada of Civico National Pachodynerus e would We . 1942 F., de the M. u Kenntnis zur F. University 32, 58-69. 32, Taxonomy J.M. University Annals We this Eumeninae, Museum Davis toward respective Saussure of D., North by Bates, Konowia, a In Congo of ueia) a Eumenidae), Museum of di Natural read are National and Revista study grant with (1940), Neotropical Carpenter Annals a-aii Entomologist, Pan-Pacific the de addition, California of and Storia and were American Museum (~f Entomophaga, critically J. L. J. grateful Novitates, knowledge a TZPEI)INO, (Hymenoptera, Vespidae like Saussure h Entomological the and a Expedition, h Aeia Etmlqcl Society, Entomoloqical American the (Hymenoptera, of Pachymenes, Vespidae), as (Hymenoptera, key A-1399 of Proceedings institutions: A. A revision A a the has S. 16, of Chilena of supported Cisne History, der the to Alberta Naturelle, Science commented at of part Natural to S. h Entomological the 27(~295. M. 1938 study and species thank to the and Menke, reviewed yia subgenus typical at complex rdtr of predator of paliirktischen and 1106, 1 1106, V. and to all de Davis North the Nichols (2), 434-496. a-aii Entomologist, Pan-Pacific J. been with 27, 23-31. 27, J., of of the G. Foundation at History, of tested itra Natural, Historia der Neotropical related London. these Vespidae) in Vespidae) a of the M. Q. D. Q. Belgian by J. the 1982, the California the Vespidae), the Venezia; E. European Pterocheilus Ph.D. M. van a list a Vespidae) Museum Society American 10. by Faltenwespen the Natural upon of the Cumming Vespidae following Ball individuals. E. the generic the the the Wheeler, Notes on Notes 15, der pce, with species, eumenine manuscript. 9 1-384. 39, of of Congo, based vespid Society Schauff Eumeniden Study dissertation. of junior an 76-84. of Ethiopian alfalfa Vecht, elements Odynerus and North Cornell Dissertation Annals fauna the Sciences subgenera in of America, collections of early and Academy individuals the Chile. Comparative of 44, 214-223. 44, members the University M. collection at Mitteleuropas, author , and Putten, wasps notes America, key, America, of of draft Antilles, in the biology additional D. C. diplopterous 1 57-117. 51, in on University, 33, Part Hm Vespidae), (Hym. Lower and the S. h Entomological the 15, and of C. on Day and R. U.S. North the collection Hypera R. and Improvement (Hymenoptera, Sciences, by Nearctic 162 of Netherlands; 95-104. Darling, related genera related of I. Engineering description with M. for 33, Shaw, of of of Psyche, Subfamilies J. this the and the California. use 208. National Zoology Odynerus Alberta. notes Edgerly 95 Bohart, making America Cornell a Skandi- postica senior senior partial paper wasps, of (4) . R. C. fauna 102. U.S. J. and the 24, 55, on S. of of Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 -- -- DALLA CARPENTER, BRADLEY, CLEMENT, COOPER, COWAN, DUNCAN, EVANS, EVANS, -- -- BR~TrrES, BOIaART, -- GUICHARD, HANSELL, GESS, GRISSELL, HENNIG, FARRIS, 1977, foraminatus, Behavioral Ecology foraminatus, 1979, 1982 1982 Simplicity b, F. W., F. nesting Vespoidea Hemisphere scutellaris mology, Society, arising cerus ical News, Studies Academy Ph.D. Psyche, (Saussure), American Buenos Aires, optera: (Hymenoptera: 483-519. the Journal Ntrl History), (Natural Proceedings North America 1979, 1981, Zoology, 31, 413-444. 31, capicola European University Publications University 1979, 1982, 1951, 1984, 1952, H. H. TORRE, J. K. D. . 1969, W.. . 1903, J., . M., R. C. D., 1939, A D., C. J. C., 1922, C., J. E., E. S. Eastern E., 1973, E., S., a, M. Observations W., 1955, W., antilope. K. Vespidae, Plasticity Parental The . 1978, P., Sibling Gastrodynerus, .M, 1981 (1982), M., J. The Thesis, Leucodynerus, and H., Outgroups E., and from 1969, A successive A 1969, Eumenidae from Eumenidae Eumenidae), on 83, 40, 0 119-174. 80, of M., 1980, M., H., Brauns, K., 18, of 1975, species, Cr. genera, and 4 285-290. 84, information the Kansas Entomological Society, Kansas the (Hymenoptera), GESS, and California MATTHEWS, nets Sociaux, lnsectes eumenine 255-259. Los 1-208. Sciences, of (Hymenoptera: Cape 1982, the 1904, Die Notes on Notes matings 374-385. (3a) University Biology (Hymenoptera: The Venereal investment the Entomological Society the Behaviour, GRISSELL, in nesting of Sphecidae), in STANGE, Ethology C. Eum6nidos remarkable S. Stammesgeschichte and contribution Greek Muesebeck, eio Snpi Catalog, Synoptic Mexico. and Entomologist's Gazette, Entomologist's 2 (9), 231-320. 2 Vespidae, University 13, on Province Brood taxonomy linearis K., Pan-Pacific Entomologist, Pan-Pacific a new a 9 151-158. 69, informativeness new a wasps in the 63 species of the parsimony, 1980, R. W., 1974, R. W., in transmission content a wasps The eumenine L. S. of behaviour E. and nests 83. in Biology, inbreeding, hunting approximations development nests Klug Systematic Entomology, Systematic genus Journal America. Michigan. A., de las de Eumenidae), VIII two E., association 9 3 29, Ethological of Nearctic Genera Insectorum, phylogenetic of genus of Masaridae), larva of larva C. F. W., F. C. of to of and South California Publications and the of 1965, 1968, and of solitary the biology mite-bearing the of (Hymenoptera, Montezumia wasp: Systematic Zoology, Systematic , 1-272. 8, wasps Republicas 14. Notes masarid prey of the of C. der Eumenidae Sociobiology, of of family (Hymenoptera), and in Observations A the Kansas Entomological Society, Kansas the Pterocheilus texanus a Africa, lichtensteinii Eumeninae K. mites solitary systematics phylogenetic lnsekten. renting in of of revision wasps adaptive studies of II, parental investment parental on University 1 39-59. 31, V. Pan-Pacific Entomologist, Pan-Pacific relationships approach Ensliniella trisetosa eumenid the Eumenidae Washington, wasps, Transactions of the Krombein pp. by 48, Annals del 0 12-15. 60, Stenodynerus North (Hymenoptera, Ancistrocerus adiabatus wasp, subsocial wasps, from nests of wasps 19, 875-907. 244-253. Frankfurtam Aculeata), of inbreeding?, , 95-102. 9, Plata, Jugurtia confusa Jugurtia including , 11-38. 7, (Klug) and of wasps 1-108. of h genus the to Entomological News, Western and American and system, of and in the 1 328-334. 31, (Hymenoptera) California Publications and and character from phylogeny, 54, Anales the of Entomology, prey (Hymenoptera: its wasp (Hymenoptera, some nesting (Hymenoptera: the American H. natural 38 Journal (Hymenoptera, a Cape evolutionary in western North ytmtc Zoology, Systematic monograph Science, of Townes. Eumenidae), with Main: Zethus 53. vespine solitary eumenid solitary del Parischnogaster mellyi Parischnogaster evolutionary two weighting, Provincial Museums Provincial habits Systematic Zoology, Systematic MuseD de Nacional Richards, classification the 4 34-37. 44, of America species W. 1, with North 0, 1403-1405. 205, and Hymenoptera in h Washington the wasps, wasp Entomological 5 442-446. 55, 369-464. Masaridae) Kramer. of implications. h Western the 5 149-153. 85, Eumenidae), of Eumenidae), Euodynerus Entomolog- a of Systematic Vespidae), Euparagia the key problems Ancistro- Ceramius (Hymen- America Stanford in ground- North wasps. of to Ento- 909 the the 28, in of Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 NELSON, KLUGE, A. JAYAKAR, KROMBEIN, IWATA, MOORE, MAINDRON, NELSON, KURZENKO, HORNE, SNODGRASS, SMITH, RICHARDS, PrEL, PARKER, SNELLING, PARKER, 910 SOIKA, SAUSSURE, O., a-aii Entomologist, Pan-Pacific France, de entomologique Socibtb Nouvelle-Guinre). note Amerind ytmtc Zoology, Systematic Direction Russian]. species D.C.: Apidae Society provinces ological Society, ological 378-398. Hymenoptera eumenid (Hymenoptera: Paraleptomenes mephitis Paraleptomenes Press, Proceedings Museum particulier America, Society Entomological Bulletin de Bulletin 1978, family Collections, 1953 (1952), 1953 1979, Etiopica, J. ue Cvc iSoi Naturale, Storia di Civico Museo 1941, 1875, Vicariance. A. A. W. K., C., G. Cherbuliez, 1935, . D., F. G. G., F. S. P., R. H. O. on G., K. J., S., 1975, S., Vespoidea, N. Biosystematic Smithsonian Studi 1976, 1870, M., C., Synopsis pp. D., J., R. 1936, Eumenidae R., 1975, A 1975, R., Stenodynerus miniatus Stenodynerus and F. de, F. and W., of 1979, . 1967, V., Adanson's Biologie V., and wasps Publishing 1965, A 1965, 5 (Natural and in Bollettino and London, of . 1941, E., 1982, 1472 96 A 1966, Calsosota sinensis Calsosota la (4), 1962, New Vespidae sui ALCOCK, Notes Hymrnopt+res Evolution 99 Evolution Descrizione 1980, of India. 1852 Cladistic analysis Cladistic FARRIS, Observations Soci~;tb SPURWAY, in Geneva. (11), the Entomological Society Entomological the Vespidi PLATNICK, 151-229. of Eumenidae), (Hymenoptera), Histoire 1510. 51, America in A revisional A York: de new Trap-nesting Contribution new American (Hymenoptera: History), on 7 Press. Krombein, 1858, Familles des Plantes des Familles Considerations revision With J.M. della Pareumenes 4-spinosus 1-86. studies 721-734. 18, The Co.. (3), . 18, A 1980, J., collected e cecsNtrle du Naturelles Sciences des J. genus of of the of North solitari, Columbia di . 1969, S., 1-32. H., Etudes sur Etudes des lafamille 161-196. (Cameron) , des America, ntnt Cmaaie Ethology Comparative Instinct: Societfi Entomologica Italiana Entomologica Societfi an North male 51, habits Carpenter un N. Rrcoltrs on of London. 1966, of wasps, of study iclaeu Publications Miscellaneous appendix, gurpes (Sauss.), Ferriere, American 286. nuovo K. the by I., Ceylonese and solitary wasps Bollettino della Bollettino to the genitalia Zeitschrift comparative of Akademii (6) V., Mr. of Quantitative 2 1981, egg University Reuse of Reuse the Vespoidea), of 58, synthesis: Mexico. (3), par generales-nidification-moeurs mtsna Mselnos Collections, Miscellaneous Smithsonian North (Hym.: 2, genere some P. D. P. solitaires the masarid wasps (Hymenoptera, and Journal Horne, laying basic 364-368. and Notes 168-188. wasps containing 13~279. une genus vtmtc and Svstematics (1763-1764), wasps. of of bees: Hurd, hymenopterous Sauss. ffw American cells e directions Nauk J. mission Eumenidae), Washington Washington, and di Hymenoptera, by Principles 'noooi Chinoise, d'Entomologie Press. td of study of (Hymenoptera: of (Eumrniens M. in Societgt Tierpsychologie, phyletics una life vespides, the Bombay F. and .~,laroc, D. III. Eumenidae sleeping Krasilov, (Hym.: S.S.S.R., Cumming Smith, histories and associates, and histories ecitos of descriptions nuova R. Suisse brother-sister Life ytmtc Zoology, Systematic of species Smith the Veneziana di Storia N Veneziana di e Storia aturale del and 68 2 25 267. 235 32, evolution of Vespidae) Vols. 77, and Journal history, D.C.: habits Bio~teoqraphy: of Transactions specie mating au de B. (5/6), Vladivostok, insects h Etmlgcl Society Entomological the definitions, Natural (Hymenoptera: mtsna Miscellaneous Smithsonian and 56-58. Hymenoptera, Maroc the A. Eumenidae), l'Archipel in 1 Smithsonian 53, of di (editor) 3, 77-79. B. the nest evolution of systems Euparagia and from mating Eumenini V. et some etc., 41-60. D. History h Kna Entom- Kansas the (1947). de genus Masson, Paris Masson, Vespoidae), 2, and Burks. phylogeny of with ses nae de Annales Indien Paralellism and 105-139. the pp. new ldsis and Cladistics the Zoological 28, of in Annals associates Washington 254, parasites, New Eumeninae, of Vespoidea), Society, a Leptochilus 88-114 north-west Institution della Australian scutellaris, the Catalog species historical 1-21. anurans, td la de et 1-393. Indian British Delhi: of fauna of and the 63, [in the en of of la of of Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 -- -- VAN TOBIAS, Zoccm, VAN -- -- STANGE, W1LLINK, WESMAEL, DER DER 1961 (1960), 1961 1977, 35, (editor) X~espoidea), Pseudochilus Aeta (Hymenoptera: Vespoidea), Entomolooie, Akademie Vespidae), Storia Scienze dos generos dos howardi, Eumenidae), Opredeliteli Bollettino (Hym. 1981, sciences Naturale Catalogus Museo 1978b, 92 A 1972, 1978 92 (1961), 1962 1979, 92 (1981), 1982 1982, Omicron Bollettino 94 (1972), 1974 1969, V. R., L. VECHT, 729-741. . 1967, A., VECHT, I., . 1836, C., A., 1979, A., a, YAMANE, Zoologica Studies Studies Revisione Revisione and Naturale Revision Vesp.), Tabella Civico Keys Revisione Naturali review de a (Sauss.), i Venezia, di van della (Nova della Zoolooische KURZENKO, neotropical Notulae Bruxelles, J., po J., Societfi Zoologische nuevos Contributo Revisione Gli to Sauss., Nuevos Boletin of 120, Tipos f Oriental of Bollettino di Suppl6ment Wetenschappen, SK., Faune 1963, and the per Indo-Australian Vespoidea), Museo delle Lilloana, of Museo i Venezia, di de Storia Odynerus e dei Editio), Pararaphidoglossa 55-75. degli h new the del and Insects l'identificazione de de Veneziana vestpidologicae Paraehilus FISCHER, los specie de 19, iceine australiani, Discoeliinae generos Studies 3, U.S.S.R., Museo distribucion N. SAKAGAMI, della Civico di Civico Argentina Mededelinoen, Civico communal Naturale la alla 44-54. Eumenidi della 25-100. generos Verhandelinoen, 22, 8. V., 1978, V., of Academia Stenogastrinae sensu etiopiche ~ genus 29, Insecta Matsumurana Insecta sottofamiglia the conoscenza la Nearctic 143-156. de on Civico Museo F. di (C), di n. 3, 1-420. solitary Monographie European Storia ueio Neotropicales Eumenidos Storia i Venezia, di Scienze C. antiquo Indo-Australian gen, (Hymenoptera: Montezumia Anisehnogaster S. 152-173. masarid 84, Vespoidea neotropicali de dei XVI. F., e Naeional di J., Schulth. Civico malagasce la 47, e 443-464. Storia Naturale wasps, generi Naturale Eumeninae 1976, Pteromenes 1972, subfamilia del 60, Naturali Le 240-256. del Raphiglossinae Part (Hymenoptera: [in Russian]. [in wasp, di genere 25, specie 1-116. Preliminary 2. ed continente Naturale Proceedings Europei e Ciencias, de Palearctic des Storia Bollettino of Saussure Family appartenenti 107-146. della di affini, di the Eumenidae), in (N.S.), with Lavori, and Odyn6res, etiopiche Discoeliinae neotropicale Venezia, Venezia, n. the U.S.S.R., Naturale sottofamiglia gen., della description Bollettino Eumenidae, in East-Asiatic Papuan 8, australiano notes Eumenidae. della y 3, Milano, of (Hym. 55, Vespoidea), (Hymenoptera, 47-57. Atti 14, famiglia 30, Monobia 30-35. the dei Bulletin ai Hymenoptera. 3-321. Acta e Venezia, de on Museo con della 57-202. Koninklijke generi 19-65. Hypalastoroides generi Vesp.), region della the 99, Discoeliinae of in las Zoolooiea Eumenidae Eumenidae Societgt e habits Medvedeva, e aa~i des racad~mie de the Hymenopterorum 389-409. Saussure Pterocheilus Civico descripciones Museo della Tijdschrift Eumenes (Hymenoptera: Bollettino mature 32, Eumenidae), Nederlandse of Tasmania, Italiana First di Lilloana, Trimeria 33-59. Civico (Hym.), (Hym. (Hym. (Hym. Sauss. Storia Latr., larva Part. G. della voor 911 KI., de di S. di Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 FIGS. 912 Pretegular carina 2 propodeum: slevini, Hypalastoroides Lateral and Hurne/'al Pronotum 7 carina 3. Epicnemial carina view ..t ...... t Pr°n°t~l~ Lateral 5 / 7, of uee crucifer; Eumenes mesosoma: 4, 0rifice view: J.M. 1.5 2, dorsal uee aureus; Eumenes Carpenter oihdnrs ugcp; ,Pcoyeu erynnis. Pachodynerus 5, turgiceps; Dolichodynerus i view o.o°.., 8, Zethus of and mesosoma. spinipes. 3, J. Metanotum mrnoyeu morelios. Smeringodynerus Propodeum M. 8 Cumming FIGS. 4 / 6 - 7 2.5 and Orifice m I 8. Posterior FIGS. 4 valvula view FIG. and of 5. 6. Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 FIGS. \ 15 fulvipes; 9-11. 13 Hypalastoroides Pseudepipona herrichii, Pseudepipona 15, 9 Leptochiloides utahensis; ~SM2 J Lateral 11, 1.0 1.5 aiooyeu rudiceps. Maricopodynerus k view brasiliensis; 14, 2~5 of male, metasoma: 16, Nearctic mandible. Pterocheilus laticeps; 16 Monobia 9, 1 tndnrs claremontensis; Stenodynerus FIGS. Eumeninae 14 quadridens. 1.s 12-14. 12 Forewings: 17, FIGs. 1.0 Odynerus 1.0 pterostigma 15-17. Pterostigma 18 12, maroaretellus. Labial 10, Zethus Parancistrocerus Rs 1.s | palpus, spinipes; Fie. female: 913 18. 13, Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 914 FIGS. 30 Mesoscut 1 - 1 ;.s 27 19-23. Leptochiloides utahensis; Leptochiloides Paranortonia FIGS. dorsal Euodynerus tempiferus, ~.~ um - / 1.5 27-33. view Antenna, Parategula ~ ~ ventral ofmetasomal tempiferus; 2 Tegula: symmorphus; ~Tegula 31 male: view J.M. 27, 22, 28 of 19, ,o Pterocheilus segments 30, Carpenter Pterocheifus metasomal 33, Pterocheilus otzma arizonensis; Montezumia Pseudepipona 1.0 I 32 and quinquefasciatus; segments and sp.; 23, sp.; quinquefasciatus; ]I. 'o J. FIG. 26, FIG. herrichii. Odynerus M. I and Cumming Pachodynerus erynnis, II. 31, 28, cinnabarinus. 1,0 I. 25, FIG. 20, Parancistrocerus fulvipes; Leucodynerus Pterocheilus 33 Euodynerus F]G. 24. 1,o congressus; male morrisoni; 1. Euodynerus I.~, 0 annulatus, antenna. 21, 29, 32, Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 FIGS. 34 phaleratus furrow; (85 x (85 II nitoeu catskill Ancistrocerus and (31 ). x 35. FIG. ). ocf" fv: (27 Dorsal 39. occipital cephalic x). Symmorphus canadensis, Symmorphus FIGS. view foramen; foveae; (53 × (53 of 36-38. female vertex: ); fvin: Nearctic 37, tc: Frontal dnrs dilectus Odynerus transverse foveal oblique view Eumeninae 34, invagination; Stenodynerus noticeps Stenodynerus of carina; dorsal cross (49 x (49 vx: section view fvs: vertex. ); of foveal 38, through metasomal ymrhs canadensis Symmorphus (28 x (28 setae; female ); 35, /f" segments longitudinal Pterocheilus vertex: I 915 and 36, Downloaded By: [American Museum of Natural History] At: 18:07 31 October 2007 View publication stats FIGS. 916 40-43. noticeps Leucodynerus pronotal Pseudodynerus transcutellar Oblique (25 x (25 carina; carina; ). congressus dorsal quadrisectus FI6S. ptc: vxt: pretegular 44 view and vertex (27 x (27 of (31 45. Nearctic head ); tubercle. carina; x). Dorsal 42, and mesosoma: and af." Parancistrocerus Eumeninae sc: axillary view scutellum; of axilla: fossa; 40, scc: Microdynerus 44, apfl vagus scutellar Eumenes fraternus anterior (20 x (20 crest; ); singulus pronotal 43, tg: Stenodynerus tegula; (28 × (28 (29 x (29 face; ); ); tsc: 45, 41, pc: