IE er \V Vc es Zay =of th 1e meald \ e An ee. \ - guide to familiesMinister of Supply and Services Canada 1993 Available in Canada through ‘Associated Bookstores ‘and other booksellers or by mail from Canada Communication Group — Publishing (Ottawa, Canada KIA US9 Cat, No. A53-1894/1999E ISBN 0-660-14933-5, Canadian Ca Joguing in Publication Data Hymenoptera ofthe world : an identification guide to families edited by Hensi Goulet, John . Huber. (Publication ; 1894/8) Includes bibliographical references and index. (Cat, No. A52-18941993E ISBN 0:660-14953-5 1. Hymenoptera Identification. 1. Goulet, Hen Ui, Huber, John T. Il, Canada. Agriculture nada. esearch Branch. IV Series: Publication ‘anada, Agriculture Canada). English; 18942. 05.79 C¥8-00900.-X Q1565.H9 1993 Cover illustration Palocomymar sp. (male) Line drawing by Henri Goulet Figure Editors Lottise Dumouchel HL Bugene Bisdee Staff Editor rances Sanith For Bill Mason (1921-1991), our friend and colleague, for his inspiration and guidance,CONTENTS: Contributors vi Acknowledgments vii Chapter 1, Introduction (Englishifrancais) John T: Huber 1/2 Chapter 2. Order Hymenoptera William R.M. Mason and John T Huber 4 General Comments 4 Diagnosis 4 Order sketch 5 Suborder Symphyta 5 Suborder Apocrita 6 Higher classification 6 Literature on Hymenoptera 7 Newsletters for Hymenopterists 8 References 10 Chapter 3. Structure John T: Huber and Michael J. Sharkey 13 General comments 13 Morphology 14 References 18 Glossary 34 Chapter 4. Use of keys Henri Goulet and William RM. Mason 60 General comments 60 Preamble to superfamily key 61 Flowchart for superfamily key 62 References 64 Chapter 5. Key to superfamilies of Hymenoptera William R.M. Mason 65 Chapter 6, Superfamilies Cephoidea, Megalodontoidea, Orussoidea, Siricoidea, ‘Tenthredinoidea, and Xyeloidea Henri Goulet 101 Key to families of Megalodontoidea 102 Key to families of Tenthredinoidea 105 References to Symphyta 112 Habitus drawings of Symphyta 116 Chapter 7. Superfamily Chrysidoidea Albert T Finnamore and Denis J Brothers 130 Key to families of Chrysidoidea 131 Key to subfamilies of Bethylidae 134 Key to subfamilies of Chrysididae 137 Key to subfamilies of Dryinidae 140 References to Chrysidoidea 146 Habitus drawings of Chrysidoidea 150 Chapter 8 Superfamily Vespoidea Denis J. Brothers and Albert T Finnamore 161 Key to families of Vespoidea 162 Key to subfamilies of Tiphiidae 178 Key to subfamilies of Sapygidae 186 Key to subfamilies of Mutillidae 188 Key to subfamilies of Pompilidae 203 Key to subfamilies of Bradynobaenidae 207 Key to subfamilies of Scoliidac 211 Key to subfamilies of Vespidae 213 Key to subfamilies of Formicidae 218 References to Vespoidea 225 ‘Habitus drawings of Vespoidea 232 Chapter 9. Chapter 10. Chapter 11. Chapter 12, Chapter 13. Chapter 14. Chapter 18, Chapter 16. Appendix 1 Index 660 Superfamily Apoidea Albert T: Finnamore and Charles D. Michener 279 Key to series of Apoidea 279 Key to families of Spheciformes 280 Key to subfamilies of Ampulicidae 291 Key to subfamilies of Sphecidae 292 Key to subfamilies of Pemphredonidae 294 Key to subfamilies of Astatidae 295 Key to subfamilies of Crabronidae 297 Key to subfamilies of Mellinidae 298 Key to subfamilies of Nyssonidae 300 Key to subfamilies of Philanthidae 304 Key to families of Apiformes 308 References to Apoidea 321 Habitus drawings of Apoidea 326 Superfamily Ichneumonoidea David B. Wahl and Michael J. Sharkey 358 Key to families of Ichneumonoidea 359 Key to subfamilies of Braconidae 363 Key to subfamilies of Holarctic and Neotropical Ichneumonidae 396 References to Ichneumonoidea 442 Habitus drawings of Ichneumonoidea 449 Superfamilies Evanioidea, Stephanoidea, Megalyroidea, and Trigonalyoidea William R.M. Mason 510 Key to families of Evanioidea $10 References to Evanioidea, Stephanoidea, Megalyroidea, and Trigonalyoidea 514 Habitus drawings of Evanioidea, Stephanoidea, Megalyroidea, and Trigonalyoidea 515 Superfamily Cynipoidea Alasdair. Ritchie 521 Key to families of Cynipoidea $22 References to Cynipoidea 529 Habitus drawings of Cynipoidea $31 ‘Superfamily Proctotrupoidea Lubomir Masner $37 Key to families of Proctotrupoidea 538 References to Proctotrupoidea 547 Habitus drawings of Proctotrupoidea 549 Superfamily Platygastroidea Lubomir Masner 558 Key to families of Platygastroidea 559 References to Platygastroidea 562 Habitus drawings of Platygastroidea 564 ‘Superfamily Ceraphronoidea Lubomir Masner 566 Key to families of Ceraphronoidea 566 References to Ceraphronoidea 567 Habitus drawings of Ceraphronoidea 568 Superfamilies Mymarommatoidea and Chalcidoidea Gary A.P Gibson S70 Key to families of Chalcidoidea (including Mymarommatidae) $73 References to Mymarommatoidea and Chalcidoidea 627 Habitus drawings of Mymarommatoidea and Chalcidoidea 635 List of habitus drawings, with generic names 656Contributors D.J. Brothers, Ph.D. Department of Zoology and Entomology University of Natal Pietermaritzburg, 3200 Republic of South Africa ACT. Finnamore, Ph.D, Provincial Museum of Alberta 12845 102nd Street Edmonton, AB Canada rN OM6 GAP. Gibson, Ph.D. Agriculture Canada Centre for Land and Biological Resources Research K.W. Neatby Building Central Experimental Farm Ottawa, ON KIA 0C6 Canada H, Goulet, Ph.D. Agriculture Canada Centre for Land and Biological Resources Research K.W.Neatby Building Central Experimental Farm Ottawa, ON KIA 0C6 Canada JT. Huber, Ph.D. Forestry Canada Se Centre for Land and Biological Resources Research, KW. Neatby Building Central Experimental Farm Ottawa, ON KIA 0C6 Canada » Masner, Ph.D. Agriculture Canada Centre for Land and Biological Resources Research KW. Neatby Building Central Experimental Farm Ottawa, ON KIA 0C6 Canada 1WR.M. Mason, Ph.D. Agriculture Canada Centre for Land and Biological Resources Research KW Neatby Building Central Experimental Farm Ottawa, ON KIA 0C6 Canada C.D. Michener, Ph.D. ‘Snow Entomological Museum, Snow Hall University of Kansas Lawrence, KS 66045 USA A.J. Ritchie, Ph.D. Gordon & Breach Science Publishers 270 8th Avenue New York, NY 10011 USA, MJ. Sharkey, Ph.D. Agriculture Canada Centre for Land and Biological Resources Research KW. Neatby Building Central Experimental Farm Ottawa, ON KIA 0C5 Canada D.B. Wahl, Ph.D. American Entomological Institute 3005 S.W. 56th Avenue Gainesville, FL 32608 USA Acknowledgments This publication was produced with support from the Centre for Land and Biological Resources Research, Agriculture Canada, Ottawa, As editors of the project, we sincerely thank the contributing authors for their prompt attention to the questions that arose during editing ofthe text and their goodwill in agreeing to changes that were needed to Produce a relatively uniform set of keys and family sketches. lp in completing the project was efficiently and cheerfully provided by both technical and professional staff. S. Righy drew many ofthe illustrations. L. Dumouchel and HE. Bisdee performed the exacting and time-consuming task of arranging the numerous ilustrations for each key couplet and preparing and rhumbering most ofthe plates. We extend special thanks to them for the quality of their work, for their ‘thoughtful relationship with each collaborator, and for their patience in dealing with what must have seemed endless changes requested by the authors and editors J, Denis and two summer students, M. MeKenzie and S. Grant, provided capable assistance in preparing many ofthe plates, and D. Tiemey prepared a few of the illustrations, J.D. Read carefully proofread the entire text, checked many of the references for accuracy of citation, prepared the computer- generated flowchart, and organized the illustrations for the glossary. J. MeWilliams typed much of the text and patiently made the countless corrections requested by authors and editors to several drafts of the ‘manuscript.The thousands of prints required for publication were done by B. Wollenschlager under the direction of B. Edwards and R. St.John of the Informations Systems and Cartography Unit, Centre for Land and Biological Resources Research, A. Lutes and J. McCarthy of Research Program Service are thanked for willingly permit- ting access to the equipment for plate production in their unit and freely giving instruction in its use. In addition to contributing to the text of three superfamilies, AT: Finnamore prepared the illustrations for the Keys to Chrysidoidea, Vespoidea, and Spheciformes, large additional task. C.D. Michener critically reviewed Chapter 3 “The various sections were reviewed by several specialists. We thank A. Austin, Waite Agricultural Research Institute, Adelaide, Australia; . Boutek, CAB. International Institute of Entomology, London, England; N.D.M. Fergusson and JS. Noyes, The Natural History Museum, London, England; L. Kimsoy, University of California, Davis, California; M. Sanborne, MacDonald College, Sainte-Anne-de-Bellevue, Quebec; D.R, Smith, United States National Museum, Washington, D.C; HLK. Townes, American Entomological Institute, Gainesville, Florida; J Whitfield, University of Missouri, St. Louis, Missouri; F. Ronquist, University of Uppsala, ‘Sweden, as well as the contributors to this project for their fast and accurate reviews of earlier drafts, ‘of the manuscript. RS. Anderson, Canadian Museum of Nature, Ottawa, Ontario, reviewed Chapters 1~5. As a coleopterist he provided a fresh perspective on the publication and made several useful suggestions for improving its clarity. We specially thank R. Wharton, Texas A & M University, College Station, for critically reviewing the entire publication, ‘The manual was initially developed for a course on identification of Hymenoptera given on three ‘occasions to a total of 45 students from across ‘Canada and several foreign countries, The students tested early versions of the keys and pointed out inaccuracies or lack of clarity in couplets and illustrations that needed improvement. We thank all of them for their suggestions In this respect, we want to thank M. Sarazin, who organized the courses, Several figures were redrawn with permission and are acknowledged below. Chapters 12 and 16, Cynipoidea and Chalcidoidea Figs. 192-196; from The Insects of Australia, Figs. 37.24A,B,D,EI; 37.25A; 37.261, respectively, published in 1970 by the Commonwealth Scientific and Industrial Research Organization, Canberra, Australia, ‘Chapters 9, Apoidea Figs. 95, 100, 101, 102, 104, 106, 113; from R.M. Bohart and A.S. Menke Specid Wasps of the World: A Generic Revision, Figs. 8, 42, 56, 69, 136, 151, 190, respectively, published 1976 by the Regents of the University of ifornia, Berkeley, California, Chapter 16, Chalcidoides figures for couplet ‘numbers 1a (right), 30a (1op), Abb (lef top), Abb (left bottom), 312, 31b, 174 (left), 29a, 27bb (right), 4e (top), 8aa, See, 27bb, 24a (left), 24a (right), 24b, aa, Tb (right); from Z. Bouek, Australasian Chalcidoidea (Hymenoptera), Figs. 67 in part), 161, 164, 215, 256, 275, 313, 372, 394, 497, 615, 919, 920, 923, 1298, respectively, published in 1988 by the CAB. International. Chapter 12, Cynipoidea figures for couplet numbers 1c, 1d, lee, ld left botwom, lec, Idd right top, lec, 1d right bottom, Ice, Ldd right middle, Ac top right, 4b, Tee, Te; irom J. Quinlan, A Revisionary Classification of the Cynipoidea (Hymenoptera) of the Ethiopian Zoogeographical Region. Aspicerinae (Figitidae) and Oberthuerellinae (Liopteridae), Figs 1, 2,3, 6,8, 33, 34, 38, 40, respectively), Bulletin of the British Museum (Natural History) 39(2): 85—133, published in 1979 by the Trustees of the British Museum (Natural History).Chapter 1 Introduction (English) John T. Huber Since the early 1980s, application of biological control methods against insect pests has had a major resurgence. Parasite Hymenoptera are among the most important agents for biological control and are extremely abundant in most terrestrial habitats. Despite their importance, their biology and taxonomy remain poorly known and their identification is difficult This publication is the result of a course on identification of Hymenoptera given three times since 1985 at the Centre for Land and Biological Resources Research (then known as the Biosystematics Research Centre). The initial impetus for the course came from an annual course sponsored since 1980 by the Systematic Entomology Laboratory, United States Department of Agriculture, Washington, D.C. A similar course offered jointly since 1989 by The Natural History Museurn, London, and by Sheffield University has also been very successful. The considerable interest in these courses indicated the need for a comprehensive identification guide to all extant families of Hymenoptera Keys to families of Hymenoptera in standard textbooks tend to repeat those published earlier, usually using the same characters and format, and ‘often permitting only the more characteristic ‘members of a family to be identified. Rare groups ‘or groups difficult to key out because of unusual attributes often were not included. The keys presented here were written by specialists who tried to include as many exceptional groups as possible to make the keys comprehensive. Therefore, although the guide was written primarily for nontaxonomists, ‘and for taxonomy students identifying Hymenoptera for the first time, it should also be useful to more experienced Hymenoptera taxonomists. Ninety-nine families and 20 superfamilies are treated (Table 1), although further rescarch on relationships will almost certainly result in changes to these numbers. At Teast one new family is known but is not included here, pending its formal description. Because this guide is intended for a wide group of users, several features were incorporated into the keys to make them easy to use while maintaining. accuracy and completeness: each distinct attribute is listed alphabetically in the alternates of the couplets; figures illustrating Key attributes are placed with the respective couplets; and terms are standardized. One chapter on structure, including an illustrated glossary of terms, and another on the use of the keys are included to help the first-time user of identification keys to Hymenoptera. The main emphasis is on family identification using the keys, which are complemented by family sketches, The sketches include a taxonomic diagnosis to supplement the keys, a summary of the biology, the size and distribution, and important (mostly taxonomic) literature references. For several families we have provided sketches and keys for the constituent subfamilies. Also included is at least one habitus drawing of a member of each family and many subfamilies. Literature to specific genera and species is usually excluded. Literature of a more general nature is cited in Chapters 2 and 3, For convenience, in this text “North America” refers to Canada and the United States only Although much effort has gone into key construction, there is always room for improvement. Largely because of differences in size of the various families, the level to which such groups are keyed is variable. Although all extant families are keyed, keys to subfamilies may or may not be included, For example, we have presented keys to the subfamilies of Braconidae and Ichneumonidae because of the large size of each of these two families. When included, not all the subfamily keys give complete world coverage, but North America, at least, is covered, Using the Keys will likely lead to discovering exceptions to them. It is hoped that suggestions for improving the keys will be made to the editors or to the authors of the appropriate section. Eleven authors contributed to the publication. Because each specialist had his own style of key ‘writing, some editorial changes had to be made to provide a reasonably uniform work. This included compromises and choices in use of morphological terms where several equally meaningful or useful terms could be used, The editors tried to maintain a fairly simple and uniform set of terms, in the keys at east, without losing accuracy or clarity If this guide results in greater interest in taxonomy of Hymenoptera and encourages biologists, teachers, students, biological control officers, and other nontaxonomists to identify their ‘own specimens of Hymenoptera to family, then the goal of the authors will have been fulfilledIntroduction (frangais) Depuis Je début de la demnigre décennic, application des techniques de lutte biologique contre les insectes nuisibles connait un grand essor. Les hyménoptéres parasites comptent parmi les agents les plus importants de cette lutte biologique. En dépit de leur importance, leur biologic et leur ie demeurent peu connues et leur tion est difficile ‘une demande accrue de cours d'introduction aux hyménoptéres et en absence de publications traitant des familles d'hyménoptéres ‘au niveau mondial, plusieurs hyménoptéristes Conjointement avec le “Systematic Entomology Laboratory” & Washington ont créé un cours. portant sur les familles d’hyménoptéres parasites. La popularité et le suceés de ce cours encourages lune orientation semblable au les Centre de recherches sur les terres et les ressources biologiques au Canada od plusieurs hyménopteristes ont coopéré a la préparation d'un cours annuel sur toutes les familles d hyménopteres. ‘Apres avoir présenté ce cours & trois reprises, il devint clair qu'un guide détaillé d'identification des familles ’hyménoptéres serait un outil indispensable & tout étudiant. Aprés plusieurs années d'efforts et avec la collaboration de plusieurs scientifiques de ce Centre et d'ailleurs, ce guide voit finalement le jour. Les quelques guides sur les familles dhyménoptéres se ressemblent dans la construction des clefs d'identification, et ne permettent la reconnaissance que des familles majeures. Certains ‘groupes demeurent difficiles a caractériser & cause G'attributs inhabituels, ou sont simplement ignorés car ils semblent trop peu communs (particuligrement les groupes basés sur de petites especes qui sont représentées par peu de spécimens dans les collections malgré leur grande abondance cen nature). Ce guide-ci se veut Pun des plus ccomplets car les experts qui Pont rédigé y ont inclus tous les groupes exceptionnels conus. Ainsi, 99 familles et de 20 super-familles y sont traitée (Tableau 1), ces nombres étant appelés a changer & Pavenir avec la poursuite de la recherche sur la classification des hyménoptres. Ce guide s‘adresse A tout biologiste ayant un intérét dans la lutte biologique et & tout débutant désireux de se familiariser avec les hyménopt@res, De plus, il sera particuligrement utile a tous les étudiants de la taxonomic, de méme qu’aux spécialistes de la systématique des hyménoptéres. ‘Comme ce travail est desting aussi bien aux débutants qu’aux experts, nous avons simplifié 1a présentation et les illustrations des clef identification sans pour autant en réduire Vintégralité et la précision. Premigrement, nous naré en paragraphes et dénoté alphabétiquement chaque attribut particulier pour chacune des deux composantes d'un couplet. ‘Deuxiemement, les abondantes figures illustrant divers attributs apparaissent simultanément avec les couplets correspondants. Troisiémement, seul un vocabulaire uniforme et clair est employé. Enfin, pour aider les personnes peu familiéres avec les hyménopteres, nous avons préparé des chapitres sur la morphologic ct sur utilisation des clets identification ainsi qu’un glossaire illustré. Bien que lemphase de ce guide soit mise sur {es clefs d'identification, les familles qui y sont identifiges sont caractérisées davantage dans de brefs exposés. On y présentent des diagnoses ‘morphologiques qui completent les clefs, un sommaire sur la biologie, la diversité et la distribution de chaque famille et dans bien des cas, de leurs sous-familles, des références pertinentes et ‘au moins une illustration complete d'une espece typique & chaque famille. Les références de nature spécifique et générique y sont exclues, cependant sous les chapitres 2 et 3 on mentionne plusicurs travaux d'ordre général. Par souci de commodité, « Amérique du Nord » dans cet ouvrage correspond au Canada et aux Etats-Unis seulement. Malgré tous les efforts apportés a une definition claire de toutes les families, certaines demeurent néanmoins obscures. De telles ifférences nous ont contraints de détinir les familles & des niveaux différents. Pour certaines grandes familles (e.g,, Ichneumonidae et Braconidac), il est beaucoup plus utile et significatif