Catalog of Hymenoptera in America North of Mexico CarlF. W. Muesebeck This catalog is dedicated to our cherished colleague with affectionate regard for his kindliness and with admiration for his distinguished scholarly contributions to our knowledge of North American Hymenoptera for more than half a century. Catalog of Hymenoptera in America North of Mexico Prepared cooperatively by specialists on the various groups of Hymenoptera under the direction of Karl V. Krombein and Paul D. Hurd, Jr. Smithsonian Institution and David R. Smith and B. D. Burks Systematic Entomology Laboratory Insect Identification and Beneficial Insect Introduction Institute Science and Education Administration, United States Department of Agriculture VOLUME 1 Symphyta and Apocrita (Parasitica) SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1979 Library of Congress Cataloging in Publicatimt Data Main entry under title: Catalog of Hymenoptera in American north of Mexico. "An outgrowth of . . . Hymenoptera of America north of Mexico, synoptic catalog (1951) in- cluding the first and second supplements (1958, 1967)" Includes index. CONTENTS: v.l. Symphytaand Apocrita (Parasitica). — v. 2. Apocrita (Aculeata) 1. Hymenoptera — North America. 2. Insects — North America. I. Krombein, Karl V. QL567.1.A1C37 595.r9'097 78-606008 Contents Page Introduction by Karl V. Krombein, Paul D. Hurd, and David R. Smith v Hymenoptera by Karl V. Krombein and Paul D. Hurd, Jr 1 Symphyta by David R. Smith 3 Megalodontoidea 7 Xyelidae 7 Pamphiliidae 10 Tenthredinoidea 19 Pergidae 19 Argidae 20 Cimbicidae 26 Diprionidae 29 Tenthredinidae 39 Siricoidea 125 Syntexidae 125 Siricidae 125 Xiphydriidae 130 Orussidae 131 Cephoidea 133 Cephidae 133 Apocrita by Karl V. Krombein : 139 Parasitica by Paul M. Marsh and Robert W. Carlson 141 Ichneumonoidea by Paul M. Marsh and Robert W. Carlson 143 Braconidae by Paul M. Marsh 144 Aphidiidae by Paul M. Marsh 295 Hybrizontidae by Paul M. Marsh 313 Ichneumonidae by Robert W. Carlson 315 Stephanidae by Robert W. Carlson 740 Chalcidoidea by Gordon Gordh 743 Torymidae (except Agaoninae) by E. E. Grissell 748 Pteromalidae by B. D. Burks 768 Eurytomidae by B. D. Burks 835 Chalcididae by B. D. Burks 860 Leucospididae by B. D. Burks 874 Eucharitidae by B. D. Burks 875 Eupelmidae by B. D. Burks 878 Encyrtidae by Gordon Gordh 890 Eulophidae by B. D. Burks 967 Mymaridae by B. D. Burks 1022 Trichogrammatidae by B. D. Burks 1033 Unplaced Taxa in Chalcidoidea by B. D. Burks 1042 Cynipoidea by B. D. Burks 1045 Ibaliidae 1045 Liopteridae 1046 Figitidae 1047 Eucoilidae 1052 Alloxystidae 1058 Cynipidae 1060 Evanioidea by Robert W. Carlson 1109 Evaniidae 1109 Aulacidae 1111 Gasteruptiidae 1115 Pelecinoidea by Carl F. W. Muesebeck 1119 Pelecinidae 1119 Proctotrupoidea by Carl F. W. Muesebeck 1121 Vanhorniidae 1122 Roproniidae 1122 Heloridae 1122 Proctotrupidae 1123 Diapriidae 1127 Scelionidae 1150 Platygastridae 1171 Ceraphronoidea by Carl F. W. Muesebeck 1187 Ceraphronidae 1187 Megaspilidae 1191 Trigonaloidea by Robert W. Carlson 1197 Trigonalidae 1197 Aculeata by Karl V. Krombein 1199 Bethyloidea by Karl V. Krombein 1203 Bethylidae 1203 Sclerogibbidae 1219 Chrysididae 1220 Dryinidae 1240 Embolemidae 1251 Scolioidea by Karl V. Krombein 1253 Tiphiidae 1253 Sierolomorphidae 1276 Mutillidae 1276 Scoliidae 1314 Sapygidae 1319 Formicoidea by David R. Smith 1323 Formicidae 1323 Vespoidea by Karl V. Krombein 1469 Masaridae 1469 Eumenidae 1472 Vespidae 1510 VI Pompiloidea by Karl V. Krombein 1523 Pompilidae 1523 Rhopalosomatidae 1570 Sphecoidea by Karl V. Krombein 1573 Ampulicidae 1574 Sphecidae 1575 Pemphredonidae 1595 Larridae 1617 Crabronidae 1650 Mellinidae 1683 Nyssonidae 1684 Philanthidae 1720 Apoidea by Paul D. Hurd, Jr 1741 Colletidae 1748 Oxaeidae 1770 Andrenidae 1772 Halicitidae 1932 Melittidae 1978 Megachilidae 1981 Anthophoridae 2081 Apidae 2188 Institutional Affiliation of Contributing Authors Smithsonian Institution; Paul D. Hurd, Jr., Karl V. Krombein, Carl F. Muesebeck U.S. Department of Agriculture; B. D. Burks, Robert W. Carlson, E. E. Grissell, Paul M. Marsh, David R. Smith University of California, Riverside: Gordon Gordh vu Introduction By Karl V. Krombein, Paul D. Hurd, Jr., and David R. Smith This catalog, which is an outgrowth of the highly successful Hymenoptera of America North of Mexico Synoptic Catalog (1951), including the first and second supplements (1958, 1967), provides simultaneously, by use of computer technology (Krombein, Mello and Crockett, 1974. Ent. Soc. Amer., Ann. 20 :24-29), a printed version as well as a computer-queriable data base of the basic systematic, biological, and morphological in- formation on the order Hymenoptera as it occurs in America north of Mexico. While the format of the present catalog closely parallels that of the original catalog, every effort has been made to increase the information con- tent to fulfill better the needs of the biological community. This has been accomplished chief- ly by the introduction of textbook prose at most higher category levels, by the presenta- tion of explanatory or descriptive comments as appropriate, and by the inclusion of more complete listings of citations to taxonomic, biological, and morphological literature at all hierarchical levels together with parentheti- cal annotations concerning the content for many of these citations. Similarly, the data about hosts, parasites, prey, predators, and pollen sources are covered more fully than in the original catalog. Since these data have been reported in various ways and under dif- fering names in the primary literature, an attempt has been made to organize this in- formation and present it, usually, alphabeti- cally even though sometimes such data have been recorded in the literature with scien- tific or vernacular names or both. Although many of these names have been checked for accuracy, no consistent attempt has been made to verify that the names agree with current usage or status. While those authors of zoological names are usually abbreviated in the citation of hosts, parasites, and the like, the authors of the scientific botanical names are not included. Elsewhere in the catalog, the last name of an author is cited in full, but without initials. Since the tech- nology employed in the preparation of this catalog precluded the use of diacritical marks, these have been omitted and consequently, as in the case of the umlaut, a variant spell- ing has been substituted as appropriate. The International Code of Zoological No- menclature (1961) including the intent and preamble of the Code and of any pertinent opinions usually has been followed. Thus a name in current usage as deterimned by the code is employed, but mention is made, as appropriate, that an application is pending before the International Commission of Zoological Nomenclature. An attempt has been made to record all the recent taxa, with their synonyms, described Vlll from, or known to occur in, the political di- visions of the conterminous United States, Canada, Alaska, and Greenland. Insofar as known, all species introduced from other countries and liberated in America north of Mexico for biological control purposes have been listed. Some of these have never been recovered and, apparently, were unsuccessful in establishing themselves. For each such species a statement has been included, follow- ing the distribution, that the species was liberated but did not become established. As in the original catalog, the arrangement is systematic for species-groups and higher categories insofar as our present knowledge and the limitations of a linear arrangement permit. The generic and subgeneric concepts represented in this arrangement are based upon what are believed to be the correct type- species. In each instance the type-species is cited together with the authority for the selection. Where designations of type-species have been found to be invalid under the Inter- national Code of Zoological Nomenclature, new type-species designations, believed to be valid, are given in the catalog. Generic syn- onymy is included under the generic head- ings except where subgenera are recognized, in which cases such synonymy is given under the subgeneric names, and references to re- visional or other papers are listed under the appropriate higher category. The arrangement of species within genera, subgenera, and species groups is alphabetical. Where subspecies are recognized, the sub- specific names are placed in alphabetical or- der under the species to which they belong; and varieties are listed under the particular species or subspecies in which they were described. In each case the specific, subspeci- fic, or infrasubspecific, name is followed by an indication of the known distribution, and by brief statements, as appropriate, of pre- ferred habitats or the like, hosts, parasites, prey, predators, or pollen sources. Much of this information on synonymy, distribution, ecology, hosts, parasites, prey, and so forth has not been published previously. The type localities are usually recorded for those forms that are known only from the localities where the type specimens were obtained. Otherwise the distribution is usually shown by states and provinces, or by other means such as life zones. Since unquestionably a catalog is indispen- sable in the support of systematic and other biological research, no effort has been spared toward making this catalog as useful as pos- sible to all of the scientific community inter- ested in these fascinating insects. LITERATURE COVERAGE All authors have attempted to include all pertinent refei'ences to synonyms, revisions, taxonomy, biology, and morphology begin- ning with 1758, the publication date of the 10th edition of "Systema Naturae" by Lin- naeus. The cut-off dates vary for the several sections of the catalog and are as follows: Symphyta through 1974 ; Ichneumonoidea through 1976; Chalcidoidea — Torymidae (except Agaoninae) and Encyrtidae through 1976; other fam- ilies and Agaoninae through 1972 ; Cynipoidea through 1972 ; Evanioidea through 1976 ; Pelecinoidea, Proctotrupoidea, and Ceraphro- noidea through 1972; Trigonaloidea through 1976; Bethyloidea and Scolioidea through 1975 ; Formicoidea through mid-1975 ; Vespoidea, Pompiloidea, and. Sphecoidea through 1975; and Apoidea through 1976. All authors have included some references subsequent to the dates listed above. IX Abbreviations and Symbols LITERATURE CITATIONS The source for journal abbreviations is Whit- lock, C, 1939, Abbreviations used in the De- partment of Agriculture for titles of 'publica- tions. United States Department of Agricul- ture Miscellaneous Publication No. 337, 278 pages. Abbreviations for other titles and journals not found in Whitlock essentially follow the same format and abbreviations that she recommends. Book titles are usually shortened to omit irrelevant adjectives and include abbreviations, e.g. : Wheeler and Wheeler, 1963. The Ants of North Dakota, p. — , is cited as Wheeler and Wheeler, 1963. Ants of N. Dak., p. — . Certain lengthy non- serial titles are also abbreviated, e.g. : Say, 1824. /;; Keating, Narr. Long's 2nd Exped., V. 2 (App.), p. — , rather than Say, 1824. In Keating, Narrative of an Expedition to the Source of St. Peter's River, Lake Winnepeek . . ., etc. The titles are intended to be uni- form throughout the catalog, but, in a work of this magnitude, there will naturally be some deviations. The abbreviations should be adequate to find the cited publication. SYMBOLS AND ABBREVIATIONS Certain symbols and abbreviations are fre- quently used in this catalog. Though there may be slight variations in some, they are generally as follows : (!) — lapsus or misspelling of a scientific name. " 9 " = $ or " 5 " = 9 — incorrect sex deter- mination. 9 {$ misdet.) or i (9 misdet.) — only one of the sexes described belongs to the spe- cies cataloged. 9 — female. <5 — male. 5 — worker. 'H — soldier. ab. — aberration. app. — appendix. cent. — central. changed status — used after a species-group name to indicate a rank different from that previously accorded to it; not necessarily the same as new status. desig. — designated; e.g., in type-species de- signation, "Desig. by Rohwer, 1911." e., east. — east, eastern. emend.- — emendation. fasc. — fascicle. fig., figs. — figure, figures. h.— heft. n. comb. — new combination ; used after a species-group name to indicate a new ge- neric assignment. n. name — new name; used after a genus- group or species-group name to indicate a substitute name for a homonym. N. name — New name ; used after a biblio- graphic citation to indicate a previously proposed name. Nom. nud. — Nomen nudum. n. s. — new series. n. status — new status ; used where a taxon is here accorded a rank different from that which it had previously. N. syn. — New synonymy; used to indicate a synonym newly proposed in this catalog. n., no., north. — north, northern. n.e., northeast. — northeast, northeastern. n.w., northwest. — northwest, northwestern. orig. desig. — original designation; used to indicate type-species designation. p., pp. — page, pages. pi., pis. — plate, plates. preocc. — preoccupied ; used after a genus- group or species-group name to indicate a homonym. pt. — part. revised status — revised status; used to de- note a taxon that has been removed from synonymy. ser. — series. s., so., south. — south, southern. s.e., southeast. — southeast, southeastern. S.W., southwest. — southwest, southwestern. sp., spp. — species. ssp., sspp. — subspecies. subg. — subgenus. transcont. — transcontinental. v., vol. — volume. var. — variety. w., vi'est. — west, western. FAUNAL ZONES Bor. Boreal Canad. Canadian Huds. Hudsonian Transit. Transition Austr. Austral U. Austr. Upper Austral L. Austr. Lower Austral Alleghan. Alleghanian Austrorip. Austroriparian Carol. Carolinian Sonor. Sonoran U. Sonor. Upper Sonoran L. Sonor. Lower Sonoran GEOGRAPHICAL NAMES Abbreviation Political Unit Ala. Alabama Alaska Alaska Alta. Alberta Ariz. Arizona Ark. Arkansas B.C. British Columbia Calif. California Canada Canada Colo. Colorado Conn. Connecticut Del. Delaware D.C. District of Columbia Fla. Florida Ga. Georgia Greenland Greenland Idaho Idaho 111. Illinois Ind. Indiana Iowa Iowa Kans. Kansas Ky. Kentucky La. Louisiana Labrador Labrador, Newfoundland (Labrador) Maine Maine Man. Manitoba Mass. Massachusetts Md. Maryland Mexico Mexico Mich. Michigan Minn. Minnesota Miss. Mississippi Mo. Missouri Mont. Montana N. B. New Brunswick N. C. North Carolina N. Dak. North Dakota Nebr. Nebraska Nev. Nevada Newfoundland Newfoundland, Newfoundland (insular) N. H. New Hampshire N.J. New Jersey N. Mex. New Mexico N. S. Nova Scotia N. W. T. Northwest Territories N. Y. New York Ohio Ohio Okla. Oklahoma Ont. Ontario Oreg. Oregon Pa. Pennsylvania P. E. I. Prince Edward Island Que. Quebec R. I. Rhode Island Sask. Saskatchewan S. C. South Carolina S. Dak. South Dakota Tenn. Tennessee Tex. Texas U. S. United States Utah Utah Vt. Vermont Va. Virginia Wash. Washington (state) W. Va. West Virginia Wis. Wisconsin Wyo. Wyoming Yukon Yukon Territory XI TAXONOMIC AND NOMENCLATURAL CHANGES The catalog contains one undiagnosed new genus also in the Ichneumonidae, Woldsted- species in the Ichneumonidae, Pterocormus tins Cai'lson, type-species Bass^is bigidtatus clasma Carlson, p. 521, proposed for the Gravenhorst, p. 719, proposed for Syrphoc- taxon misidentified as Ichneumon canadensis tonus Foerster sensu Dasch (1964). Cresson by Heinrich (1961). The following new names are proposed to The catalog contains one undiagnosed new replace preoccupied names : Tenthredinidae Pachynematus gamus Smith for Pachynematus graminis Marlatt (1896) p. 58 Nematus attus Smith for Amauronematus dyari Marlatt (1896) p. 68 Amauronematus peralus Smith for Nematus pectoralis Cresson (1880) p. 80 Ichneumonidae Oedomopsis davisi Carlson for Trophon ? iiasutns Cresson (1868) p. 366 Gelis cushmani Carlson for Hemiteles apantelis Cushman (1927) p. 405 Oresbiiis shimiaginensis Carlson for Stiboscopus ferrugineus Ashmead (1902) ... .p. 438 Pterocormus dionymus Carlson for Ichneumon anonymus Heinrich (1961) p. 522 Casinaria affinisima Carlson for Casinaria affinis Walley (1947) p. 635 Pteromalidae Mesopolobiis fuscipedes Burks for Platyterma fuscipes Ashmead (1896) p. 816 Eurytomidae Harmolita ovatella Burks for Harmolita ovata Phillips and Emery (1919) p. 840 Eulophidae Syntomosphyrum orgyiazele Burks for Tetrastichomyia orgyiae Girault (1916) . .p. 1005 Diapriidae Trichopria kiefferi Muesebeck for Diapria montana Kieffer (1906) p.ll47 Scelionidae Trimorus contractus Muesebeck for Gryo)i flavipes Ashmead (1893) p. 1162 Ceraphronidae Aphanogmus harringtoni Muesebeck for Aphanogmus salicicola Harrington (1899) p.ll90 Megaspilidae Dendrocerus obscurellus Muesebeck for Atritomus califortiicus Kieffer (1906) ..p.ll94 Masaridae Euparagia richardsi Bohart for Psiloglossa simplicipes Rohwer (1909) p. 1470 Philanthidae Cerceris bolingeriana Krombein for Cerceris bolingeri ScuUen (1972) p. 1730 Anthophoridae Triepeolus mitchelli Hurd for Triepeolus sublunatus Mitchell (1962) p.2094 There are a number of other nomenclatural valid genera and species for each family and and taxonomic changes. These are considered higher category. These nomenclatural and of lesser bibliographic importance than the taxonomic changes are as follows : new names, so tabulations of them are de- A number of generic transfers are made, ferred to Volume 3 which will also contain They are usually cited in the text as xanti- the indexes and a table of the number of anum (Saussure), n. comb. The authority xu responsible for the transfer is the author of that section unless the name of another specialist is included. There are also a number of instances where a taxon formerly considered to be a species is treated here as a subspecies of another taxon, or where a taxon formerly considered to be a subspecies is now raised to specific rank. The authority responsible for the change is the author of that section unless the name of another specialist is included. These are usu- ally cited in the text as clavatum johannis (Richards), n. status or alba Rohwer, n. status. The words — changed status — occasionally follow the author of a species-group name. This indicates that the taxon has a rank different from that accorded it elsewhere. It is not the same as new status for it reflects a change which has already been published. In a few taxa, the words — revised status — follow the author of a species-group name. This denotes a taxon which has been removed from synonymy. There are a few new synonyms at the genus-group level and numerous new syn- onyms at the species-group level. These are indicated by the abbreviation N. syn. follow- ing the bibliographic citation of the new syn- onym. As noted above, the synonymy is to be attributed to the author of the section unless the name of another specialist appears in parentheses following the abbreviation N. syn. Volume 3 It is intended that Volume 3 will contain separate indexes to the taxa of Hymenoptera, and to their hosts, parasites, prey, predators, and pollen and nectar sources. Preparation of the indexes has already begun, and we antici- pate that the tapes for Linotron production will be sent to the Government Printing Office during 1978. We will also include in Volume 3 a tabulation of the number of valid genera and species for each family and higher cate- gory, and lists of the nomenclatural and taxonomic changes other than the new names which are listed above. Acknowledgments Preparation of the catalog and funding for its publication have had the enthusiastic support of Porter M. Kier, Director, National Museum of Natural History (NMNH), Smithsonian Institution (SI) and of Lloyd V. Knutson, Chairman, Insect Identification and Beneficial Insect Introduction Institute, U.S. Depart- ment of Agriculture. The catalog in its printed form could not have been achieved without their help, and we are most grateful that their assistance was available whenever we required it. We are indebted to a host of cooperating hymenopterists for generous assistance which xui has greatly enhanced the content and quality of the catalog. The aid furnished has involved such diverse activities as reviewing prelimi- nary drafts of various sections, and providing new information on taxonomy, synonymy, distribution, and biology. In the Symphyta, H. E. Milliron, formerly of the Biosystematics Research Institute (BRI), Agriculture Canada, Ottawa, re- viewed parts of the manuscript and provided information, as did H. R. Wong, Northern Forest Research Centre, Edmonton, Alberta, for Piistiphoia Latr. H. Greenbaum, Uni- versity of Florida, Gainesville, furnished data on Florida sawflies. The accuracy of host names in the Para- sitica was checked by the following special- ists : Smithsonian Institution — J. F. G. Clarke, D. R. Davis, W. D. Duckworth, T. L. Erwin, W. D. Field, and R. C. Froeschner; Systematic Entomology Laboratory (SEL), U. S. Department of Agriculture — D. C. Ferguson, R. J. Gagne, R. D. Gordon, A. B. Gurney, J. L. Herring, R. W. Hodges, J. M. Kingsolver, J. P. Kramer, A. S. Menke, D. M. Miller, L. M. Russell, C. W. Sabrosky, D. R. Smith, T. J. Spilman, M. B. Stoetzel, E. L. Todd, R. E. Warner, and R. E. White. D. M. Weisman (SEL) identified the remains of some lepidopterous larvae which served as hosts. Specialists who were helpful in the Ichneu- monoidea were : L. E. Caltagirone, University of California, Albany, and C. C. Loan (BRI), who reviewed parts of the manuscript on Braconidae ; C. van Achterberg, Waarder, The Netherlands, W. R. M. Mason (BRI), R. D. Shenefelt, University of Wisconsin, Madison, and R. Wharton, University of California, Berkeley, who provided advice and informa- tion on Braconidae; H. K. and M. C. Townes, American Entomological Institute, Ann Ar- bor, Michigan, who provided information and advice on Ichneumonidae; and P. M. Marsh (SEL) , who proofed the first-phase computer printouts for Mesochorinae, Diplazontinae, Oxytorinae and Orthocentrinae, and the third- phase edit for Ichneumoninae. Z. Boucek, Commonwealth Institute of En- tomology, London, and M. Graham, Oxford University, provided much information on Chalcidoidea, and D. P. Annecke, Plant Pro- tection Research Institute, Pretoria, South Africa, advised on the placement of some species assigned erroneously to Aphycus Mayr. A special debt of gratitude is due C. F. W. Muesebeck who painstakingly proofed all edit phases of the computer printouts for Encyrtidae and Torymidae and all but the first-phase edits of all other families of Chalcidoidea. D. B. Krombein provided welcome assist- ance by helping to proof the manuscripts and printouts for all superfamilies of aculeate wasps. F. D. Parker, Bee Biology and Syste- matics Laboratory, Utah State University, Logan, furnished biological data for a number of Utah wasps. We are grateful to R. M. Bohart, Univer- sity of California, Davis, for reviewing the manuscript on Chrysididae and for consider- able other assistance which included informa- tion on new synonyms and distribution in the Elampinae and Chrysididinae, and the assign- ment of taxa to species groups in Chrysis L. H. E. Evans, Colorado State University, Fort Collins, reviewed the section on Bethyli- dae. In the Scolioidea, the late J. C. Bradley, Cornell University, Ithaca, New York, and J. G. Betrem, Deventer, The Netherlands, reviewed the manuscript on Scoliidae, and H. W. Allen, Moorestown, New Jersey, that on Tiphiinae. W. E. Ferguson, San Jose State University, California, and C. E. Mickel, Uni- versity of Arizona, Tucson, furnished infor- mation on synonymy and taxonomy of some Mutillidae. R. R. Snelling, Los Angeles County Museum, California, and M. R. Smith, Arlington, Virginia, reviewed the section on Formicoi- dea. J. F. Watkins II, Baylor University, Waco, Texas, reviewed the section on Dory- linae, and A. C. Cole, University of Tennessee, Knoxville, that on Pogonomyrmex Mayr. A. Francoeur, University of Quebec, Chicou- timi, provided information on the fusca group of Formica L. 0. W. Richards, British Museum (Natural History) , London, and R. M. Bohart reviewed the entire manuscript for Vespoidea. J. van der Vecht, Putten, The Netherlands, recom- mended the systematic sequence adopted in the Eumenidae and reviewed the manuscript. XIV M. J. West-Eberhard, Universidad del Valle, Call, Colombia, and R. R. Snelling reviewed the manuscript on Vespidae and supplied data on taxonomy and biology. J. E. Gillaspy, Texas A & I University, Kingsville, provided information on PoHsfes Latr. H. E. Evans reviewed the manuscript on Pompilidae, and F. E. Kurczewski, University of Syracuse, N. Y., contributed some prey records. R. M. Bohart and A. S. Menke generously made available a copy of their manuscript, "Sphecid Wasps of the World," which was most helpful in assembling the section on Sphecoidea ; Menke, in addition, reviewed the catalog manuscript for this superfamily. Other specialists contributed data on the groups mentioned after their names : R. E. Coville, University of California, Berkeley (taxonomy and distribution of Trypoxyloni- nae) ; J. E. Gillaspy (Bembicinae) ; F. E. Kurczewski (biology of Larridae and Cra- bronidae) ; R. C. Miller, Cornell University (taxonomy, distribution, and biology of Crabronidae) ; W. J. Pulawski, Wroclaw, Poland (taxonomy and distribution of Tachij- sphe.r Kohl) ; and D. Vincent. University of Maryland, College Park (taxonomy and dis- tribution of Passaloeciis Shuck.). E. G. Linsley, University of California, Berkeley, reviewed the entire manuscript on Apoidea and offered valuable suggestions. C. D. Michener, University of Kansas, Law- rence, discussed with the author of that sec- tion the biology and systematics of bees; many of his suggestions have been incorpo- rated in the classification adopted. The fol- lowing specialists have cooperated by con- tributing data on the groups specified after their names: G. E. Bohart, Bee Biology and Systematics Laboratory, Utah State Univer- sity, Logan (taxonomy and biology of Apoidea) ; W. E. LaBerge, Illinois Natural History Survey, Urbana (taxonomy of An- drenidae and Anthophoridae) ; U. N. Lanham, University of Colorado, Boulder (taxonomy of Apoidea) ; M. A. Lieftinck, Rhenen, The Netherlands (taxonomy of Anthophoridae) ; A. L0ken, University of Bergen, Norway (taxonomy of Apidae) ; T. B. Mitchell, North Carolina State University, Raleigh (taxon- omy of Apoidea) ; J. S. Moure, Universidade Federal do Parana, Curitiba, Brazil (taxon- omy of Apoidea) ; F. D. Parker (taxonomy and biology of Apoidea) ; J. G. Rozen, Jr., American Museum of Natural History, New York, New York (taxonomy and biology of Apoidea) ; R. R. Snelling (taxonomy of Hylaeus F.) ; R. W. Thorp, University of California, Davis (taxonomy and biology of Apidae) ; P. H. Timberlake, University of California, Riverside (taxonomy of Apoidea) ; and T. J. Zavortink, University of San Francisco, California (taxonomy of Antho- phoridae) . C. W. Sabrosky (SEL) has been helpful to all of us in the discussion of abstruse nomen- clatural problems. J. F. Gates Clarke (SI) participated patiently in philosophical dis- cussions of a wide variety of subjects per- taining to the catalog. G. C. Steyskal (SEL) was the resource person for the grammar of scientific names and other linguistic matters. Computerization of the catalog and its pro- duction by the computer-driven Linotron re- quired the highly skilled technical expertise of specialists in computer storage and appli- cations. James F. Hello, formerly Chief of the Data Processing (ADP) Program at NMNH, carefully analyzed the 1951 Hymenoptera Catalog and developed the data analysis matrix which governed entry of information into the computer. R. Creighton, Manager, Information Retrieval and Indexing Division, Office of Computer Services (OCS), SI, de- vised the programs for editing, arranging, querying, and displaying data from the man- uscripts. J. J. Crockett, Manager, Software Systems and Program Maintenance (OCS), developed the program for conversion of the computerized data to special magnetic tapes capable of driving the Photo Typesetting Unit, the Mergenthaler Linotron 1010, in the Government Printing Office. T. G. Gautier, Chief (ADP), and D. Bridge, Operations Manager (ADP), maintained daily collabora- tion with the Editorial Board in assignment of clerical assistance, and production of the edit phases, merge files, and SELGPO print- outs. Finally, we are most grateful for the care- ful, accurate typing of the manuscripts for computer entry by the corps of dedicated clerk typists. R. M. Garlick served with the XV program as principal typist from its inception during the six years of manuscript produc- until his reassignment in mid-1976 as a com- tion and computer entry were : L. E. Back, puter technician; he also trained the other L. M. Bybell, R. Cloyed, L. E. Hatton, M. assigned typists. P. R. Brown is currently the Monahan, L. G. Oliver, J. Peabody, P. A. only typist assigned full time. Other typists Sunkel, and M. F. Ward. who worked for varying lengths of time XVI Order HYMENOPTERA By Karl V. Krombein and Paul D. Hurd, Jr. The Hymenoptera is one of the largest orders of insects with more than 20,000 species in America north of Mexico, most of which are beneficial and many of which are of considerable economic importance to agriculture and forestry either as parasites or predators of pests or as pollinators of more than 100 commercially grown crops. Among the relatively few injurious Hymenoptera are the sawflies, some of which are serious defoliators or stem-borers of trees or crops. Taxonomy: Fabricius, 1804. Systema Piezatorum, 439 pp., 1 p. of errata, 3 pp. of index. — Panzer, 1806. Kritische Revision der Insektenfauna Deutschlands, v. 2, Hymenoptera, 271 pp. 2 pis. Cited in text as Krit. Rev. Insektenf. Deutschlands. —Andre, 1879-1913, Species des Hymenopteres d'Europe et d'Algerie, vols. 1-11. Cited in text as Spec. Hym. Eur. Alg. —Cameron, 1883-1900. In Godman and Salvin, Biologia Centrali-Americana. Hymenoptera; vol. 1, pp. 1-487, 20 pis., 1883-1900; vol. 2, pp. 413, 13 pis., 1888-1900. Cited in text as Cameron, Biol. Cent.-Amer., Hym. — Cresson, 1887. Amer. Ent. Soc, Trans., Sup. Vol., pp. 1-350 (key to No. Amer. families, genera and catalog of spp.). — Dalla Torre, 1892-1902. Cat. Hym., 10 vols, (world spp.). — Schulz, 1906. Spolia Hymenopterologica, 355 pp., 11 figs., 1 pi. Cited in text as Spolia Hym. — Viereck, et ai, 1917 (1916). Conn. State Geol. and Nat. Hist. Survey, Bui. 22: 1-824, 10 pis., 15 text figs, (keys to Conn. spp.). — Boerner, 1919. Biol. Zentbl. 39: 145-186, 6 figs, (phylogeny). — Schroeder, 1925. Handb. d. Ent. 3: 712-825, figs. 593-705. — Tillyard, 1926. The Insects of Australia and New Zealand, pp. 252-307, 53 figs., 2 pis. — Schmiedeknecht, 1930. Die Hymenopteren Nord- und Mitteleuropas, 2nd ed., 1062 pp., 127 figs. Cited in text as Hym. Mitteleuropas. — Brues and Melander, 1932. Mus. Compar. Zool., Bui. 73: 471-526, figs. 887-971 (keys to families, adults, and larvae). — Handlirsch and Meixner, 1933. In Krumbach, Handb. d. Zool., v. 4, Insecta 2, pp. 895-1036. — Comstock, 1940. An Introduction to Entomology, pp. 884-1007. — Essig, 1942. College Entomology, pp. 619-727. — Imms, 1948. A General Textbook of Entomology, ed. 7, pp. 544-615. — Lanham, 1951. Ent. Soc. Amer., Ann. 44: 614-628, 27 figs, (phylogeny based on wing venation). — Muesebeck, Krombein, Townes et a/., 1951. U. S. Dept. Agr., Agr. Monog. 2, 1,420 pp., 1 map (catalog of species north of Mexico). — Michener, 1953. Kans. Univ. Sci. Bui. 35: 993-995 (larvae, key to certain families). — Richards, 1956. Hymenoptera, Introduction and Keys to Families. Handbooks Ident. Brit. Ins., Roy. Ent. Soc, London 6, pt. 1, 94 pp., 197 text figs., 11 pis. — Boucek et ai, 1957. In Klic Zvireny CSR (keys to the fauna of Czechoslovakia), ed. by Kratochvil, pp. 35-406, 1108 figs. —Bradley, 1958 (1956). Tenth Internatl. Congr. Ent. Proc. 1: 265-269 (phylogeny). —Krombein et al., 1958. U. S. Dept. Agr., Agr. Monog. 2, Sup. 1, 305 pp. Hymenoptera in America North of Mexico (supplement to catalog of species north of Mexico). — Berland, 1958. Atlas des Hymenopteres de France. I, Tenthredes, Parasites, Porte-aiguillon (Bethyloides, pars), 155 pp., 14 pis. II, Porte-aiguillon (Bethyloides (fin), Scolioides, Formicoides, Pompiloides, Vespoides, Sphecoides, Apoides), 184 pp., 14 pis. — Oeser, 1961. Zool. Mus. Berlin, Mitt. 37: 75-85, figs. 106-107 (phylogeny based on ovipositor). — Krombein, Burks et al., 1967. U. S. Dept. Agr., Agr. Monog. 2, Sup. 2, 584 pp. (supplement to catalog of species north of Mexico). — Malyshev, 1968. Genesis of Hymenoptera and phases of their evolution 319 pp. (English translation by Richards and Uvarov). — Borror and DeLong, 1970. An Introduction to the Study of Insects, ed. 3, pp. 537-616, figs. 495-577. — Riek et ai, 1970. /)( The Insects of Australia, C. S. I. R. 0., pp. 867-959, 40 figs. -Richards, 1972 (1971). Ent. Essays to Commemorate Retirement of Prof. K. Yasumatsu, pp. 1-13, 10 figs, (thoracic spiracles in classification). — Rasnitsyn, 1971. XIII Internatl. Congr. Ent., Proc. 1: 289 (origin). -Koenigsmann, 1976. Deut. Ent. Ztschr. (n.f. 23) 4-5: 253-279, 4 figs, (phylogeny). Biology: Bishoff, 1927. Biologie de Hymenopteren, 598 pp., 224 figs. — Fulmek, 1957. Naturhist. Mus. Wien, Ann. 61: 110-227 (aphid parasites and predators). — iWata, 1972 (1971). Evolution of Instinct: comparative Studies of Hymenoptera Behavior, 503 pp. (Japanese edition). — Farish, 1972. Anim. Behavior 20: 662-676 (grooming behavior). — Iwata, 1976. Evolution of Instinct: Comparative Ethology of Hymenoptera, 535 pp., frontisp., 50 figs. (English translation, Natl. Tech. Inform. Serv., PB 257052). Morphology: Snodgrass, 1911. U. S. Natl. Mus., Proc. 39: 37-91, 16 pis., 19 text figs, (thorax). — rohwer and Gahan, 1916. Ent. Soc. Wash., Proc. 18: 20-76, 3 pis. (wing venation). -Comstock, 1918. The Wings of Insects, pp. 362-381, pi. 10, text figs. 380-405. —Snodgrass, 1935. Principles of Insect Morphology, 667 pp., 319 figs. —Ross, 1936. Ent. Soc. Amer., Ann. 29: 99-111, 2 pis. (wing venation). —Snodgrass, 1941. Smithsn. Inst., Misc. Collect. 99 (14): 1-86, 33 pis., 6 text figs, (male genitalia). — Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 367-446, 81 figs, (thorax of wingless and brachypterous spp.). —Richards, 1956. Roy Ent. Soc. London, Proc. (A) 31: 99-104, 7 figs, (interpretation of thoracic venter). -Short, 1959. Roy. Ent. Soc, London, Trans. Ill: 175-203, 11 figs, (abdominal musculature). —Oeser, 1961. Zool. Mus. Berlin, Mitt. 37: 3-119, 107 figs, (female ovipositor). —Daly, 1963. Ent. Soc. Amer., Ann. 56: 295-306 (thoracic musculature). —Smith, 1970. Ent. Soc. Amer., Ann. 63: 1-27 (evolutionary morphology of external genitalia). — Eady, 1974. Jour. Ent., Ser. B, 48: 63-72 (wing venation). Suborder SYMPHYTA By David R. Smith The suborder Symphyta, commonly known as sawflies and horntails, has also been recorded in the early literature under the names Chalastogastra, Sessiliventria, or Phyllophaga and Xylophaga. The suborder includes about 10,000 world species grouped into about 1,000 genera and 12 families. Representatives are found on all continents except for Antarctica, and they are also absent on many of the more isolated islands of the world such as Hawaii and many other Pacific islands. The most northern record is that for Pachynematus parvilabris (Thomson) on Ward Hunt Island in Canada, 83" 05' N., and several species are found as far south as Tierra del Fuego. The North American fauna consists of around 1,000 species in 10 families. The two fami- lies that have no living representatives in North America are the Megalodontidae and Blasticotomidae,both of which are Palearctic, though the Blasticotomidae is represented in North America by the fossil species Paremphytus ostentus Brues from the Miocene of Florissant, Colorado. The Blasticotomidae contains only several species, very secretive and rarely found, the larvae of which bore in the stems of ferns. Living forms may yet be discovered on this con tinent. Other than the families discussed here, a number of fossil families have been described, most all fi im the Old World. These are Anaxyelidae, Gigasiricidae, Karatavitidae, Myrmiciidae, Parapamphiliidae, Pararchexyelidae, Paroryssidae, Pseudosiricidae, Sepulcidae, Xyelotomidae, and Xyelydidae. The common name sawfly, applied to members of most families, is derived from their flylike appearance and the sawlike female ovipositor which is used to cut open plant tissue for insertion of eggs. The name horntail is usually applied to members of the family Siricidae, the females of which have a long slender ovipositor. Adults of the suborder may be distinguished from other Hymenoptera by the abdomen which is broadly joined to the thorax, the trochanters which are always two-segmented and the hindwing which usually has three closed basal cells. The larvae of most sawflies are entirely different from other Hymenoptera larvae and are most often con- fused with those of Lepidoptera, though sawfly larvae lack crochets on the prolegs, have only one pair of ocelli, and normally have more than five pairs of prolegs. Some larvae, especially those modified for an internal existence, resemble the grublike larvae of other Hymenoptera but normally have a projection at the apex of the abdomen and vestiges of thoracic legs. The higher classification of the Symphyta in most universal use at present is that proposed by Ross (1937) and Benson (1938). The suborder is divided into two major series, the Orthandria and Strophandria basically separated by the male genitalia which remains normal in the former but is turned 180° prior to eclosion in the latter. The Strophandria is expressed as the super- family Tenthredinoidea. The Orthandria have been further divided on the basis of characters of the mesosternum and head capsule into three superfamilies, the Megalodontoidea, Siricoidea, and Cephoidea. Some authors have recognized the superfamily Xyeloidea including only the family Xyelidae. This has some merit, as both positions of male genitalia are found, the orthan- 4 Hymenoptera in America North of Mexico drous in the Macroxyelinae and strophandrous in the Xyelinae. All members of this suborder are phytophagous except for the family Orussidae which is parasitic on other wood-boring insects. Host plants and habits are various. Many larvae are ex- ternal feeders on the foliage of the host, but others are leaf miners, gall formers on twigs, petioles, leaves, or buds, leaf-edge rollers, stem borers, wood borers, or petiole miners. Because of their plant feeding habits, many sawflies are pests of ornamental plants, agricultural crops, or forest stands. Their occurrence is normally local, but outbreaks in forests may be extensive, covering many thousands of acres and resulting in considerable growth loss or death of trees. Appearance in epidemic proportions is commonly sporadic, suddenly appearing for a year or more then practically disappearing for several years, probably being kept under control by their natural parasite-predator-disease complex. Some of the most destructive species are aliens which were accidentally introduced from abroad and found a vacuum for development without natural enemies. Many parasites have been introduced to help combat these pests. Some of the more important sawflies are those that feed on timber species, roses, apples, pears, peaches, and wheat and other grain crops. On the other hand, a few sawflies have been used beneficially, or at least have been attempted for use in the biological control of weeds. One species, Ucona acaenae Smith (=Antholcus varinervis of authors), was introduced into New Zealand from Chile in the early 1930's to help control the spread of Acaena. This attempt was only partially successful. Another species, Priophorus morio (Lepeletier), was released in Hawaii from the western United States to help control the spread of Rubus on the islands, but this attempt failed. The life cycle of many sawflies follows a similar pattern, though with many variations. Adults of most species fly in spring and early summer and are very short-lived. It is difficult to keep in- dividual adults alive in captivity for much more than a few days although the actual flight of a species may extend over several weeks or months. Some adults may not feed but some may feed on moisture, nectar and pollen of flowers, leaf pubescence, or other insects. The most productive areas for collecting are in vegetation on edges of rivers or streams, marshes, open and scrubby woodland, or undisturbed meadows. The catkins of willows and alders are especially productive for many groups. Oviposition is in the foliage, stems, twigs, or wood, and the eggs are most al- ways inserted in the plant tissue though some Pamphiliidae may glue their eggs to the leaf sur- face. Larval feeding time varies, but usually lasts about two weeks. Many larvae, especially Diprionidae, feed gregariously at first but later disperse to other parts of the host. After feed- ing is completed the larva molts into a non-feeding stage called the prepupa or resting stage. The prepupa normally leaves the host in search of a site for pupation. This may be in a cell in the ground or litter, in a papery cocoon in the ground, in some other substance such as wood, stems of the host or nearby plants, or fruits of other plants. The prepupal stage differs morphologically from the feeding stages, especially in the shape of the mandibles and sometimes color pattern. Some larvae with spines loose these in the final molt. If there is a single generation, the prepupa will remain in its cocoon or cell the rest of the summer and pupate the following spring. How- ever, depending on the species, latitude, or diapause requirements, there may be several genera- tions a year, or it may take several years to complete the cycle. The following references are to general articles on the Symphyta. Some, such as Maxwell, 1955, in morphology, pertain to many species and the reference is not repeated under each spe- cies. Under some species, such as Pristiphora erichsonii (Hartig), a listing of all biological references would take many more pages, and the references are selected or pertain to some recent synopsis which contains a good bibliography. Some of the new distribution records, hosts, and taxonomic changes are taken from unpublished notes. Only those taxonomic changes cited as being new have not before appeared in the literature. I am indebted to several colleagues for reviewing parts of this section and or providing some of the information contained herein: H. E. Milliron, formerly with the Biosystematics Research Institute, Agriculture Canada, Ottawa and other authorities of the Institute for allowing study of their sawfly collection; H. R. Wong, Northern Forest Research Centre, Canadian Forestry Service Edmonton, Alberta, Canada; and H. Greenbaum, Department of Entomology, University of Arkansas Fayetteville. Taxonomy: Norton, 1867-1869. Amer. Ent. Soc, Trans. 1: 31-84, 193-280; 2: 211-242, 321-368 (N. Amer. catalog). — Cresson, 1880. Amer. Ent. Soc, Trans. 8: 53-68 (N. Amer. catalog). — Dalla Torre, 1894. Cat. Hym., v. 1, 459 pp. (world catalog). — Ashmead, 1898. Canad. Ent. 30: 141-145, 177-183, 205-213, 225-232, 249-257, 281-287, 305-316 (classification, keys to Suborder SYMPHYTA 5 genera). — Konow, 1905. In Wytsman, Gen. Ins., fasc. 27, 27 pp.; fasc. 28, 14 pp.; fasc. 29, 176 pp. (world catalog). -Dyar, 1893-1895. Canad. Ent. 25: 244-248; 26: 42-45, 185-189; 27: 191-196, 208-212 (larvae). —Dyar, 1895 Amer. Ent. Soc, Trans. 22: 301-312 (larvae) -Dyar, 1897. N. Y. Ent. Soc, Jour. 5: 18-30, 190-201 (larvae) -Dyar, 1898. Canad. Ent. 30: 173-176 (larvae) -Dyar, 1898. N. Y. Ent. Soc, Jour. 6: 121-138 (larvae) -Dyar, 1900. N. Y. Ent. Soc, Jour. 8: 26-31 (larvae). -Rohwer, 1911. Ent. Soc. Wash., Proc. 13: 215-224 (classification). -Rohwer, 1911. U. S. Dept. Agr., Bur. Ent., Tech. Ser. 20, pt. 2, pp. 69-109 (genotypes and work of Ashmead). —Rohwer, 1911 Ent. News 22: 218-219 (additions and correction to "genotypes"). — Enslin, 1912-1918. Deut. Ent. Ztschr., Beih. 790 pp. (Tenthredinoidea of middle Europe). — MacGillivray, 1913. Ent. Soc Ont., Ann. Rpt. 44: 54-75 (larvae). —MacGillivray, 1916. Conn. State Geol. and Nat. Hist. Survey Bui. 22: 25-175. -Yuasa, 1923. 111. Biol. Monog. 7(4): 1-168 (N. Amer. larvae). —Ross, 1937. 111. Biol. Monog. 15(2): 1-173 (generic revision, N. Amer.). —Benson, 1938. Roy. Ent. Soc. London, Trans. 87: 353-384 (world classification). —Benson, 1951. Handb. for Ident. of Brit. Ins., v. 6, pt. 2(a): 1-49; 1952, v. 6, pt. 2(b): 51-137; 1958, v. 6, pt. 2(C): 139-252. -Lorenz and Kraus, 1957. Die Larvalsystematik der Blattwespen, 339 pp. (larvae of European species). —Ross, 1960. Amer. Ent. Soc, Trans. 85: 315-321 (early history of sawfly study in N. Amer.). —Benson, 1962. Brit. Mus. (Nat. Hist.) Ent., Bui. 12: 381-409 (Holarctic sawflies). — Rasnitsyn, 1969. Origin and evolution of the lower Hymenoptera, 187 pp. (fossil). Biology: Rohwer, 1915. Ent. Soc. Wash., Proc 17: 195-198 (mating habits). —Benson, 1950. Soc. Brit. Ent., Trans. 10: 45-142 (natural history of Brit, sawflies). —Wong, 1954. Canad. Ent. 86: 154-158 (sawflies on white birch in Man., Sask.). — Neilson, 1958. Canad. Ent. 90: 229-234 (life histories of sawflies in low-bush blueberry fields in N. B. ). — Raizenne, 1957. Canada Dept. Agr. Pub. 1009, 45 pp. (forest sawflies of s. Ont. and their parasites). — Lindquist, 1959. Canad. Ent. 91: 625-627 (leaf mining sawflies on birch). — Lindquist and Miller, 1970. Ent. Soc. Ont., Proc. 100: 117-123 (free-feeding sawflies on birch and alder in Ont.). -Lindquist and Miller, 1971. Ent. Soc. Ont., Proc. 102: 118-122 (larvae feeding on spruce and balsam fir in Ont.). Morphology: Marlatt, 1891. Ent. Soc. Wash., Proc. 2: 115-117 (final molting of larvae). — Marlatt, 1894. Ent. Soc. Wash., Proc. 3: 78-82 (neuration of wings). —MacGillivray, 1906. U. S. Natl. Mus., Proc 29: 569-654 (wings). —Van Dine, 1906. Hawaii. Ent. Soc, Proc. 1: 19-22 (comparative anatomical study of mouthparts). — Crampton, 1919. Ent. Soc. Wash., Proc 21: 129-155 (genitaHa and terminal abdominal structure of males and larvae). — Middleton, 1921. Ent. Soc. Wash., Proc. 23: 139-144 (terminal abdominal structures). — Middleton, 1921. Ent. Soc. Wash., Proc 23: 173-192 (suggested homologies between larvae and adults). — Boulange, 1924. Mem. et Travaux des Facultes Catholiques de Lille 28: 1-444 (genitalia). —Taylor, 1931. Roy. Phys. Soc, Proc. 22: 41-70 (morphology of the tenthredinid head). -Taylor, 1931. Ent. Soc. Amer., Ann. 24: 451-466 ("Dyar's Rule", its application to sawfly larvae). — Malloch, 1936. Inaugural- Dissertation, Erlangung der Doktorwurde, Philosophischen Fakultat der Friedrich- Wilhelms-Universitat zu Berlin, 55 pp. (thorax and cenchri). —Parker, 1934. Bol. Lab. Zool. Gen. e Agr. Portici 28: 159-191 (anatomy of larvae, with special reference to head). —Ross, 1936 Ent. Soc. Amer., Ann. 29: 99-111 (ancestry of wing venation). —Ross, 1945. Ent. News 56: 261-268 (genitalia: terminology and study techniques). —Maxwell, 1955. Canad. Ent. 87 (sup. 1): 1-132 (comparative internal larval anatomy). — Arora, 1956. Panjab Univ., Res. Bui., Zool. 90: 85-119 (relationship of Symphyta to other insect orders based on adult external morphology). — Togashi, 1965. Kontyu 33: 230-234 (rectal papillae). — Togashi, 1970. Mushi 43 (sup.): 1-114 (comparative morphology of internal reproductive organs). — Kenchington, 1972. Jour. Ent. 46: 111-116 (variations in silk gland morphology among larvae). Superfamily MEGALODONTOIDEA Family XYELIDAE A small family of primitive sawflies better represented in North America than in other parts of the northern hemisphere. Most of the family is associated with conifers; only members of the tribe Macroxyelini are associated with deciduous trees. Revision: Ross, 1932. Ent. Soc. Amer., Ann. 25: 153-169 (Nearctic). Taxonomy: Klima, 1937. In Hedicke, Hym. Cat. pt. 4, 12 pp. (catalog). —Benson, 1945. Roy. Ent. Soc. London, Proc, Ser. B: Taxonomy 14: 34-37 (generic classification, world). — Burdick, 1959. Kans. Ent. Soc, Jour. 32: 120-122 (fossil). — Rasnitsyn, 1964. Paleontol. Jour. No. 1, pp. 88-96 (fossil). -Rasnitsyn, 1965. Polski Pismo Ent. 35: 483-519 (Xyelinae of world). —Rasnitsyn, 1966. Paleontol. Jour. No. 4, pp. 69-85 (fossil). —Smith, 1967. Ent. Soc. Amer., Ann. 60: 376-384 (larvae). Subfamily XYELINAE Tribe XYELINI Genus XYELA Dalman Pinicola Brebisson, 1818. Soc. Philomath, de Paris, Bui. des Sci., p. 116. Preocc. Type-species: Pinicola julii Brebisson. Desig. by Rohwer, 1911. Xyela Dalman, 1819. Svenska Vetensk. Akad., Handl. 40: 122. Type-species: Xyela pusilla Dalman. Desig. by Curtis, 1824. Mastigocera Klug, 1827. In Berthold, Latreille's Nat. Fam. Thierr. p.442. Tritokreion Schilling, 1825. Schles. Gesell. f. Vaterland. Kult. Ubers., Arb. 2: 43. No included species. Neoxyela Curran, 1923. Canad. Ent. 55: 20. Type-species: Neoxyela alberta Curran. Monotypic. Xyelatana Benson, 1938. Roy. Ent. Soc. London, Proc, Ser. B: Taxonomy 7: 35. Type-species: Xyela longula Dalman. Orig. desig. Larvae of most species of Xyela live and feed in the developing staminate cones of Pinus spe- cies. One species, gallicaulis, forms shoot galls. Adults fly early in the spring and are commonly found in large numbers on the catkins of Salix or Ahius near their host plants. Revision: Rohwer, 1913. U. S. Natl. Mus., Proc 45: 265-281. -Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 285-353 (also biology). alberta (Curran). Alta., Mont., Wyo., Calif. Host: Piyms contorta. Neoxyela alberta Curran, 1923. Canad. Ent. 55: 20. 9, J. 8 Hymenoptera in America North of Mexico alpigena (Strobl). Que. and Maine to Md. w. to Utah and N. Mex.; Holarctic. Host: Pinus strobus. Pinkola alpigena Strobl, 1895. Wien. Ent. Ztg. 14: 277. 9,6. Xyela bnnmeiceps Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 269. 9. Xyela kamtshatica Gussakovskij, 1935. Faune URSS, Ins. Hym., T. 2, 1: 133. $. Xyela middlekauffi Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 343. 9. Taxonomy: Benson, 1962. Brit. Mus. (Nat. Hist.) Bui. Ent. 12: 385. bakeri Konow. Que. to Fla. w. to B. C. and Calif. Ecology: Widespread. Host: Pmus elliottii, P. palustris, P. ponderosa, P. sabiniana, P. virginiana. Xyela bakeri Konow, 1898. Ent. Nachr. 24: 328. Xyela negundinis Cockerell. 1907. Canad. Ent. 39: 324. Xyela salicis Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 271. 6. Xyela intrabilis MacGillivray, 1923. Brooklyn Ent. Soc, Bui. 18: 53. 6. Biology: Ebel, 1966. Ent. Soc. Amer., Ann. 59: 127-129. californica Rohwer. Calif. Xyela californica Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 272. 6. cheloma Burdick. Idaho, Nev., B. C, Wash., Oreg. Host: Pinus ponderosa. Xyela cheloma Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 336. 9,6. concava Burdick. Utah, Nev., Calif. Host: Pinus monophylla, P. ponderosa. Xyela concava Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 342. 9,6. deserti Burdick. Nev., Calif. Host: Pinus monophylla. Xyela deserti Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 347. 9,6. dodgei Greenbaum. Fla. (Cedar Key). Xyela dodgei Greenbaum, 1973. Fla. Ent. 57: 104-106. 6. gallicaulis Smith. Va., Ga. Ecology: Larva feeds inside and forms galls on shoots. Host: Pinus echinata, P. elliottii, P. taeda. Xyela gallicaulis Smith, 1970. Ga. Ent. Soc, Jour. 5: 69-72. 9,6. larva. linsleyi Burdick. Idaho, B. C, Wash., Calif. Host: Pinus ponderosai^). Xyela li^isleyi Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 344. 9,6. lunata Burdick. Calif. Host: Pinus coulteri, P. sabiniana. Xyela lunata Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 334. 9,6. minor Norton. Que. to Fla. w. to B. C. and Calif. Ecology: Widespread. Host: Pinus coulteri, P. elliottii, P. muricata, P. pahistris, P. ponderosa, P. sabiniana, P. taeda, P. virginiayia. Xyela 7ni7ior Norton, 1868. Amer. Ent. Soc, Trans. 2: 349. 9. Xyela luteopicta Cockerell, 1902. Canad. Ent. 34: 194. 6. Xyela alni Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 268. 9,6. Xyela winnemanae Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 268. 9. Xyela errans Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 269. 9. Xyela dissimilis Rohwer, 1913. U. S. Natl. Mus., Proc 45: 270. 9. Xyela nevadensis Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 271. 9. Xyela coloradensis Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 272. 6. N. syn. Biology: Ebel, 1966. Ent. Soc. Amer., Ann. 59: 127-129. Morphology: Daly, 1963. Ent. Soc Amer., Ann. 56: 299. obscura (Strobl). Newfoundland to Fla. w. to N. W. T., B. C. and Calif.; Holarctic. Ecology: Widespread. Host: Pinus banksiayia, P. palustris, P. ponderosa, P. virginiana. Piyiicola julii var. obscura Strobl, 1895. Wien. Ent. Ztg. 14: 277. 9. Xyela japonica Rohwer, 1910. U. S. Natl. Mus., Proc. 39: 99. 9. Xyela pini Rohwer, 1913. U. S. Natl. Mus., Proc 45: 267. 9,6. Xyela slossonae Rohwer, 1913. U. S. Natl. Mus., Proc. 45: 270. 9,6. Xyela similis Rohwer, 1913. U. S. Natl. Mus., Proc 45: 271. 6. Taxonomy: Benson, 1962. Brit. Mus. (Nat. Hist.) Bui. Ent. 12: 385. Biology: Ebel, 1966. Ent. Soc. Amer., Ann. 59: 127-129. Superfamily MEGALODONTOIDEA priceae Burdick. Calif. Host: Pinus coulteri. Xyela prkeae Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 339. 9, 6. radiatae Burdick. Calif. Host: Pinus radiata(^) Xyela radiatae Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 337. 9,6. serrata Burdick. Calif. Host: Pinus muricata. Xyela serrata Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 341. 9. styrax Burdick. D. C, Ga., Md., N. Y., Va. Host: Pinus virginiana. Xyela styrax Burdick, 1961. Calif. Univ., Pubs., Ent. 17: 345 9, 6. Tribe PLERONEURINI Genus PLERONEURA Konow Pleronenra Konow, 1897. Ent. Nachr. 23: 56. Type-species: Xyela dahli Hartig. Desig. by Rohwer, 1911. Pleuroneura Ashmead, 1898. Psyche 8: 214. Emend. Manoxyela Ashmead, 1898. Psyche 8: 214. Type-species: Manoxyela californica Ashmead. Orig. desig. Larvae of this genus live in the developing shoots and buds of Abies. Revision: Rohwer, 1910. Canad. Ent. 42: 88-93. —Ross, 1932. Ent. Soc. Amer., Ann. 25: 156-159. Taxonomy: Smith, 1967. Ent. Soc. Amer., Ann. 60: 382-383 (larvae). —Smith, 1967. Ent. Soc. Wash., Proc. 69: 182-183 (species synonymy). aldrichi Ross. Idaho, Oreg., Nev., Calif. Host: Abies. Pleroneura aldrichi Ross, 1932. Ent. Soc. Amer., Ann. 25: 158-159. 9,6. brunneicornis Rohwer. N. S., N. B., Maine, Que., Ont., N. Y., Minn. Host: Abies balsamea, A. concolor. Pleroneura bruneiconiisV.) Rohwer, 1910. Canad. Ent. 42: 89. 9. Pleroneura bmyineicomis Rohwer, 1910. Canad. Ent. 42: 220. Pleroneura borealis Felt, 1917. Canad. Ent. 49: 220. 9,6. Biology: Webb and Forbes, 1951. Canad. Ent. 83: 181-183. californica (Ashmead). Wyo., Alta., Utah, B. C, Oreg., Calif. Host: Abies spp. Manoxyela califoi-nica Ashmead, 1898. Psyche 8: 214. 6. Pleroneura fulvicornis Rohwer, 1910. Canad. Ent. 42: 89. 6. Pleroneura sclucarzi Rohwer, 1910. Canad. Ent. 42: 220. 9. koebelei Rohwer. Idaho, Oreg., Calif. Host: Abies spp. Pleroneura koebelei Rohwer, 1910. Canad. Ent. 42: 90. 6. lutea Rohwer. Idaho, Oreg., Nev., Calif. Pleroneura lutea Rohwer, 1910. Canad. Ent. 42: 90. 6. Subfamily MACROXYELINAE Tribe XYELECIINI Genus XYELECIA Ross Xyelecia Ross, 1932. Ent. Soc. Amer., Ann. 25: 159. Type-species: Xyelecia nearctica Ross. Orig. desig. nearctica Ross. "Mont, and Colo.", Idaho, Oreg., Calif. Ecology: Larvae live in shoots. Host: Abies spp. Xyelecia nearctica Ross, 1932. Ent. Soc. Amer., Ann. 25: 160. 9. Taxonomy: Smith, 1964. Pan-Pacific Ent. 40: 54-56 (male). —Smith, 1967. Ent. Soc. Amer., Ann. 60: 381-382 (larva). 10 Hymenoptera in America North of Mexico Tribe MACROXYELINI Genus MACROXYELA Kirby Macroxyela Kirby, 1882. List Hym. Brit. Mus. v. 1, p. 351. Type-species: Xyela ferruginea Say. Orig. desig. Protoxyela MacGillivray, 1912. Canad. Ent. 44: 295. Type-species: Xyela aeniad) Norton. Orig. desig. Larvae of this genus feed on the leaves of deciduous trees. Revision: Ross, 1932. Ent. Soc. Amer., Ann. 25: 164-166. aenea (Norton). La., Mo., Tex. Xyela aenea Norton, 1872. Amer. Ent. Soc, Trans. 4: 86.