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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 494

A Catalog and Review of Immature Apoidea (Hymenoptera)

Ronald J. McGinley

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1989

ABSTRACT

McGinley, Ronald J. A Catalog and Review of Immature Apoidea (Hymenoptera). Smithsonian Contributions to Zoology, number 494, 24 pages, 2 figures, 1 table, 1989.—The systematic literature covering the larvae and pupae of all bees is reviewed and presented in catalog form. Historical aspects of the study of immature apoids are reviewed with emphasis placed on what yet needs to be accomplished and which taxa should be considered primary targets for collection.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

McGinley, Ronald J.

A catalog and review of immature Apoidea (Hymenoptera).

(Smithsonian contributions to zoology ; no. 494)

Bibliography: p.

Supt. of Docs. no.: SI 1.27:494

1. Bees—Classification. 2. Bees—Development. 3. Bees—Larvae. I. Title. II. Series.

QL1.S54   no. 494   [QL566]   591 s   89-600106   [595.79′9]

A Catalog and Review of Immature Apoidea (Hymenoptera)

Ronald J. McGinley

____________

Ronald J. McGinley, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Introduction

The immature stages of holometabolous insects provide systematists with character sets that are too often overlooked in taxonomic and phylogenetic studies. The most obvious reason for this neglect is that the immatures of most species are unknown; this because they are often difficult to locate, collect and correctly associate with adults. Another reason for ignoring immature stages is that many systematists are reluctant to utilize character sets that are unfamiliar to them. In the case of apoid “paedophobia,” the problem is exacerbated by a seemingly unorganized literature that leaves many workers bewildered. Because the publication of Immature Insects (Stehr, 1987) will hopefully stimulate greater interest in larval studies, this seems an appropriate time to summarize the immature-bee literature on a global basis.

The present paper presents a listing of those bee species that are known from the larval and/or pupal stages and provides the associated bibliographic information. It is hoped that this information will encourage bee biologists to preserve immature stages for future study, especially the immatures of those taxa that are unknown or poorly represented in the literature and collections. Immatures should be killed and preserved in alcohol (75 to 80 percent) or Kahle's Solution (a mixture of ethyl alcohol, formaldehyde, glacial acetic acid and water); techniques of specimen preparation and examination are presented in Rozen and McGinley (1974a) and Stephen, Bohart, and Torchio (1969).

ACKNOWLEDGMENTS.—The manuscript was greatly improved by the comments and suggestions provided by Robert W. Brooks, Connal D. Eardley, George C. Eickwort, Oliver S. Flint, Jr., Terry L. Griswold, Terry F. Houston, Wallace E. LaBerge, Charles D. Michener, Beth B. Norden, Frank D. Parker, Yuriy A. Pesenko, Jerome G. Rozen, Jr., Shoichi F. Sakagami, and Philip F. Torchio. Beth Norden also assisted with the literature search and figure lettering, and Philip Torchio kindly provided a prepublication copy of his 1987 paper (with B. Burwell) on colletid pupae. Radclyffe B. Roberts compiled the initial listing of bee taxa that was the basis for the general classification scheme followed herein.

Historical Review

As mentioned above, our knowledge of apoid diversity is understandably based primarily on studies of adults. The discrepancy in our knowledge of immature forms versus adults is well illustrated by the history of bee taxonomy in the United States and Canada, a fauna of approximately 3,589 species and subspecies (Figure 1). Study of this fauna (based on adults) from 1758 through the 1840s resulted in the description of 90 currently recognized species. During this early period, Thomas Say contributed significantly with descriptions of 33 species. Exploration of the North American bee fauna really began during the 1850–1870 period with the work of F. Smith and E.T. Cresson. T.D.A. Cockerell and P.H. Timberlake contributed most significantly to our knowledge of North American species with descriptions of 1138 and 735 species and subspecies, respectively; nine additional workers, listed in figure 1, have each contributed over 75 species descriptions to the North American fauna. Among these most prolific contributors, only Michener and LaBerge have published descriptions of immature forms and have utilized this information in assessments of phylogenetic relationships and classification. Mature larvae of only 174 (5.0%) of the approximately 3,589 species and subspecies in this area are known.

Among the earliest known larval descriptions of Apoidea are those of Hoplitis parvula and H. tridentata by Dufour and Perris (1840). Earlier depictions of the honey bee are so generalized as to be virtually useless for comparative purposes. Through the 1940s, several workers contributed scattered larval treatments (most notably F. Claude-Joseph and G. Grandi), but it wasn't until Michener's publication in 1953 that the study of larval bees became truly comparative and organized for the first time. In this publication, Michener described or redescribed the larvae of 135 species and subspecies, provided summary descriptions for the bee families based on mature larvae, presented a key to the larvae of the major hymenopteran taxa and discussed evolutionary trends among bee larvae. Michener followed this landmark paper with a similar treatment of apoid pupae in 1954. These papers have done much to stimulate other bee systematists to collect and study immature bees. Most notably, from 1954 to the present, J.G. Rozen, Jr., has contributed a large body of work that has greatly advanced our understanding of the immatures of virtually all bee families and has emphasized their value in phylogenetic considerations (literature cited in bibliography). Also contributing significantly during this period were G.C. Eickwort (primarily halictine immatures), B. Lucas de Oliveira (Neotropical species), W.P. Stephen and T. Koontz (Bombus larvae) and P.F. Torchio (various taxa). Stephen et al., (1969) provided an overview of apoid larval and pupal morphology and included an account of preservation and examination techniques. McGinley (1981) published a key to the known mature larvae throughout the Apoidea and described 96 genera in matrix form (126 characters). An illustrated key to North American bee taxa based on mature larvae, family treatments and introduction to the associated literature was published by McGinley, 1987. Torchio and Burwell (1987) have contributed descriptions (in tabular form) of several colletid pupae and discuss some general problems in studying apoid pupae.

FIGURE 1.—History of the bee fauna in America north of Mexico: number of adult and larval descriptions per decade (including larval treatments outside North America) and associated systematists who have described over 75 species from this area (most adult data from Hurd, 1979).

Systematic Contribution of Immature Apoid Studies

The few studies published that consider the immature forms of any significant number of congeneric species indicate that larvae and pupae, for some taxa, appear to be of limited value at the alpha-taxonomic level. In his study of adults and larvae of Nomadopsis, Rozen (1958) states, “there appears to be no differentiation of the larvae with respect to species groups or subgenera.” Eickwort (1981) writes, “I have examined in detail larvae and pupae of 6 species of Agapostemon and they are all very similar in even the most minute morphological characters. This suggests that immature stages are unlikely to provide good characters to distinguish closely related halictine species but will prove valuable in generic studies.” However, closely related species in other bee taxa can be distinguished by larval characters. For example, Torchio and Torchio (1975) were able to differentiate four closely related species of Apis on the basis of larval features and in an unpublished study they found it easy to distinguish the larvae of nine species of Diadasia (P.F. Torchio, pers. comm.). Rozen and McGinley (1974a) were able to provide a key to the mature larvae of eight species of Hesperapis. In his study of Perdita, Eickwort (1977 and pers. comm.) shows that larval characters might be valuable in this taxon at the subgeneric and species group level. As stated by P.F. Torchio (pers. comm.), at the species level “the value of larval characteristics may, therefore, vary from one taxon to another within Apoidea.”

That larval and pupal characters can be extremely valuable in systematic studies at the generic and higher categorical levels is well illustrated by Rozen's various studies on the Nomadinae (1966c, 1977c), on the Panurginae (1966a, 1968, 1970a, 1971), and the Fideliinae (1970b, 1973b, 1977b) and Lithurginae (1973c). In contrast to the ordinary cell-reared bee larvae, allodapine larvae which do not live in cells have excellent characters at both generic and specific levels. Michener (1977) showed that allodapine larvae were extremely valuable in generic studies; analysis of larval characters alone gave the closest approximation of the topology of a general cladogram based on cladistic analysis of adult, pupal, larval and egg characters.

That apoid larvae do not seem to be highly informative at the higher apoid categorical levels was suggested by Michener and Greenberg (1980). An attempt to illustrate this apparent lack of resolution is presented in Figure 2, where the highest known categorical levels supported by larval synapomorphies are presented (characters listed in Appendix) and contrasted with the greater clustering detected on the basis of adult morphology (Michener and Greenberg, 1980). Among the larger bee families, only the Andrenidae is supported by larval characters alone. This does not argue against the monophyly of the other apoid families (supported by other character sets), but illustrates the lack of phylogenetic resolution detected at this categorical level.

Immature Apoid Collecting/Research Priorities

The primary goal of this paper is not to review what has been accomplished in the study of immature apoids, but rather to identify what still needs to be done. The figures presented in Table 1 review the percentage of genera in each apoid family and subfamily that are known as mature larvae. Of the 480 genera listed in this compilation only 151 (31%) genera are known in the larval stage. The following listing points out the major synthetic papers within each family and emphasizes which taxa might be considered primary targets for future research efforts.

COLLETIDEA.—Although the mature larvae of this family were reviewed by McGinley (1981), only 26 percent of the currently recognized colletid genera are known as larvae. Most work is needed on the many genera of Neotropical and Australasian Paracolletini (Colletinae) and the Australasian Euryglossinae. Of special interest, would be the location of immature representatives of the Neotropical diphaglossine tribe Dissoglottini (Mydrosoma, Mydrosomella, Ptiloglossidia). In the United States and neotropics, immature forms of Eulonchopria (Paracolletini) remain unknown. Known colletid pupae have been reviewed by Torchio and Burwell (1987).

FIGURE 2.—Highest known categorical levels supported by apoid larval synapomorphies (left-hand side) contrasted with higher resolution detected from study of adult characters (Michener and Greenberg, 1980). Apparent larval synapomorphies are listed in Appendix.

TABLE 1.—Percentage of genera known as mature larvae among apoid families and subfamilies.

STENOTRITIDAE.—Houston (1975) located and described larvae of Stenotritus pubescens and has recently collected the larvae of S. greavesi and larval representatives of two species of Ctenocolletes (McGinley, in prep.).

HALICTIDAE.—Despite the significant contributions by G.C. and K.R. Eickwort (literature cited in bibliography), the larvae and pupae of this family remain virtually unknown with only 20 percent of the recognized genera represented by immatures. Given the high level of diversity exhibited by adult nomiines, the study of additional immatures of this subfamily should prove most valuable. The immatures of Nomioidini (Ceyalictus, Nomioides) in the Halictinae are unknown.

ANDRENIDAE.—Among andrenines, immatures of only a few of the numerous Andrena subgenera are known; as more representatives become known they may contribute significantly to our understanding of species group relationships. In the United States, immatures of Ancylandrena and Megandrena are unknown.

Panurgine immatures are relatively well known, largely due the efforts of J.G. Rozen. Rozen (1966a) contributed a synthetic account of panurgine larvae and additional treatments of the immatures of Meliturgula (Rozen, 1968), Hypomacrotera and Protandrena (as Psaenythia) (Rozen, 1970a), and Melitturga and Panurgus (Rozen, 1971). Despite this progress, the immatures of 19 genera of panurgines remain unknown, including Metapsaenythia and Xenopanurgus from the United States. Yager and Rozen (1966) published a preliminary account of andrenid pupae.

OXAEIDAE.—The known larvae of Oxaea (Roberts, 1973) and Protoxaea (Rozen, 1964a) are very similar. Immatures of Mesoxaea and Notoxaea are unknown.

MELITTIDAE.—The known immatures of this family were reviewed by Rozen and McGinley (1974b). Rozen has contributed additional papers on the immatures of Meganomia (Rozen, 1977a) and Macropis (Rozen and Jacobson, 1980). Among those genera for which immature representatives are unknown, most interesting to locate would be immatures of the dasypodine tribes Promelittini (Promelitta) and Sambini (Samba).

CTENOPLECTRIDAE.—The mature larva of Ctenoplectra armata was described by Rozen (1978). At that time, the Ctenoplectrinae were classified as a melittid subfamily, and Rozen's phylogenetic considerations were made within that context. Larval characters should be reanalyzed in light of recent information presented by Michener and Greenberg (1980) that indicates ctenoplectrids are the sister group of the long-tongued bee families (Figure 2).

MEGACHILIDAE.—In a series of papers, Rozen (1970b, 1973b,c, 1977b) has contributed most significantly to our knowledge of fideliine and lithurgine megachilids. Immature representatives of all genera in these two subfamilies are known, with the exception of pupal Parafidelia. This progress contrasts sharply with our understanding of megachiline immatures, which have never been comprehensively reviewed. In the Anthidiini alone, 64 genera remain unknown in immature form.

ANTHOPHORIDAE.—Within the Anthophorinae, Rozen has contributed comprehensive papers on the Eucerini and Centridini (Rozen, 1965b) and the Melectini, Ericrocini, and Rhathymini (Rozen, 1969). Lucas de Oliveira (1962, 1966a) has published preliminary comparative studies of exomalopsine larvae. Within the Anthophorini, Michener (1953) compared a number of Anthophora larvae and Torchio and Stephen (1961) described the immatures of Emphoropsis. In spite of the above work, our understanding of larval/pupal diversity in this subfamily remains meager, for example, the immatures of 21 genera of Eucerini are unknown including those of Gaesischia, Martinapis, Melissoptila, Simanthedon, and Xenoglossodes, all of which occur within the United States. The immatures of four anthophorine tribes await discovery: Ancylini (Ancyla, Tarsalia), Canephorulini (Canephorula), Eucerinodini (Eucerinoda), and Tetrapediini (Tetrapedia). In the United States, immatures of Ericrocis, apparent parasites of Centris, should be considered primary targets.

Known mature larvae of the Nomadinae were reviewed by Rozen (1966c, 1977c) and those of the included Ammobatini by Rozen and McGinley (1974b). Because nomadines are cleptoparasitic in the nests of other bees, their immature forms are especially difficult to locate and to correctly associate with adult forms. Most exciting finds would include the discovery of immature Epeoloidini (Epeoloides; apparent parasites of Macropis) and Townsendiellini (Townsendiella, tentatively associated with Hesperapis), both of which occur within the United States. Other nomadine tribes unknown as immatures are the Ammobatoidini (Ammobatoides), Caenoprosopidini (Caenoprosopidis) and Osirini (Osiris). Important contributions could be made with the discovery of the many unknown genera of the Ammobatini and the Ericrocidini.

Within the Xylocopinae, study of immature allodapines include the early efforts of Rayment (1949a,b, 1951) and Syed (1963). This group, which exhibits the greatest morphological diversity known among apoid larvae, was the focus of a series of comprehensive papers by Michener (1964, 1975, 1976, 1977). The known immatures of Xylocopa have been studied by Lucas de Oliveira (1974). Immatures of Lestis, and Proxylocopa have not been treated in the literature.

APIDAE.—The known larvae of Apis were thoroughly studied by Torchio and Torchio (1975). Their study is especially notable for documenting considerable levels of intraspecific morphological variability which should be considered in future larval work. Larval Bombini were subject to early study by Ritcher (1933) and more recently by Stephen and Koontz (1973). Surprisingly, immatures of only two genera of the popularly studied Euglossini are known; immatures of Eufriesea, Exaerete, and Aglae remain unknown and should be considered primary targets. What we know about the immatures of the Meliponinae comes mostly from the contributions of Lucas de Oliveira (1958, 1960, 1965, 1966a, 1968, 1970). Many more larval/pupal representatives of this subfamily need to be collected; immatures of Dactylurina and Meliponula are unknown.

Catalog of Immature Apoidea

The information presented below was obtained from a personal card catalog maintained since 1977. A computer search of the literature was carried out in August 1988; the data bases searched were the Biosis Previews Database (1969–1988), the Zoological Record Database (1978–1983) and the Life Sciences Collection Database (1978–1985). Only systematic treatments of immatures are listed; a large body of literature dealing with immature development, physiology and biology exists but is not included here (see for example the paper on embryogenesis by Torchio and Trostle, 1986). Some larvae are known to have been collected but as of yet have not been treated in the literature; these are herein listed as being unknown.

The classification utilized largely follows an unpublished listing of bee taxa at and above the subgeneric level compiled by Radclyffe B. Roberts; this listing is not intended to be exhaustive, especially for Old World taxa, but is most adequate for cataloging immature studies.

Papers dealing only with larvae are cited without annotation; papers treating both larvae and pupae are cited with the annotation “[+ PUPA]”; citations followed by “[PUPA]” indicate that only pupae were treated. The numbers in parentheses after the higher categories indicate the percentage of bee genera in that taxon known as mature larvae. Taxa enclosed by brackets are known to have been collected, but as yet have not been treated taxonomically (listed depositories are: the University of Kansas (KU), the Bee Biology and Systematics Laboratory, Utah State University (UTAH), and the Smithsonian Institution (USNM; Karl V. Krombein trap-nesting material). The American Museum of Natural History, New York, is a major depository for immature apoids and houses many unstudied specimens not listed herein; interested biologists should contact Jerome G. Rozen, Jr., for information concerning loan and deposition policies.

COLLETIDEA (26%)

COLLETINAE (27%)

COLLETINI (25%)

Colletes araucariae Friese.—Claude-Joseph, 1926; Michener, 1953; McGinley, 1981.

Colletes ciliatoides Stephen.—Nielsen and Bohart, 1967 [sex characters]; Torchio and Burwell, 1987 [PUPA].

Colletes ciliatus Friese.—Claude-Joseph, 1926; Michener, 1953.

Colletes cognatus Spinola.—Torchio and Burwell, 1987 [PUPA].

Colletes daviesanus Smith.—Semichon, 1936.

Colletes fulgidus Swenk.—Michener, 1953, 1954 [PUPA]; McGinley, 1981.

Colletes kincaidii Cockerell.—Torchio and Burwell, 1987 [PUPA].

Colletes laticeps Friese.—Claude-Joseph, 1926; Michener, 1953; McGinley, 1981.

Colletes michenerianus Moure.—McGinley, 1981.

Colletes succinctus (Linnaeus).—Mayet, 1875 [+ PUPA]; Michener, 1953.

Colletes thoracicus Smith.—McGinley, 1981.

Colletes validus Cresson.—Rajotte, 1979.

Unknown: Hemicotelles, Mourecotelles, Xanthocotelles.

PARACOLLETINI (27%)

Callomelitta species A.—McGinley, 1981.

Leioproctus cingulatus (Moure).—McGinley, 1981.

Leioproctus zonalis (Reed).—Claude-Joseph, 1926 [as Lonchopria marginata]; Michener, 1953; McGinley, 1981.

Leioproctus zonatus (Moure).—McGinley, 1981.

Leioproctus species near semipurpureus (Cockerell).—McGinley, 1981.

Scrapter longula (Friese).—McGinley, 1981.

Unknown: Anthoglossa, Eulonchopria, Hesperocolletes, Hexantheda, Neopasiphae, Paracolletes, Tetraglossula, Trichocolletes.

DIPHAGLOSSINAE (44%)

CAUPOLICANINI (100%)

Caupolicana gayi Spinola.—Claude-Joseph, 1926; Michener, 1953; McGinley, 1981.

Crawfordapis luctuosa (Smith).—Otis et al., 1982; Roubik and Michener, 1985 [PUPA]; Torchio and Burwell, 1987 [PUPA].

Ptiloglossa fulvopilosa (Cameron).—McGinley, 1981.

Ptiloglossa species A.—McGinley, 1981.

Ptiloglossa species B.—McGinley, 1981.

DIPHAGLOSSINI (33%)

Cadeguala occidentalis (Haliday).—Claude-Joseph, 1926 [as Policana occidentalis]; Michener, 1953; McGinley, 1981; Torchio and Burwell, 1987 [PUPA].

Cadeguala albopilosa (Spinola).—Claude-Joseph, 1926; Michener, 1953, 1954 [PUPA]; McGinley, 1981 [all above as Policana herbsti]; Torchio and Burwell, 1987 [PUPA].

Unknown: Cadegualina, Diphaglossa.

DISSOGLOTTINI (0)

Unknown: Mydrosoma, Mydrosomella, Ptiloglossidia.

EURYGLOSSINAE (11%)

Eurygloss fasciatella Cockerell.—Michener, 1953.

Euryglossa subsericea Cockerell.—McGinley, 1981.

Euryglossa species A.—McGinley, 1981.

Pachyprosopis angophorae Cockerell.—McGinley, 1981.

Pachyprosopis indicans (Cockerell).—McGinley, 1981.

Unknown: Argohesma, Brachyhesma, Dasyhesma, Euryglossina, Euryglossella, Euryglossula, Heterohesma, Hyphesma, Melittosmithia, Parapachyprosopis, Quasihesma, Sericogaster, Stenohesma, Stilpnosoma, Turnerella, Xanthesma.

HYLAEINAE (33%)

Amphylaeus morosus (Smith).—McGinley, 1981.

Hylaeus alcyoneus (Erichson).—McGinley, 1981.

Hylaeus asininas (Cockerell and Casad).—McGinley, 1981.

Hylaeus bisinuatus Forster.—Torchio and Burwell, 1987 [PUPA].

Hylaeus brevicornis (Nylander).—Danks, 1970.

Hylaeus cressoni (Cockerell).—Nielsen and Bohart, 1967 [sex characters].

Hylaeus modestus Say.—McGinley, 1981.

Hylaeus nigritus Fabricius.—Janvier, 1969.

Hylaeus parvulus Janvier.—Janvier, 1972.

Hylaeus perhumilis (Cockerell).—McGinley, 1981.

Hylaeus pictus Smith.—Janvier, 1969.

Hylaeus pubescens (Perkins).—Daly and Coville, 1982.

Hylaeus variegata (Fabricius).—Stockhert, 1922; Michener, 1953.

Hylaeus species—Michener, 1953.

[Hylaeus species, UTAH.]

Hyleoides concinna (Fabricius).—McGinley, 1981.

Meroglossa species A, near torrida (Smith).—McGinley, 1981.

[Palaeorhiza gratiosa Cheesman; Y. Hirashima material, on loan to RJ. McGinley.]

[Palaeorhiza gressittorum Hirashima; Y. Hirashima material, on loan to R.J. McGinley.]

Unknown: Agogenohylaeus, Gephyrohylaeus, Hemirhiza, Heterapoides, Hylaeorhiza, Nothylaeus, Pharohylaeus.

XEROMELISSINAE (25%)

Chilicola ashmeadi (Crawford).—Eickwort, 1967 [+ PUPA]; McGinley, 1981; Torchio and Burwell, 1987 [PUPA].

Chilicola friesei Herbst—Claude-Joseph, 1926; Michener, 1953.

Chilicola inermis Friese.—Claude-Joseph, 1926; Michener, 1953.

Xeromelissine species A.—McGinley, 1981.

Unknown: Chilimelissa, Xenochilicola, Xeromelissa.

STENOTRITIDAE (50%)

[Stenotritus greavesi (Rayment); T.F. Houston material, on loan to RJ. McGinley.]

Stenotritus pubescens (Smith).—Houston, 1975; McGinley, 1981.

[Ctenocolletes nicholsoni (Cockerell); T.F. Houston material, on loan to RJ. McGinley.]

[Ctenocolletes ordensis Michener; TF. Houston material, on loan to RJ. McGinley.]

HALICTIDAE (20%)

DUFOUREINAE (15%)

Dufourea mulleri (Cockerell).—Torchio et al., 1967 [partial description]; McGinley, 1981.

Dufourea novaeangliae (Robertson).—Eickwort et al., 1986.

Rophitoides canus (Eversm.).—Enslin, 1921 [as Rhophites canus].

[Systropha punjabensis Batra and Michener.—Batra and Michener, 1966 (immature larva)].

Unknown: Conanthalictus, Michenerula, Micralictoides, Morawitzia, Morawitzeiella, Penapis, Protodufourea, Rophites, Sphecodosoma, [Systropha, UTAH], Xeralictus.

HALICTINAE (20%)

AUGOCHLORINI (17%)

Augochlora cordiaefloris Cockerell.—Eickwort and Eickwort, 1972a [+ PUPA].

Augochlora hallinani Michener.—Eickwort and Eickwort, 1973a [+ PUPA].

Augochlora nominata Michener.—Eickwort and Eickwort, 1972a [+ PUPA].

Augochlora pura (Say).—Michener, 1953, 1954 [PUPA]; McGinley, 1981.

Augochlora sidaefoliae Cockerell.—Eickwort and Eickwort, 1973a [+ PUPA].

Corynura chloris (Spinola).—Claude-Joseph, 1926 [as Halictus chloris].

Corynura cristata (Smith).—Claude-Joseph, 1926 [as Halictus cristatus].

Neocorynura colombiana Eickwort.—Eickwort, 1979 [+ PUPA].

[Neocorynura species, KU.]

Pereirapis edentata Michener.—Eickwort and Eickwort, 1973b [+ PUPA, as Augochlorella]; McGinley, 1981.

Unknown: Andinaugochlora, Ariphanarthra, Augochlorella, Augochlorodes, [Augochloropsis, KU], Caenaugochlora, Ceratalictus, Chlerogas, Chlerogella, Corynurella, Halictillus, Megalopta, Megaloptidia, Megommation, Paroxystoglossa, [Pseudaugochloropsis, KU], Rhectomia, Rhinocorynura, Temnosoma, Thectochlora.

HALICTINI (24%)

Agapostemon angelicus Cockerell.—Eickwort, 1981 [+PUPA].

Agapostemon cockerelli Crawford.—Eickwort, 1981 [+PUPA].

Agapostemon nasutus Smith.—Eickwort and Eickwort, 1969 [+ PUPA].

Agapostemon sericeus (Forster).—LaBerge and Ribble, 1966a [as A. radiatus]; Eickwort, 1981 [+ PUPA]; McGinley, 1981 [as A. radiatus].

Agapostemon texanus Cresson.—Eickwort, 1981 [+ PUPA].

Agapostemon virescens (Fabricius),—Abrams and Eickwort, 1980 [+ PUPA].

[Halictus (Halictus) farinosus Smith, UTAH.]

Halictus (Halictus) ligatus Say.—Packard, 1897; Michener, 1953; McGinley, 1981.

Halictus (Halictus) parallelus Say.—Packard, 1897; Michener, 1953.

Halictus (Halictus) scabiosae Rossi.—Grandi, 1954, 1961.

Halictus (Halictus) sexcinctus (Fabricius).—Grandi, 1937,1961; Michener, 1953.

Halictus (Seladonia) tripartitus Cockerell.—Michener, 1953, 1954 [PUPA].

Lasioglossum (Dialictus) spinolae (Friese).—Claude-Joseph, 1926.

Lasioglossum (Ctenonomia) albescens sepulchralis Cameron.—Sakagami, 1968 [+ PUPA].

Lasioglossum (Evylaeus) duplex Dalla Torre.—Sakagami and Hayashida, 1960 [+ PUPA].

Lasioglossum (Dialictus) herbstiellum (Friese).—Claude-Joseph, 1926 [as Halictus herbstiellus].

Lasioglossum (Dialictus) imitatum (Smith).—Michener, 1953 [as L. sparsum], 1954 [PUPA, as L. sparsum].

Lasioglossum (Evylaeus) kincaidii (Cockerell).—Michener, 1953, 1954 [PUPA].

Lasioglossum (Hemihalictus) lustrons (Cockerell).—Daly, 1961.

Lasioglossum (Evylaeus) malachurum (Kirby).—Soika, 1934 [+PUPA]; Michener, 1953.

Lasioglossum (Evylaeus) ohei Hiroshima and Sakagami.—Sakagami, Hiroshima, and Ohe, 1966.

Lasioglossum (Dialictus) umbripenne (Ellis).—Eickwort and Eickwort, 1971 [+PUPA].

Lasioglossum (Dialictus) zephyrum (Smith).—(McGinley, 1981).

Lasioglossum (Dialictus) species.—Michener, 1953.

Microsphecodes kathleenae Eickwort.—Eickwort and Eickwort, 1972b [+ PUPA].

Ruizantheda (Ruizanthedella) mutabilis (Spinola).—Claude-Joseph, 1926 [as Halictus mutabilis].

Sphecodes albilabris (Kirby).—Rozen, 1965a [+ PUPA];McGinley, 1981.

Sphecodes species.—Michener, 1953,1954 [+ PUPA; reported as Neopasites species?].

Sphecodes species, near fragariae Cockerell.—Torchio, 1975 [+ PUPA].

Unknown: Archihalictus, [Caenoha lie tus, KU], Dinagapostemon, Echthralictus, Eupetersia, [Habralictus, KU], Homalictus, Madagalictus, Mexalictus, Pachyhalictus, Paragapostemon, Parathrincostoma, Patellapis, [Pseudagapostemon, KU], Ptilocleptis, Rhinetula, Thrincohalictus, Thrinchostoma, Zonalictus.

NOMIOIDINI (0)

Unknown: Ceyalictus, Nomioides.

NOMIINAE (100%)

(Many taxa in this group have variously been treated as genera or subgenera; undoubtedly, there are several valid nomiine genera but the following conservative scheme, i.e., Nomia (S.I.), is used here to avoid additional confusion).

Nomia australica Smith.—Rayment, 1957.

Nomia esakii Hiroshima.—Hiroshima, 1961 [as Rhopalomelissa],

[Nomia heteropoda Say, UTAH.]

Nomia melanderi Cockerell.—Michener, 1953, 1954 [PUPA]; Nielsen and Bohart, 1967 [sex characters]; McGinley, 1981.

Nomia n. nevadensis Cresson.—Michener, 1953; Nielsen and Bohart, 1967 [sex characters].

Nomia punctulata Dalla Torre.—Masuda, 1943; Hiroshima, 1961.

Nomia ruficornis Spinola.—Soika, 1932; Michener, 1953.

Nomia triangulifera Vachal.—Nielsen and Bohart, 1967 [sex characters].

Nomia yasumatsui Hiroshima.—Hiroshima, 1961 [as Rhopalomelissa].

Nomia species.—LaBerge and Ribble, 1966a [as Agapostemon splendens].

ANDRENIDAE (41%)

ANDRENINAE (17%)

Andrena (Callandrena) accepta Viereck.—Rozen, 1973a; McGinley, 1981.

Andrena (Leucandrena) barbilabris (Kirby).—Thorp and Stage, 1968 [+ PUPA; as A. placida Smith].

Andrena (Thysandrena) bisalicis Viereck.—Yager and Rozen, 1966 [PUPA]; Rozen, 1973a.

Andrena (Euandrena) caerulea Smith.—Michener, 1953 [as A. complexa Viereck].

Andrena (Thysandrena) candida Smith.—Youssef and Bohart, 1968 [+ PUPA].

Andrena (Melandrena) carlini Cockerell.—Schrader and LaBerge, 1978.

Andrena (Diandrena) chalybaea (Cresson).—Thorp, 1969.

Andrena (Melandrena) dunningi Cockerell.—Johnson, 1981.

Andrena (Ptilandrena) erigeniae Robertson.—Davis and LaBerge, 1975.

Andrena (Leucandrena) erythronii Robertson.—Michener, 1953; Yager and Rozen, 1966 [PUPA].

Andrena (Scrapteropsis) imitatrix Cresson.—Yager and Rozen, 1966 [PUPA]; Rozen, 1973a.

Andrena (Larandrena) miserabilis Cresson.—Norden and Scarbrough, 1979.

Andrena (Scrapteropsis) morrisonella Viereck.—Rozen, 1973a.

Andrena (Tylandrena) perplexa Smith.—Stephen, 1966 [+ PUPA; as A. viburnella].

Andrena (Melandrena) regidaris Malloch.—Schräder and LaBerge, 1978.

Andrena (Melandrena) vicina Smith.—Packard, 1897 [+ PUPA]; Michener, 1953.

Andrena species.—Michener, 1953.

Unknown: Alocandrena, Ancylandrena, Euherbstia, Megandrena, Orphana.

PANURGINAE (45%)

Acamptopoeum submettallicum (Spinola).—Claude-Joseph, 1926 [as Camptopaeum]; Michener, 1953.

Calliopsis andreniformis Smith.—Michener, 1953; Rozen, 1966a; Yager and Rozen, 1966 [PUPA].

Calliopsis crypta Shinn.—Rozen, 1966a; McGinley, 1981.

Calliopsis rozeni Shinn.—Rozen, 1966a.

Camptopoeum bakeri Rozen.—Rozen, 1988.

Heterosarus boylei (Cockerell).—Rozen, 1966a; McGinley, 1981.

Hypomacrotera callops callops Cockerell and Porter.—Rozen, 1970a [+PUPA]; McGinley, 1981.

Melitturga caudata Perez.—Rozen, 1971.

Melitturga clavicornis (Latreille).—Rozen, 1965a; Yager and Rozen, 1966 [PUPA]; McGinley, 1981.

Meliturgula braunsi Friese.—Rozen, 1968 [+ PUPA]; McGinley, 1981.

Nomadopsis anthidia anthidia (Fowler).—Rozen, 1958; Yager and Rozen, 1966 [PUPA]; Nielsen and Bohart, 1967 [sex characters].

Nomadopsis anthidia lutea Rozen.—Rozen, 1958.

Nomadopsis australior (Cockerell).—Custer, 1929; Michener, 1953; Rozen, 1958.

Nomadopsis barbata Timberlake.—Rozen, 1958.

Nomadopsis boharti Rozen.—Rozen, 1958; Yager and Rozen, 1966 [PUPA].

Nomadopsis comptula (Cockerell).—Rozen, 1958.

Nomadopsis edwardsii (Cresson).—Rozen, 1958.

Nomadopsis filiorum Rozen.—Rozen, 1963.

Nomadopsis fracta Rozen.—Michener, 1953; Rozen, 1958.

Nomadopsis helianthi (Swenk and Cockerell).—Michener, 1953,1954 [PUPA, described as N. euphorbiae (Cockerell)]; Rozen, 1958; Yager and Rozen, 1966 [PUPA].

Nomadopsis hesperia equina (Cockerell).—Rozen, 1958; Yager and Rozen, 1966 [PUPA].

Nomadopsis linsleyi Rozen.—Rozen, 1958.

Nomadopsis micheneri Rozen.—Rozen, 1958.

Nomadopsis personata (Cockerell).—Yager and Rozen, 1966 [PUPA].

Nomadopsis puellae (Cockerell).—Rozen, 1958; Yager and Rozen, 1966 [PUPA]; McGinley, 1981.

Nomadopsis scitula (Cresson).—Yager and Rozen, 1966 [PUPA].

Nomadopsis scutellaris scutellaris (Fowler).—Rozen, 1958; Yager and Rozen, 1966 [PUPA]; Nielsen and Bohart, 1967 [sex characters].

Nomadopsis trifolii Timberlake.—Rozen, 1958.

Nomadopsis zonalis sierrae Rozen.—Rozen, 1963; Yager and Rozen, 1966 [PUPA].

Panurginus albopilosus Lucas.—Rozen, 1971 [+ PUPA].

Panurginus atriceps (Cresson).—Rust, 1976.

Panurginus melanocephalus (Cockerell).—Michener, 1953; Rozen, 1966a.

Panurginus potentillae (Crawford).—Rozen, 1966a; McGinley, 1981.

Panurginus species A.—Rozen, 1966a; Yager and Rozen, 1966 [PUPA].

Panurgus banksianus (Kirby).—Micheli, 1931; Michener, 1953.

Panurgus calcaratus Scopoli.—Micheli, 1936; Michener, 1953.

Panurgus dentipes Latreille.—Rozen and Rozen, 1966; McGinley, 1981.

Panurgus intermedius Rozen.—Rozen, 1971 [+ PUPA].

Panurgus oraniensis Perez.—Rozen, 1971 [+ PUPA].

Panurgus podagricus Perez.—Rozen, 1971.

[Perdita (Cockerellia) albipennis Cresson, UTAH.]

[Perdita (Perdita) bohartorum Timberlake, UTAH.]

Perdita (Perdita) confusa Timberlake.—Rozen, 1966a.

Perdita (Perdita) halictoides Smith.—Eickwort, 1977 [+ PUPA].

Perdita (Perdita) lenis Timberlake.—Rozen, 1966a.

Perdita (Cockerellia) lingualis Cockerell.—Michener, 1963 [+ PUPA]; Rozen, 1966a; Yager and Rozen, 1966 [PUPA]; McGinley, 1981.

Perdita (Perdita) maculigera maculipennis Graenicher.—Michener and Ordway, 1963 [+ PUPA]; Rozen, 1966a; Yager and Rozen, 1966 [PUPA].

Perdita (Perdita) nuda Cockerell.—Torchio, 1975 [+ PUPA].

Perdita (Perdita) octomaculata (Say).—Eickwort, 1977 [+ PUPA].

Perdita (Cockerellula) opuntiae Cockerell.—Custer, 1929; Michener, 1953.

Perdita (Perdita) sexmaculata Cockerell.—Rozen, 1966a.

Perdita (Perdita) zebrata zebrata Cresson.—Custer, 1929; Michener, 1953; Rozen, 1966a.

Protandrena bicolor (Timberlake).—Rozen, 1970 [+ PUPA];McGinley, 1981.

Pseudopanurgus aethiops (Cresson).—Rozen, 1966a.

Pseudopanurgus verticalis Timberlake.—Rozen, 1966a.

Pterosarus occidus (Timberlake).—Rozen, 1966a.

Spinoliella herbsti (Friese).—Claude-Joseph, 1926 [as Camptopaeum]; Michener, 1953.

Spinoliella maculata (Spinola).—Claude-Joseph, 1926 [as Camptopaeum]; Michener, 1953.

Unknown: Anthemurgus, Anthrenoides, Austropanurgus, Callonychium, Cephalurgus, Epimethea, Liopoeum, Liphanthus, Mermiglossa, Metapsaenythia, Parafriesea, Parapsaenythia, Plesiopanurgus, Poecilomelitta, Protomeliturga, Pseudosarus, Rhophitulus, Xenopanurgus.

OXAEIDAE (50%)

Oxaea flavescens Klug.—Roberts, 1973 [+ PUPA]; Torchio and Burwell, 1987 [PUPA].

Protoxaea gloriosa (Fox).—Rozen, 1964a; McGinley, 1981.

Unknown: Mesoxaea, Notoxaea.

MELITTIDAE (29%)

DASYPODINAE (38%)

DASYPODINI (60%)

Capicola braunsiana Friese.—Rozen and McGinley, 1974a; McGinley, 1981.

Dasypoda plumipes Panzer.—Rozen and McGinley, 1974a; McGinley, 1981.

Hesperapis carinata Stevens.—Rozen and McGinley, 1974a.

Hesperapis ilicifoliae (Cockerell).—Rozen and McGinley, 1974a.

Hesperapis nitidula Cockerell.—Rozen and McGinley, 1974a.

Hesperapis pellucida Cockerell.—Rozen and McGinley, 1974a.

Hesperapis regularis (Cresson).—Burdick and Torchio, 1959; Rozen and McGinley, 1974a.

Hesperapis rufipes (Ashmead).—Michener, 1953; Rozen and McGinley, 1974a.

Hesperapis trochanterata Snelling.—Rozen, 1987a.

Hesperapis species A.—Rozen and McGinley, 1974a.

Hesperapis species B.—Rozen and McGinley, 1974a.

Unknown: Eremaphanta, Xeralictoides.

PROMELITTINI (0)

Unknown: Promelitta.

SAMBINI (0)

Unknown: Haplomelitta, Samba.

MEGANOMIINAE (25%)

Meganomia gigas Michener.—Rozen, 1977a [+ PUPA; as M. binghami (Cockerell)]; McGinley, 1981.

Unknown: Agemmonia, Ceratomonia, Uromonia.

MELITTINAE (40%)

Macropis europaea Warncke.—Rozen and McGinley, 1974a [+PUPA]; McGinley, 1981.

Macropis nuda (Provancher).—Rozen and Jacobson, 1980 [+ PUPA].

Melitta leporina (Panzer).—Rozen and McGinley, 1974a [+ PUPA]; McGinley, 1981.

Unknown: Dolichochile, Rediviva, Redivivoides.

CTENOPLECTRIDAE (50%)

Ctenoplectra armata Magretti.—Rozen, 1978; McGinley, 1981.

Unknown: Ctenoplectrina.

MEGACHILIDAE (22%)

FIDELIINAE (100%)

Fidelia villosa Brauns.—Rozen, 1970b [+ PUPA]; McGinley, 1981.

Neofidelia profuga Moure and Michener.—Rozen, 1973b [+ PUPA]; McGinley, 1981.

Parafidelia pallidula Cockerell.—Rozen, 1977b; McGinley, 1981.

LITHURGINAE (66%)

Lithurge apicalis (Cresson).—Rozen, 1973c.

Lithurge atratiformis Cockerell.—Rozen, 1973c; McGinley, 1981.

Lithurge chrysurus Fonscolombe.—Roberts, 1978 [+ PUPA].

Lithurge echinocacti (Cockerell).—Rozen, 1973c [compared to L. apicalis].

Trichothurgus dubius (Herbst).—Claude-Joseph, 1926 [as Lithurge]; Michener, 1953; Rozen, 1973c [+ PUPA].

Trichothurgus echinocacti (Cockerell).—Rozen, 1973c [as Lithurge].

Trichothurgus species—Rozen, 1973c [as Lithurge].

Unknown: Microthurge.

MEGACHILINAE (18%)

ANTHIDIINI (12%)

Anthidium chilense Spinola.—Claude-Joseph, 1926; Michener, 1953.

Anthidium oblongatum Latreille.—Xambeu, 1896; Maneval, 1937; Michener, 1953.

Anthidium 7-dentatum Lepeletier.—Grandi, 1961.

Anthidium species—McGinley, 1981.

Anthidiellum ehrhorni (Cockerell).—Michener, 1953 [as Anthidium ehrhorni].

Anthidiellum perplexum (Smith).—Baker et al., 1985 [PUPA].

Anthidiellum notatum robertsoni (Cockerell).—Michener, 1953.

Dianthidium curvatum sayi Cockerell.—Michener, 1953.

Dianthidium heterulkei heterulkei Schwarz.—Clement, 1976.

Dianthidium species—Michener, 1953; McGinley, 1981.

[Dianthidium spp., UTAH.]

Heterostelis hurdi Thorp.—Thorp, 1966 [+ PUPA].

Odontostelis bilineolata (Spinola).—Rozen, 1966b [+ PUPA; as Stelis]; McGinley, 1981.

Paranthidiellum lituratum (Panzer).—Enslin, 1923; Micheli, 1934; Michener, 1953.

Paranthidium caturigense (Giraud).—Micheli, 1935; Maneval, 1936; Michener, 1953.

Paranthidium septemdentatum (Latreille).—Xambeu, 1896; Grandi, 1934; Michener, 1953.

Stelis chlorocyanea (Cockerell).—Rust and Thorp, 1973 [+ PUPA].

Stelis elongativentris Parker.—Rozen, 1987b.

Stelis lateralis Cresson.—Michener, 1953; Rozen, 1966b.

Stelis minuta Lepeletier.—Enslin, 1925; Michener, 1953.

Stelis nasuta (Latreille).—Maneval, 1937; Michener, 1953.

Stelis ornatula Nylander.—Micheli, 1935; Michener, 1953.

[Stelis spp., UTAH.]

Trachusa perdita Cockerell.—Michener, 1953; McGinley, 1981.

Unknown: Adanthidium, Afranthidium, Allanthidium, Anthidioma, Anthodioctes, Apianthidium, Archianthidium, Asianthidium, Aspidosmia, Atropium, Ausanthidium, Axillanthidium, Aztecanthidium, Bathanthidium, Bellanthidium, Benanthis, Branthidium, Bothranthidium, [Callanthidium, UTAH], Capanthidium, Carinanthidium, Cyphanthidium, [Dolichostelis, UTAH], Domanthidium, Doxanthidium, Epanthidium, Euaspis, Exanthidium, Gnathanthidium, Heteranthidium, Honanthidium, Hypanthidioides, Hypanthidium, Icteranthidium, Immanthidium, Malanthidium, Manthidium, Meganthidium, Melanthidium, Mesanthidiellum, Mesanthidium, Nananthidium, Neanthidium, Nigranthidium, Notanthidium, Oranthidium, Oxyanthidium, Pachyanthidium, Paraanthidium, [Parevaspis, UTAH], Plesianthidium, Pseudoanthidium, Pyctanthidium, Reanthidium, Rhodanthidium, Royanthidium, Serapista, Spinanthidium, Trianthidiellum, Trianthidium, Trichanthidium, Tuberanthidium, Ulanthidium, Xenanthidium.

DIOXINI (100%)

Dioxys cincta (Jurine).—Micheli, 1936; Michener, 1953.

Dioxys pomonae pomonae Cockerell.—Rozen, 1967 [+ PUPA]; McGinley, 1981.

Dioxys productus productus (Cresson).—Rozen, 1967.

MEGACHILINI (75%)

Chalicodoma (Chelostomoides) campanulae campanulae (Robertson).—Baker, Kuhn, and Bambara, 1985.

Chalichodoma (Pseudomegachile) flavipes Spinola.—Moustafa and Mazeed, 1977 [+ PUPA].

Chalicodoma (Callomegachile) mystaceana Michener.—King, 1984 [+ PUPA].

Chalicodoma (Chelostomoides) species.—Michener, 1953, 1954 [PUPA].

[Chalicodoma spp., UTAH.]

Coelioxys elongata Lepeletier.—Iwata, 1939.

[Coelioxys funeraria Smith, UTAH.]

Coelioxys modesta Smith.—Baker et al., 1985 [additional material in USNM].

Coelioxys octodentata Say.—Michener, 1953; Baker, 1971.

Coelioxys sayi Robertson.—Baker, 1971; Baker et al., 1985 [PUPA]; [first instar in USNM].

Coelioxys species.—McGinley, 1981.

Megachile albisecta Klug.—Grandi, 1931; Michener, 1953; Grandi, 1961.

Megachile argentata Fabricius.—Grandi, 1931; Michener, 1953; Grandi, 1961.

Megachile brevis Say.—Michener, 1953, 1954 [PUPA]; McGinley, 1981.

Megachile buyssonii Perez.—Buysson, 1902; Michener, 1953.

Megachile centuncularis (Linnaeus).—Packard, 1897; Buysson, 1902; Grandi, 1934; Michener, 1953; Grandi, 1961; Romasenko, 1983b.

Megachile euzona Perez.—Claude-Joseph, 1926; Michener, 1953.

Megachile lagopoda (Linnaeus).—Buysson, 1902; Michener, 1953; Marikovskaja, 1984.

Megachile ligniseca Kirby.—Romasenko, 1983a.

Megachile macularis Dalla Torre.—King, 1984.

Megachile maritima (Kirby).—Buysson, 1902; Michener, 1953.

Megachile mendica Cresson.—Baker et al., 1985 [+ PUPA].

Megachile montivaga Cresson.—Baker et al., 1985.

Megachile muraria Fabricius.—Grandi, 1934; Michener, 1953; Grandi, 1961.

Megachile nigriventris Schenck.—Micheli, 1937; Michener, 1953.

Megachile policaris Say.—Baker et al., 1985 [+ PUPA; as Eumegachile].

Megachile pollinosa Spinola.—Claude-Joseph, 1926; Michener, 1953.

Megachile pyrenaica pyrenaica Lepeletier.—Buysson, 1902; Michener, 1953.

Megachile pyrenaica alpina Morawitz.—Micheli, 1935; Michener, 1953.

Megachile rancaguensis Friese.—Claude-Joseph, 1926; Michener, 1953.

Megachile rotundata (Fabricius).—Nielsen and Bohart, 1967 [sex characters].

Megachile saulcyi Guerin-Meneville.—Claude-Joseph, 1926; Michener, 1953.

Megachile versicolor Smith.—Danks, 1970.

Megachile species.—King, 1984.

Megachile species.—Michener, 1953.

[Megachile spp., USNM.]

Unknown: Creightoniella.

OSMIINI (23%)

Many unstudied taxa, including nearly all species of Ashmeadiella, are in the collections of the Bee Biology and Systematics Laboratory, Utah State University and the Smithsonian Institution.

Ashmeadiella species.—Michener, 1953; McGinley, 1981.

Heriades carinatus Cresson.—Matthews, 1965.

Heriades crenulatus Nylander.—Grandi, 1934; Grandi, 1961; Michener, 1953.

Hoplitis adunca (Panzer).—Grandi, 1935 [as Osmia]; Michener, 1953; Grandi, 1961 [as Osmia].

Hoplitis anthocopoides (Schenck).—Eickwort, 1973 [+ PUPA].

Hoplitis cylindrica (Cresson).—Baker, Kuhn, and Bambara, 1985.

Hoplitis lepeletieri (Perez).—Micheli, 1933; Michener, 1953.

Hoplitis leucomelanena (Kirby).—Enslin, 1925; Micheli, 1930; Michener, 1953.

Hoplitis loti (Morawitz).—Micheli, 1931; Michener, 1953.

Hoplitis parvula (Dufour and Perris).—Dufour and Perris, 1840; Michener, 1953.

Hoplitis tridentata (Dufour and Perris).—Dufour and Perris, 1840; Michener, 1953.

Hoplitis species—Michener, 1953; McGinley, 1981.

Osmia aurulenta (Panzer).—Marechal, 1926; Michener, 1953.

Osmia lignaria lignaria Say.—Baker, Kuhn, and Bambara, 1985.

Osmia lignaria propinqua Cresson.—Michener, 1953; McGinley, 1981.

Osmia rufa (Linnaeus).—Losinski, 1911; Michener, 1953.

Osmia submicans (Morawitz).—Maneval, 1939; Michener, 1953; Wafa and El Berry, 1972 [+ PUPA]; Moustafa and El Berry, 1976 [+ PUPA].

Proteriades bunocephala (Michener).—Thorp, 1968.

Proteriades xerophila (Cockerell).—Michener, 1953.

Unknown: Archeriades, Bytinskia, [Chelostoma, UTAH], Hofferia, Jaxartinula, Kumobia, Megaloheriades, Metallinella, Noteriades, Ochreriades, Othinosmia, [Prochelostoma, USNM], [Protosmia, UTAH], Pseudoheriades, Stenosmia, Wainia, Xeroheriades.

ANTHOPHORIDAE (39%)

ANTHOPHORINAE (38%)

ANCYLINI (0)

Unknown: Ancyla, Tarsalia.

ANTHOPHORINI (33%)

Amegilla crinipes (Smith).—Radovic and Krunic, 1977.

Amegilla parietina fulvocinerea (Dours).—Radovic and Krunic, 1977.

Amegilla pulchra (Smith).—Cardale, 1968 [+ PUPA].

Amegilla species.—Cardale, 1968 [+ PUPA].

Anthophora abrupta Say.—Michener, 1953; Norden, 1984 [+ PUPA].

Anthophora bomboides stanfordiana Cockerell.—Michener, 1953 [as A. stanfordiana]; McGinley, 1981; Brooks, 1983 [PUPA].

Anthophora borealis Morawitz.—Popova, 1984.

Anthophora edwardsii Cresson.—Michener, 1953.

Anthophora furcata syringae (Cockerell).—Michener, 1953, 1954 [+ PUPA].

Anthophora incerta Spinola.—Claude-Joseph, 1926 [as Podalirius]; Michener, 1953.

Anthophora linsleyi Timberlake.—Michener, 1953, 1954 [+ PUPA].

Anthophora occidentalis Cresson.—Nielsen and Bohart, 1967 [sex characters].

Anthophora personata (Illiger).—Semichon, 1922 [+ PUPA]; Michener, 1953, 1954 [PUPA].

Anthophora urbana Cresson.—Michener, 1953.

Anthophora villosula Smith.—Torikata, 1931; Michener, 1953.

Emphoropsis miserabilis (Cresson).—Torchio and Stephen, 1961 [+ PUPA]; McGinley, 1981.

Unknown: Deltoptila, Elaphropoda, Habrophorula, Habropoda, Heliophila, Pachymelus.

CANEPHORULINI (0)

Unknown: Canephorula.

CENTRIDINI (66%)

Centris aenea Lepeletier.—Rozen, 1965b.

Centris cineraria (Smith).—Claude-Joseph, 1926; Michener, 1953.

Centris derasa Lepeletier.—Rozen, 1965b.

Centris lanipes (Fabricius).—Rozen, 1965b; McGinley, 1981.

Centris rufosuffusa Cockerell.—Rozen, 1965b.

[Centris spp., UTAH.]

Epicharis fasciata Lepeletier and Serville.—Rozen, 1965b; McGinley, 1981.

Epicharis rustica (Olivier).—Rozen, 1965b.

Epicharis rustica flava (Olivier).—Camargo, Zucchi, and Sakagami, 1975 [+ PUPA].

Unknown: Ptilotopus.

ERICROCIDINI (15%)

Acanthopus splendidus urichi Cockerell.—Rozen, 1969; McGinley, 1981.

Mesoplia rufipes (Perty).—Rozen, 1969.

Unknown: Abromelissa, Aglaomelissa, Ctenioschelus, Cyphomelissa, Epiclopus, Ericrocis, Eurytis, Hopliphora, Mesocheira, Mesonychium, Oxynedys.

EUCERINI (31%)

Alloscirtetica gayi (Spinola).—Claude-Joseph, 1926 [as Tetralonia chilensis Herbst]; Michener, 1953.

Alloscirtetica tristrigata (Spinola).—Claude-Joseph, 1926 [as Tetralonia]; Michener, 1953.

Eucera sociabilis Smith.—Masuda, 1946 [+ PUPA; as E. difficilis Perez].

Florilegus condignus (Cresson).—LaBerge and Ribble, 1966b.

[Melissodes agilis Cresson, UTAH.]

Melissodes pallidisignata Cockerell.—Rozen, 1965b; McGinley, 1981.

Melissodes persimilis Cockerell.—Buchmann and Jones, 1980.

Melissodes robustior Cockerell.—Rozen, 1965b.

Melissodes rustica (Say).—Clement, 1973.

Melissodes species.—Michener, 1953.

Peponapis fervens (Smith).—Rozen, 1965b; McGinley, 1981.

Svastra obliqua obliqua (Say).—Rozen, 1964b; McGinley, 1981.

Svastrides melanura (Spinola).—Claude-Joseph, 1926; Michener, 1953.

Tetralonia hamata Bradley.—Miliczky, 1985.

Tetralonia lanuginosa Klug.—Mohamed, 1974 [+ PUPA].

Tetralonia malvae Rossi.—Grandi, 1954, 1961.

Thygater species.—Packer, 1987.

Xenoglossa angustior Cockerell.—Rozen, 1965b; McGinley, 1981.

Xenoglossa fulva Smith.—Rozen, 1965b.

Xenoglossa strenua (Cresson).—Bohart, 1964; Rozen, 1965b.

Unknown: Anthedonia, Cemolobus, Dithygater, Eucara, Gaesischia, Gaesochira, Lophothygater, Loxoptilus, Martinapis, Melissina, Melissoptila, Megascirtetica, Micronychapis, Pachysvastra, Pectenapis, Simanthedon, Svastrina, Syntrichalonia, Tetraloniella, Thygatina, Trichocerapis, Xenoglossodes.

EUCERINODINI (0)

Unknown: Eucerinoda.

EXOMALOPSINI (40%)

Exomalopsis caerulea Friese.—Claude-Joseph, 1926; Michener, 1953; Rozen, 1957.

Exomalopsis chionura Cockerell.—Rozen, 1957; McGinley, 1981.

Lanthanomelissa species—Lucas de Oliveira, 1966a.

Paratetrapedia gigantea (Schrottky).—Lucas de Oliveira, 1962.

[Paratetrapedia swainsonae (Cockerell), KU.]

Tapinotaspis caerulea (Friese).—Lucas de Oliveira, 1962.

Unknown: [Ancyloscelis, UTAH], Caenonomada, Chalepogenus, Chilimalopsis. [Eremapis, UTAH], Monoeca.

MELECTINI (57%)

Melecta armata Panzer.—Semichon, 1922 [+ PUPA]; Michener, 1953, 1954 [PUPA].

Melecta luctuosa Sopoli.—Soika, 1936 [first instar].

Melecta pacifica Cresson.—Torchio and Youssef, 1968.

Melecta separata callura (Cockerell).—Rozen, 1969; McGinley, 1981.

Thyreus caeruleopunctatus (Blanchard).—Cardale, 1968 [PUPA].

Thyreus lieftincki Rozen.—Rozen, 1969; McGinley, 1981.

Thyreus lugubrus (Smith).—Cardale, 1968.

Thyreus species—Rozen, 1969.

Xeromelecta californica (Cresson).—Michener, 1953, 1954 [PUPA]; Rozen, 1969.

Zacosmia maculata (Cresson).—Torchio and Youssef, 1968 [+ PUPA]; Rozen, 1969; McGinley, 1981.

Unknown: Brachymelecta, Nesomelecta, Tetralonoidiella.

MELITOMINI (100%)

Diadasia chilensis Spinola.—Claude-Joseph, 1926 [as Melitoma]; Michener, 1953.

[Diadasia consociata Timberlake, UTAH.]

Diadasia diminuta (Cresson).—McGinley, 1981.

Diadasia enevata (Cresson).—Michener, 1953, 1954 [+ PUPA]; Nielsen and Bohart, 1967 [sex characters].

Melitoma euglossoides Lepeletier and Serville.—McGinley, 1981.

Ptilothrix bombiformis (Cresson).—Michener, 1953 [as Emphor bombiformis], 1954 [+ PUPA].

PARARHOPHITINI (100%)

Pararhophites orobinus (Morawitz).—McGinley and Rozen, 1987.

RHATHYMINI (100%)

Rhathymus bicolor Lepeletier.—Rozen, 1969; McGinley, 1981.

Rhathymus trinitatis Cockerell.—Rozen, 1969.

Rhathymus species—Camargo, Zucchi, and Sakagami, 1975 [+ PUPA].

TETRAPEDIINI (0)

[Tetrapedia, UTAH.]

NOMADINAE (40%)

AMMOBATINI (33%)

Ammobates carinatus Morawitz.—Rozen and McGinley, 1974b; McGinley, 1981.

Morgania histrio transvaalensis Bischoff.—Rozen and McGinley, 1974b [+ PUPA]; McGinley, 1981.

Oreopasites vanduzeei Cockerell.—Rozen, 1954, 1966c; Rozen and McGinley, 1974b [+ PUPA]; McGinley, 1981.

Pseudodichroa fumipennis Bischoff.—Rozen and McGinley, 1974b; McGinley, 1981.

Unknown: Caesarea, Melanempis, Omachthes, Parammobatodes, Pasites, Pasitomachthes, Pseudopasites, Sphecodopsis.

AMMOBATOIDINI (0)

Unknown: Ammobatoides.

BIASTINI (50%)

Neopasites cressoni Crawford.—Rozen, 1966c; McGinley, 1981.

Unknown: [Biastes, UTAH].

CAENOPROSOPIDINI (0)

Unknown: Caenoprosopidis.

EPEOLINI (66%)

Doeringiella gayi (Spinola).—Claude-Joseph, 1926 [as Epeolus]; Michener, 1953.

Epeolus pusillus Cresson.—Rozen, 1966c.

Epeolus tristis Smith.—Mayet, 1875 [+ PUPA]; Michener, 1953.

Epeolus species A.—Michener, 1953; Rozen, 1966c; McGinley, 1981.

Odyneropsis apicalis Ducke.—Rozen, 1966c.

Triepeolus dacotensis (Stevens).—Nielsen and Bohart, 1967 [sex characters].

[Triepeolus helianthi (Robertson), UTAH.]

Triepeolus mesillae Cockerell.—Rozen, 1966c.

Triepeolus remigatus (Fabricius).—Rozen, 1966c.

Triepeolus species A.—Rozen, 1966c; McGinley, 1981.

Triepeolus species B.—Rozen, 1966c.

Triepeolus species C.—Michener, 1953; Rozen, 1966c.

Unknown: Thalestria, Trophocleptria.

EPEOLOIDINI (0)

Unknown: Epeoloides.

HOLCOPASITINI (50%)

Holcopasites calliopsidis (Linsley).—Rozen, 1966c.

Holcopasites insoletus (Linsley).—Rozen, 1966c; McGinley, 1981.

Holcopasites species A.—Rozen, 1966c.

Unknown: Schmiedeknechtia.

ISEPEOLINI (100%)

Isepeolus luctuosus (Spinola).—Claude-Joseph, 1926; Michener, 1953.

Isepeolus viperinus (Holmberg).—Michener, 1957; Rozen, 1966c; Lucas de Oliveira, 1966b; McGinley, 1981.

NEOLARRINI (100%)

Neolarra pruinosa Ashmead.—Rozen, 1966c; McGinley, 1981.

NOMADINI (71%)

Kelita tuberculata Ehrenfeld and Rozen.—Ehrenfeld and Rozen, 1977.

Melanomada sidaefloris (Cockerell).—Rozen, 1977c [+ PUPA]; McGinley, 1981.

Melanomada annectens Snelling and Rozen.—Snelling and Rozen, 1987.

Nomada obscurella Fowler.—Michener, 1953 [as N. fowleri]; Rozen, 1966c; McGinley, 1981.

Nomada imbricata Smith.—Packard, 1897 [+ PUPA]; Michener, 1953.

Nomada japonica Smith.—Masuda, 1946 [+ PUPA].

Nomada suavis Cresson.—Rozen, 1966c.

Nomada species A.—Rozen, 1966c.

Nomada species B.—Rozen, 1966c.

Nomada species C.—Rozen, 1966c.

Paranomada velutina Linsley.—Rozen, 1977c [+ PUPA]; McGinley, 1981.

Triopasites penniger (Cockerell).—Rozen, 1977c.

Unknown: Brachynomada, Hexepeolus.

OSIRINI (0)

Unknown: Osiris.

PROTEPEOLINI (50%)

Protepeolus singularis Linsley and Michener.—Rozen, Eickwort, and Eickwort, 1978 [+ PUPA]; McGinley, 1981.

Unknown: Leiopodus.

TOWNSENDIELLINI (0)

Unknown: Townsendiella.

XYLOCOPINAE (74%)

CERATININI (81%)

Allodape ceratinoides Gribodo.—Brauns, 1926; Michener, 1953, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape collaris Vachal.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape dapa Strand.—Michener, 1975.

Allodape exoloma Strand.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape friesei Strand.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape interrupta Vachal.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape mucronata Smith.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape panurgoides Smith.—Michener, 1975; McGinley, 1981 [as A. crinita Brauns MS].

Allodape pernix (Bingham).—Michener, 1975.

Allodape pictifrons Smith.—Michener, 1975.

Allodape quadrilineata (Cameron).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape rufogastra Lepeletier and Serville.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape stellarum Cockerell.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodape species.—Friese, 1914; Brauns, 1926; Michener, 1953.

Allodapula acutigera Cockerell.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodapula dichroa (Strand).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodapula hessei Michener.—Michener, 1975.

Allodapula melanopus (Cameron).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Allodapula ornaticeps Michener.—Michener, 1975.

Allodapula rozeni Michener.—Michener, 1975.

Allodapula variegata (Smith).—Brauns, 1926 [as Allodape pringlei]; Michener, 1953, 1975; Michener and Scheiring, 1976 [PUPA].

Allodapula xerica Michetner.—Michenesr, 1975.

Braunsapis acuticauda Michener.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapins albitarsis (Friese).—Michener, 1975.

Braunsapis bouyssoui (Vachal).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis draconis Michener.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis elizabethana (Strand).—Michener, 1975.

Braunsapis facialis (Gerstaecker).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis foveata (Smith).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis ghanae Michener.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis gorillarum (Cockerell).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis iwatai (Sakagami and Yoshikawa).—Sakagami and Yoshikawa, 1961 [as Allodape iwatai].

Braunsapis leptozonia (Vachal).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis luapulana (Cockerell).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis natalica Michener.—Michener, 1975.

Braunsapis nitida (Smith).—Michener and Syed, 1962 [as Allodapula].

Braunsapis pallida Michener.—Michener, 1975.

Braunsapis paradoxa (Strand).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis perkinsiella (Michener and Syed).—Michener and Syed, 1962 [as Allodapula].

Braunsapis plebeia (Cockerell).—Michener and Syed, 1962 [as Allodapula].

Braunsapis rhodesi (Cockerell).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis rolini (Vachal).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis sauteriella Cockerell.—Yasumatsu, 1938 [as Allodape marginata Cockerell]; Michener, 1953 [as A. marginata]; Sakagami and Yoshikawa, 1961 [as Allodape sauteriella].

Braunsapis simillima (Smith).—Michener and Syed, 1962 [as Allodapula].

Braunsapis simplicipes Michener.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis strandi (Masi).—Masi, 1930 [as Allodape]; Michener, 1953.

Braunsapis stuckenbergorum Michener.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis trochanterata (Gerstaecker).—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Braunsapis unicolor (Smith).—Michener and Syed, 1962 [as Allodapula]; Michener, 1962 [+ PUPA].

Braunsapis vitrea (Vachal).—Michener, 1975.

Ceratina callosa (Fabricius).—Micheli, 1936; Michener, 1953.

Ceratina chalcites Latreille.—Grandi, 1957.

Ceratina cucurbitina (Rossi).—Grandi, 1935; Michener, 1953.

Ceratina dupla Say.—Packard, 1897; Michener, 1953.

Ceratina species.—McGinley, 1981.

[Ceratina spp., UTAH.]

Compsomelissa stigmoides (Michener).—Michener, 1976; Michener and Scheiring, 1976 [PUPA].

Eucondylops konowi Brauns.—Michener, 1975.

Eucondylops reducta Michener.—Michener and Scheiring, 1976 [PUPA].

Exoneura apposita Rayment.—Rayment, 1949b; Michener, 1953.

Exoneura concava Rayment.—Rayment, 1949b; Michener, 1953.

Exoneura concinnula Cockerell.—Syed, 1963; Michener and Scheiring, 1976 [PUPA].

Exoneura frogattii Cockerell.—Rayment, 1949a; Michener, 1953.

Exoneura fultoni Cockerell.—Rayment, 1949a; Michener, 1953.

Exoneura hamulata Cockerell.—Rayment, 1951; Michener, 1953; Syed, 1963; Michener and Scheiring, 1976 [PUPA].

Exoneura illustris Erickson and Rayment.—Erickson and Rayment, 1951; Michener, 1953.

Exoneura montana Rayment.—Rayment, 1949b; Michener, 1953.

Exoneura obliterata Cockerell.—Rayment, 1949a; Michener, 1953.

Exoneura obscuripes Michener.—Syed, 1963.

Exoneura pictifrons Alfken.—Erickson and Rayment, 1951; Michener, 1953.

Exoneura richardsoni Rayment.—Rayment, 1951; Michener, 1953.

Exoneura roddi Rayment.—Rayment, 1949b; Michener, 1953.

Exoneura rufitarsis Rayment.—Rayment, 1951 [+ PUPA]; Michener, 1953.

Exoneura setosa Houston.—Houston, 1976 [+ PUPA].

Exoneura simillima Rayment.—Rayment, 1949a; Michener, 1953.

Exoneura subbaculifera Rayment.—Rayment, 1951; Michener, 1953; Syed, 1963.

Exoneura subholmesi Rayment.—Rayment, 1949b; Michener, 1953.

Exoneura variabilis Rayment.—Rayment, 1949b; Michener, 1953; Syed, 1963; Michener and Scheiring, 1976 [PUPA].

Exoneurella eremophila (Houston).—Houston, 1976 [+ PUPA; as Exoneura].

Exoneurella lawsoni (Rayment).—Syed, 1963 [this and following citations as Exoneura]; Michener, 1964; Michener and Scheiring, 1976 [PUPA].

Exoneurella tridentata (Houston).—Houston, 1976 [+ PUPA; as Exoneura].

Halterapis angustula (Cockerell).—Michener, 1976; Michener and Scheiring, 1976 [PUPA].

Halterapis nigrinervis (Cameron).—Michener, 1976; Michener and Scheiring, 1976 [PUPA].

Inquilina excavata (Cockerell).—Rayment, 1949b [also included as Exoneura concava]; Michener, 1953 [also as E. concava]; Syed, 1963; Michener and Scheiring, 1976 [PUPA].

Macrogalea candida (Smith).—Michener, 1976; Michener and Scheiring, 1976 [PUPA].

Manuelia gayatina (Spinola).—Claude-Joseph, 1926; Michener, 1953.

Nasutapis straussorum Michener.—Michener, 1975; Michener and Scheiring, 1976 [PUPA].

Unknown: Effractapis, Exoneuridia, Pithitis.

XYLOCOPINI (33%)

Xylocopa aestuans (Linnaeus).—Dover, 1924 [+ PUPA]; Michener, 1953.

Xylocopa augusti Lepeletier.—Lucas de Oliveira, 1974.

Xylocopa brasilianorum Linnaeus.—Lucas de Oliveira, 1974.

Xylocopa ciliata Burmaister.—Lucas de Oliveira, 1974.

Xylocopa frontalis Oliver.—Lucas de Oliveira, 1974; McGinley, 1981.

Xylocopa hirsutissima Maidl.—Lucas de Oliveira, 1974.

Xylocopa iris Christ.—Grandi, 1957, 1961.

Xylocopa macrops Lepeletier.—Lucas de Oliveira, 1974.

Xylocopa mordax Smith.—Lucas de Oliveira, 1974.

Xylocopa nogueirai Hurd and Moure.—Lucas de Oliveira, 1974.

Xylocopa subcyanea Perez.—Lucas de Oliveira, 1974.

Xylocopa submordax Cockerell.—Lucas de Oliveira, 1974.

Xylocopa violacea (Linnaeus).—Lucas, 1868, 1888; Grandi, 1934; Michener, 1953; Janvier, 1977.

Xylocopa virecens Lepeletier.—Lucas de Oliveira, 1974.

Xylocopa virginica (Linnaeus).—Packard, 1897; Michener, 1953, 1954 [PUPA].

Unknown: Lestis, Proxylocopa.

APIDAE (72%)

APINAE (100%)

Apis cerana Fabricius.—Torchio and Torchio, 1975.

Apis dorsata Fabricius.—Torchio and Torchio, 1975.

Apis florea Fabricius.—Torchio and Torchio, 1975.

Apis mellifera Linnaeus.—Nelson, 1924; Grandi, 1934; Michener, 1953, 1954 [PUPA]; Grandi, 1961; Dade, 1962; Nielsen and Bohart, 1967 [sex characters]; Torchio and Torchio, 1975; McGinley, 1981.

BOMBINAE (62%)

BOMBINI (100%)

Bombus agrorum (Fabricius).—Stephen and Koontz, 1973; Cumber, 1949.

Bombus agrorum pascuorum Scopoli.—Grandi, 1937; Michener, 1953.

Bombus appositus Cresson.—Stephen and Koontz, 1973.

Bombus atratus Franklin.—Lucas de Oliveira, 1979 [+ PUPA].

Bombus bimaculatus Cresson.—Ritcher, 1933; Michener, 1953.

Bombus brasiliensis Lepeletier.—Lucas de Oliveira, 1979 [+ PUPA].

Bombus californicus Smith.—Stephen and Koontz, 1973.

Bombus caliginosus (Frison).—Stephen and Koontz, 1973.

Bombus crotchii Cresson.—Stephen and Koontz, 1973.

Bombus deuteronynus Schulz.—Sakagami, 1951 [as B. senilis Smith]; Michener, 1953 [as B. senilis].

Bombus fervidus (Fabricius).—Packard, 1866, 1897; Ritcher, 1933; Michener, 1953.

Bombus griseocollis (Degeer).—Ritcher, 1933; Michener, 1953; Stephen and Koontz, 1973.

Bombus hortorum (Linnaeus).—Cumber, 1949.

Bombus hypnorum (Linnaeus).—Stephen and Koontz, 1973.

Bombus hyperboreus Schonherr.—Stephen and Koontz, 1973.

Bombus impatiens Cresson.—Ritcher, 1933; Michener, 1953; Stephen and Koontz, 1973.

Bombus lapidarius (Linnaeus).—Cumber, 1949.

Bombus lucorum (Linnaeus).—Cumber, 1949.

Bombus mastrucatus Gerstaecker.—Moczar, 1938; Michener, 1953.

Bombus morio (Swederus).—Lucas de Oliveira, 1979.

Bombus morrisoni Cresson.—Stephen and Koontz, 1973.

Bombus nevadensis Cresson.—Stephen and Koontz, 1973.

Bombus nevadensis auricomus (Robertson).—Ritcher, 1933; Michener, 1953; Stephen and Koontz, 1973.

Bombus pennsylvanicus (Degeer).—Ritcher, 1933; Michener, 1953, 1954 [PUPA; as B. americanorum]; Stephen and Koontz, 1973; McGinley, 1981.

Bombus pennsylvanicus sonorus Say.—Stephen and Koontz, 1973.

Bombus perplexus Cresson.—Ritcher, 1933; Michener, 1953.

Bombus pratorum (Linnaeus).—Cumber, 1949.

Bombus ruderarius (Muller).—Cumber, 1949.

Bombus silvarum Linnaeus.—Grandi, 1934; Michener, 1953.

Bombus sylvicola Kirby.—Stephen and Koontz, 1973.

Bombus terrestris (Linnaeus).—Cumber, 1949.

Bombus terricola Kirby.—Ritcher, 1933; Michener, 1953; Stephen and Koontz, 1973.

Bombus terricola occidentalis Greene.—Stephen and Koontz, 1973.

Bombus vagans Smith.—Ritcher, 1933; Michener, 1953.

Bombus vandykei (Frison).—Stephen and Koontz, 1973.

Bombus vosnesenskii Radoszkowski.—Ritcher, 1933; Michener, 1953, 1954 [PUPA]; Stephen and Koontz, 1973.

Psithyrus rupestris (Fabricius).—Cumber, 1949.

Psithyrus variabilis (Cresson).—Ritcher, 1933; Michener, 1953.

[Psithyrus suckleyi (Greene), UTAH.]

EUGLOSSINI (50%)

Euglossa intersecta Latreille.—Zucchi, de Oliveira, and de Camargo, 1969 [+ PUPA].

Euglossa imperialis Cockerell.—Roberts and Dodson, 1967.

Eulaema nigrita Lepeletier.—Zucchi, Sakagami, and de Camargo, 1970.

Euplusia violacea (Blanchard).—Michener, 1953 [as Centris (Euplusia) violacea].

Unknown: Eufriesea, [Exaerete, UTAH], Aglae.

MELIPONINAE (78%)

Hypotrigona muelleri (Friese).—Lucas de Oliveira, 1970.

Lestrimelitta ehrhardti Friese.—Lucas de Oliveira, 1968 [+ PUPA].

Lestrimelitta limao (Smith).—Lucas de Oliveira, 1968 [+ PUPA].

Melipona fasciata trinitatis Cockerell.—McGinley, 1981.

Melipona marginata Lepeletier.—Michener, 1953.

Melipona nigra schencki Gribodo.—Lucas de Oliveira, 1960.

Melipona quadrifasciata quadrifasciata Lepeletier.—Michener, 1953.

Melipona variegatipes Gribodo.—Michener, 1953.

Nannotrigona postica (Latreille).—Lucas, 1958.

Partamona cupira Smith.—Michener, 1953 [as Trigona], 1954 [PUPA].

Partamona testacea (Klug).—McGinley, 1981 [as Trigona].

Plebeia droryana (Friese).—Lucas de Oliveira, 1965.

Trigona corvina Cockerell.—Michener, 1953.

Unknown: Dactylurina, Meliponula.

Appendix

This section lists the apparent larval synapomorphies supporting the taxa listed in Figure 2 (in upper case, below). Many of these taxa, while clearly defined by adult characters, are only tenuously supported by presumed larval synapomorphies; below, these characters are followed by a question mark. Future research may result in greater resolution based on larval features; this treatment should be considered preliminary.

COLLETIDEA.—DIPHAGLOSSINAE: elongate, cylindrical salivary lips (McGinley, 1981, figs. 2, 17); COLLETINI: extremely broad, unproduced spiracular atria? (McGinley, 1981, fig. 63); PARACOLLETINI: maxilla greatly exceeding labium in lateral view (McGinley, 1981, figs. 29, 33, 38, 45); Scrapter: genal area conspicuously expanded (McGinley, 1981, fig. 51); EURYGLOSSINAE: labiomaxillary region strongly recessed? (McGinley, 1981, figs. 66, 74); XEROMELISSINAE: frontal swellings above antennae present, maxillary palpi enlarged (McGinley, 1981, fig. 80); HYLAEINAE: salivary lips absent with labiomaxillary region produced? (McGinley, 1981, figs. 86, 87).

STENOTRITIDAE.—Mandibular apex obliquely truncate? (Houston, 1975, fig. 15).

HALICTIDAE.—DUFOUREINAE: venter of abdominal segment IX strongly protuberant (Eickwort et al., 1986, fig. 10); NOMIINAE: dorsal tubercles well developed, conical (Michener, 1953, fig. 68), mandibular apex usually bifid? (Michener, 1953, fig. 71; Hirashima, 1961, figs. 61–66); HALICTINAE: maxillary palpi apparently absent (Michener, 1953, fig. 51).

OXAEIDAE.—Primary tracheal opening slit-like, not circular (Rozen, 1964a, fig. 6), labral apex cleft (Rozen, 1964a, fig. 2).

ANDRENIDAE.—Salivary opening appearing U-shaped in frontal view (Rozen, 1966a, fig. 5; Rozen, 1973a, fig. 5; see Torchio (1975) for discussion).

MELITTIDAE.—MEGANOMIINAE: area immediately behind posterior mandibular articulation projecting (Rozen, 1977a, figs. 18, 19), spiracular subatrial wall spinose (Rozen, 1977a, fig. 20); MELITTINAE: salivary plate present (Rozen and McGinley, 1974a, figs. 10, 21); DASYPODINAE: salivary lips absent? (Rozen and McGinley, 1974a, figs. 29, 35, 46).

CTENOPLECTRIDAE.—Monophyly based on larval characters alone is questionable (see Rozen, 1978).

MEGACHILIDAE.—MEGACHILINAE: conspicuous body setae present? (see Rozen, 1978:644); LITHURGINAE: large, conspicuous tooth at the base of the apical mandibular concavity (Rozen, 1973c, figs. 7, 19, 21); FIDELIINAE: monophyly based on larval characters alone is questionable (two of three known generic representatives share with the Lithurginae a conspicuous, adoral tooth; see Rozen, 1977b).

ANTHOPHORIDAE.—PARARHOPHITINI: the traditional classification of this taxon as an anthophorid is probably incorrect; more likely associated with the megachiloid lineage (McGinley and Rozen, 1987). EXOMALOPSINI: body slender, venter of abdominal segment IX protuberant? (Rozen, 1957, fig. 4), maxillary apices not bent mesiad? (Rozen, 1957, fig. 1); MELITOMINI: body extremely slender, venter of abdominal segment IX protuberant (Michener, 1953, figs. 206, 213), maxillary apices strongly bent mesiad (Michener, 1953, figs. 208, 215; most likely plesiomorphic); at present, this tribe is difficult to differentiate from the Exomalopsini on the basis of larval synapomorphies. EUCERINI: larvae extremely uniform but difficult to define cladistically? (see Rozen, 1965b); CENTRITINI: anus situated dorsally on segement X, mandibular concavity extremely broad and scoop-shaped? (Rozen, 1965b); ANTHOPHORINI: monophyly based on larval characters questionable? (salivary lips absent, mandibular concavity scoop-shaped, Michener, 1953, figs. 220, 223); MELECTINI: labiomaxillary region recessed (Rozen, 1969, figs. 4, 12, 26); ERICROCIDINI and RATHYMINI: labiomaxillary region greatly enlarged (Rozen, 1969, figs. 33, 40, 47); NOMADINAE: position of posterior tentorial pits and/or posterior arms modified, mandibles with simple apex and reduced cusp (Rozen et al., 1978); XYLOCOPINAE: salivary opening on dorsal surface of labium (hidden from view, Michener, 1953, figs. 177, 178).

APIDAE.—APINAE (except Apis): conical, often highly sclerotic and darkly pigmented, dorsal papillae on thorax (Michener, 1953, figs. 248, 266).

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1968.   Immature Stages of Australian Anthophorinae (Hymenoptera: Apoidea). Journal of the Australian Entomological Society, 7:35–41.

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1926.   Recherches Biologiques sur les Hymenopteres du Chili (Melliferes). Annales des Sciences Naturelles, Zoologie, series 10, 9:113–268.

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1973.   The Nesting Biology of Melissodes (Eumelissodes) rustica (Say), with a Description of the Larva (Hymenoptera: Anthophoridae). 46:516–525.

1976.   The Biology of Dianthidium heterulkei Schwarz, with a Description of the Larva (Hymenoptera, Megachilidae). The Wasmann Journal of Biology, 34:9–22.

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1949.   Larval Specific Characters and Instars of English Bombidae. Proceedings of the Royal Entomological Society of London, 24:14–19.

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1929.   Habits of Perdita zebrata with Description of Larva. Canadian Entomologist, 61:49–51.

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1962.   Anatomy and Dissection of the Honey Bee. 158 pages. Buckinghamshire, England: Bee Research Association Publishers.

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1961.   Biological Observations on Hemihalictus lustrans, with a Description of the Larva (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 34:134–141.

Daly, H.V., and R.E. Coville

1982.   Hylaeus pubescens and Associated Arthropods at Kilauea, Hawaii Volcanoes National Park (Hymenoptera: Apoidea and Chalcidoidea; Mesostigmata: Ameroseiidae). Proceedings of the Hawaiian Entomological Society, 24:75–81.

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1970.   Biology of Some Stem-Nesting Aculeate Hymenoptera. Transactions of the Royal Entomological Society of London, 122:323–399.

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1975.   The Nest Biology of the Bee Andrena (Ptilandrena) erigeniae Robertson (Hymenoptera: Andrenidae). Illinois Natural History Survey, Biological Notes, 95:1–16.

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1924.   Some Observations on the Bionomics of Xylocopa aestuans Linn. (Apidae). Transactions of the Entomological Society of London, 73:144–150.

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1840.   Sur les Insectes Hymenopteres qui nichent dans l'interieur de tiges seches de la ronce. Annales de la Société Entomologique de France, 9:5–53.

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1977.   The Cuckoo Bee Genus Kelita, Its Systematics, Biology, and Larvae. American Museum Novitates, 2631: 24 pages.

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1967.   Aspects of the Biology of Chilicola ashmeadi in Costa Rica (Hymenoptera: Colletidae). Journal of the Kansas Entomological Society, 40:42–73.

1973.   Biology of the European Mason Bee, Hoplitis anthocopoides (Hymenoptera: Megachilidae), in New York State. Search: Agriculture, 3:1–31.

1977.   Aspects of the Nesting Biology and Descriptions of Immature Stages of Perdita octomaculata and P. halictoides (Hymenoptera: Andrenidae). Journal of the Kansas Entomological Society, 50:577–599.

1979.   A New Species of Wood-Dwelling Sweat Bee in the Genus Neocorynura, with Description of Its Larva and Pupa (Hymenoptera: Halictidae). Entomologia Generalis, 5:143–148.

1981.   Aspects of the Nesting Biology of Five Nearctic Species of Agapostemon (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 54:337–351.

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1969.   Aspects of the Biology of Costa Rican Halictine Bees, I: Agapostemon nasutus (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 42:421–452.

1971.   Aspects of the Biology of Costa Rican Halictine Bees, II: Dialictus umbripennis and Adaptations of Its Caste Structure to Different Climates. Journal of the Kansas Entomological Society, 44:343–373.

1972a. Aspects of the Biology of Costa Rican Halictine Bees, IV: Augochlora (Oxystoglossela) (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 45:18–45.

1972b. Aspects of the Biology of Costa Rican Halictine Bees, III: Sphecodes kathleenae, a Social Cleptoparasite of Dialictus umbripennis (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 45:529–541.

1973a. Notes on the Nests of Three Wood-Dwelling Species of Augochlora from Costa Rica (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 46:17–22.

1973b. Aspects of the Biology of Costa Rican Halictine Bees, V: Augochlorella edentata (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 46:3–16.

Eickwort, G.C., P.F. Kukuk, and F.R. Wesley

1986.   The Nesting Biology of Dufourea novaeangliae (Hymenoptera: Halictidae) and the Systematic Position of the Dufoureinae Based on Behavior and Development. Journal of the Kansas Entomological Society, 59:103–120.

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1921.   Beitrage zur Kenntnis der Hymenopteren, I: Biologie von Rhophites canus Evers. Deutsche Entomologische Zeitschrift, 1921:59–65.

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1925.   Beitrage zur Kenntnis der Hymenopteren, IV. Deutsche Entomologische Zeitschrift, 1925:177–210.

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1951.   Simple Social Bees of Western Australia. Western Australian Naturalist, 3:45–59.

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1914.   Bienenlarven mit Pseudopodien und neue Arten der Gattung Allodape (Hym.). Deutsche Entomologische Zeitschrift, 1914:144–150.

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1931.   Contributi alla conoscenza biologica e morfologica degli imenotteri melliferi e predatori XII. Bollettino del Laboratorio Entomologia del R. Istituto Superiore Agrario Bologna, 4:19–72.

1934.   Contributi alla conoscenza degli imenotteri melliferi e predatori XIII. Bolettino del Laboratorio Entomologia del R. Istituto Superiore Agrario Bologna, 7:1–144.

1935.   Contributi alla conoscenza degli imenotteri aculeati XV. Bollettino dell'Istituto di Entomologia della Universita di Bologna, 8:27–121.

1937.   Contributi alla conoscenza degli imenotteri aculeati XVI. Bollettino dell'Istituto di Entomologia della Universita di Bologna, 9:253–348.

1954.   Contributi alla conoscenza degli imenotteri aculeati XXVI. Bollettino dell'Istituto di Entomologia della Universita di Bologna, 20:81–255.

1957.   Contributi alla conoscenza degli imenotteri aculeati XXVII. Bollettino dell'Istituto di Entomologia della Universita di Bologna, 22:307–398.

1961.   Studi di un entomologo sugli imenotteri superiori. Bollettino dell'Istituto di Entomologia della Universita di Bologna, 25:1–661.

Hirashima, Y.

1961.   Monographic Study of the Subfamily Nomiinae of Japan. Acta Hymenopterologica, 1:241–303.

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1975.   Nests, Behaviour and Larvae of the Bee Stenotritus pubescens (Smith) and Behaviour of some Related Species (Hymenoptera: Apoidea: Stenotritinae). Journal of the Australian Entomological Society, 14:145–154.

1976.   New Australian Allodapine Bees (Subgenus Exoneurella Michener) and their Immatures (Hymenoptera: Anthophoridae). Transactions of the Royal Society of Australia, 100:15–28.

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1939.   Bionomics and Ecology of Coelioxys elongata Lepeletier. Mushi, 12:34–40.

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1969.   Apuntes sobre las Hylaeus F., 1793, de Espana. Graellsia, 25:217–227.

1972.   La Nidification des Hylaeus parvulus Janvier. Entomologiste, 28:178–189.

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1981.   Observations on the Biology of Andrena (Melandrena) dunningi Cockerell (Hymenoptera: Andrenidae). Journal of the Kansas Entomological Society, 54:32–40.

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1984.   Immature Stages of Some Megachilidae (Hymenoptera: Apoidea). Journal of the Australian Entomological Society, 23:51–57.

LaBerge, W.E., and D.W. Ribble

1966a. The Nests and Larvae of Two Species of Agapostemon (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 39:467–472.

1966b. Biology of Florilegus condignus (Hymenoptera: Anthophoridae), with a Description of Its Larva, and Remarks on Its Importance in Alfalfa Pollination. Annals of the Entomological Society of America, 59:944–950.

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1911.   Ueber einen Eigentumlichen Nestbau von Osmia bicornis L., Zeitschrift für wissenschaftliche Insektenbiologie, 7:223–230, 316–322.

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1868.   Etude pour servir a l'histoire naturelle de la vie evolutive de la Xylocopa violacea, Hymenoptere Perce-Bois de la tribu des Apiens. Annales de la Société Entomologique de France, series 4, 8:727–736.

1888.   [Note on a Nymph of Xylocopa violacea.] Annales de la Société Entomologique de France, series 6, 8:158.

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1958.   Mudas ontogeneticas em larvas de larvae de Meliponinae (Hyemoptera, Apoidea). Dusenia, 8:37–40.

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1960.   Mudas ontogeneticas em larvas de Melipona nigra schencki Gribodo (Hymenoptera: Apoidea). Boletim da Universidade Federal do Parana, Zoologia, 2:1–16.

1962.   Morfologia externa da larva de Paratetrapedia (Paratetrapedia) gigantea (Schrottky, 1909) e consideracoes filogeneticas sobre algumas larvas de Anthophoridae (Hymenoptera: Apoidea). Boletim da Universidade Federal do Parana, Zoologia, 12:1–21.

1965.   Observacoes em larvas e pupas de Plebia (Plebeia) droryana (Friese, 1900) (Hymenoptera, Apoidea). Papéis Avulsos do Departamento de Zoologia, Secretariada Agricultura, Sao Paulo, 18(3): 21–27.

1966a. Descrição de estadios imaturos de Lanthanomelissa sp., com inferencia sobre a posição filogenetica deste genero entre os Exomalopsinae (Hymenoptera, Apoidea). Studia Entomologica, 9:429–440.

1966b. Descrição de estadios imaturos de Isepeolus viperinus (Holmberg) e confrontacoes com outras larvas de Anthophoridae parasitas conhecidas (Hym. — Apoidea). Boletim da Universidade Federal do Parana, Zoologia, II, 11:163–176.

1968.   Estadios imaturos de Lestrimelitta neotropicais (Hymenoptera-Apoidea). Boletim da Universidade Federal do Parana, Zoologia, 3:1–12.

1970.   Estadios imaturos de Hypotrigona (Leurotrigona) muelleri (Friese, 1900) e caracteristicas de algumas larvas de Apidae (Hymenoptera—Apoidea). Boletim da Universidade Federal do Parana, Zoologia, 3:233–244.

1974.   Estadios imaturos de algumas Xylocopa neotropicais (Hymenoptera—Apoidea). Acta Biologica Parana, 3:93–112.

1979.   Estadios imaturos de Bombus neotropicais (Hymenoptera, Apoidea). Dusenia, 11:83–89.

Maneval, H.

1936.   L'Anthidium Caturigense, son nid et sa Larve (Hym. Apidae). Annales de la Société Entomologique de France, 105:1–5.

1937.   Notes sur les Hymenopteres (5e serie), Revue Francaise d'Entomologique, 4:162–181.

1939.   Notes sur les Hymenopteres (6e serie), Annales de la Société Entomologique de France, 108:49–108.

Marechal, P

1926.   Etude biologique de l'“Osmia aurulenta” Panz. Bulletin Biologique de la France et de la Belgique, 60:561–592.

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1984.   [Contributions to Morphology and Ecology of the Megachile-Bees of the Subgenus Xanthosarus Robertson (Hymenoptera, Megachilidae).] In Yu. A. Pesenko, editor, Systematics and Ecology of Bees, Proceedings of the Leningrad Zoological Institute, 128:64–73. [In Russian.]

Masi, L.

1930.   Descrizione di un'Allodape vivente nelle spine di un'acacia nella Somalia Italiana. Memoire della Societa Entomologica Italiana, 9:67–75.

Masuda, II.

1943.   Biological Notes on Nomia punctata Westwood (Hymenoptera, Andrenidae). Mushi, 15:16–28.

1946.   Biological Notes on Eucera difficilis Perez and Whose [sic] Parasitic Bee, Nomada japonica Smith, with Description of a Gynandromorphic Eucera, Kontyu, 14:45–60.

Matthews, R.W.

1965.   The Biology of IIeriades carinata Cresson (Hymenoptera, Megachilidae). Contributions of the American Entomological Institute, 1:1–33.

Mayet, V.

1875.   Memoire sur les moeurs et les metamorphoses d'une nouvelle espece de Coleoptere de la famille des vesicants le Sitaris colletis. Annales de la Société Entomologique de France, series 5, 5:65–94.

McGinley, R.J.

1981.   Systematics of the Colletidae Based on Mature Larvae with Phenetic Analysis of Apoid Larvae (Hymenoptera: Apoidea). University of California Publications in Entomology, 91:1–307.

1987.   [Apoidea Section.] In F.W. Stchr, editor, Immature Insects, 1:602–617 (key), 689–704. Dubuque, Iowa: Kendall/Hunt Publishing Co.

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1987.   Nesting Biology, Immature Stages, and Phylogenetic Placement of the Palacarctic Bee Pararhophites (Hymenoptera: Apoidea). American Museum Novitates, 2903: 21 pages.

Micheli, L.

1930.   Note biologiche e morfologiche sugli immenotteri (contributo 2). Memoire della Società Entomologica Italiana, 9:46–66.

1931.   Note biologiche e morfologiche sugli imenotteri (contributo 3). Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 70:19–28.

1933.   Note biologiche e morfologiche sugli imenotteri (contributo 4). Memoire della Società Entomologica Italiana, 12:5–15.

1934.   Note biologiche e morfologiche sugli imenotteri (contributo 5). Bollettino della Società Entomologica Italiana, 66:246–252.

1935.   Note biologiche e morfologiche sugli imenotteri. Bollettino della Società Veneziana de Storia Naturale, series 7, 1:126–134.

1936.   Note biologiche e morfologiche sugli imenotteri. Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, series 6, 75:1–16.

1937.   Note biologiche e morfologiche sugli imenotteri. Memorie della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, series 8, 76:280–290.

Michener, C.D.

1953.   Comparative Morphological and Systematic Studies of Bee Larvae with a Key to the Families of Hymenopterous Larvae. University of Kansas Science Bulletin, 35:987–1102.

1954.   Observations on the Pupae of Bees. Pan-Pacific Entomologist 30:63–70.

1957.   Notes on the Biology of a Parasitic Bee, Isepeolus viperinus (Hymenoptera, Anthophorinae). Entomological News, 68:141–146.

1962.   Biological Observations on the Primitively Social Bees of the Genus Allodapula in the Australian Region (Hymenoptera, Xylocopinae). Insectes Sociaux, 9:355–373.

1963.   Observations on the Bionomics of a Colonials Bee of the Genus Perdita (Hymenoptera: Apoidea, Panurginae). Journal of the Kansas Entomological Society, 36:114–118.

1964.   The Bionomics of Exoneurella, a Solitary Relative of Exoneura. Pacific Insects, 6:411–426.

1975.   Larvae of African Allodapine Bees, 1: The Genus Allodape; 2: Braunsapis, and Nasutapis; 3: The Genera Allodapula and Eucondylops. Journal of the Entomological Society of Southern Africa, 38:1–12, 223–242, 243–250.

1976.   Larvae of African Allodapine Bees, 4: Halterapis, Compsomelissa, Macrogalea, and a Key to African Genera. Journal of the Entomological Society of Southern Africa, 39:33–37.

1977.   Discordant Evolution and the Classification of Allodapine Bees. Systematic Zoology, 26:32–56.

Michener, C.D., and L. Greenberg

1980.   Ctenoplectridae and the Origin of Long-Tongued Bees. Zoological Journal of the Linnean Society, 69:183–203.

Michener, C.D., and E. Ordway

1963.   The Life History of Perdita maculigera maculipennis (Hymenoptera: Andrenidae). Journal of the Kansas Entomological Society, 36:34–45.

Michener, C.D., and J.F. Scheiring

1976.   Pupae of Allodapine Bees. Journal of the Australian Entomological Society, 15:63–70.

Michener, C.D., and I.H. Syed

1962.   Specific Characters of the Larvae and Adults of Allodapula in the Australian Region (Hymenoptera: Ceratinini). Journal of the Entomological Society of Queensland, 1:30–41.

Miliczky, E.R.

1985.   Observations on the Nesting Biology of Tetralonia hamata Bradley with a Description of Its Mature Larva (Hymenoptera: Anthophoridae). Journal of the Kansas Entomological Society, 58:686–700.

Moczar, L.

1938.   Zur Okologie zweier Apiden. Zoologischer Anzeiger, 123:90–95.

Mohamed, M.I.

1974.   Seasonal Distribution of Tetralonia lanuginosa Klug with a Taxonomic Description of Different Stages (Hym., Anthophoridae). Deutsche Entomologische Zeitschrift, 21:167–178.

Moustafa, M.A., and A.A. El Berry

1976.   Morphological Studies on Osmia submicans Mor. Mitteilungen aus dem Zoologischen Museum Berlin, 52(2):209–225.

Moustafa, M.A., and M.M. Mazeed

1977.   Morphological Studies on Chalichodoma (Pseudomegachile) flavipes Spinola. Mitteilungen aus dem Zoologischen Museum Berlin, 53(1):161–171.

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1924.   Morphology of the Honeybee Larva. Journal of Agricultural Research, 28:1167–1213.

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1967.   Sex Characters of Larval Bees (Hymenoptera: Apoidea). Annals of the Entomological Society of America, 60:414–419.

Norden, B.B.

1984.   Nesting Biology of Anthophora abrupta (Hymenoptera: Anthophoridae). Journal of the Kansas Entomological Society, 57:243–262.

Norden, B.B., and A.G. Scarbrough

1979.   Nesting Biology of Andrena (Larandrena) miserabilis Cresson and Description of the Prepupa (Hymenoptera: Andrenidae). Brimleyana, 1:141–146.

Otis, G.W., R.J. McGinley, L. Garling, and L. Malaret

1982.   Biology and Systematics of the Bee Genus Crawfordapis (Colletidae, Diphaglossinae). Psyche, 89:279–296.

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1866.   Observations on the Development and Position of the Hymenoptera, with Notes on the Morphology of Insects. Proceedings of the Boston Society of Natural History, 10:279–296.

1897.   Notes on the Transformations of the Higher Hymenoptera, II and III. Journal of the New York Entomological Society, 5:77–87, 109–120.

Packer, L.

1987.   A Description of the Mature Larva and Cocoon of the Bee Thygater (Hymenoptera; Anthophoridae). Journal of the New York Entomological Society, 95:23–27.

Popova, L.M.

1984.   [Nesting of the Three Bee-Species of the Genus Anthophora Latr. (Hymenoptera, Anthophoridae).] In Yu. A. Pesenko, editor, Systematics and Ecology of Bees, Proceedings of the Leningrad Zoological Institute, 128:74–81. [In Russian.]

Radovic, I., and M.D. Krunic

1977.   External Morphology of the Larvae Anthophohora [sic] (Amegilla) parietina (F) var. fulvocinerea (Dours) and Anthophora (Amegilla) crinipes (Sm.) (Hymenoptera: Apoidea). Arhiv Bioloskih Nauka, Beograd, 27:137–144.

Rajotte, E.G.

1979.   Nesting, Foraging, and Pheromone Response of the Bee Colletes validus Cresson and Its Association with Lowbush Blueberries. (Hymenoptera: Colletidae) (Ericaceae: Vaccinium). Journal of the Kansas Entomological Society, 52:349–361.

Rayment, T.

1949a. New Bees and Wasps, Part VIII. Victoria Naturalist, 65:208–212.

1949b. New Bees and Wasps, Part IX. Victoria Naturalist, 65:247–254.

1951.   Biology of the Reed-Bees with Descriptions of Three New Species and Two Allotypes of Exoneura. The Australian Zoologist, 11:285–313.

1957.   The Nomia australica Smith Complex; Its Taxonomy, Morphology, and Biology, with the Description of a New Mutillid Wasp. The Australian Zoology, 12:176–200.

Ritcher, P.O.

1933.   The External Morphology of Larval Bremidae and Key to Certain Species (Hymenoptera). Annals of the Entomological Society of America, 26:53–63.

Roberts, R.B.

1973.   Nest Architecture and Immature Stages of the Bee Oxaea flavescens and the Status of Oxaeidae (Hymenoptera). Journal of the Kansas Entomological Society, 46:437–446.

1978.   The Nesting Biology, Behavior and Immature Stages of Lithurga chrysurus, an Adventitious Wood-Boring Bee in New Jersey (Hymenoptera: Megachilidae). Journal of the Kansas Entomological Society, 51:735–745.

Roberts, R.B., and C.H. Dodson

1967.   Nesting Biology of Two Communal Bees, Euglossa imperialis and Euglossa ignita (Hymenoptera: Apidae), Including Description of Larvae. Annals of the Entomological Society of America, 60:1007 1014.

Romasenko, L.P.

1983a. [Peculiarities of Megachile ligniseca (Hymenoptera, Megachilidae).] Vestnik Zoologii (Kiev) 2:76–78. [In Russian.]

1983b. [A Study of Megachile centuncularis Biology.] Vestnik Zoologii (Kiev) 4:31–34. [In Russian.]

Roubik, D.W., and C.D. Michener

1985.   Nesting Biology of Crawfordapis in Panama (Hymenoptera, Colletidae). Journal of the Kansas Entomological Society, 57:662–671.

Rozen, J.G., Jr.

1954.   Morphological Description of the Larva of Oreopasites vanduzeei Cockerell (Hymenoptera: Anthophoridae). Pan-Pacific Entomologist, 30:203–207.

1957.   External Morphological Description of the Larva of Exomalopsis chionura Cockerell, Including a Comparison with Other An-thophorids (Hymenoptera: Apoidea). Annals of the Entomological Society of America, 50:469–475.

1958.   Monographic Study of the Genus Nomadopsis Ashmead (Hymenoptera: Andrenidae). University of California Publications in Entomology, 15:1–202.

1963.   Notes on the Biology of Nomadopsis, with Descriptions of Four New Species (Apoidea, Andrenidae), American Museum Novitates, 2142: 17 pages.

1964a. Phylogenetic-Taxonomic Significance of Last Instar of Protoxaea gloriosa Fox, with Descriptions of First and Last Instars (Hymenoptera: Apoidea). Journal of the New York Entomological Society, 72:223–230.

1964b. The Biology of Svastra obliqua obliqua (Say), with a Taxonomic Description of Its Larvae (Apoidea, Anthophoridae). American Museum Novitates, 2170: 13 pages.

1965a. The Biology and Immature Stages of Melitturga clavicornis (Latreille) and of Sphecodes albilabris (Kirby) and the Recognition of the Oxaeidae at the Family Level (Hymenoptera, Apoidea). American Museum Novitates, 2224: 18 pages.

1965b. The Larvae of the Anthophoridae (Hymenoptera, Apoidea), Part 1: Introduction, Eucerini, and Centridini (Anthophorinae). American Museum Novitates, 2233: 27 pages.

1966a. Systematics of the Larvae of North American Panurgine Bees (Hymenoptera, Apoidea). American Museum Novitates, 2259: 22 pages.

1966b. Taxonomic Descriptions of the Immature Stages of the Parasitic Bee, Stelis (Odontostelis) bilineolata (Spinola) (Hymenoptera: Apoidea: Megachilidae). Journal of the New York Entomological Society, 74:84–91.

1966c. The Larvae of the Anthophoridae (Hymenoptera, Apoidea), Part 2: The Nomadinae. American Museum Novitates, 2244: 38 pages.

1967.   The Immature Instars of the Cleptoparasitic Genus Dioxys (Hymenoptera: Megachilidae). Journal of the New York Entomological Society, 75:236–248.

1968.   Biology and Immature Stages of the Aberrant Bee Genus Meliturgula (Hymenoptera, Andrenidae). American Museum Novitates, 2331: 18 pages.

1969.   The Larvae of the Anthophoridae (Hymenoptera, Apoidea), Part 3: The Melectini, Ericrocini, and Rhathymini. American Museum Novitates, 2382: 24 pages.

1970a. Biology and Immature Stages of the Panurgine Bee Genera Hypomacrotera and Psaenythia (Hymenoptera, Apoidea). American Museum Novitates, 2416: 16 pages.

1970b. Biology, Immature Stages, and Phylogenetic Relationships of Fideliine Bees, with the Description of a New Species of Neofidelia (Hymenoptera, Apoidea). American Museum Novitates, 2427 : 25 pages.

1971.   Biology and Immature Stages of Moroccan Panurgine Bees (Hymenoptera, Apoidea). American Museum Novitates, 2457: 37 pages.

1973a. Biology Notes on the Bee Andrena accepta Viereck (Hymenoptera, Andrenidae). Journal of the New York Entomological Society, 81:54–61.

1973b. Life History and Immature Stages of the Bee, Neofidelia (Hymenoptera, Fideliidae). American Museum Novitates, 2519: 14 pages.

1973c. Immature Stages of Lithurgine Bees with Descriptions of the Megachilidae and Fideliidae based on Mature Larvae (Hymenoptera, Apoidea), American Museum Novitates, 2527: 14 pages.

1977a. Biology and Immature Stages of the Bee Genus Meganomia (Hymenoptera, Melittidae). American Museum Novitates, 2630: 14 pages.

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1968.   Ecology of a Proteriades and Its Chrysura Parasite, with Larval Descriptions (Hymenoptera: Megachilidae: Chrysididae). Journal of the Kansas Entomological Society, 41:324–331.

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