The History of Bee Classifications

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The species of bees have been classified in many different ways. An exhaustive historical treatment of the classifications proposed would occupy a great deal of space. The following pages briefly summarize the classifications found in some of the major publications on the subject. To facilitate comparison of these classifications, I have sometimes rearranged the groups so that the sequence does not inhibit comparisons. The genera I list are not necessarily all those listed by the authors, but are representatives of the various currently accepted tribes, subfamilies, or families. Thus a reader can determine what groups were intended by the authors. For example, where for brevity I have written Megachile after Megachilidae, most authors also included Osmia, Chalicodoma, Che-lostoma, Heriades, etc. Of course, when authors placed these genera in different tribes or higher categories, this fact has been indicated. The genera listed vary, largely for geographical reasons; for example, a classification of European bees cannot be expected to include taxa not found in Europe. Misplaced genera that are not types of higher taxa are often ignored; in some cases the authors had not seen authentic specimens and had therefore placed the genera on the basis of descriptions. In other cases they were simply careless. Modern generic names and accepted spellings have been introduced as necessary to permit easy understanding.

Because the small Nomadinae have been included as a block in most classifications, but the genera and tribes cited often differ among authors, I have used the word "pasitines" to indicate this group instead of listing particular genera or tribes. The word "pasitines" has the advantage that it is familiar, yet no currently recognized tribe is called Pasitini. Therefore the word "pasitines" can reasonably stand for the entire group of tribes, Ammobatoi-dini to Caenoprosopidini in Table 16-1.

Kirby (1802), in the first major account of the bees of any area (Britain), placed all bees in two genera, Melitta for the S-T bees and Apis for the L-T bees. In the same year Latreille (1802b) recognized the same two groups as families, with certain supergeneric subdivisions as shown in Table 18-1. The idea of recognizing numerous genera was quickly established; Klug (1807b), only five years after Kirby's monograph, gave a summary listing 32 bee genera.

Lepeletier (1835, 1841) not only gave a later classification by Latreille that is similar to the above, but also presented his own classification. He separated the social Ap-inae (Bombus, Melipona, Apis) in a separate account (1835). Other bees (1841) were divided into two major groups, the solitary nesters and the parasites, the latter including some that we now know to be nonparasitic: Cer-atina, Allodape, and Hylaeus. Each of the two major groups was divided into three families (Table 18-2). The recognition of parasitic bees as families separate from nonparasitic bees affected bee classifications for years, parasites like Psithyrus being placed in families different from those of closely related nonparasites, in this case

Table 18-1. Classification of Bees Based on Latreille (1802b).

Family Andrenetes (Andrenetae)

Division I. Tongue blunt (Colletes, Hylaeus) Division II. Tongue pointed (Andrena, Dasypoda) Family Apiares (Apiariae) Megachiles (Megachile) Nomades (Epeolus, Melecta, Nomada) Euceres (Eucera)

Podaliries (Centris, Podalirius = Anthophora)

Claviceres (Clavicera = Ceratina)

Xilocopes (Xylocopa)

Euglosses (Euglossa)

Bourdons (Bombus)

Apiares domestiques (Apis)

other Bombus. It was not until the time of Robertson (1904) that such parasitic forms were regularly placed with their nonparasitic relatives in bee classifications, although Thomson (1872) had also correctly placed them. A century later, Tkalciu (1972), as noted below, again postulated that in their phylogenetic history the parasitic bees never possessed a scopa but evolved from wasps, independently from other bees.

Some features of Lepeletier's classification that seem extraordinary to modern melittologists are the placement of Hylaeus and Ceratina among the parasitic bees, of Rhathymus in the same tribe with Sphecodes, of Melitta in the Xylocopites, and ofpasitines and parasitic megachilids in the same tribe.

Schenck (1861, 1869), in accounts of the bees ofNas-sau, Germany, provided the classification summarized in Table 18-3, using the 1869 version with interpretation as necessary from that of 1861. All subfamily names (with family name endings by modern standards) were based on generic roots. The placement of Ceratina in the An-thophoridae removed it from the parasitic groups, where it does not belong, and Melitta was removed from association with Xylocopa. Segregation of the parasitic mega-chilids from other parasites was an important step. A noteworthy problem, which extended through many later classifications, concerned "short-tongued" bees that have a somewhat elongate glossa, e.g., Dasypoda, Du-fourea, Melitturga, Panurginus, Panurgus, and Systropha. The classification contained enough taxa (subfamilies), but these genera were scattered in an almost random way; for example, the three of these genera placed in the Pa-nurgidae fall in three families by current standards, but Dufourea,, Halictoides, Rophites, and Systropha, scattered among three subfamilies by Schenck, belong in one.

Thomson (1872) made a classification that in various ways is more modern than those of earlier writers, although he called the main named divisions tribes. His classification is summarized in Table 18-4. Except for the association of Epeolus, Nomada, and the pasitines with

Solitary-nesting Bees Family Podilegides

Tribe Eulmites (Euglossa, Eulaema) Tribe Anthophorites (Anthophora, Eucera, Melitturga, Systropha)

Tribe Xylocopites (Centris, Epicharis, Melitta, Xylocopa) Family Gastrilegides (Anthidium, Chelostoma, Lithurgus,

Megachile) Family Merilegides

Tribe Andrenites (Andrena, Halictus, Nomia) Tribe Panurgites (Dasypoda, Dufourea, Panurgus) Tribe Colletides (Colletes)

Social Bees

Family Apiarides Tribe Apiarites (Apis) Tribe Meliponites (Melipona) Family Bombides (Bombus)

Parasitic Bees

Family Psithyrides (Psithyrus) Family Dimorphides

Tribe Melectites (Aglae, Ceratina, Epeolus, Melecta,

Mesoplia, Nomada) Tribe Phileremides (Coelioxys, Dioxys, pasitines, Stelis) Family Monomorphides Tribe Prosopites (Hylaeus) Tribe Rhathymites (Rhathymus, Sphecodes)

Melecta, Thomson put the parasitic bees where they belong—Sphecodes in the halictids, the megachilid parasites in the Megachilidae, and Psithyrus with Bombus. For the first time, Halictus appeared in a major taxon different from that of Andrena; Colletes and Hylaeus were in the

Solitary Bees

Tribe Halictina [ Colletes, Halictus, Hylaeus, Rophites (including Dufourea and Halictoides), Sphecodes] Tribe Andrenina (Andrena, Panurgus) Tribe Megachilina (Anthidium, Coelioxys, Dioxys, Megachile, Stelis)

Tribe Megillina (Ceratina, Cilissa = Melitta, Dasypoda,

Eucera, Macropis, Megilla = Anthophora) Tribe Nomadina (Epeolus, Melecta, Nomada, pasitines)

Social Bees

Tribe Bombina (Apathus = Psithyrus, Bombus) Tribe Apina (Apis)

Table 18-5. Classification of Bees Based on Schmiedeknecht (1882).

I. Solitary bees

A. Podilegidae (Scopulipedes, leg collectors)

a. Femorilegidae (femur collectors) Andrenidae (Andrena, Colletes, Halictus, Nomia) Panurgidae (Biareolina, Dasypoda, Panurgus, Rophites) Xylocopidae (Ceratina, Xylocopa)

b. Crurilegidae (tibia collectors) Melittidae (Macropis, Melitta)

Anthophoridae (Ancyla, Anthophora, Eucera, Systropha)

B. Gastrilegidae (Dasygastrae, belly collectors) Megachilidae (Anthidium, Lithurgus, Megachile, Osmia)

C. Pseudoparasitae (nonparasitic bees without a scopa) Prosopidae = Hylaeidae

Sphecodidae (of course this is now known to be parasitic)

II. Social bees

Apidae (Apis) Bombidae (Bombus)

III. Parasitic bees

Psithyridae (Psithyrus)

Melectidae (Melecta, Nomada, pasitines)

Stelidae (Coelioxys, Dioxys, Stelis)

same tribe, along with Rophites, and the melittids were among what are frequently called the anthophorines, i.e., the noncorbiculate Apidae.

In spite of Thomson's finding that most parasitic taxa do not belong in their own separate families but can be associated with their nonparasitic relatives, Schmiedeknecht (1882), Friese (1895), and subsequent works as late as Schmiedeknecht (1930) reverted to a system similar to that of Lepeletier. They recognized three sections, the solitary nest-making bees, the social bees, and the parasitic bees. In the first section these authors provided considerable classificatory structure, as seen in Schmiedeknecht's 1882 version, which is summarized in Table 18-5. Schmiedeknecht's families agreed with Schenck's (1861) in uniting Panurgidae and Rophitidae under the former name. Ceratina was included in the Xy-locopidae. Bombidae was recognized as separate from the Apidae.

Table 18-3. Classification of Bees Based on Schenck (1861, 1869).

Contrary to current practice, Schenck used the -idae ending for subfamilies.

Subfamily Andrenidae (Andrena, Colletes, Hylaeus = Halictus, Nomia)

Subfamily Prosopidae (Prosopis = Hylaeus) Subfamily Sphecodidae (= Rhathymidae in 1869) (Sphecodes) Subfamily Panurgidae (Dasypoda, Dufourea, Panurgus) Subfamily Rophitidae (Halictoides, Rophites) (In 1869 these were included in the Panurgidae.) Subfamily Melittidae (Macropis, Melitta, Panurginus) Subfamily Megachilidae (Anthidium, Lithurgus, Megachile) Subfamily Anthophoridae (Anthophora, Ceratina, Eucera,

Melitturga, Systropha) Subfamily Xylocopidae (Xylocopa) Subfamily Apidae (Apis, Bombus) Subfamily Psithyridae (Psithyrus)

Subfamily Melectidae (Epeolus, Melecta, Nomada, pasitines) (In 1869 the pasitine bees were put in a separate subfamily, the Phileremidae.) Subfamily Stelidae (Coelioxys, Dioxys, Stelis)

Family Colletidae (Colletes, Diphaglossa, Paracolletes) Family Prosopidae (Euryglossa, Prosopis = Hylaeus) Family Andrenidae

Subfamily Andreninae (Ancyla, Andrena, Melitta, Nomia, Stenotritus)

Subfamily Halictinae (Augochlora, Halictus, Systropha) Subfamily Sphecodinae (Sphecodes, Temnosoma) Family Panurgidae (Dasypoda, Dufourea, Hylaeosoma =

Chilicola, Macropis, Panurgus, Rophites) Family Megachilidae

Subfamily Osmiinae (Heriades, Osmia) Subfamily Megachilinae (Ctenoplectra, Lithurgus, Megachile) Subfamily Anthidiinae (Anthidium) Family Stelidae

Subfamily Stelidinae (Stelis)

Subfamily Coelioxinae (Allodape, Chilicola, Coelioxys, Dioxys, pasitines) Family Nomadidae (Aglae, Epeolus, Exaerete, Melecta, Nomada, Osiris)

Family Anthophoridae (Anthophora, Centris, Emphor =

Ptilothrix, Eucera, Exomalopsis, Melitturga) Family Ceratinidae (Ceratina) Family Xylocopidae (Oxaea, Xylocopa) Family Euglossidae (Euglossa) Family Bombidae (Bombus) Family Psithyridae (Psithyrus) Family Apidae

Subfamily Meliponinae (Melipona) Subfamily Apinae (Apis)

Ashmead (1899a) greatly modified Schmiedeknecht's (1882) system and included genera from all parts of the world. His classification, summarized in Table 18-6, placed all parasitic bees in families of their own, except for the Sphecodinae; he presumably did not realize that his Sphecodinae consisted of parasitic forms. Anomalies, in view of our present knowledge, were the placement of parasitic euglossines in the Nomadidae, the placement of the melittids and dufoureines in the Panurgidae, and the positions of such genera as Chilicola (two places), Oxaea, Ctenoplectra, and Allodape. Melitturga was equally out of place; its position in the Anthophoridae, although traditional, was incorrect. I have ignored some of Ashmead's careless placements of genera that were little known to him.

Robertson (1904) thoughtfully developed a new classification for bees; his families were widely accepted by North American hymenopterists such as Viereck (1916) and by American textbook writers. Table 18-7 summarizes it, with some interpretation based on Robertson's 1903 papers. A noteworthy feature of Robertson's classification is recognition of the two large groups, Pygidialia and Apygidialia. As stated elsewhere, the pygidial plate has been lost repeatedly and independently. Robertson was not familiar with the numerous pygidialate colletids or the remnants of such plates in many of his Cerati-noidea and in the megachilid tribe Lithurgini. Like

Apygidialia Colletoidea

Family Colletidae (Colletes) Family Prosopididae (Prosopis = Hylaeus) Trypetoidea

Family Megachilidae Subfamily Osmiinae Tribe Osmiini (Osmia) Tribe Trypetini (Trypetes = Heriades) Subfamily Megachilinae

Tribe Megachilini (Megachile) Tribe Coelioxyini (Coelioxys) Family Stelidae

Subfamily Trachusinae (Trachusa) Subfamily Anthidiinae Tribe Stelidini (Stelis) Tribe Anthidiini (Anthidium) Ceratinoidea

Family Ceratinidae (Ceratina) Family Exoneuridae (Allodape, Exoneura) Family Xylocopidae (Xylocopa) Apoidea

Family Apidae (Apis, Bombus, Psithyrus) Pygidialia Andrenoidea

Family Andrenidae (Andrena) Family Panurgidae

Subfamily Panurginae (Panurgus) Subfamily Protandreninae (Protandrena) Family Halictidae (Augochlora, Halictus, Sphecodes) Family Nomiidae (Paranomia = Nomia) Family Dufoureidae (Dufourea, Halictoides, Rophites) Family Macropididae (Macropis) Anthophoroidea

Family Anthophoridae (Anthophora) Family Euceridae (Eucera)

Family Emphoridae (Emphor = Ptilothrix, Melitoma) Family Melectidae (Melecta, pasitines, and presumably Nomada and Epeolus)

Thomson, but probably independently, Robertson placed the parasitic forms appropriately except for the association of Nomada and its relatives with Melecta. He was the first to properly recognize the limits of his Dufoureidae ( = Rophitinae); Schenck had named the family but had somehow put Dufourea itself in the Panurgidae. Unfortunately, it is not clear where Robertson would have placed Melitta (he omitted it because it does not occur in his area), although he said that it would not be near Macropis.

Börner (1919) constructed a classification that, for ST bees, foreshadowed some later classifications. The parasitic bees were appropriately placed, as shown by Grütte (1935), who followed Börners system. Some taxa, such as Macropis, Melitta, and the Ceratinini, were misplaced, and the Nomiinae and Halictinae were mixed, as were the Osmiini and Megachilini. Börner did not mention the pasitine bees, but probably he intended them to be in

Table 18-8. Classification of Bees Based on Börner (1919).

Family Colletidae

Subfamily Prosopinae (Hylaeus) Subfamily Colletinae (Caupolicana, Colletes) Family Andrenidae

Subfamily Andreninae (Andrena) Subfamily Panurginae (Macropis, Melitta, Panurgus) Family Halictidae Subfamily Halictinae

Tribe Nomiini (Agapostemon, Augochlora, Nomia) Tribe Halictini (Halictus, Paragapostemon, Sphecodes) Tribe Nomioidini (Nomioides) Subfamily Halictoidini (Dufourea, Rophites) Family Megachilidae

Subfamily Osmiinae (Osmia, Stelis) Subfamily Megachilinae (Anthidium, Coelioxys, Megachile) Family Nomadidae

Subfamily Ceratininae (Allodape, Ceratina) Subfamily Nomadinae (Nomada) Family Apidae

Subfamily Anthophorinae

Tribe Eucerini (Centris, Eucera, Exomalopsis, Melissodes,

Tetrapedia) Tribe Anthophorini (Anthophora) Tribe Xylocopini (Xylocopa) Subfamily Apinae

Tribe Bombini (Bombus, Euglossa, Psithyrus)

Tribe Apini (Apis)

Tribe Meliponini (Melipona)

the Nomadinae. Borner's classification is summarized in Table 18-8.

The older classifications of bees were based largely on various characters of mouthparts, wings, legs, and scopa. Bischoff (1934) was among the first to call attention to various little-used characters of the body, such as suban-tennal sutures and the episternal groove, as well as to the jugal lobe of the hind wing. Grutte (1935) made use of Bischoff's findings in a study of parasitic bees, and subsequent classifications, such as that of Michener (1944), utilized the same characters.

A classification for the bees of the world developed by Michener (1944) is summarized in Table 18-9. Some principal features of this classification were the placement of Melitturga in the Panurginae, of the Nomiinae and Du-foureinae in the Halictidae, of Chilicola in the Colletidae, and of Oxaea in the Andrenidae. Note also the enlarged Melittidae and the recognition of the Lithurginae as a distinct subfamily of the Megachilidae. The broad Apidae, including all L-T bees except the Megachilidae, was also novel. The enormous number of tribes in the An-thophorinae had not been anticipated by previous general bee classifications, although most of the tribes had been named before 1944.

Michener (1965b) summarized the bee classification and modified that of 1944 as follows: The tribe Caupoli-canini was transferred to the Diphaglossinae. The subfamily Euherbstiinae was added to the Andrenidae.

Family Colletidae

Subfamily Euryglossinae (Euryglossa) Subfamily Hylaeinae (Hylaeus) Subfamily Chilicolinae (Chilicola, Xeromelissa) Subfamily Colletinae

Tribe Paracolletini (Paracolletes) Tribe Colletini (Colletes) Tribe Caupolicanini (Caupolicana) Subfamily Stenotritinae (Stenotritus) Subfamily Diphaglossinae (Diphaglossa) Family Andrenidae

Subfamily Andreninae (Andrena) Subfamily Panurginae

Tribe Panurgini (Panurgus, Protandrena) Tribe Melitturgini (Melitturga) Subfamily Oxaeinae (Oxaea) Family Halictidae

Subfamily Dufoureinae (Dufourea, Rophites, Systropha) Subfamily Nomiinae (Nomia)

Subfamily Halictinae (Augochlora, Halictus, Sphecodes, Temnosoma) Family Melittidae

Subfamily Melittinae (Melitta) Subfamily Macropidinae (Macropis) Subfamily Dasypodinae (Dasypoda) Subfamily Ctenoplectrinae (Ctenoplectra) Family Megachilidae

Subfamily Lithurginae (Lithurgus) Subfamily Megachilinae

Tribe Megachilini (Coelioxys, Heriades, Megachile, Osmia)

Tribe Anthidiini (Anthidium, Dioxys, Stelis) Family Apidae

Subfamily Fideliinae (Fidelia) Subfamily Anthophorinae

Tribe Exomalopsini (Exomalopsis) Tribe Ancylini (Ancyla) Tribe Nomadini (Nomada) Tribe Epeolini (Epeolus) Tribe Osirini (Osiris)

Tribe Protepeolini (Protepeolus = Leiopodus) Tribe Epeoloidini (Epeoloides) Seven tribes of pasitine bees Tribe Emphorini (Melitoma, Ptilothrix) Tribe Eucerini (Eucera) Tribe Anthophorini (Anthophora) Tribe Hemisiini (Centris, Epicharis) Tribe Melectini (Melecta) Tribe Rhathymini (Rhathymus) Tribe Ericrocini (Ctenioschelus, Ericrocis, Mesoplia) Subfamily Xylocopinae

Tribe Ceratinini (Allodape, Ceratina, Exoneura) Tribe Xylocopini (Xylocopa) Subfamily Apinae

Tribe Euglossini (Aglae, Euglossa, Eulaema, Exaerete) Tribe Bombini (Bombus, Psithyrus) Tribe Meliponini (Melipona) Tribe Apini (Apis)

Family Andrenidae Subfamily Colletinae a. (Colletes, Hylaeus)

b. (Caupolicana) Subfamily Andreninae a. (Andrena)

b. (Melitturga, Oxaea, Panurgus) Subfamily Halictinae a. (Rophites, Systropha)

b. (Halictus, Nomia, Nomioides, Sphecodes) Family Apidae

Subfamily Melittinae a. (Dasypoda, Pararhophites)

b. (Ctenoplectra, Macropis, Melitta) Subfamily Megachilinae a. (Lithurgus)

b. (Anthidium, Stelis; Dioxys, Osmia; Coelioxys, Megachile)

Subfamily Ceratinae (sic)

a. (Exomalopsis, Fidelia)

b. (Allodape, Ceratina) Subfamily Anthophorinae a. (Ancyla, Manuelia, Xylocopa)

b. (Dasiapis = Diadasia, Eucera, Lanthamelissa (sic), Tapinotaspis, Tetrapedia)

c. (Ancyloscelis, Anthophora, Caenonomada, Epeoloides, Melecta)

Subfamily Nomadinae a. (Biastes, Epeolus)

b. (Nomada, most pasitines) Subfamily Apinae a. Melipona b. Apis, Bombus c. Euglossa

Fideliinae was raised to family rank. Likewise, Anthophorinae was raised to family rank with the following subfamilies: Exomalopsinae (for Exomalopsini and An-cylini), the Nomadinae (for tribes Nomadini through the pasitine bees in the 1944 classification), the Anthophorinae (for the Emphorini to Ericrocini), and the Xylo-copinae (for Ceratinini and Xylocopini). Of course, the Apinae of 1944 were also raised to family rank, and two subfamilies were recognized, Bombinae for the tribes Euglossini and Bombini and Apinae for the tribes Meliponini and Apini. Name changes were Emphorini to Melitomini, Hemisiini to Centridini, and Ericrocini to Ctenioschelini.

Some of these changes resulted from the tradition of recognizing numerous families and the hesitation of others to accept a broad family Apidae. Recognition of families like Bombidae, Meliponidae, and Xylocop-idae was widespread, and the 1965 classification was a compromise. I now consider that raising the Anthophorinae to family level, and the related changes, were mistakes.

A modified world classification can be extracted from Michener (1979a: 297-323). It incorporated the changes that appeared in 1965 except that the Euherbstiinae was incorporated into the Andreninae. Additionally, Oxaea and its relatives were placed in the Oxaeidae, Halictinae was divided into three tribes (Augochlorini, Halictini, Nomioidini), Dioxys was placed in a separate megachilid tribe, Ancyla was tentatively put in the Exomalopsini, additional tribes of Anthophoridae (Eucerinodini, Tetra-pediini, Canephorulini, Pararhophitini, Isepeolini) were recognized, and the Meliponini was raised to subfamily rank.

In general, based on the classification above but with more categories are the classifications of Engel (2001b, 2005). Another version by Melo and Gongalves (2005) is similar but with all the Apiformes included in a single family, Apidae, which therefore equals the Apiformes.

Table 16-1 shows the classification accepted for the present book.

Sustera (1958), reacting to many of the same findings that led to the development of my classification of 1979, had proposed a similar classification 21 years earlier. It placed the Ceratininae and Xylocopinae in a separate family, the Xylocopidae. Its most unusual feature was the placement of the Nomadinae (Nomadini and Ammoba-tini) in the Andrenidae while Epeolini, Epeoloidini, and the remaining pasitines were placed in a subfamily Bi-astinae in the family Anthophoridae. This division and placement of Nomadinae had been indicated earlier in a diagram by Pittioni and Schmidt (1942, pl. I).

Although the classifications by Michener outlined above have been widely used, other authors have proposed very different classifications. The samples discussed below demonstrate the persistent lack of agreement about bee classification and phylogeny. A principal reason for the preparation of the present work is to present cladistic patterns and classifications that best represent our current knowledge of these insects.

Table 18-10 summarizes a classification, limited largely to European genera, by Warncke (1977a); tribal names were not used, but the genera were nonetheless grouped into units (here lettered a, b, or c) falling below the subfamily level. Some unusual features of this classification include the placement of Pararhophites in the Melittinae, of the Melittinae in the Apidae (L-T bees), and of Fidelia and Exomalopsis close together and in the Ceratininae, the wide separation of Ceratina from Manuelia and Xylocopa, and the placement of Ancyla with the latter. The genera included by Warncke in each group ofAnthophorinae are so diverse that I have listed them all rather than merely presenting a representative of each group.

Finally, to further demonstrate the lack of general agreement on bee evolution and classification, I note that Tkalcil (1972, 1974a) proposed that all parasitic bees arose, not from pollen-collecting ancestors, but from nonpollen-collecting ancestors of the pollen-collecting groups. If true, this would greatly change accepted classifications, for it implies different nonpollen-collecting wasp ancestors for major groups of bees. I believe, on the contrary, that there is abundant morphological evidence for the relations of parasitic bees to different nonparasitic taxa; see Section 8.

The status of the corbiculate Apidae (Euglossini,

Bombini, Meliponini, and Apini) is of special historical interest, partly because honey bees and bumble bees are included, but also because the group has often been recognized, in the above classifications and by Michener (1990a), as a taxon of family rank. The four tribes consistently came out as a single clade in analyses by Roig-Alsina and Michener (1993), who included them as part of a large subfamily Apinae but called them the "apine clade" (the apine line of Silveira, 1993b). To recognize the corbiculate Apidae as a family would require the recogni tion of numerous other families, a procedure that does not seem helpful or appropriate. Yet because of their relationship to one another, as well as because of history, it is convenient to have a term for the four tribes considered collectively. The name "corbiculate Apidae" is appropriate. Additional characters showing their monophyletic relationship are enumerated in Section 102. The difficulty of giving them a technical name within our system is somewhat troubling.

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