Classification of the Bee Sphecoid Clade

Lomholdt (1982), Brothers (1975), Michener (1944, 1979a), Gauld and Bolton (1988), and others, including older authors such as Comstock (1924), have advocated placing the sphecoid wasps and the bees in the same superfamily. The custom of separating them as the su-perfamilies Sphecoidea and Apoidea obscures their close relationship to one another, as compared to other super-families of aculeate Hymenoptera. It has been common to recognize only one family of sphecoid wasps in spite of their...

The Identification of Bees

For the practical problems of identification of bees to the genus level, it may be easier to use a regional key than to go through the worldwide treatment below, even though many of the keys herein are regional. Identification to species is of course more difficult. Throughout Sections 36 to 121, I have included references to works containing keys or revisions of species. For some taxa and some areas, no such treatments exist. Table 32-1 lists some works containing keys to bee genera of certain...

Diversity and Abundance

Michener (1979a) published on the biogeography of bees, and the following material is in part derived from that paper. Bees appear to attain their greatest abundance, greatest numbers of species, and probably greatest numbers of genera and subgenera, not in the tropics, but in various warm-temperate, xeric regions of the world. It is easy to make statements like this, but to provide suitable documentation for them is difficult. The taxonomic literature is a poor guide because of the different...

Practical Key to Family Group Taxa Based on Females

Because the key to families (Sec. 33) depends heavily on characters that are difficult to see in dry specimens with the mouthparts in repose, a key based on more readily observable characters seems worthwhile. This key does not usually lead to families, but rather directs the user to tribes or subfamilies. It is based on females for males, it is best to make the necessary examinations of mouthparts and use the key to families, Section 33. The tibial hairs of Pararhophites (Megachilidae) look...

Solitary versus Social Life

Many works treat aspects of behavior of diverse kinds of bees. Specialized papers are cited throughout this book some more general treatments are the books by Friese (1923), with its interesting colored plates of nests of European bees Iwata (1976), with its review of previous work on the behavior of bees and other Hymenoptera and O'Toole and Raw (1991), which offers readable accounts and fine illustrations of bees worldwide. A major aspect of behavior involves intraspecific interactions, i.e.,...

Info

S8 of male with two flat, delicate, apical processes, longer than body of sternum (Fig. 39-4b) supraclypeal area with longitudinal, strongly elevated, impunctate, shining carina or broad ridge extending from frontal carina down to upper margin of clypeus distal three an-tennal segments of male modified (Fig. 39-4d) (Australia) Glossurocolletes . S8 of male ending in the usual single, commonly heavily sclerotized, apical process supraclypeal area broadly convex, median part sometimes...

Fossil Bees

Fossil bees are rare, and with some exceptions only those in amber are well enough preserved to shed light on phy-logeny. The fossils so preserved are probably biased toward bees that used resin in nest construction, and therefore sometimes became mired and trapped as they collected the resin. Bees are prone to the development of convergent features in the wing venation, legs, and other parts often visible in fossils, whereas the mouthparts and male genitalia so often important in phylogenetic...

Key to the Families Based on Adults

The key to families below is intended to work for the great majority of bees, but exceptions to some of the characters of some couplets exist, usually in rare or geographically limited taxa. These problems are addressed in the notes in Section 34, each note keyed to the pertinent couplet number. Some general attributes of the families are discussed in Section 21, and the bases for the recognition of families are discussed in Sections 18 to 21, as well as in the main systematic text, Sections 36...

Reduction or Loss of Structures

Systematists devote a great deal of time and energy to finding shared characters that have not evolved conver-gently and that therefore characterize taxa and are useful in developing phylogenetic hypotheses. Some of the most interesting characters that organisms have, however, are those that are convergent and that therefore suggest some common behavior or common environmental challenge perhaps working through behavior. The following are some examples of characters that have arisen two or more...

Structures and Terminology of Immature Stages

Eggs of bees are described in Sections 4 and 8. They are not rich in external characters, although size varies as described in those sections, and the chorion of some clep-toparasitic bees is variously modified (Sec. 8). Rozen (2003a) and Rozen and Ozbek (2003) have investigated Figure 11-1. Diagrammatic lateral view of a bee larva. Figure 11-1. Diagrammatic lateral view of a bee larva. eggs of numerous, especially parasitic, bees and in addition to characters of the chorion in general, have...

The Higher Classification of Bees

The following paragraphs serve to identify those groups recognized as families, as well as the relationships among the families. They concern only extant families the extinct groups that have been given family-group status are ignored. They add detail to the information already provided by the dendrograms (Figs. 20-1, 20-2) and Table 16-1. They are not intended as descriptions of the families, but rather as indications of some of the reasons for decisions about the family-level classification....

About Bees and This Book

Since ancient times, people have been drawn to the study of bees. Bees are spritely creatures that move about on pleasant bright days and visit pretty flowers. Anyone studying their behavior should find them attractive, partly because they work in warm sunny places, during pleasant seasons and times of day. The sights and odors of the fieldwork ambience contribute to the well-being of any researcher. Moreover, bees are important pollinators of both natural vegetation and crops, and certain...

Subfamily Colletinae

This subfamily is by far the largest and most abundant group ofhairy colletids. It consists of small to rather large, generally hairy andreniform bees, most of them superficially resembling species of the genera Andrena or Halictus. A few small species are only sparsely hairy and are almost hylaeiform, especially in males. The first flagellar segment is shorter than the scape and not recognizably petiolate. The glossa usually has two short lobes (it is thus weakly bilobed, Fig. 19-2b, as in...

New and Modified Structures

In contrast to loss characters, new or newly modified structures are more likely to be unique synapomorphies that can be used to recognize clades in phylogenetic studies. Nonetheless, remarkable cases of convergence exist, provided that our phylogenetic hypotheses are correct. If our hypotheses are considered to be incorrect, worse problems of understanding phylogenies usually arise. The following are some examples of apparently new structures that seem to have arisen independently in different...

Bee Taxa and Categories

Classifications, of course, are based in large part on phy-logeny. Some specialists cladists base classification entirely on phylogeny others consider also information from diverse sources in developing a classification. No one, however, should presume to make a classification without having all available phylogenetic information. For practical purposes, I present here some information on bee classification, prior to the section on phylogeny, because use of the family-group names makes...

The History of Bee Classifications

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...

Bee Wing Diagram

Diagram Bee Vein

Basal area of propodeum propodeal spiracle hind wing base propodeal spiracle scutellum -bj metanotum- parapsidal line tegula mesepisternum is not true, as careful examination shows nevertheless the middle coxa, which in many bees is expanded upward to form a vertically elongated cylinder see Michener, 1981b , displaces or is partly hidden beneath the lower part of the metepisternum. The form and subdivisions of the propodeum are important systematically. Many bees have a pair of impressed lines...

Family Stenotritidae

This family comprises two Australian genera of moderate-sized to large, robust, euceriform, hairy bees Fig. 362 . Superficially, these bees closely resemble those of the American tribe Caupolicanini of the Diphaglossinae although one stenotritid species, Cenocollees smaragdinus Smith , is bright metallic green . Unlike colletids, they have a well-developed tibial scopa and a reduced femoral scopa, and pollen is accordingly carried principally on the tibia, as shown by Houston 1984 . The first...

Body Form Tagmata and Sex Differences

Bees ofmany genera can be identified to genus at a glance, or at least very promptly, by a person familiar with the bees of the relevant region. This is possible in part because of the diverse body shapes of bees. I have indicated general body shape in the text by a series of terms such that with a single word a person who knows a few common genera of bees can get an idea of what bees of an otherwise unknown genus look like. These terms, following Michener, McGinley, and Danforth 1994 , are...

Short Tongued versus Long Tongued Bees

At least since Kirby 1802 , it has been the custom of specialists on bees, unlike other hymenopterists, to devote a great deal of attention to mouthparts. Continuing this practice, recent phylogenetic studies Roig-Alsina and Michener, 1993 Alexander and Michener, 1995 have been heavily weighted by numerous characters of mouth-parts. An old atlas of proboscides of bees, that of Saun-ders 1891 , shows many of the characters discussed below. From at least as early as Kirby's monograph 1802 to the...

Some Problematic Taxa

Scattered through the families of bees are several large species complexes for which the current generic classification is arbitrary and will probably be revised in the near future. I am not referring here to the many differences of opinion on rank should a given group of species be regarded as a species group or be given subgeneric status, should a taxon be a subgenus or a separate genus, or should a taxon be regarded as a tribe or a subfamily Such problems are perpetual they have no right or...

Tribe Paracolletini

The Paracolletini, in a superficial way, replaces the holarc-tic genus Andrena in the Australian region and in temperate South America. It contains numerous andreni-form bees, some of them with special features relevant to the flowers where they collect pollen. As noted in Section 20, these bees exhibit a series of characters listed as probably ancestral for bees in an old work Michener, 1944 because of similarity to characters of wasps. Yet the para-colletines are not at all wasplike in...

Family Colletidae

As noted in Section 21, the Colletidae are morphologically diverse bees, such that one could easily justify recognizing several families among them, as some authors have done. These bees, however, have synapomorphies as indicated below , and it seems reasonable to retain them as a single, large, worldwide family. It is most abundant and most diversified in temperate parts of Australia and South America. In the holarctic region there are only two common genera, Colletes and Hylaeus neotropical...

Floral Relationships of Bees

Outer Surface Tibia Bee

Wind and bees are the world's most important pollinating agents. Bees are either beneficial or actually essential for the pollination, and therefore for the sexual reproduction, of much of the natural vegetation of the world, as well as for many agricultural crops see Sec. 3 . The pollinators are primarily female bees, which collect pollen as the principal protein source in their own food and especially to feed their larvae. Flowers produce not only nectar and sometimes oil but also excess...

Nests and Food Storage

Food Stored Stem Diagram

The nests of bees are the places where their young are reared. They are always to some degree made by the mother, or, in social bees, by the workers. Nests and especially cells, their provisions, and larval behavior are full of meaningful details of importance not only for bee survival but also for our understanding of adaptations and of phylogeny. Malyshev's many papers were among the most important and detailed early studies of these matters, culminating in his summary work Malyshev, 1935 ....

Structures and Anatomical Terminology of Adults

The illustrations and text in this section provide the names for the external parts of adult bees. Many structures are simply labeled in the illustrations. Those that do not appear in the figures, or for which discussion or explanation is needed, are treated in the text, with the preferred terms appearing in boldface type at the places where they are defined or explained. The hope is that the illustrations plus the text will encourage uniformity ofus-age in future work. The emphasis is on...

Development and Reproduction

Pre Defecating

As in all insects that undergo complete metamorphosis, each bee passes through egg, larval, pupal, and adult stages Fig. 4-1 . The haplodiploid system of sex determination has had a major influence on the evolution of the Hymenoptera. As in most Hymenoptera, eggs of bees that have been fertilized develop into females those that are unfertilized develop into males. Sex is controlled by alleles at one or a few loci heterozygosity at the sex-determining locus or loci produces females. Development...

Parasitic and Robber Bees

In many groups of organisms that store food for themselves or their young, parasitic or robber individuals, species, or genera can be found. Such forms steal or feed upon the stored food, often starving or more directly killing the hosts. Bees well illustrate such tendencies reviews were by Bischoff 1927 397-401 , Grutte 1935 , Bohart 1970 , and Iwata 1976 . This section consists of four parts, as follows first, on nest usurpation and robbing second, on social parasites that live in the nests...

Scopa Colletidae

1st m-cu 2nd m-cu 2nd abscissa of Rs 1st r-m 2nd r-m 1st abscissa of R1 2nd abscissa of R1 r and Rs Rs M, 2nd and following abscissae of M It has the drawback that the first is not technically a crossvein, but is thought to be a transverse section of a longitudinal vein, Rs. Louis 1973 reviewed prior alar terminologies and proposed a new nomenclature for veins, attempting to avoid considerations of homology and phylogeny. He called the submarginal crossveins the first to third RM, or...

Family Level Phylogeny and the Proto Bee

Stenotritidae

Phylogeny, presumably, was at least in the back of the minds of the proponents of some of the classifications and their modifications summarized in Section 18. Recent studies, directed toward understanding the relationships of major groups like families, subfamilies, and tribes, provide a more overt and detailed look at bee phy-logeny than was available in the past. These studies Roig-Alsina and Michener, 1993 Alexander and Michener, 1995, and Danforth et al., 2006 , however, leave various...