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6(5). 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 impunctate; apical an-tennal segments of male unmodified or rarely crenulate, or last one rarely broadened and flattened 7

7(6). Claws (at least ofmale, those offemale unknown) each with broad, flat inner ramus arising near base (Fig. 39-4g); episternal groove below scrobal groove represented only by weak, short depression; strong carina just behind posterior orbit (Australia) Hesperocolletes

—. Claws cleft apically in male and usually in female, inner ramus pointed like outer ramus or sometimes in female reduced to a tooth or absent (in one undescribed species of Leioproctus somewhat like those of Hesperocolletes); episternal groove distinct below scrobal groove; no strong carina behind posterior orbit 8

8(7). S8 of male with median apical process slender and hairy at apex, pale, not exposed (Fig. 39-3b); volsella large, produced posteriorly, reaching beyond apex of articulated gonostylus (Fig. 39-3a); stigma less than one-half as long as marginal cell, measured on wing margin (Fig. 39-5a); apex of marginal cell bent gradually from wing margin for about one-sixth length of cell and bearing long appendage; profile of propodeum nearly vertical (Australia) Chrysocolletes

—. S8 of male with median apical process robust, heavily sclerotized (Fig. 39-6c, g), its apex suggesting a pygidial plate and commonly exposed in repose, or (in some species of subgenus Goniocolletes) process broadened and appearing as extension of elongated disc (Fig. 39-7e); volsella not reaching gonostylus or reaching only its basal part, gonostylus fused to gonocoxite; stigma usually larger; apex of marginal cell only minutely bent away from wing margin; profile of propodeum usually with sloping or subhorizontal basal zone (Australian area) Leioproctus

Genus Brachyglossula Hedicke

Brachyglossa Friese, 1922a: 577, not Boisduval, 1829. Type species: Brachyglossa rufocaerulea Friese, 1922, monobasic. Brachyglossula Hedicke, 1922: 427, replacement for Brachy-glossa Friese, 1922. Type species: Brachyglossa rufocaerulea Friese, 1922, autobasic.

This genus of large (body length 12-16 mm), dark-haired, unbanded bees is distinctive in appearance, except for the superficially similar Leioproctus (Cephalocolletes) laticeps (Friese). In some major features, such as the vesti-ture and form of the hind basitarsus of the female and the shape of the process of S8 of the male, Brachyglossula resembles Leioproctus. The other characters are sufficiently marked and unique, however, to support recognition at

Figure 39-1. Brachyglossula bouvieri (Vachal). a, b, Dorsoventral and lateral views of male genitalia; c, d, Dorsoventral views of S8 and S7 of male; e, f, Mandibles of male and female. In the divided drawings, dorsal views are at the left. From Michener, 1989.

the genus level, even though the result, at least for the time being, is a probably paraphyletic genus Leioproctus. The integument is black or metallic blue, the metasoma in some species red. The inner orbits diverge below; the facial fovea is as described for Lonchopria. The preapical tooth of the mandible of the male is broad and weakly to strongly emarginate, the mandible thus at least weakly tri-dentate (Fig. 39-1e). All scopal hairs have strong axes and numerous fine side branches at more or less right angles to the axes. T1 is markedly narrower than T2. S2-S5 of the female support a well-developed, dense, plumose scopa. For illustrations of the male genitalia and sterna, see Figure 39-1a-d and Michener (1989).

Brachyglossula is found in Bolivia and Argentina (provinces of Misiones to Catamarca). There are four named species.

Brachyglossula may collect pollen principally from cactus flowers.

Genus Callomelitta Smith

Callomelitta Smith, 1853: 85. Type species: Callomelitta picta

Smith, 1853, monobasic. Binghamiella Cockerell, 1907c: 235. Type species: Sphecodes antipodes Smith, 1853, monobasic. Binghamiella (Pachyodonta) Rayment, 1954: 48. Type species: Binghamiella fulvicornis Rayment, 1954, monobasic.

Figure 39-1. Brachyglossula bouvieri (Vachal). a, b, Dorsoventral and lateral views of male genitalia; c, d, Dorsoventral views of S8 and S7 of male; e, f, Mandibles of male and female. In the divided drawings, dorsal views are at the left. From Michener, 1989.

This is a genus ofblack or metallic blue-black bees with the head and thorax coarsely punctate, often with the metasoma and sometimes with parts of the thorax red. The body length is 6 to 10 mm. There are no conspicuous metasomal hair bands, and the prepygidial and py-gidial fimbriae are of rather dense but short hairs. The ju-gal lobe ofthe hind wing is nearly five-sixths as long as the vannal lobe (Fig. 39-2) (one-fourth to three-fourths as long in most other Australian Colletinae). There are three submarginal cells. The inner hind tibial spur of the female is finely ciliate (Fig. 39-2) and the basitibial plate is one-third as long as the tibia or nearly so, the margins sometimes nodulose; in the male the basitibial plate is one-fifth to one-sixth as long as the tibia. For figures of genitalic and sternal characters as well as wing venation, see Mich-ener (1965b).

Callomelitta ranges from Tasmania to northern Queensland, west to Western Australia. Most species are from eastern Australia. The 11 species were listed by Michener (1965b) and Cardale (1993).

The body form, coarse punctation, nonmetallic coloration, and red metasoma characteristic of Callomelitta

Figure 39-2. Callomelittapicta Smith. a, Wings; b, Inner hind tibial spur of female. From Michener, 1965b.

antipodes (Smith) produce a superficial similarity to a species of Sphecodes (Halictidae), and F. Smith accordingly described the species in that genus. The other ten species, however, because of their usually metallic coloration and more elongate body form, do not resemble Sphecodes.

Genus Chrysocolletes Michener

Leioproctus (Chrysocolletes) Michener, 1965b: 71. Type species: Paracolletes moretonianus Cockerell, 1905, by original designation.

This genus suggests Phenacollees in its occipital development, the median ocellus being about midway between the antennal bases and the posterior edge of the vertex. The appressed, short pubescence of the metasoma also suggests Phenacolletes, but the rest of the body and the legs have the ordinary plumose hairs characteristic of Leioproctus. Body length is 9 to 13 mm. The marginal cell is bent away from the costa for a greater distance than in most Leioproctus (Fig. 39-5a). The male is unique in having the inner prongs of the hind claws much nearer the bases of the claws than is the case on the other legs. The facial foveae are absent. There are three submarginal cells; the stigma is subparallel-sided, less than half as long as the costal edge of the marginal cell (Fig. 39-5a). The inner hind tibial spur of the female is finely pectinate; that of the male, ciliate. S7 and S8 and the enormous volsellae are also distinctive; see Figure 39-3a-c and Michener (1965b) and Maynard (1996).

Chrysocolletes, which has been found in Queensland, New South Wales, Northern Territory, and Western Australia, comprises at least five species. The genus was revised by Maynard (1996).

Chrysocolletes moretonianus (Cockerell) is probably restricted in pollen collecting to species of Goodeniaceae.

Although originally described as a subgenus of Leio-proctus, Chrysocolletes differs from Leioproctus in many characters, and its phylogenetic position is obscure. Leio-proctus crenulatus Michener was placed in Chrysocolletes by Michener (1965b) and was thought to be intermediate between Leioproctus s. str. and Chrysocolletes. It now seems that the characters in which it resembles Chryso-

Figure 39-3. Male terminalia of Colletinae. a-c, Genitalia, S8, and S7 of Chrysocolletes moretonianus (Cockerell); d-f, Genitalia, S8, and S7 of Phenacolletes mimus Cockerell. (Dorsal views are at the left.) From Michener, 1965b.

colletes (e.g., the crenulate flagellum ofthe male) are probably convergent. Leioproctus crenulatus is transferred to the subgenus Leioproctus s. str., at least until that group is properly revised.

Genus Eulonchopria Brethes

This is a genus of coarsely sculptured, nonmetallic bees 8 to 11 mm long with yellow apical integumental bands on at least some of the metasomal terga, but without hair fasciae on the terga, and often with plaited (longitudinally folded) forewings with darkened costal margins (Dan-forth and Michener, 1988), the whole thus yielding a superficial resemblance to eumenine wasps. The pubescence is short; in Eulonchopria s. str., at the anterior and posterior scutal angles and often elsewhere, the hairs are so short that each fits inside a puncture and is broadly plumose. Facial foveae are absent or deeply impressed and well defined in both sexes; when present, they are elongate and low on the face, that is, not reaching the summits of the eyes. The propodeal triangle has large, deep pits; some of the ridges margining the pits are produced, lamella-like or toothlike. The horizontal and vertical surfaces of the propodeum are separated by a sharp angle or lamella. The front basitarsus of the female in Eulon-chopria s. str. ends with an outer apical process from which a comb extends basad on the outer edge of the ba-sitarsus; in Ethalonchopria the comb is present but the apical process is absent. The inner hind tibial spur of the female is coarsely pectinate (three to five teeth); that of the male is coarsely toothed or ciliate, or the hind tibial spurs are completely absent. The hairs of the outer surface of the hind tibia of the female are short and not scopa-like, especially on the distal half of the tibia. The basitib-ial plate of the male ends in a carina extending to the apex of the tibia. The stigma is nearly parallel-sided, vein r arising near the apex, and the margin within the marginal cell is oblique, not convex. There are three submarginal cells; the apex of the marginal cell is obliquely bent away from the wing margin or obliquely truncate. The genitalia and hidden sterna of males were illustrated by Michener (1963a), the wing venation by Danforth and Michener (1988).

Typical members of this genus possess various probable apomorphies that are unusual among bees and not shared by related groups (Leioproctus and other Collet-inae). The deep, rather slender, bare, well-defined facial foveae (of one subgenus), however, are shared with the Australian genus Callomelitta and with some species of the African genus Scrapter, as well as with the Euryglossi-nae and Hylaeinae. This character is probably a ple-siomorphy. Likewise, the distinct, slender male gonostyli of some species, unique for the Colletidae, are a possible plesiomorphy. The apparently disjunct distribution of Eulonchopria (Americas but absent in wet tropics), combined with its unusual characters and the morphological diversity of its species, suggests that Eulonchopria is an archaic group possessing many derived features.

Leioproctus simplicicrus Michener (1989) from Peru, originally incorrectly placed as an unusual species of L. (Nomiocolletes), to which it runs in the keys to genera and subgenera, seems to connect Eulonchopria and Leioproc-tus, suggesting that Eulonchopria is a specialized deriva tive of the large paraphyletic genus Leioproctus. G. Melo pointed out to me that L. simplicicrus resembles Eulon-chopria in its short pubescence, the obliquely truncate apex of the marginal cell, the apical yellow tergal bands, the carina along the hind tibia of the male, and (especially) its genitalia and hidden sterna (compare Michener, 1963a and 1989). (The female is unknown, and the characters of the scopa cannot be determined.) On the other hand, L. simplicicrus differs from Eulonchopria and resembles Leioproctus in the absence of a preoccipital carina, the absence of a lamella from the posterior pronotal lobe to the dorsum of the pronotum, and the relatively large stigma, which is broadest at the base of vein r and convex within the marginal cell. The propodeal triangle has a few weak rugae basally but lacks the sharp carinae or lamellae characteristic of Eulonchopria s. str. When both sexes are known, L. simplicicrus may well be placed as a distinct genus or subgenus; females of an unnamed species from Brazil may fall in the same group.

Key to the Subgenera of Eulonchopria

1. Facial fovea absent; omaulus not carinate

E. (Ethalonchopria)

—. Facial fovea distinct; omaulus carinate

Eulonchopria / Subgenus Ethalonchopria Michener

Eulonchopria (Ethalonchopria)Michener, 1989: 670. Type species: ApistagaulleiVachal, 1909, by original designation.

This subgenus differs in many features from the other subgenus. In nearly all of the subgeneric characters it is less strange than Eulonchopria s. str., that is, more like other colletines. Noteworthy are the punctate and only slightly concave foveal areas on the face, such that distinct foveae are absent; the simple axillae; and the jugal lobe of the hind wing, which extends little more than halfway from the wing base to the level of vein cu-v. Although most of the subgeneric characters are plesiomorphic relative to Eulonchopria s. str., the small jugal lobe and the small second submarginal cell are apomorphic. This subgenus is probably the sister group to Eulonchopria s. str.

The subgenus is known from Bolivia, southern Brazil, and eastern Colombia. The two species names, both dating from Vachal (1909), may represent only one species.

Eulonchopria / Subgenus Eulonchopria Brethes s. str.

Eulonchopria Brethes, 1909a: 247. Type species: Eulonchopria psaenythioides Brethes, 1909, monobasic.

This subgenus contains the more ornate and extraordinary members of the genus. The carinate omaulus and produced, angulate axillae are especially unusual. The preoccipital ridge is expanded as a strong lamella. The carina on the upper margin of the hind tibia of the male is toothed. The jugal lobe of the hind wing nearly attains the level of cu-v. The apex of T7 of the male is bilobed or bidentate.

This subgenus ranges from Paraguay, Argentina (Salta province), and Brazil (Santa Catarina to Minas Gerais)

northward to Venezuela, Colombia, Nicaragua, Mexico (Oaxaca to Sonora), and the USA (southern Arizona). The three named species, only one of them from South America, were revised by Michener (1963a); there are additional (undescribed) species in South America.

The species of Eulonchopria may be oligolectic visitors to flowers of Acacia; at least the North and Central American species appear to collect pollen regularly from that plant.

Eulonchopria s. str. contains two species groups. In the South American E. psaenythioides Brethes and its unde-scribed relatives, the hind tibial spurs of the male are present, S8 of the male lacks an apical process, and there are no slender male gonostyli. In the two North and Central American species, E. punctatissima Michener and oaxa-cana Michener, the hind tibial spurs of the male are absent, S8 of the male has a small, apically expanded process, and there is a distinct, slender, hairy gonostylus.

Genus Glossurocolletes Michener

Leioproctus (Glossurocolletes)Michener, 1965b: 60. Type species: Leioproctus bilobatusMichener, 1965, by original designation.

This genus includes species that have hitherto been included in Leioproctus, but seem so different as to justify generic status. They agree in various features with Leioproctus (Protomorpha), although the species are larger (about 7.5 mm long) than the average of that group and more robust. In these respects, the lack of metasomal fasciae, and the reduced apical lobes of S7, this genus also resembles Leioproctus (Odontocolletes). Noteworthy distin guishing features of the male of Glossurocolletes are the apically modified antenna (Fig. 39-4d) [differing in type of modification from those of Leioproctus (Ceratocol-letes)], the swollen antennal scape, the broad mandible (in males three times as long as the basal width, less than four times as long as the minimum width), and the strong longitudinal ridge on the supraclypeal area. The really striking feature of the group, however, is the enormously elongate and biligulate apex ofS8 of the male (Fig. 39-4b), the apices of the two lobes being exposed in undissected specimens. S7 of the male does not have a narrow neck where the apodemes join the small (almost insignificant) disc of the sternum, as is usual in colletids. Instead, the apo-demes are broadly joined to one another, forming the transverse body of the sternum (Fig. 39-4c). Thus, the S7 of this genus does not look like the S7 of a colletid.

The following are additional characters of Glossurocol-letes. The facial fovea is about three times as long as broad, impunctate but minutely roughened, and distinctly impressed. The eyes bear scattered hairs [very short and inconspicuous in G. xenoceratus (Michener)]. Of the three submarginal cells, the second is unusually small (Fig. 395 c); the stigma is small, not parallel-sided, with vein r arising near its middle. The hind tibial spurs of both sexes are thick and strongly curved; both the outer margin of the outer spur and the inner margin of the inner spur are pectinate with coarse teeth. For illustrations of the genitalia, sterna, and antennae, see Figure 39-4 and Michener (1965b).

Ghssurocolletes is found in Western Australia. The named species are G. bilobatus (Michener) and xenoceratus (Michener), the two distinguished by Michener (1965b).

Figure 39-4. Details of para-colletine structures. a-c, Male genitalia, S8, and S7 of Glossurocolletes bilobatus (Michener); d, Male antennal flagellum of G. xenoceratus (Michener); e-j, Hesperocol-letes douglasiMichener, male, portion of wing, claws and arolium, side view of claw, genitalia, S8, and S7. (In the divided drawings dorsal views are at the left. From Michener, 1965b.

Figure 39-4. Details of para-colletine structures. a-c, Male genitalia, S8, and S7 of Glossurocolletes bilobatus (Michener); d, Male antennal flagellum of G. xenoceratus (Michener); e-j, Hesperocol-letes douglasiMichener, male, portion of wing, claws and arolium, side view of claw, genitalia, S8, and S7. (In the divided drawings dorsal views are at the left. From Michener, 1965b.

Genus Hesperocolletes Michener

Hesperocolletes Michener, 1965b: 75. Type species: Hesperocolletes douglasi Michener, 1965, by original designation.

This genus, known only in the male, consists a moderate-sized (body length 12 mm), nonmetallic species similar in appearance to Trichocolletes and Paracolletes (especially P. crassipes Smith), as well as to Leioproctus. The stigma (Fig. 39-4e) is like that in those groups of Leioproctus having a slender stigma, such as the subgenus Go-niocolletes. There are three submarginal cells, the second and third equal in length and together slightly longer than the first. The wholly vertical propodeum, as seen in profile, is almost matched in a few groups of Leioproctus, but the lack of a defined basitibial plate in the male separates Hesperocolletes from nearly all Leioproctus. The rather protuberant and yellowish clypeus of the male suggests that of Paracolletes and Trichocolletes. The most distinctive generic characters (male only) are (1) the strong carina around, and especially behind, each eye; (2) the near absence of the episternal groove (only a shallow concavity) below the scrobal groove; and (3) the deeply cleft claws, their inner prongs broad, flat, and directed more or less ventrally (Fig. 39-4g), suggesting the claws of many clep-toparasitic bees such as some female Melectini (An-thophorinae). For illustrations of structures including male genitalia and sterna, see Figure 39-4 and Michener (1965b).

This genus is known from a single specimen reportedly from Rottnest Island, Western Australia. Additional material of this genus has not been found at the type locality. The specimen had been handled by T. Rayment, who is known to have been careless about switching locality labels on specimens. The specimen certainly came from somewhere in Western Australia, however.

Genus Leioproctus Smith

This huge genus, with its many subgenera, is found both in the Australian area and in South America, principally in temperate parts of that continent. In the appearance, abundance, and diversity of its species it is equivalent to the genus Andrena of the holarctic region. The integument is black, metallic blue or green, or sometimes red on the metasoma; metasomal hair bands and colored integumental bands may be present or absent. The clypeus is usually rather uniformly punctured, with or without a weak, upper-median, depressed, densely punctate area, or if such an area is strongly evident, as in the subgenus Kylopasiphae and a few species of the subgenus Perditomorpha, then this area is often surrounded by extensive impunctate, usually convex areas, as in the genus Lonchopria. The facial foveae are usually not recognizable or are indicated by broad, weakly defined areas of slightly different texture, or (in a few Australian subgenera) the foveae are rather well defined, depressed, and rather broad (not groovelike). The mandible of females and most males has a preapical tooth on the upper margin; the expansion or tooth found on the lower margin in many male Lonchopria is absent. The labrum is usually more than three times as wide as long. The front basitar-sus of the female usually lacks a well-formed comb of hairs on the outer margin, but such a row of hairs is present in the subgenera Cephalocolletes, Nomiocolletes, Reedapis, and Spinolapis. The inner hind tibial spur of the female is ciliate to coarsely pectinate, the bases of the teeth forming a uniform series, not crowded and diverging from one another as in females of Lonchopria and Trichocolletes. The tibial scopa, unlike that of Lonchopria, is not so dense as to completely obscure the tibial surface; the hairs of the inner surface of the hind tibia of the females are moderately long, simple (at least in a limited area), and do not form a zone of short keirotrichia. The hind basitarsus of the female tapers slightly, but the apex is more than half as wide as the maximum width near the base; the outer surface is flat or convex, its vestiture being superficially rather similar to that of the tibia, and its hairs are longer than those of the inner surface. The basitibial plate of the female is usually well defined, pointed or rounded, and sometimes hidden by hair, but it is sometimes absent in the subgenus Excolletes. The basitibial plate of the male is defined, but sometimes one margin is missing. There may be either two or three submarginal cells; if three, the second is usually much shorter than the third on the posterior margin. The stigma is moderate-sized, vein r arising at or slightly beyond its middle, and the margin within the marginal cell is usually convex (Fig. 39-5d-l). The sternal scopa is present or absent. The genitalia and sterna of many species and nearly all subgenera were illustrated by Michener (1965b, 1989) and in other works cited therein; see Figures 39-6 and 39-7.

A feature of this genus is that, although membership in the genus is indicated by a complex of characters, no one of these is invariable, and indeed characters that elsewhere among bees are often regarded as of generic importance vary within this genus. Thus arolia are absent from the subgenus Urocolletes; claws are simple instead of cleft in the females of several groups and all intermediate conditions are found in the platycephalus group of the subgenus Leioproctuss. str. and in the subgenus Euryglos-sidia; basitibial plates are partly gone in one species of the subgenus Lamprocolletes and wholly gone in females of some species of Excolletes; and the apical lobes of S7 of the male, characteristically present in this family, vary from four in number (subgenera Andrenopsis, Ceratocolletes, and Euryglossidia and the advenus group of Leioproctus s. str.) to two in most forms, these lobes very much reduced in the subgenera Nesocolletes and Odontocolletes.

A character that has often been considered as of generic or subgeneric importance, namely the inner hind tibial spur of the female (ciliate or pectinate, Fig. 39-8) is not always even a subgeneric character. It varies among species of the subgenera Leioproctus s. str., Euryglossidia, and Perditomorpha. Likewise, the number of submarginal cells, though usually a good subgeneric character (i.e., constant among species that closely resemble one another in other features), varies among species of the subgenera Sarocolletes and Nomiocolletes and even among individuals of an Australian species of Leioproctus s. str., L. ab-normis (Cockerell).

The subgenera in South America are mostly well-defined taxa; those in Australia are less well understood. In Australia there is partial intergradation among some of the taxa that are here called subgenera; at least some of the group characters break down. The problem confronting

(Cockerell); g, L. (Andrenopsis) flavorufus (Cockerell); h, L. (Baeo-colletes) calcaratus Michener; i, L. (Nesocolletes) fulvescens (Smith); j, L. (Urocolletes) rhodurus Michener; k, L. (Leioproctus) unguidentatus Michener; l, L. (Excolletes) impatellatus Michener. From Michener, 1989.

Figure 39-5 Wings and parts of forewings of Colletinae of the Australian region. a, Chrysocolletes moretonianus (Cockerell); b, Phenacolletes mimus Cockerell; c, Glossurocolletesxenoceratus Michener; d, Leioproctus (Protomorpha) tarsalis (Rayment); e, L. (Filiglossa) filamentosus (Rayment); f, L. (Nodocolletes) dentiger

Figure 39-5 Wings and parts of forewings of Colletinae of the Australian region. a, Chrysocolletes moretonianus (Cockerell); b, Phenacolletes mimus Cockerell; c, Glossurocolletesxenoceratus Michener; d, Leioproctus (Protomorpha) tarsalis (Rayment); e, L. (Filiglossa) filamentosus (Rayment); f, L. (Nodocolletes) dentiger

(Cockerell); g, L. (Andrenopsis) flavorufus (Cockerell); h, L. (Baeo-colletes) calcaratus Michener; i, L. (Nesocolletes) fulvescens (Smith); j, L. (Urocolletes) rhodurus Michener; k, L. (Leioproctus) unguidentatus Michener; l, L. (Excolletes) impatellatus Michener. From Michener, 1989.

a comprehensive review of the situation at this time is that there are many unusual undescribed species in Australia, and many of the described species are known in only one sex. I believe that Australian workers, particularly G. V. Maynard, are rectifying this situation, and in fact her as yet unpublished thesis will help greatly, but I can present only the currently published information and indicate some of these problems in the discussions of various subgenera.

Leioproctus occurs in Australia (north to New Guinea and Misool, an island west of New Guinea), Tasmania, New Caledonia, New Zealand, and temperate South America. It is probably a paraphyletic taxon from which Bmchyglossula, Eulonchopria, Lonchopria, and perhaps Lonchorhyncha and Niltonia were derived in South America and from which Glossurocolletes, Neopasiphae, Para-

colletes, Phenacolletes, and Trichocolletes and perhaps Chrysocolletes and Hesperocolletes were derived in Australia. Leioproctus is nonetheless definable and generally useful taxonomically, in spite of intergrading through the one species of the subgenus I^nchoprel^la to the genus Lonchopria. Until more species are known and cladogeny is better studied, especially in the diverse Australian fauna, there is no point in attempting a new classification, for new combinations that would last only until the next revision would be the result. Table 39-1 lists the subgenera from the two areas where Leioproctus exists.

There is no known character that distinguishes the Australian and South American groups of the genus. In the absence of a phylogenetic analysis I would not use this fact in any biogeographical sense, for Leioproctus s. str. simply consists of Leioproctus species not readily at-

Figure 39-6. Male genitalia and hidden sterna of Leioproc-tus, showing similarity of certain subgenera. a-d, Dorsoven-tral and lateral views of genitalia, S8, and S7 of L. (Perditomorpha) eulonchopri-odes Michener; e-h, Same structures of L. (Halictan-threna) malpighiacearum (Ducke). (Dorsal views are at the left.) From Michener, 1989.

Figure 39-6. Male genitalia and hidden sterna of Leioproc-tus, showing similarity of certain subgenera. a-d, Dorsoven-tral and lateral views of genitalia, S8, and S7 of L. (Perditomorpha) eulonchopri-odes Michener; e-h, Same structures of L. (Halictan-threna) malpighiacearum (Ducke). (Dorsal views are at the left.) From Michener, 1989.

Figure 39-7. Male terminalia of Leioproctus, showing variation in S7 among diverse subgenera. a, L. (Perditomorpha) brunerii (Ash-mead); b, L. (Holmbergeria) rubriventris (Friese); c, L. (Ky-lopasiphae) pruinosus Michener. d-f, Male genitalia, S8, and S7 of L. (Goniocolletes) dolosus Michener. a-c, from Michener, 1989; d-f, from Michener, 1965b.

Figure 39-7. Male terminalia of Leioproctus, showing variation in S7 among diverse subgenera. a, L. (Perditomorpha) brunerii (Ash-mead); b, L. (Holmbergeria) rubriventris (Friese); c, L. (Ky-lopasiphae) pruinosus Michener. d-f, Male genitalia, S8, and S7 of L. (Goniocolletes) dolosus Michener. a-c, from Michener, 1989; d-f, from Michener, 1965b.

Figure 39-8. Variation in inner hind tibial spurs of females of Colleti-nae. a, Leioproctus (Spinolapis) caerulescens (Spinola); b, L. (Perditomorpha) brunerii(Ashmead); c, L. (P.) inconspicuus Mich-ener; d, Brachyglossula bouvieri (Vachal); e, Lonchopria (Lon-choprella) annectens Michener; f, Trichocolletes venustus (Smith). Both a and b are considered ciliate, the others pectinate. a-e, from Michener, 1989; f, from Michner, 1965b.

Figure 39-8. Variation in inner hind tibial spurs of females of Colleti-nae. a, Leioproctus (Spinolapis) caerulescens (Spinola); b, L. (Perditomorpha) brunerii(Ashmead); c, L. (P.) inconspicuus Mich-ener; d, Brachyglossula bouvieri (Vachal); e, Lonchopria (Lon-choprella) annectens Michener; f, Trichocolletes venustus (Smith). Both a and b are considered ciliate, the others pectinate. a-e, from Michener, 1989; f, from Michner, 1965b.

tributable to any other subgenus. It is almost certainly pa-raphyletic. The lone South American species included in the subgenus by Michener (2000) and now placed in Actenosigynes shows no special affinity at the subgenus level with the Australian members of the genus.

Table 39-1. Subgenera of Leioproctus Segregated by Geographical Area.

South America

Australian Region

Actenosigynes

Andrenopsis

Albinapis

Baeocolletes

Cephalocolletes

Ceratocolletes

Chilicolletes

Cladocerapis

Glossopasiphae

Colletellus

Halictanthrena

Colletopsis

Hexantheda

Euryglossidia

Holmbergeria

Excolletes

Hoplocolletes

Filiglossa

Kylopasiphae

Goniocolletes

Nomiocolletes

Lamprocolletes

Perditomorpha

Leioproctus s. str.

Protodiscelis

Nesocolletes

Pygopasiphae

Odontocolletes

Reedapis

Protomorpha

Sarocolletes

Urocolletes

Spinolapis

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