Protandrena bachue Gonzalez & Ruz, 2007.

The new subgenus can be recognized easily by the following combination of characters: body predominantly dark brown to black with reduced yellow maculations; forewing with two submarginal cells (e.g., Figs 1, 13); propodeum glabrous basally; mesoscutum finely punctate; dorsal surface of propodeum longer than metanotum; anterior tentorial pit at outer subantennal sulcus, just above intersection between outer subantennal and epistomal sulci; female metatibial scopa with sparse, mostly simple setae; and male SVI with broad U- or V-shaped midapical emargination (Figs 8, 16); TVII with distal margin straight or medially emarginate (Fig. 6); and gonostylus simple, without apical lobes or projections, without long apical setae, and completely fused to gonocoxite (Figs 11, 12, 19, 20).

Female:Small to moderate-sized bees (4–12 mm in length); color mainly dark brown to black, nonmetallic, without yellow maculations except on pronotal lip; integument dull to weakly shiny, distinctly imbricate to granular between punctures, especially on dorsal surface of mesosoma and posterior surface of mesofemur; punctures coarser, denser on head than on meso- and metasoma; pubescence predominantly dark brown to black, short, and sparse; pubescence longer and denser on head and mesosoma; metatibial scopa consisting of sparse, long, mostly simple setae; metasomal terga and sterna, except on apical segments, with distal margins glabrous; SVI with dense patch of branched setae laterally (cf. Gonzalez and Ruz 2007: Figs 41–44). Head broader than long, about as wide as mesosoma; mandible edentate, pointed; labrum with strong ridge bordering glabrous, impunctate basal area; clypeus less than three times broader than long; supraclypeal area usually more distinctly convex than clypeus in profile; lower mesal paraocular area gently convex; anterior tentorial pit at outer subantennal sulcus, just above epistomal sulcus; antennal toruli about at middle of face; antennal scape unmodified, not surpassing lower tangent of median ocellus in repose; antennal flagellum about as long as head width or longer, unmodified or crenulate basally on posterior surface; facial fovea well-marked, elongate; compound eyes subparallel; lower margin of median ocellus coinciding with upper orbital tangent; vertex gently convex; gena about as wide as or slightly wider than compound eye in profile, widest dorsally, narrower ventrally; labiomaxillary complex of moderate length, not distinctly elongate; maxillary palpus with six equally long palpomeres; labial palpus with four palpomeres, first palpomere about as long as combined lengths of remaining palpomeres, second palpomere about as long as third and fourth palpomeres individually; glossa about one-half length of prementum; galeal comb composed of 20 bristles. Pronotal collar rounded, not carinate; dorsal surface of propodeum gently sloping to subhorizontal, longer than metanotum, weakly striate. Forewing with pterostigma more than three times longer than broad, about twice as wide as prestigma, margin basal to vein r-rs diverging from costa, that within marginal cell slightly convex; marginal cell obliquely and broadly truncate at apex, appendiculate, slightly longer than distance from its apex to wing tip; two submarginal cells (i.e., 1rs-m absent), first submarginal cell longer than second; basal vein gently curved to nearly straight; 1m-cu distal to 2Rs (second free abscissa Rs, or first submarginal crossvein sensu Michener 2007); 2m-cu basal to 2rs-m (second submarginal crossvein sensu Michener 2007); jugal lobe about three-fourths length of vannal lobe; hind wing with second abscissa of M+Cu more than three times length of cu-a; 6–8 distal hamuli. Legs unmodified; mesofemur without well-developed comb on ventral margin basally; mesotibial spur slightly shorter than mesobasitarsus, straight or nearly so, with coarse branches (sensu Engel 2009) (cf. Gonzalez and Ruz 2007: Figs 39, 40); metatibia about twice as long as metabasitarsus, keirotrichia on inner surface except on anterior and posterior margins; metabasitibial plate carinate, with semierect, short, stiff setae on disc; metatibial spurs slightly curved apically to nearly straight; metabasitarsus strongly projecting on posterodistal margin; pretarsal claws cleft, inner ramus shorter than outer. Metasomal TII with well-marked lateral fovea; pygidial plate subtriangular, well-defined, medially elevated; SVI with distal margin rounded or truncate.

Male: As in female except longer, sparser body pubescence, clypeus often maculate, metabasitibial plate glabrous, and the following: antennal flagellum unmodified (Figs 13, 27) to weakly (Fig. 2) or strongly (Fig. 26) crenulate on posterior surface; compound eyes subparallel to slightly convergent ventrally. Outer surfaces of pro- and mesotibiae apically with small spine; metatibia with posterior marginal carina weakly toothed basally; metabasitarsus with posterodistal margin not distinctly projecting as in female; pretarsal claws symmetrical or with inner ramus shorter than outer. Metasoma usually more elongate than in female, sometimes petiolate; TVII without pygidial plate, distal margin straight or with V-shaped median emargination (Fig. 6); sterna with distal margins straight or convex, except SVI with distinct U- or V-shaped median emargination (Figs 8, 16); SVII with apical lobes attached to small discal area, not distinctly constricted basally, distally retrorse (Figs 9, 17, 30); SVIII longer than broad, midapical projection rather short (about one-half length of disc body), not distinctly constricted basally, broadly rounded apically (Figs 10, 18); genital capsule slightly longer than broad, gonobase absent; gonostylus about as long as penis valves or slightly longer, simple, fully fused to gonocoxite, gently or strongly curved in profile, without long, branched setae on apex; volsella clearly differentiated in medial digitus and lateral cuspis, denticulate, digitus elongate; penis valves simple, narrow, basally fused; penis membranous, bilobed, apically wide, about as long as penis valves (Figs 11, 12, 19, 20).

The new genus-group name is a combination of Andes, referring to the Andean distribution of this group of bees, and Panurgus, type genus of the Panurginae. The name is masculine.

In addition to the type species, Protandrena bachue Gonzalez & Ruz, the subgenus includes the following taxa: Protandrena amyae sp. n., Protandrena femoralis sp. n., Protandrena guarnensis Gonzalez & Ruz, Protandrena maximina Gonzalez & Ruz, Protandrena rangeli Gonzalez & Ruz, and Protandrena wayruronga Gonzalez & Ruz.

The subgenus occurs at mid- and high elevations (1100–3400 m) in the Andes from Venezuela to Peru. Two species groups (one consisting of Protandrena guarnensis and Protandrena femoralis, the other including the remaining species) can be recognized within Andinopanurgus by the characters indicated in the key to species (infra) (Table 2).

The general habitus of Andinopanurgus as well as the shape of the male sixth and seventh sterna suggest species of Rhophitulus Ducke and Heterosarus Robertson but the propodeum is basally pubescent in Rhophitulus and, in both taxa, TVII is gently or strongly projected medially on the distal margin, SVII has lobes with much broader apex, and the gonostylus has long branched setae apically and is partially fused to the gonocoxite, at least ventrally. In addition, Andinopanurgus lacks the distinctive dorsal remnant of the gonobase of Rhophitulus. The rather narrow and distally retrorse apical lobes of SVII of Andinopanurgus (Figs 9, 17) resemble those of Protandrena s.str. and Metapsaenythia Timberlake, but the apex of these lobes lack the distinctive spatulate setae present in the latter. Metapsaenythia has also a propodeum basally pubescent and a metasoma that is frequently red. If future studies demonstrate that Protandrena s.l. is paraphylectic, perhaps some of its subgenera, including Andinopanurgus, may well be recognized at the generic level.

♂ (Fig. 1), Ecuador: Napo. Past. Road from Baeza to Papallacta, km. 188, 13-IV-1977 [13 April 1977], Elaine R. Hodges (USNM).

♂, Ecuador: Napo, Baeza (22 Kms. W.), 15 May 1975, elev. 8000ft. / Collected by sweeping net above damp road bed / Collected by Ashley B. Gurney (SEMC).

The male of this species can easily be recognized by the antennal flagellum weakly crenulate on the posterior surface (Fig. 2), the mandible distinctly broad apically (Figs 4, 5), and the posterior hypostomal carina strongly projecting into a tooth (Fig. 4).

Male: Body length 8.70 mm (8.50 mm); forewing length 6.50 mm (6.70 mm); head width 2.40 mm (2.48 mm). Head 1.4× wider than long; inner orbits of compound eyes subparallel (Fig. 3); intertorular distance 1.7× OD, 0.7× length of torulorbital distance; torulus diameter equal to OD; ocellocular distance 3.4× OD, 2.0× greater than ocelloccipital distance; interocellar distance 1.3× OD; compound eye 1.8× longer than wide; clypeus 2.6× broader than long, projecting about 0.4× compound eye width in lateral view; gena 1.2× broader than compound eye in profile; supraclypeal area, just below inferior torular tangent, distinctly protuberant; frontal line elevated just above antennal toruli to one-half distance between antennal toruli and median ocellus, ending at that point; inner subantennal sulcus about 0.7× length of outer subantennal sulcus; facial fovea 1.7× longer than broad, about one-half length of scape; scape 2.1× longer than broad, antennal flagellum slightly longer than head width, F1–F6 weakly crenulate on posterior surface, not forming deep concavity between flagellomeres (Fig. 2); pedicel about one-third length of F1, about as long as broad, F1 1.8× longer than broad, about 1.5× longer than F2 and F3 individually, remaining flagellomeres about as long as broad, except last flagellomere longer than broad; mandible distinctly broad apically (Figs 4, 5); posterior hypostomal carina strongly projecting into a tooth (Fig. 4). Forewing pterostigma 4.0× longer than broad; prestigma 3.1× longer than broad (prestigma width measured to its margin). Mesosoma slightly narrower than head width; mesoscutum 1.3× wider than long, 2.3× longer than mesoscutellum, 4.5× longer than metanotum; propodeum with basal part about three-fourths of mesoscutellum length in dorsal view; protibial spur with apical portion of rachis long, about three-fourths of malus length, with distinct row of 10 elongate branches (not including apical portion of rachis); mesotibial spur gently curved apically, with coarse branches, less than one-half of mesobasitarsus length; metatibia with posterior marginal carina weakly toothed on upper two-thirds; metatibial spurs slightly curved apically, inner spur slightly longer than outer; pretarsal claws cleft, inner ramus slightly shorter than the outer. Lateral fovea of TII ellipsoid, about 2.0× longer than broad; TVII with V-shaped median emargination on distal margin (Fig. 6); SV–SVIII, and genital capsule as in figures 7–12.

Color dark brown to black, except apex of mandible reddish brown and clypeus with yellow maculation as in figure 3. Wing membranes brownish, veins and pterostigma dark brown.

Head with sparse, long (2.5–3.0× OD), semierect, poorly-branched, black setae except brownish setae on condylar and outer grooves of mandible, gena posteriorly, and hypostomal area; scape with long setae, 2× as long as maximum scape diameter. Pronotum with short (0.5–1.0× OD), dense, brownish setae along dorsal margin and pronotal lobe; mesoscutum, mesoscutellum, and metanotum with two types of setae: sparse, long (2.5–3.0× OD), erect, poorly-branched, black setae, and dense, short (0.5× OD), brownish setae; mesepisternum and lateral and posterior areas of propodeum with mostly sparse, long (2.5–3.0× OD), erect, branched, brownish setae; legs with setae mostly brownish, longer and denser on coxae, trochanters, and profemur. Metasoma with terga mostly bare, with minute (≤ 0.3× OD), semierect, sparse ferruginous setae on discs, laterally with denser and longer setae; TVI with long (2× OD), semierect, dark brown setae on disc, setae denser on TVII; sterna with sparse, short (1× OD), semierect setae, denser and longer on sides of each sternum.

Outer surface of mandible and basal area of labrum smooth and shiny, impunctate; clypeus with sparse (1–1.5× PW), faint punctures, integument between punctures imbricate; supraclypeal area with scattered punctures laterally, weakly imbricate, shinier than on clypeus medially; subantennal area and inferior paraocular area with punctures separated by a puncture width or less, integument strongly imbricate to nearly granular (as on remainder of face); remaining areas of face with coarse punctures, contiguous, smaller than on clypeus; gena strongly imbricate with faint punctures. Mesoscutum, mesoscutellum, and metanotum with small, dense punctures (≤ 1× PW), integument granular between punctures; mesepisternum strongly imbricate with scattered (1–2.0× PW), faint punctures, punctures coarser and denser dorsally; metepisternum transversely weakly striate near wing base, otherwise strongly imbricate. Propodeum strongly imbricate with fine and weak striae basally, lateral and posterior surfaces with faint, scattered punctures. Metasomal terga and sterna shiny, weakly imbricate with minute, scattered (2–3.0× PW) punctures on discs, punctures coarser and denser on TVII; distal margins of terga shiny, weakly imbricate, impunctate except on TVII.

Female: Unknown.

The specific epithet is a matronym honoring Mrs. Amy Comfort de Gonzalez, loving and supporting wife of the senior author.

♂, Peru: Pasco Dept. [Departamento] San Miguel Eneñas, NW Villa Rica-Puerto Bermudas Rd., 1780 m, 10°44'0"S, 75°11'54"W, 16 Oct 1999; R. Brooks, PERU1B99 037, ex: yellow composite / SM0149359, KUNHM-ENT [barcode label] (SEMC).

(n = 3♀♀, 12♂♂) 2♂♂ with same date as the holotype; 1♂, Pasco Dept. San Juan, Villa Rica-Puerto Bermudas Rd., Rio Cacazu, 1100 m, 10°39'12"S, 75°6'54"W, 16 Oct 1999; R. Brooks, PERU1B99 034A, ex: on flowering tree; 3♀♀, 9♂♂, Pasco Dept. Villa Rica Rd., 1475 m, 10°47'6"S, 75°18'54"W, 15 Oct 1999; R. Brooks, D. Brzoska, PERU1B99 030 (SEMC).

Both sexes of Protandrena femoralis are most similar to Protandrena guarnensis from northwestern Colombia in their small body size (4.5–6.0 mm in body length), F1 about as long as F2, absence of maculations on the male clypeus, male TVII with a straight distal margin, male SV without spines on the midapical margin, and general shape of the genitalia and hidden sterna. The male can easily be separated by the structure of SVII, which has broader apical lobes (cf. Figs 17 and 30), and the gonostylus, which is more robust in profile than that of Protandrena guarnensis, about the same width across its length, and basally not protuberant on the medial margin in dorsal view (Figs 19, 20). The female can be recognized by the posterior surface of the mesofemur and anterior and posterior surfaces of the metafemur distinctly depressed (Figs 23–25). In Protandrena guarnensis the apical lobes of SVII are narrow, parallel-sided, with the retrorse section of the apex comma-shaped (Fig. 30); the gonostylus is slender in profile, slightly tapering towards the apex, and strongly protuberant basally on the medial margin in dorsal view (cf. Gonzalez and Ruz 2007: Figs 28, 32). The meso- and metafemora of the female of Protandrena guarnensis are unmodified, not distinctly depressed.

Male: Body length 5.0 mm (4.73–5.33 mm); forewing length 4.47 mm (4.47–4.60 mm); head width 1.50 mm (1.50–1.60 mm). Head 1.4× wider than long; inner orbits of compound eyes converging below (Figs 14); intertorular distance 2.3× OD, 0.9× length of torulorbital distance; torulus diameter equal to OD; ocellocular distance 3.4× OD, 2.8× greater than ocelloccipital distance; interocellar distance 1.3× OD; compound eye 1.8× longer than wide; clypeus 2.4× broader than long, projecting about 0.3× compound eye width in lateral view; gena 0.8× width of compound eye in profile; supraclypeal area, just below inferior torular tangent, distinctly protuberant; frontal line weakly elevated just above antennal toruli to one-half distance between antennal toruli and median ocellus, ending at that point; inner subantennal sulcus about 0.7× length of outer subantennal sulcus; facial fovea about 2.0× longer than broad, 0.4× length of scape; scape 2.1× longer than broad, antennal flagellum unmodified, slightly longer than head width (Figs 13–14); pedicel slightly shorter than F1, about as long as broad, F1 about as long as broad, subequal to F2 and F3 individually, remaining flagellomeres about as long as broad, except last flagellomere longer than broad; mandible pointed. Forewing prestigma 3.2× longer than broad (prestigma width measured to its margin); pterostigma 3.6× longer than broad. Mesosoma narrower than head width; mesoscutum 1.3× wider than long, 2.7× longer than mesoscutellum, 5.7× longer than metanotum; propodeum with basal part about three-fourths of mesoscutellum length in dorsal view; protibial spur with apical portion of rachis long, about one-half length of malus, with distinct row of about 10 elongate branches (not including apical portion of rachis); mesotibial spur straight or nearly so, with coarse branches, slightly more than one-half mesobasitarsus length; metatibia with posterior marginal carina weakly toothed on upper third; metatibial spurs of similar length, slightly curved apically; pretarsal claws with rami of similar length. Lateral fovea of TII elongate, about 4.0× longer than broad; TVII with distal margin straight or nearly so; SV–SVIII, and genital capsule as in figures 15–20.

Color dark reddish brown to black, without yellow maculations. Wing membranes subhyaline, slightly brownish, veins and pterostigma dark brown.

Head with sparse, long (2.5–3.0× OD), semierect, poorly-branched, black setae except brownish setae on condylar and outer grooves of mandible, gena posteriorly, and hypostomal area; scape with long setae, 2× as long as maximum scape diameter. Pronotum with short (0.5–1.0× OD), dense, brownish setae along dorsal margin and pronotal lobe; mesoscutum and mesoscutellum with two types of dark brown setae: sparse, long (1.5–2.0× OD), erect, poorly-branched setae, and short (0.5–1.0× OD), slightly denser setae; metanotum with short setae as on mesoscutellum; mesepisternum and lateral and posterior areas of propodeum with very sparse, long (1.5–2.0× OD), erect, branched, brownish setae; legs with setae mostly brownish, longer and denser on coxae, trochanters, and profemur. Metasomal terga with minute (≤ 0.3× OD), semierect, dense ferruginous setae on discs, laterally with denser and longer setae; TVI with long (1.5–2× OD), semierect, dark brown setae on disc, setae denser on TVII; sterna with sparse, short, semierect setae (1.5× OD) on discs, denser and longer laterally; preapical margin of SIV with few, semierect thick setae, each seta consisting of short rachis with three or four long branches, resembling scales or bundles of several setae at low magnifications.

Outer surface of mandible and basal area of labrum smooth and shiny, impunctate; clypeus with sparse (1–1.5× PW), faint punctures, integument between punctures weakly imbricate basally, becoming nearly smooth and shiny toward apex; supraclypeal area with scattered punctures laterally, weakly imbricate, medially shiny as on clypeus; remaining areas of face with coarse punctures separated by a puncture width or less, integument strongly imbricate to nearly granular, punctures becoming weaker and sparser on vertex; gena strongly imbricate with faint punctures. Mesoscutum, mesoscutellum, and metanotum with small, sparse punctures, integument granular between punctures; mesepisternum strongly imbricate with scattered (1–2.0× PW), faint punctures, punctures coarser and denser dorsally; metepisternum transversely weakly striate near wing base, otherwise strongly imbricate. Propodeum strongly imbricate with few fine, weak striae basally (barely visible), lateral and posterior surfaces with faint, scattered punctures. Metasomal terga and sterna shiny, lineolate-imbricate with minute punctures separated by about two puncture widths on discs, punctures coarser and denser on TVII, sparser on sterna; distal margins of terga shiny, weakly imbricate, impunctate except on TVII.

Female: As in male except shorter body pubescence, lighter and shinier integument (Fig. 21), and the following: Body length 5.53–5.73 mm; forewing length 4.87–5.0 mm; head width 1.60–1.67 mm. Inner orbits of compound eyes subparallel (Fig. 22); intertorular distance 2.6× OD, about as long as torulorbital distance; torulus diameter subequal to OD; ocellocular distance 3.5× OD, 2.5× greater than ocelloccipital distance; interocellar distance 1.6× OD; compound eye 2.1× longer than wide; clypeus 2.9× broader than long, projecting about 0.4× compound eye width in lateral view; supraclypeal area gently convex, not distinctly protuberant medially; facial fovea 3.3× times longer than broad, 0.7× length of scape; scape 2.7× longer than broad, antennal flagellum about as long as head width. Forewing prestigma 5.0× longer than broad (prestigma width measured to its margin); pterostigma 4.5× longer than broad. Mesoscutum about 5.0× longer than metanotum; protibial spur with apical portion of rachis about three-fourths length of malus, with about five branches (not including apical portion of rachis); mesotibial spur about 0.7× mesobasitarsus length; mesofemur with posterior surface and metafemur with anterior and posterior surfaces distinctly depressed (Figs 23–25); inner metatibial spur slightly shorter than outer; pretarsal claws with inner ramus shorter than the outer. Lateral fovea of TII about 3.0× longer than broad.

The specific epithet refers to the distinctly depressed meso- and metafemora of the female of the species.

1♂, Ecuador, Pich. [Pichincha], Quito (48 KmS), 6 May 1975, Ashley Gurney (USNM).

This species was previously known from the male holotype collected in Papallacta, Napo (Ecuador). Gonzalez and Ruz (2007) tentatively identified a single specimen from Cauca, Colombia, as the female of this species. The putative female is very similar to another one from Valle, Colombia, except in the length of the apex of the protibial spur; in the specimen from Cauca the apex is long, about one-half of the malus length, whereas in the female from Valle it is short, about one-third of the malus length, as it is in the holotype. Such a difference in both females suggests that they might not be conspecific, although we do not know how variable this character is within and among species. The complete label data of the female from Valle are as follows: “Colombia: Valle, Tenerife, Paramo at 12, 000’ R. E. Dietz, Sept. 15, 1970” (USNM).