Andinopanurgus, a new Andean subgenus of Protandrena (Hymenoptera, Andrenidae)

Abstract A new subgenus of Protandrena Cockerell (Panurginae: Protandrenini) from South America, Andinopanurgus Gonzalez & Engel, subgen. n., is described and figured for distinctive species of the genus occurring at mid- and high elevations in the Andes from Venezuela to Peru (1100–3400 m). In addition to the distribution, the subgenus is easily distinguished from other subgenera by a unique combination of morphological characters in both sexes, especially in the hidden sterna and genitalia of the male. Protandrena amyae sp. n., and Protandrena femoralis sp. n., are also described and figured from the Ecuadorian and Peruvian Andes. New geographical records and a key to the species are also provided.


Introduction
Panurgine bees of the tribe Protandrenini (sensu Michener 2007) are restricted to the Western Hemisphere where they are diverse and abundant in temperate areas of the Americas table 1. Summary of generic and subgeneric classification of Protandrenini (sensu Michener 2007), with the new subgenus included. Those subgenera still recognized at the generic level by some authors are indicated with an asterisk; note that Neffapis has been segregated into a separate tribe, Neffapini (Engel 2005). The distribution and approximate number of species are based on Michener (2007), Moure et al. (2008), and Ascher and Pickering (2011). NA = North America; CA = Central America; SA = South America.
Description. 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.

Etymology.
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.
Included species. In addition to the type species, P. bachue Gonzalez & Ruz, the subgenus includes the following taxa: P. amyae sp. n., P. femoralis sp. n., P. guarnensis Gonzalez & Ruz, P. maximina Gonzalez & Ruz, P. rangeli Gonzalez & Ruz, and P. wayruronga Gonzalez & Ruz. Comments. The subgenus occurs at mid-and high elevations (1100-3400 m) in the Andes from Venezuela to Peru. Two species groups (one consisting of P. guarnensis and P. 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.
Diagnosis. 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).
Description. 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 table 2. Summary of currently included species in Andinopanurgus with information on the known sexes, distribution, and some morphological characters. Plus (+) and dash (-) symbols indicate presence and absence of a particular character, ? = unknown. 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.
Female: Unknown. Etymology. The specific epithet is a matronym honoring Mrs. Amy Comfort de Gonzalez, loving and supporting wife of the senior author. Diagnosis. Both sexes of P. femoralis are most similar to P. 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 P. 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 P. guarnensis the apical lobes of SVII are narrow, parallel-sided, with the retrorse section of the apex commashaped (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 P. guarnensis are unmodified, not distinctly depressed.
Description. 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 ; 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.
Etymology. The specific epithet refers to the distinctly depressed meso-and metafemora of the female of the species.

Protandrena (Andinopanurgus) wayruronga Gonzalez & Ruz http://species-id.net/wiki/Protandrena_(Andinopanurgus)_wayruronga
Protandrena wayruronga Gonzalez and Ruz, 2007: 400 [♂]. Rhophitulus wayruronga (Gonzalez and Ruz) Comments. 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). SVII with apical lobes not parallel-sided, much broader apically (apex about twice as broad as base), retrorse section of apex not comma-shaped (Fig. 17); gonostylus more robust in profile, about same width across its length, basally not protuberant on medial margin in dorsal view (Peru) ... P. femoralis sp. n.