Ancylini Michener, 1944: 273. Type genus: Ancyla Lepeletier de Saint Fargeau, 1841.

Ancylaini Michener, nomen emendatum; Engel et al., 2008: 199; ICZN, 2011: 116.

Diagnosis: Bees of small to moderate size (5–10 mm in length); head about as wide as long to wider than long and face comparatively narrow, interocular distance less than compound eye length; vertex comparatively straight, not uniformly convex nor excavated between compound eyes and ocelli in facial view. Lower face often without maculation (see sexes below); clypeus flat to weakly protuberant in profile, scarcely bent back at side of labrum. Labrum broader than long. Malar space linear, nearly lacking with mandibular base abutting lower compound eye margin. Mandible simple, without distinct subapical tooth; articulations equidistant from compound eye, posterior articulation behind mediolongitudinal axis of compound eye. Postpalpal portion of galea broad proximally, then attenuate and with weakly sclerotized strip extending to apex; maxillary palpus with six palpomeres. Glossa and labial palpus much shorter than prementum; submental spine present; paraglossa short; labial palpus with four palpomeres, first two palpomeres similar in shape to apical two palpomeres, not flattened; submental spine absent. Pronotum without transverse carina. Propodeum basally declivitous, basal area impunctate, not covered by setae. Protibial calcar of strigilis with anterior carina or thin lamella on rachis bordering primary velum but lacking anterior, secondary velum; primary velum rectangular with inner margin rounded, with inner apical corner projected as acutely pointed process; malus short, often shorter than velum, with apex straight, inner margin minutely ciliate. Mesotibial spur long, serrate, apex slightly incurved; mesobasitarsal comb absent. Metabasitibial plate short, wider than long, apically weakly and broadly rounded; metatibial spurs long, serrate, apices straight or slightly incurved. Pretarsal arolium present. Forewing with pterostigma small, longer than wide, scarcely wider than prestigma; pterostigmal margin inside marginal cell convex, sometimes weakly so; marginal cell apex not truncate, bent away from anterior wing margin; three submarginal cells present; membrane with setae throughout. Hind wing with 2M+Cu about one-half as long as M, or slightly more; cu-a transverse, one-half length or more of M; jugal lobe short, slightly less than one-half length of vannal lobe. Metasomal tergum I without carina at angle of anterior- and dorsal-facing surfaces, at most sharply angulate in some males.

Female: Face without maculation; probasitarsus without anterior or posterior combs; metatibial and metabasitarsal scopa large, dense, composed of long, plumose setae, without oil-collecting structures; metabasitibial plate surface bare and smooth; pretarsal claws with small basal tooth; metasomal tergum VI with narrow pygidial plate; gradulus of metasomal sternum II simple; sterna II–V with scattered long, subdecumbent setae apically, but not forming scopa.

Male: Clypeus, supraclypeal area, paraocular area, and antennal scape with maculation; apical margin of clypeus with linear patch of distinctive setae; ventral surface of mandible and usually postgena near mandibular base with characteristic patch of dense setae; flagellum crenulate; mesobasitarsus, metatibia, metabasitarsus, and metatibial spurs frequently greatly modified (e.g., mesobasitarsus swollen with dense setae; metatibia swollen, with modified setae; metabasitarsus thickened and arched, with concave inner surface and modified; metatibial spurs elongate and broadly curved); metabasitibial plate covered with appressed setae; pretarsal claws deeply cleft; tergum VII setose, with pygidial plate present; sternum V with subapical, paramedial setal patches and apical, paramedial, thumblike processes bordering medial concavity; sternum VI with medial tubercle extending between processes of sternum V, and variously with broadly concave areas laterally and medially, with mediolongitudinal ridge or carina and medioapical margin various modified; sternum VII with lateral and apical lobes; sternum VIII with apical lobes; gonostylus without setigerous parapenial lobe; spatha present.

Comments: The presently included species, all in the genus Ancyla (= Plistotrichia Morawitz, 1874), and their distributions are summarized in table 1. The nesting biology and immature stages have been reported for two species of Ancyla (Straka and Rozen, 2012), and the species are generally considered specialists of Apiaceae and has mouthparts similar to short-tongued bees (Silveira, 1993b; Straka and Rozen, 2012).

Tarsaliini Engel, 2015: 4. Type genus: Tarsalia Morawitz, 1895.

Diagnosis (modified and expanded from Engel, 2015): Bees of small to moderate size (5.5–13 mm in length); head wider than long, but face comparatively narrow, interocular distance less than compound eye length; vertex comparatively straight, not uniformly convex nor excavated between compound eyes and ocelli in facial view (fig. 1D). Lower face often with maculation (see sexes below); clypeus weakly to moderately protuberant in profile, strongly bent back at side of labrum. Labrum broader than long. Malar space linear, nearly lacking with mandibular base abutting lower compound eye margin. Mandible simple, without distinct subapical tooth; articulations equidistant from compound eye, posterior articulation behind mediolongitudinal axis of compound eye. Postpalpal portion of galea broad proximally, then attenuate and with weakly sclerotized strip extending to apex; maxillary palpus with six palpomeres. Glossa and labial palpus longer than prementum; submental spine present; paraglossa short, not exceeding first labial palpomere; labial palpus with four palpomeres, first two palpomeres long, sheathlike (contrasting with condition in Ancylaini where first two palpomeres are similar in shape to apical two palpomeres); submental spine present. Pronotum without transverse carina. Propodeum basally with subhorizontal area, subhorizontal area punctate and setose. Protibial calcar of strigilis with anterior carina or thin lamella on rachis bordering primary velum but lacking anterior, secondary velum; primary velum rectangular with inner margin blunt, straight, with inner apical corner projected as acutely pointed process (as in most Eucerini, Ancylaini, Exomalopsini, and Emphorini: Schönitzer, 1986); malus elongate, as long as or longer than velum, with apex straight or slightly incurved, inner margin minutely ciliate. Mesotibial spur long, serrate, apex slightly incurved; mesobasitarsal comb absent. Metabasitibial plate short, wider than long, apically weakly and broadly rounded; metatibial spurs long, serrate, apices slightly incurved. Pretarsal arolium present. Forewing with pterostigma small (fig. 1A), as long as wide, scarcely wider than prestigma; pterostigmal margin inside marginal cell weakly or not convex; marginal cell apex not truncate, bent away from anterior wing margin; three submarginal cells present; membrane with setae throughout. Hind wing with 2M+Cu one-half as long as M or less; cu-a transverse, more than one-half length of M; jugal lobe short, slightly less than one-half length of vannal lobe. Metasomal tergum I with or without carina at angle of anterior- and dorsalfacing surfaces (absent in two species).

Female: Labrum, clypeus, and supraclypeal area usually with maculation (fig. 1D) (absent in one species); paraocular area usually without maculation (present in one species); antennal scape usually without maculation (present in one species); probasitarsus without anterior or posterior combs; metatibial and metabasitarsal scopa large, dense, composed of long, plumose setae, without oil-collecting structures; metabasitibial plate surface bare and smooth; pretarsal claws with minute basal tooth; metasomal tergum VI with narrow pygidial plate; gradulus of metasomal sternum II weakly birecurved (similar to condition in some Thygater Holmberg); sterna II–V with scopa composed of long, dense setae, such setae often apically wavy or sinuous (similar to type IV of Pasteels and Pasteels, 1974).

Male: Labrum with maculation (reduced to a medioapical spot in one species); clypeus and supraclypeal area usually with maculation (absent in one species); paraocular area with maculation; antennal scape usually without maculation (absent in one species), flagellum not crenulate nor elongate; metabasitibial plate often shorter than in female, covered with appressed setae; pretarsal claws deeply cleft; tergum VII setose, with true pygidial plate absent (no raised pygidial plate, instead surface continuous with disc and uniformly covered with setae), instead with distinct lateral carina and apicomedially produced and truncate to weakly bilobate (broadly and shallowly emarginate), medial projected area rounded and deflexed in two species; apical margin sternum V straight to medially lobate; sternum VI narrowed apically, apical margin usually characteristic (truncate, rounded, or medially emarginate), with lateral marginal areas broadly concave and typically smooth, producing narrow medial area usually bearing a medial tubercle or carina; sternum VII with lateral and apical lobes; sternum VIII with apical lobes; gonostylus with parapenial lobe bearing thickened setae; spatha present (absent in one species); hidden sterna and genitalia asymmetrical (sternum VII usually strongly asymmetrical; sternum VIII less dramatically so; sometimes with penis valves strongly asymmetrical).

Comments: Recently, the tribe was proposed as “new” again in Plant and Paulus (2016), despite having already been made available a year prior to this. The name as employed by these authors has no nomenclatural standing.

Baker (1998) speculated that the characteristic sternal setae functioned as a scopa. However, conclusive evidence demonstrating the use of the sternal setae for pollen transport is lacking, and both Michener (2007) and Plant and Paulus (2016) indicated that specimens with copious pollen in the hind-leg scopae lacked pollen on the sterna. However, we've followed Baker (1998) in considering the sternal setae as a metasomal scopa as the female of the new species described below has appreciable pollen in only two places on the body: the hind-leg scopae and the sternal setae. It is hoped that biological studies will someday be possible to permit an evaluation of the true functional significance of the dense sternal setae of Tarsalia.

The chiral form of the male terminalia is unique among bees. The asymmetry of the male sternum VII, and to a lesser degree sternum VIII, along with the sometimes hypertrophy of one penis valve, is immediately distinctive and peculiar. The purpose for this consistent chirality is unclear and, as noted by Baker (1998), does not correspond to any asymmetry in the female terminal sclerites. Until such time as the mating behavior of these bees is discovered, the function of these asymmetries shall remain a mystery, but in the meantime they represent a strong synapomorphy for the tribe.

We have considered the males of Tarsalia to lack a true pygidial plate. It is definitive that a pygidial plate as traditionally defined (Michener, 1944) is lacking in males of Tarsalia. However, the seventh tergum is medioapically produced with the tergal apical margin carinate, suggestive of a plate, and leading some to speculate that the majority of the tergal apical dorsum is equivalent to a pygidial plate (Michener, 2007). However, this apical surface is continuous on all sides with the remainder of the tergal disc, as well as identically sculptured and uniformly covered with setae, in stark contrast to the distinctly raised surface of a pygidial plate which also almost invariably differently sculptured and either lacking in setae or with such setae distinctly differing from those of the surrounding disc. Thus, from a strictly morphological standpoint we do not consider a true pygidial plate to be present in Tarsalia, and the form of the seventh tergum in these bees should not be confused with those lineages in which such a plate is present (e.g., Ancylaini). Even if the apical protrusion of the seventh tergum of male Tarsalia should be shown later to be developmentally homologous to the pygidial plate, it still remains a fundamentally different structure of uncertain function and such a character state should not be equated or confused with the pygidial plate in other bees. Referring to the present character state as a “pygidial plate” obscures the uniqueness of the condition found among Tarsalia, as well as misleads others, we believe, as to its morphological identity (i.e., that a true pygidial plate is lacking).

Tarsalia Morawitz, 1895: 9. Type species: Tarsalia hirtipes Morawitz, 1895, by monotypy. Friese, 1896: 211; Baker, 1998: 837; Michener, 2000: 666; Michener, 2007: 686.

Ancyla (Tarsalia) Morawitz; Warncke, 1977: 58; Warncke, 1979: 192.

Diagnosis: As for tribe (above).

Comments: The genus currently includes two rather distinct groups, one with three species in northeastern Africa, Arabia, and Iran, while the other has five species in Western and Central Asia, south into India, and west on the Mediterranean islands of Cyprus and Sardinia (table 1).

Type species: Tarsalia kindahensis Engel, new species.

Diagnosis: Body length approximately 5.5–9.0 mm; integument pale castaneous to testaceous on metasoma and majority of mesosoma (figs. 3–5, 6). Female metatibial scopa densely closed (fig. 2B); outer surface of metatibia apical to metabasitibial plate without patch of distinctive setae, area similar to remainder of scopal setae. Metasomal tergum I with or without carina at angle of anterior- and dorsal-facing surfaces (all species of Tarsalia s. str. with carina, albeit sometimes weakly so).

Etymology: The new subgeneric name is a combination of the Greek astibes (meaning, “untrodden,” hence “desert”) and melissa (meaning, “bee”). The gender of the name is considered feminine.

Included species: The subgenus presently includes three species: T. persica (Warncke) from forest steppes and semidesert regions of Iran to either side of the Zagros Mountains (Warncke, 1979; Baker, 1998); T. mimetes (Cockerell) from Upper Egypt and the Sudan (Cockerell, 1933; Baker, 1998); and T. kindahensis, n. sp., from the Najd Plateau of Saudi Arabia (east of the Hejaz and immediately west of the ad-Dahna Erg or Desert), with records from near Unayzah and separated by the Thuwayrat dunes from the others southeast near al-‘Amāriah on the Tuwayq escarpment, Riyadh.

Diagnosis: Tarsalia kindahensis can be recognized by combination of largely pale castaneous to light testaceous integument on metasoma, legs, and sometimes large portions of mesosoma although nota and much of pleura darker (figs. 4, 6); outer surface of female probasitarsus with dense, long, suberect setae with sinuate or wavy apices; metasomal tergum I without transverse dorsal carina; female metasomal scopa with long, subdecumbant setae with sinuate or wavy apices; male tergum VII with medioapical produced margin weakly bilobate (fig. 5A); male sternum VI apically truncate and with medial tubercle (fig. 5B); male hidden sterna asymmetrical and of characteristic form (fig. 5C); left penis valve hypertrophied (figs. 5E, 5G); and unique possession of yellow facial maculation on female paraocular area and scape, in addition to labrum, clypeus, and supraclypeal area (fig. 6C).

Description: Male: Total body length 6.72 mm (7.14 mm); forewing length 4.70 mm (5.18 mm). Head wider than long, length 1.69 mm (1.94 mm), width 2.28 mm (2.34 mm); inner margins of compound eyes comparatively straight, parallel in lower two-thirds, slightly diverging in upper third, upper interorbital distance 1.31 mm (1.37 mm), lower interorbital distance 1.16 mm (1.22 mm); apex of clypeus slightly below lower tangent of compound eyes; clypeus weakly protuberant and convex; ocelli at upper tangent of compound eyes; lateral ocelli separated from median ocellus by about one-half median ocellar diameter; lateral ocellus separated from eye by about 1.6× lateral ocellar diameter; lateral ocellus separated from posterior of head by about its diameter; scape short, extending at most to lower border of median ocellus; FI slightly longer than apical width, longer than FII. Intertegular distance 1.41 mm (1.53 mm). Basal area of propodeum weakly defined, about as long as metanotum. Legs without special modifications; mesobasitarsus elongate, straight, spur extending to slightly basad basitarsal midlength; metafemur slightly thicker than mesofemur; metabasitarsus elongate. Forewing with basal vein basad cu-a by 2–3 times vein width; 1m-cu entering second submarginal cell; second submarginal cell shortest, narrowed toward anterior wing margin; 2rs-m arcuate. Hind wing with nine distal hamuli. Metasomal tergum I without carina at angle of anterior- and dorsal-facing surfaces (similar to T. mimetes); tergum VII gradulus medially pointed posteriorly, without lateral gradular teeth (sensu Baker, 1998), but lateral area instead angulate, apical margin projected, deflexed medially, margin shallowly and broadly emarginate (fig. 5A); sternum V with medioapical margin produced with broad, apically blunt lobe; sternum VI narrowed apically, medioapical margin truncate, with broadly concave, smooth ovoid areas laterally along margin producing narrow medial disc bearing a medial tubercle (fig. 5B); hidden sterna VII and VIII asymmetrical and as in figures 5C and 5D; genitalia as in figures 5E–G, with penis valves markedly asymmetrical (figs. 5E, 5G).

Integument somewhat shining. Labrum with coarse, shallow, contiguous punctures, otherwise smooth. Clypeus with coarse, faint, shallow punctures separated by less than a puncture width, except slightly more spaced along borders and more oblique, approximately separated by a puncture width or sometimes slightly more, integument between punctures smooth. Supraclypeal area with smaller, more well-defined punctures, separated by a puncture width, except medially slightly more separated, integument otherwise smooth. Face below level of antennae with small, faint, scattered punctures, separated by less than a puncture width along border with clypeus and subantennal sulcus, then becoming sparser toward inner margin of compound eye such that paraocular margin impunctate, integument between punctures smooth. Face above level of antennae with small, well-defined punctures separated by less than a puncture width, integument between punctures faintly imbricate; punctures sparse in short, broad triangular patch immediate anterior to median ocellus and absent in paraocular margin where integument is strongly imbricate; ocellocular space medially as on upper face but with punctures absent along outer border of lateral ocellus and in paraocular margin bordering compound eye, former patch smooth, latter strongly imbricate; vertex as on upper face; gena as on upper face except punctures smaller and becoming gradually more widely spaced ventrally such that on lower gena punctures separated by 2–3 times a puncture width, integument between punctures smooth; postgena with minute punctures similar to those on lower gena except sparse, integument otherwise smooth. Pronotum with sparse, minute punctures, integument otherwise smooth. Mesoscutum with small, well-defined punctures separated by less than a puncture width, with some posteromedially separated by about a puncture width, integument between punctures; tegula generally smooth to faintly imbricate, with sparse, minute punctures; mesoscutellum with punctures as on mesoscutum except somewhat larger and coarser. Metanotum finely imbricate with scattered, shallow, faint punctures. Pleura with small, well-defined punctures, integument otherwise smooth except more imbricate in area of mesepisternal border with metepisternum; punctures on hypoepimeral area becoming oblique ventrally, nearly contiguous, except sparse along scrobal sulcus; punctures on preepisternal area smaller, becoming shallower, fainter, and oblique toward anterior margin; punctures on mesepisternum below scrobal sulcus becoming fainter along posterior border and on ventral-facing surface; metepisternum imbricate with sparse punctures. Basal area of propodeum largely obscured by setae, integument apparently imbricate and coarsely punctate; posterior and lateral surfaces of propodeum faintly imbricate to smooth, with scattered, small punctures. Metasomal terga with minute, well-defined punctures separated by less than a puncture width, except anterior-facing depression of tergum I largely smooth and narrow apical margins of terga smooth, integument between punctures smooth; sterna as on terga except punctures coarser and integument otherwise more imbricate.

Coloration of head generally testaceous to black (lighter in holotype male, darker in paratype male), except labrum entirely yellow; mandible largely yellow except dark reddish brown to black apically; clypeus and supraclypeal area entirely yellow; paraocular area yellow below level of antennae and broadly along compound eye in lower half of upper face; paraocular area in upper half of face above antennae dark brown to testaceous; scape yellow, pedicel and flagellomeres I and II yellowish brown; remainder of flagellum orange. Pronotum dark reddish brown to testaceous (testaceous in holotype male, dark testaceous along borders blending to dark reddish brown posteriorly in paratype male). Mesoscutum and mesoscutellum dark testaceous to black; tegula semitranslucent; metanotum testaceous to dark reddish brown; pleura testaceous to dark reddish brown; basal area of propodeum dark testaceous to black, remainder of propodeum testaceous to dark reddish brown. Legs testaceous. Wing membranes hyaline, faintly parchment colored; veins yellowish brown to dark brown. Metasoma testaceous.

Pubescence white and often obscuring integument on frons, vertex, gena, mesosoma, tibiae, tarsi, and metasomal terga where such setae usually appressed to decumbent and plumose (setae with numerous, minute branches along length of rachis, giving feathered appearance: similar to short-branched setal forms described by Saunders, 1878, and Braue, 1913), setae of mesoscutum not squamiform, typically as long as or slightly longer than ocellar diameter (setae abraded from large areas of mesosoma and anterior metasomal terga on paratype male, allowing view of integumental sculpture); labrum with transverse, medial patch of short, erect, branched setae; setae below level of antennae typically sparse, suberect to erect, long, and minutely branched, similar setae on postgena; mesosoma with setae less prominent on preepisternal area and posteroventrally on mesepisternum and anteroventrally on lateral propodeal surface; tergal setae largely short and appressed; sternal setae longer, suberect, and posteriorly directed.

Female: As described for male, except as follows: total body length 7.50 mm; forewing length 5.00 mm. Head wider than long, length 1.81 mm, width 2.44 mm; inner margins of compound eyes comparatively straight, slightly convergent below, upper interorbital distance 1.51 mm, lower interorbital distance 1.33 mm; lateral ocelli separated from median ocellus by about one-half median ocellar diameter; lateral ocellus separated from eye by about 2.0× lateral ocellar diameter; lateral ocellus separated from posterior of head by about its diameter; scape short, extending to slightly less than distance to lower border of median ocellus; FI elongate, length about twice apical width, about twice as long as FII. Intertegular distance 1.56 mm.

Coloration as in darker male (above) (fig. 6).

Pubescence as in male except for sex differences: probasitarsus with fine, erect to suberect setae with distinctly sinuate or wavy apices; metatibial and metabasitarsal scopa composed of dense, elongate, plumose setae; metasomal sterna II–V with rows of elongate, subdecumbent setae with sinuate or wavy apices, forming distinct scopa; sternum VI with medioapical patch of dense, short, erect setae.

Holotype: ♂, Saudi Arabia: Riyadh, al-Amariah [al-‘Amāriah, a locality about 12 km west of Riyadh], Majra [Mazra‘ah] al-Gasim [al-Qasim, farm], 23.v.2011 [23 May 2011], M.A. Hannan (KSMA).

Paratypes: 1♀, Saudi Arabia: Riyadh, al-Amariah [al-‘Amāriah], Majra [Mazra‘ah] al-Gasim [al-Qasim, farm], 19.vi.2011 [19 June 2011], M.A. Hannan & I. Naser (SEMC); 1♂, Saudi Arabia: Qassim [al-Qassim], Unizah [Unayzah], al-Watania Farm, Rowdah, 29.v.2013 [29 May 2013], M.A. Hannan; Pulicaria undulate [sic] [Pulicaria undulata] (SEMC).

Floral record: The paratype male was taken while visiting flowers of Pulicaria undulata (L.) C.A. Mey (Asteraceae: Asteroideae: Helianthodea: Inuleae), a rather common annual herb in the region, known as false fleabane or regionally as “Jethjath” and used in medicinal tea. Regional beekeepers consider P. undulata as a good source for pollen and nectar during the spring.

Etymology: The specific epithet refers to the ancient kingdom of Kindah, which occupied, among other areas, large portions of the Najd, encompassing the type localities.

Comments: There is variation in color of the head and mesosoma between the two males, the one from Qassim being darker and similar to the female from al-‘Amāriah, while the male from al-‘Amāriah is distinctly lighter. The lighter male is more testaceous in those areas where the Qassim male is dark testaceous to dark reddish brown, and dark testaceous to reddish brown in those places where the female and male from Qassim are dark reddish brown to black. However, despite these color differences, the integumental, structural, and genitalic traits are identical between the males. The male of T. kindahensis superficially resembles to some degree the male of the sympatric Tetraloniella (Tetraloniella) persiciformis Alqarni et al. (2012), a eucerine that has also been taken visiting flowers of P. undulata.