Belliturgula najdica Engel, sp. n.

The genus resembles Flavomeliturgula in the elongate and apically truncate glossa but differs in the following attributes: body almost entirely pale yellow, with scattered dark brown to black markings; outer subantennal sulcus arched; clypeus greatly protuberant and extending well in front of compound eyes; labrum as long as wide, with distinct basilateral ovoid windows of translucent integument; proximal labial palpomere greatly elongate and flattened, twice as long as combined length of remaining palpomeres, extending well past apex of paraglossa; second through fourth labial palpomeres cylindrical, not flattened; apex of glossa not broadened; galea longer than stipes; mesoscutum covered with abundant, erect, minutely spicate, white setae; forewing 1rs-m confluent with (rather than distad) 1m-cu; basal area of propodeum setose; probasitarsus short and broad, about twice as long as wide; mesotibial spur short, only about 0.5× length of mesobasitarsus; pygidial plate with margins concave apically and therefore more abruptly tapering to acutely rounded apex.

Small species (less than 8 mm in total length); integument largely yellow, with some dark brown to black markings (Figs 1–5); head broader than long; compound eyes with inner margins parallel (Fig. 5); outer subantennal sulcus greatly arched outward (Fig. 5) (not straight); facial fovea shallow, narrower than median ocellar diameter, elongate, length approximately 5.5× width (Fig. 5); clypeus greatly protuberant, extending in front of compound eye by almost as much as compound eye width (Fig. 8) (protuberance extending in front of compound eye by distance approximately 0.85–0.95× compound eye width); labrum as long as wide, with basilateral ovoid areas (‘windows’) of translucent integument, without basilateral patches of dense, elongate, branched setae, with shallow, mediolongitudinal depression; ventral surface of prementum without abundant, long, posteriorly directed, simple setae (such setae present in Khuzimelissa); first labial palpomere greatly elongate and flattened, much longer than remainder of palpus (Fig. 12) (2× as long as combined lengths of remaining palpomeres), extending greatly beyond apex of paraglossa, second palpomere not flattened, third through fourth palpomeres short, subequal in length, cylindrical (Fig. 12); glossa greatly elongate, distinctly longer than face (Fig. 8); apex truncate but not broadened (Fig. 9); galea longer than stipes (galea slightly longer than stipes in Flavomeliturgula; galea shorter than stipes in Khuzimelissa); mesoscutum with numerous scattered, erect, minutely spicate (pipecleaner-like in appearance), white setae, not obscuring integument (0.8–1.3× median ocellar diameter) (Fig. 4); basal area of propodeum setose, setae not obscuring integument; forewing with pterostigma longer than wide, approximately 3.5× as long as wide, width subequal to length of prestigma, about 3.5× as long as prestigma, margin bordering first submarginal cell slightly diverging from anterior wing margin, border inside marginal cell converging toward anterior wing margin and with convex arch; marginal cell widely and slightly obliquely truncate at apex, longer than distance from its apex to wing tip; three submarginal cells present (i.e., 1rs-m present); length of first submarginal cell slightly greater than combined lengths of second and third submarginal cells; r-rs and 3Rs (anterior border of second submarginal cell) subequal; 2Rs and 1rs-m slightly converging anteriorly, thus second submarginal cell trapezoidal; 3Rs slightly longer than 0.5× 4Rs (anterior border of third submarginal cell); 1rs-m confluent with 1m-cu (as in Gasparinhala); 1cu-a at least 2× as long as 2M+Cu (usually 2M+Cu even shorter in other genera); hind wing with six distal hamuli; probasitarsus broad, about twice as long as wide (probasitarsus more elongate slender in Flavomeliturgula, nearly four times as long as wide); protibial calcar with velum rectangular, velum with outer and apical borders comparatively straight, malus (apical portion of rachis: sensu Engel et al., 2017) ciliate and slightly shorter than velum; mesofemur ventro-proximally forming nearly orthogonal angle (as in Flavomeliturgula, Meliturgula, Khuzimelissa, and related genera); mesotibial spur serrate, about 0.5× length of mesobasitarsus; metabasitibial plate broadly rounded, with carinate margins and surface covered in appressed, minute, plumose setae; metatibial and metabasitarsal scopa composed of abundant, elongate (1.8–3.2× median ocellar diameter), simple setae; pretarsal claws simple; metasomal terga without well-developed setal bands (Fig. 4) (present in Khuzimelissa); pygidial plate with apex narrowly rounded, margins tapering more prominently in apical half (Fig. 24).

The generic name is an artificial combination of Bell, honoring explorer, archeologist, and nation-builder Gertrude Margaret Lowthian Bell (1868–1926), and a subset of Meliturgula Friese, type genus of the subtribe Meliturgulina. The name is therefore treated as a euphonious combination of letters and the gender of the name is considered to be feminine.

The genus includes at present only the type species, found in central Saudi Arabia.

As for the genus (vide supra).

♀: Total body length 7.08–7.33 mm; forewing length 4.13–4.27 mm. Head broader than long, width 2.08–2.13 mm, length 1.67–1.71 mm; compound eyes with inner orbits margins roughly parallel, upper interorbital distance 1.33–1.38 mm, lower interorbital distance 1.33–1.36 mm; distance between compound eyes slightly greater than compound eye length; compound eye length 1.25 mm, width 0.58–0.63 mm; clypeus greatly protuberant, extending in front of compound eye by nearly compound eye width; intertegular distance 1.42–1.46 mm.

Clypeus and supraclypeal area largely smooth with coarse, shallow (almost faint in most areas) punctures, such punctures separated by a puncture width or more, slightly denser along borders; labrum similar to clypeus except impunctate on transparent ovals; remainder of face similar except punctures smaller, more defined (albeit still shallow), and separated by a puncture width or frequently less; dark area between ocelli minutely and faintly imbricate and impunctate; facial fovea imbricate and impunctate; gena as on upper face except punctures becoming more sparsely separated ventrally; mesoscutum smooth to faintly and minutely imbricate with shallow punctures separated by a puncture width or less; mesoscutellum as on mesoscutum except punctures slightly denser and integument slightly more noticeably imbricate; metanotum as on mesoscutellum except shallow punctures nearly contiguous; pleura as on mesoscutum except punctures shallower and becoming sparse ventrally; basal area of propodeum as on metanotum; lateral and posterior surfaces of propodeum with small punctures separated by a puncture width or frequently less, integument between punctures smooth to faintly imbricate; metasomal terga minutely and finely imbricate with scattered shallow, faint, small punctures separated by a puncture width or less, such punctures most easily discernible on first tergum and becoming fainter on succeeding terga; pygidial plate minutely and finely imbricate; sterna as on terga except shallow, coarse punctures more prominent.

Integument generally pale yellow except marked with dark brown to black on mandibular apex, on facial fovea, around ocelli (Figs 4, 5); mesoscutum with three longitudinal dark stripes, medial stripe thin, terminating before posterior of mesoscutum, lateral stripes thicker, situated between lateral yellow borders and medial stripe, extending to axilla (Fig. 4); dark brown on lateral depression of axilla and along anterior border of mesoscutellum (slightly extended mediolongitudinally) (Fig. 4), and light brown anteriorly on basal area of propodeum (Fig. 4). Wing membranes clear, hyaline; veins pale yellow to off white (Fig. 3). Metasoma with light to dark brown marks as follows: paramedian spots on anterior-facing surface of first metasomal tergum (Fig. 4), elongate oval spots at lateral extremities of second tergum (Fig. 3), small spots on lateral extremities of third tergum (absent in some individuals); terga I–V with irregular brownish-purplish coloration (reflective of inner organs through pale integument) to varying degrees (in some individuals inner gut prominent through integument, others less so); pygidial plate brown, becoming reddish apically (Fig. 24); sterna pale brown, becoming darker on two apical sterna.

Figures 3, 4. 

Female of Belliturgula najdica Engel, gen. et sp. n., from central Saudi Arabia. 3 Lateral habitus 4 dorsal habitus.

Pubescence generally white and largely sparse (Figs 3–5); facial setae generally short to moderate (0.2–1.0× median ocellar diameter) in length and minutely spicate (i.e., minutely branched on all sides along entire length of rachis, with a pipecleaner-like appearance), such setae scattered on face and generally suberect; scape with similar moderate-length, erect, minutely spicate setae; minutely spicate setae longer (0.9–1.5× median ocellar diameter) and erect around ocelli, vertex, posterior of gena, and postgena; mandible with more sparse, elongate (1.8–3.0× median ocellar diameter), simple setae along lower margin; mesoscutum with numerous erect short to moderate-length (0.4–1.2× median ocellar diameter) minutely spicate setae, such setae not obscuring integument; mesoscutellum as on mesoscutum although setae slightly sparser; metanotum with minutely spicate setae sparse on disc, with various greatly elongate (up to 2.0× median ocellar diameter) minutely spicate setae at lateral extremities; pleura with numerous moderate-length to elongate (1.0–2.2× median ocellar diameter) minutely spicate setae, such setae not obscuring integument; basal area of propodeum with numerous fine minutely spicate setae, not obscuring integument; propodeum with lateral and posterior surfaces with pubescence similar to that of pleura; legs with minutely spicate setae except simple on tarsi and scopal setae of metatibia and metabasitarsus elongate (1.8–3.2× median ocellar diameter) and simple; metasoma with generally sparse pubescence, setae more numerous on apical terga where such setae minutely spicate and suberect to erect; setae particularly dense around pygidial plate and obscuring integument.

♂: Latet.

♀, KSA [Kingdom of Saudi Arabia], Riyadh [Region], Derab Agricultural Research Station [24°25'23"N, 46°39'01"E, 580 m elevation], 29.4.2015 [29 April 2015], M. Shebl, Carthamus [Asteraceae: Carthamus L.] (SEMC).

2♀♀, same data as holotype (SEMC, KSMA).

The specific epithet is taken from the Najd, the vast geographic central plateau portion of Saudi Arabia which encompasses the type locality which is slightly south of the city of Riyadh.

The type series was collected from an unidentified species of distaff thistle (Asteraceae: Carduoideae: Carthamus L.) growing around the Dirab Agricultural Research Station in central Saudi Arabia (just south from Riyadh). The most common species of Carthamus in Saudi Arabia are Carthamus oxyacantha M.Bieb., which grows wild in many disturbed areas, and C. tinctorius Linnaeus (safflower), the latter widely cultivated, particularly so in southern Arabia (Mandaville, 1990). It is likely that the type series of B. najdica was captured at C. oxyacantha. Species of Carthamus are spiny, dentate herbs that can be largely self-pollinated, although among insects bees are the dominant pollinators (e.g., Butler et al., 1966).

Meliturgula deserta Warncke, 1983 [1985].

The genus superficially resembles Flavomeliturgula and Belliturgula, particularly the latter in the extensive yellow coloration. However, the genus differs from both in the short, more typical labial palpomeres in which the proximal labial palpomere is elongate, flattened, longer than the combined lengths of the remaining palpomeres (approximately 1.25× length of remaining palpomeres combined), with the second palpomere flattened and slightly shorter than the length of the third and fourth palpomeres combined; and the third and fourth palpomeres subequal in length and cylindrical (Fig. 13). In addition, the glossa is acute (Figs 6, 15) (rather than broadly truncate apically), the glossa is shorter than the head length (Figs 6, 19–21) (rather than greater than the head length in the former two genera), the pretarsal claws are cleft (simple in the former two genera), the metasomal terga have distinctive broad setal bands (Figs 16–18) (lacking in the former two genera), and the pygidial plate is apically emarginate (Fig. 25) (acutely rounded in the former two genera). Other diagnostic traits include: small species, slightly more than 8 mm in length; integument largely pale yellow with few dark brown to black markings (Figs 6, 16–18); outer subantennal sulcus arched (Fig. 6); clypeus gently and weakly convex, scarcely protuberant (Figs 6, 17, 19), extending in front of compound eye by about 0.5× compound eye width; labrum wider than long, width 1.7× length, without lateral oval windows of transparent integument (Fig. 20), surface gently and weakly convex with weak subtriangular proximal medial depression; prementum ventrally with abundant, elongate (1.0–1.9× median ocellar diameter), erect to suberect, simple setae (Figs 15, 19, 21); galea shorter than stipes; mesoscutum with abundant, appressed, squamose setae (Fig. 18), intermixed with sparse, erect, minutely spicate setae (0.8–1.0× median ocellar diameter); probasitarsus elongate, slender, length approximately 3× width; mesotibial spur about as long as mesobasitarsus; forewing with r-rs distinctly shorter than 3Rs, 1rs-m distad 1m-cu, and 1cu-a about 4× as long as 2M+Cu; hind wing with 10 or more distal hamuli arranged in a dense series.

Figures 5–7. 

Female faces of representative Meliturgulina. 5 Belliturgula najdica Engel, gen. et sp. n., from Saudi Arabia (note arched outer subantennal sulci) 6 Khuzimelissa deserta (Warncke), comb. n., from Pakistan 7 Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov) from Iran (note straight outer subantennal sulci).

The new genus-group name is a combination of Khuzi, the Middle Persian name for the ancient Susian people, and from which the regional name Khuzestan derives (and encompasses the type locality of the type species), and the Greek, mélissa (μέλισσα), meaning, “bee”. The gender of the name is feminine.

The genus presently includes only the type species, Khuzimelissa deserta (Warncke), comb. n., which has been recorded from Iran (Khuzestan) and Pakistan (Balochistan).

Figures 8–11. 

Female heads and mouthparts in profile and details of glossal apexes. 8 Head profile of Belliturgula najdica Engel, gen. et sp. n., from Saudi Arabia 9 glossal apex of B. najdica 10 head profile of Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov) from Iran 11 glossal apex of F. (F.) lacrymosa.

The genus can be generally differentiated from other members of the subtribe by the following combination of features: small to medium-sized bees, 5.0–10.0 mm in length; integument dark brown to brown with extensive yellow markings (Figs 22, 23); outer subantennal sulcus straight (Fig. 7); clypeus weakly protuberant, extending in front of compound eye by about 0.35–0.5× compound eye width (Fig. 10); labrum wider than long, width 1.5× length, without lateral oval windows of transparent integument, surface with weakly raised, medial plateau, basilaterally with prominent setose patches composed of long (1.8–2.4× median ocellar diameter), sub-decumbent setae; prementum ventrally without elongate setae; proximal labial palpomere elongate (but not extending past apex of paraglossa: in Belliturgula palpomere extends well past apex of paraglossa), flattened, about as long as to slightly longer than combined lengths of remaining palpomeres; second and third palpomeres flattened and elongate, together slightly shorter than to about as long as first palpomere, fourth palpomere short, shorter than preceding palpomere, cylindrical combined (Fig. 14); glossa greatly elongate, longer than head length, apex broadly truncate and broadened (Figs 7, 10, 11); mesoscutum without appressed, squamose setae or abundant spicate setae (Fig. 22); forewing with 1rs-m distad 1m -cu (confluent in Belliturgula); probasitarsus elongate, slender, length approximately 4× width; mesotibial spur elongate, nearly 0.75× as long as mesobasitarsus; pretarsal claws simple; metasomal terga without distinctive broad setal bands; pygidial plate apically acutely rounded, with lateral margins tapering evenly toward apex, surface with mediolongitudinal ridge (Fig. 26).

Figures 12–15. 

Labial palpi of representative species (to same scale) and labiomaxillary complex of Khuzimelissa deserta (Warncke), comb. n. (not to same scale), all in ventral view. 12 Labial palpus of Belliturgula najdica Engel, gen. et sp. n. 13 labial palpus of K. deserta 14 labial palpus of Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov) 15 photograph of labiomaxillary complex of K. deserta.

The subgenus can be distinguished by the combination of: mesoscutum dark brown to black with yellow markings, surface with scattered, appressed, fine setae, such setae mostly anteriorly and laterally, and 0.5–0.75× median ocellar diameter; basal area of propodeum without setae; and hind wing with 10 or more distal hamuli arranged in a dense series along anterior wing margin.

The subgenus as here circumscribed is restricted to the type species, Flavomeliturgula lacrymosa (Popov), from Iran (Fars, Hormozgān, Khuzestan, Tehran).

Figures 16–18. 

Female of Khuzimelissa deserta (Warncke), comb. n., from Pakistan. 16 Dorsal habitus 17 lateral habitus 18 detail of mesosoma and anterior metasomal dorsum.

Meliturgula tapana Warncke, 1983 [1985].

The subgenus can be distinguished by the combination of: mesoscutum dark brown to black with yellow markings, surface with scattered, minute, fine setae, and less than 0.5× median ocellar diameter; basal area of propodeum with setae; and hind wing with 6–7 distal hamuli along anterior wing margin.

The new subgeneric name is a combination of Kóréš or Kūruš, the Hebrew and Persian names for Cyrus the Great (Cyrus II, ca. 600–530 B.C.), who first unified the ancient kingdoms of the Near East and founded the Achaemenid Empire of Persia, combined with the Greek, mélissa (μέλισσα), meaning, “bee”. The gender of the name is feminine.

The subgenus includes two species: F. (Koreshomelissa) tapana and F. (K.) centaurea (Warncke), both from northern (along the Caspian Sea) and southern costal Iran (Māzandarān and Hormozgān) and southern Pakistan (Balochistan).

Figures 19–21. 

Female of Khuzimelissa deserta (Warncke), comb. n., from Pakistan. 19 Head and mouthparts in profile 20 dorsal-frontal view of mouthparts 21 extended labiomaxillary complex in profile.

Flavomeliturgula berangeriae Patiny, 2002.

The subgenus can be distinguished by the combination of: mesoscutum entirely black without yellow maculation, surface with scattered setae of 1–1.5× median ocellar diameter; basal area of propodeum with setae; and hind wing with 6–7 distal hamuli along anterior wing margin.

The new subgeneric name is a combination of Freya, for the famed explorer of Arabia, Persia, and Afghanistan, Freya Madeline Stark (1893–1993), and the Greek, mélissa (μέλισσα), meaning, “bee”. The gender of the name is feminine.

The subgenus includes two species: F. (Freyamelissa) berangeriae and F. (F.) schwarziana Patiny, from southwestern Iran (Bushehr) and southern Pakistan (Balochistan).

Figures 22, 23. 

Female of Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov) from Iran. 22 Dorsal habitus 23 lateral habitus.

Figures 24–26. 

Female pygidial plates. 24 Belliturgula najdica Engel, gen. et sp. n. 25 Khuzimelissa deserta (Warncke), comb. n. 26 Flavomeliturgula (Flavomeliturgula) lacrymosa (Popov).