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Apolysis elbae (Efflatoun, 1945) comb. nov. Usia elbae Efflatoun, 1945: 210. Type material Lectotype (here designated) EGYPT • ♂ (pinned through circular card, in good condition); “ Gebel Elba, South Eastern Desert, 15,3 to end April 1929 /Zool. Dep. Collection, Egyptian University, Collection Tewfik/ Usia elbae Effl., Det. Efflatoun ”; TAU. Paralectotype EGYPT • 1 ♀ (pinned through circular card, in good condition, thorax partially greasy); “ Gebel Elba, South Eastern Desert, 15,3 to end April 1929 /Zool. Dep. Collection, Egyptian University, Collection Tewfik/ Usia elbae Effl., Det. Efflatoun ”; TAU. Remarks Efflatoun (1945) based his description of this species on more than 40 males and 50 females, mostly from Gebel Elba, southwestern Egypt, and two males from Wadi Feran [sic = Feiran], South Sinai. While it has not been possible to visit the collections in Egypt, where many of the syntypes still survive, two syntypes in TAU were loaned to me and are here designated as lectotype and paralectotype. A very small species, readily diagnosed as Apolysis from the distal arista in the apico-dorsal sulcus of the postpedicel, sparsely short-haired disc of mesonotum, elongate discal cell, A1 running dead straight to wing margin, A1+CuA 2 (petiole of anal cell) long, slender short-haired palps and form of the form and orientation of the male hypopygium. Neither the lectotype or paralectotype has been dissected as this would not increase confidence of generic placement in Apolysis as currently understood. Distribution Egypt (Gebel Elba, Wadi Edeib and Wadi Kansissrob, Southwestern Desert, and Wadi Feran [sic = Feiran], South Sinai)., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 149-150, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Efflatoun H. C. 1945. A monograph of Egyptian Diptera. Part VI. Family Bombyliidae. Section 1: Subfamily Bombyliidae Homoeophthalmae. Bulletin de la Societe Fouad Ier d'Entomologie 29: 1 - 483."]}

Parusia almeria gen. et sp. nov. urn:lsid:zoobank.org:act: 22C73CC7-88D1-4CE0-821A-1B3C795625DD Figs 16, 20, 32, 60 Etymology Noun in apposition, after Almeria, the region of southeast Spain where the type material was collected. Type material Holotype SPAIN • ♂; “ Almeria, Bird observatory of the Salinas del Cabo de Gata, N 36º45′39″ W 02º13′15″, dunes near salines, 9 April 2004 leg. Antonio Aguirre ”; NHMUK. Paratypes SPAIN • 1 ♀; “ Almeria, Bird observatory of the Salinas del Cabo de Gata, N 36º45′39″ W 02º13′15″, dunes near salines, 9April 2004, leg. Antonio Aguirre ”; NHMUK • 1 ♂ (in spirit); “same data”; PCMC-T • 1 ♂; “ Carboneros N36°59′39.7″ W01°56′29.6″ 100 m 19 March 2003, leg. J. Dils & J. Faes ”; PCJD • 1 ♂, 1 ♀ (in spirit); “near Torregarcía, N 36º49′58″ W 02º17′29″ zone with Ziziphus lotus and dunes, 13 April 2004 Moericke trap 1, leg. Antonio Aguirre ”; PCMC-T • 1 ♂, 1 ♀; “ Moericke Trap 3 Base del Cerro del Barronal, N 36º44′01 ′ W 02º08′29″, very sandy zone with Pitas, Rhamnus sp., and Ammophila areanaria, 12 March 2004 pitfall trap 5, leg. Antonio Aguirre ”; PCDG. Description MEASUREMENTS. Body length: 4.4–6.6 mm. Wing length: 3.7–5.8 mm. Male HEAD. Gena plus mouth margin black in ground colour, much narrower than the apical breadth of a palp, grey dusted gena very narrow, linear, more shining mouth margin a little wider. Frons black in ground colour, hind half narrowing distinctly from hind corners to just in front of anterior ocellus where eyes start to diverge before abruptly widening on anterior part before running parallel down mouth margins. Eyes separated at their narrowest by 1.5–2.5 times the diameter of the front ocellus, hind ocellus narrowly separated from the eye margin. Ocellar tubercle shining, barely dusted, narrow part of frons lightly grey dusted, subshining blackish, white dusted on front half, more thinly so medially where dark ground colour shows from some angles. Yellow-brown hairs on ocellar tubercle almost twice as long as the width of the frons at rear, narrow part of frons bare, anterior part of frons with numerous white hairs some of which are as long as scape and pedicel combined, inclined at 45°. Occiput dark in ground colour with fairly dense grey dust but ground colour still apparent, subshining, more shining on triangular area behind ocellar tubercle. Yellow to white hairs dorsally only just overtopping ocellar tubercle, tips curved anteriorly, hairs significantly longer below and rather wavy. Ommatidia uniform in size across the eyes. Antennae black, postpedicel variable but usually relatively short, less than twice length of scape and pedicel together, sensilla in subapical sulcus can be white and conspicuous. All antennal segments with short pale brown hairs above, longest just before subapical sulcus. Palps short but easily visible, strongly clavate, brown to black, the pale yellow apical setae as long as the length of the palps. Proboscis short, only about twice head length, dorsally clothed with short, adpressed brown setae basolaterally. Basoventral membrane dirty brownish-white. THORAX. Shining black cuticle dulled by relatively thin blue-grey dust except sometimes along paramedian and antehumeral vittae where the shining black cuticle can show through to a greater or lesser extent. Where paramedian vittae are clear they are approximately equal to median dusted stripe anteriorly, widening rearward where tending to fade. Antehumeral vittae interrupted at the thoracic suture. Mesonotum covered with moderately long, pale yellow hair, the longest hairs longer than those on the ocellar tubercle. Scutellum more thinly dusted centrally, long-haired on the disc, hairs fringing the apical margin longer than those on mesonotum. Pleura similar to mesonotum laterally, or a little more densely dusted, white hairs on pronotum, posterior half of the anepisternum and much of katepisternum. WING. Membrane faintly brown tinged, the veins brown. Crossvein r-m beyond basal third but before mid-point of discal cell conspicuously beyond m-cu. Anal lobe moderately developed with evenly convex margin, about as broad as the anal cell. HALTERE. Pale yellow to white, base of stem slightly brownish. LEGS. Coxae black with a coating of grey dust like that on the pleura, not entirely obscuring ground colour. Remainder of legs black or very dark brown, essentially undusted so the shining cuticle is not dulled. Coxae externally and femora ventrally covered with rather long, pale yellow hairs, on the femora longer than the greatest depth of the femora. Femora dorsally and tibia rather densely covered with both short adpressed white hairs, and on tibia a few longer, semi-erect hairs, especially at base. ABDOMEN. All tergites subshining black, dusting on disc obscure and dark, laterally, on the reflexed margin, densely grey dusted mostly obscuring the ground colour, most extensive on tergite one. Middle tergites with narrow but conspicuous and sharply demarcated yellow apical margins; on tergite one this margin disappears laterally, on remaining tergites it tapers away on reflexed margin, tergites seven and eight with yellow apices very narrow or lacking. Sternites black cuticle with grey dusting dulling but not entirely obscuring the ground colour, apical yellow margins obscure. Tergites and sternites all covered with relatively short pale yellow hairs, mostly erect but on basal tergites tending to be inclined towards mid-line. GENITALIA. Conspicuously large and globular. Gonocoxites black to dark brown, shining, composed of two rounded hemispheres separated by a deep sulcus. Epandrium shining black to dark brown. Both gonocoxite and epandrium covered in long yellow hairs, denser than on tergites, especially so on gonocoxite. Female Very similar to male except for broader frons, narrowest at rear where one fifth to one quarter head width, hind ocellus separated from eye by about one and a half times the diameter of that ocellus. Frons widening more evenly towards the front, hairs more extensive, biserial along eye margins to rear of front ocellus. Thorax very similar, hairing tending to be rather shorter. Yellow apical margins to the tergites relatively broader. Abdominal hairing a little sparser and shorter. Proximal part of genital fork more or less in line with angle of the arms, attached membranously, broadest towards pointed tip, basal spermathecal ducts normal, fusing at or just before the vaginal plate which is contiguous with the genital fork and not differentiated. Remarks Within this genus of eight species there is considerable uniformity in external characters, differences between the species being subtle and not likely to be much use in deducing affinities within the genus. The female genitalia suggest that this species is most closely allied to the other two Parusia gen. nov. that occur in Spain Pru. loewi and Pru. propinqua gen. et sp. nov. Of the African-Tyrrhenian species probably related to Pru. aurata, in which the male and female genitalia are closest to the Iberian species. Distribution Southeast Spain., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 123-125, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958

Apolysis sedophila (Brunetti, 1909) Usia sedophila Brunetti, 1909: 227. Type material Syntypes INDIA • 3 ♀♀; “ Syntype / Usia sedophila Brunetti, Cotype / Syntype Usia sedophila Brunetti. det. J.E. Chainey 1986/[leg.] Annandale, Simla, Alt. 7000ft., 16 May [19]09/On flowers of Sedum rosulatum /Recd. in exch., from, Indian Museum., Calcutta., 1910-14/ Usia sedophila Brun ♂ ♀ ”; NHMUK. Other material examined INDIA • 7 ♀♀; “ On Sedum flowers/ India, Phagu to Kufri, Simla Hills, 8-9000ft., 21 May 1916 /[leg.] Annandale & Kemp/Standing over sedophila Brunetti, in J. Bowden Coll., BMNH(E) 2003-159”; NHMUK. Remarks Although originally described in Usia this species was moved to Apolysis by Evenhuis & Greathead (1999) and this is confirmed here with some notes on the syntypes. Despite the suggestion on one label, all specimens in the NHMUK, both syntypes and non-type material, are females. While all the seven non-type specimens are conspecific, and are the same species as two of the syntypes, the third (with the cotype label) seems to be a different taxon. All 10 specimens are Apolysis as this genus is currently defined so it is reasonable to assume that any other syntypes in Indian museums will also be Apolysis. Thus, although the specific identity of A. sedophila is currently uncertain, that it belongs in Apolysis is certain. Distribution India (Himachal Pradesh), Pakistan., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on page 154, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Brunetti E. 1909. New Indian Leptidae and Bombyliidae with a note on Comastes osten Sacken, v. Heterostylum Macquart. Records of the Indian Museum 3: 211 - 230.","Evenhuis N. L. & Greathead D. J. 1999. World Catalog of Bee Flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden."]}

Parusia loewi (Becker, 1906) gen. et comb. nov. Figs 17, 31, 65 Usia loewi Becker, 1906: 219. Etymology Named after Friedrich Hermann Loew (1807–1879). Type material Lectotype (here designated) SPAIN • ♂; “ Andalusia, Rosenhauer/ Coll. H. Loew /Zool. Mus. Berlin/95719 Typus ”; ZMHB. Paralectotype SPAIN • 1 ♀ [on same pin. A second label, not on the pin when examined but associated with these specimens]; “ grata Lw. ♀ det Becker”; ZMHB. Other material examined SPAIN • 2 ♂♂, 4 ♀♀; “ Malaga Province, Marbella 29, 30 April 1983 ”; NHMUK • 6 ♂♂, 10 ♀♀; “ 2–6 May 1983 (leg. J. Bowden)[dark form]”; NHMUK • 1 ♂, 1 ♀; “ Salamanca, Villar de Ciervo, Las Coronas 23–25 May 1999 yellow pan”; PCDG • 1 ♀; “ 6 km N. of Villar de Ciervo 26.v–2.vi.1999 1 ♀ (leg. H.- P. Tschorsnig)”; PCDG • 1 ♀; “ Andalusia, Veer de la Frontera 29 March 2000 1 ♀ (leg. D.J. Greathead)”; NHMUK • 1 ♂; “ Malaga (Espagne), Don José Maria, de la Fuentes”; MNHN • 2 ♀♀; “ Museum Paris, Espagne, coll. Dufouri / Usia aurata ”; MNHN • 1 ♂; “ Madrid, Maniola [?], G. Sharma / Usia / Usia aurata Em’s/Museum Paris, Coll. J. Surcouf 1919”; MNHN • 5 ♂♂, 2 ♀♀; “ Andalusia, near Padul, 783 m, N37°01′40″ W3°37′3″ 2 May 2008, Leg. D.J. Gibbs ”; PCDG • ♂ ♀ in cop.; “ 780 m N37°01′42″ W3°37′5″ 2 May 2008, Leg. D.J. Gibbs”; PCDG • 2 ♀♀; “ 840 m N37°02′47″ W3°38′43″ 1 May 2008, Leg. D.J. Gibbs”; PCDG. Redescription MEASUREMENTS. Body length: 2.4–5.0 mm. Wing length: 2.5–5.1 mm. Male HEAD. Gena and mouth margin brownish black to black in ground colour, narrower than the apical breadth of a palp, grey dusted gena narrow, linear, more shining mouth margin broader, sometimes yellowish. Eyes separated at their narrowest by twice the diameter of the front ocellus, hind ocellus separated from the eye margin by less than half the diameter of that ocellus. Frons brownish black to black in ground colour, ocellar tubercle shining, barely dusted, narrow part of frons matt blackish, noticeably converging from hind corners to well in front of anterior ocellus, then widening relatively abruptly onto anterior of frons. Front part of frons densely blue-grey dusted, rather more thinly so centrally. White hairs on ocellar tubercle conspicuously longer than the width of the frons at rear, narrow part of frons bare, anterior part of frons with numerous white hairs which are no more than half the length of those on the ocellar tubercle. Occiput dark in ground colour densely covered with grey or brownish dust, triangular area behind ocelli more thinly dusted, subshining. The whole covered with erect pale yellow to white hairs, short above just overtopping ocellar tubercle, much longer below, silky, slightly wavy but not very dense. Ommatidia uniform in size across the eyes. Antennae blackish, tip of pedicel somewhat paler in lectotype, postpedicel variable but usually approximately twice length of scape and pedicel together, sensilla in subapical sulcus sometimes whitish so conspicuous. All antennal segments with short pale brown hairs above, longest just before subapical sulcus. Palps short but easily visible, strongly clavate, brown to black, the pale yellow apical setae as long as the length of the palps. Proboscis not very long, approximately one and a half head lengths, dorso-laterally clothed with short, adpressed brown setae. Basoventral membrane brown. THORAX. Blackish ground colour largely obscured by blue-grey dust except along the conspicuous paramedian and antehumeral vittae where the shining black cuticle shows through, dulled by a variably very thin coating of brownish dust, subshining or even shining. Paramedian vittae variable a little narrower to a little broader then acrostichal dusted stripe, extending from very front of mesonotum to three quarters the way back. Antehumeral vittae divided at thoracic suture, sometimes a vague darker spot above wing base, area in front of scutellum usually darker, more thinly dusted. Mesonotum covered with moderately long, pale yellow hair, the longest hairs as long as those on the ocellar tubercle. Scutellum relatively sparsely haired on the disc, long hairs fringing the apical margin. Pleura dusted as notopleuron, with similar hairs on pronotum, posterior half of the anepisternum and a few scattered across katepisternum. Metepimeron partly yellow in lectotype, entirely dark in other specimens, glabrous. WING. Membrane faintly brown tinged, especially in first basal cell, the veins brown. Crossvein r-m a little beyond basal third of discal cell, conspicuously beyond m-cu. Anal lobe moderately developed with evenly convex margin, about as broad as the anal cell. HALTERE. Pale yellow to white, base of stem slightly brownish. LEGS. Coxae dark with a coating of grey dust like that on the pleura. Remainder of legs black or very dark brown, only thinly dusted so not obscuring the shining cuticle. Coxae externally and femora ventrally covered with rather long, white hairs, on the femora longer than the greatest depth of the femora. Femora dorsally and tibia rather densely covered with short, white hairs. ABDOMEN. All tergites black appearing undusted but rendered matt by thin dark brown dusting, dully shining on disc, tergite seven undusted, shining. Laterally on the reflexed margin densely grey dusted obscuring the ground colour, most extensive on tergite one where it extends to the scutellum. All tergites with narrow to quite broad, conspicuous and sharply demarcated yellow apical margins, narrowing laterally but even clear on tergite one here. Sternites black and densely grey dusted obscuring the ground colour, apical yellow margins less well defined. Tergites and sternites all covered with fairly long, erect white to very pale yellow hairs, longest on sternites. GENITALIA. Conspicuously large and globular compared to most Protypusia gen. nov., but typical for Parusia gen. nov. Gonocoxites black to dark brown, shining, composed of two rounded hemispheres separated by a deep sulcus. Gonostylus sharply curved, tip at right angles to base and twisted, often visible in dried specimens. Epandrium shining black to dark brown, apico-lateral corners rounded. Both gonocoxite and epandrium covered in long whitish to pale yellow hairs, longest and densest on gonocoxite. Female Differs from the male in its more broadly separated eyes, three to three and a half times the diameter of the front ocellus, about one-fifth head width at narrowest point, hind ocelli separated from the eye margins by approximately twice the diameter of these ocelli. A few hairs on the middle part of the frons laterally. Hairs on legs and abdomen shorter, conspicuously so on the abdomen. Apical yellow margins to tergites rather broader, tergite eight not noticeably more shining than preceding tergites. Proximal part of genital fork strongly curved, sometimes at more than a right-angle when viewed laterally. Common spermathecal duct attached to distal end of main sclerotised part of genital fork at an acute angle so pointing towards base of abdomen or at right angles so pointing dorsally. Dark form A series from Malaga Province, southern Spain is significantly darker than the typical form and gives the appearance of being a different species. However, this darker appearance is due to no more than the dusting on the mesonotum being less extensive and thinner such that the ground colour shows through. In some individuals the dust also looks darker and browner but some in the series have dusting very like the typical form, just slightly less extensive. The paramedian vittae are much broader, divided by a narrow dusted acrostichal line which almost disappears posteriorly. Antehumeral vittae also more extensive, dusting on posterior part of mesonotum and scutellum inconspicuous such that these areas are subshining black. Dusting on occiput can be darker and browner. Tergites with narrower yellow apical margins. The male genitalia and female genital fork show some slight differences from the typical form but these are not sufficient to justify specific separation. Remarks This species seems to be the commonest and most widespread of the genus in Iberia, the other two Spanish species so far only known in Andalusia. The female genitalia suggest that it is closest to the other two Iberian species, with Pru. aurata showing the closest affinities in North Africa. Distribution Southern Spain north to Salamanca., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 138-141, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Becker T. 1906. Usia Latr. Berliner Entomologische Zeitschrift 50 [1905]: 193 - 228."]}

Parusia propinqua gen. et sp. nov. urn:lsid:zoobank.org:act: AE374F61-1A31-46DA-9C35-C986FD795B31 Fig. 66 Etymology From Latin ‘ propinquus ’ = ‘near’, ‘neighbouring’, ‘related’; refers to the great similarity of this species to other members of the aurata -species group. Type material Holotype SPAIN • ♂; “ Andalusia, Sierra Nevada, above Padul, 1341 m, N37°02.732′ W3°36.321′, 1 May 2008, leg. D.J. Gibbs ”; NHMUK. Paratypes SPAIN • 1 ♂; “ Andalusia, Sierra Nevada, above Padul, 1346 m, N37°02.736′ W3°36.324′, 2 May 2008, leg. D.J. Gibbs ”; PCDG • 1 ♂, 1 ♀; “ Almeria, Uleila N37°11′56″ W02°12′07″ 740 m 25 March 2001, leg. J. Dils & J. Faes ”; PCJD • 2 ♀♀; “ Baños y Mendigo, 13 km S of Murcia, Mosa Trajectum 14 May 2004 leg. C.E. Dyte ”; NHMUK • 1 ♀; “same data”; PCDG. Description MEASUREMENTS. Body length: 2.6–3.9 mm. Wing length: 2.7–4.2 mm. Male HEAD. Gena and mouth margin brownish black to black in ground colour, narrower than the apical breadth of a palp, grey dusted gena narrow, linear, more shining mouth margin broader. Eyes separated at their narrowest by two to three times the diameter of the front ocellus, hind ocellus separated from the eye margin by about half the diameter of that ocellus (on average more widely separated than in Pru. loewi). Ocellar tubercle shining, barely dusted, frons brownish black to black in ground colour, narrow part matt grey dusted, more blackish at narrowest part from some angles, noticeably converging from hind corners to well in front of anterior ocellus, then widening relatively abruptly onto anterior of frons. Front part of frons densely blue-grey dusted, rather more thinly so centrally. White hairs on ocellar tubercle conspicuously longer than the width of the frons at rear, narrow part of frons bare, anterior part of frons with numerous white hairs which are no more than half the length of those on the ocellar tubercle, mostly much shorter. Occiput dark in ground colour densely covered with grey or brownish dust, triangular area behind ocelli equally dusted. The whole covered with erect pale yellow to white hairs, short above just overtopping ocellar tubercle, much longer below, silky, slightly wavy but not very dense. Ommatidia uniform in size across the eyes. Antennae blackish, postpedicel variable but usually a little less than twice length of scape and pedicel together, sensilla in subapical sulcus white and conspicuous in the three male types. All antennal segments with short pale brown hairs above, longest just before subapical sulcus. Palps short but easily visible, strongly clavate, brown to black, the pale yellow apical setae as long as the length of the palps. Proboscis not very long, approximately one and a half head lengths, dorso-laterally clothed with short, inclined brown setae, basoventral membrane brown. THORAX. Black ground colour largely obscured by grey to brown dust except along the conspicuous paramedian and antehumeral vittae where the shining black cuticle shows through, dulled by a thin coating of brownish dust, subshining. Paramedian vittae variable a little narrower to a little broader than dusted acrostichal stripe, extending from very front of mesonotum to three quarters of the way back. Antehumeral vittae usually divided at thoracic suture, lacking darker spot above wing base, area in front of scutellum can be darker, more thinly dusted. Mesonotum covered with moderately long, pale yellow hair, the longest hairs as long as those on the ocellar tubercle. Scutellum relatively sparsely haired on the disc, long hairs fringing the apical margin. Pleura dusted as notopleuron, with similar hairs on pronotum, posterior half of the anepisternum and a few scattered across katepisternum. Metepimeron dark in specimens so far seen. WING. Membrane faintly brown tinged, the veins brown, yellower basally. Crossvein r-m a little beyond basal third of discal cell, conspicuously beyond m-cu. Anal lobe moderately developed with evenly convex margin, about as broad as the anal cell. HALTERE. Pale yellow to white, base of stem slightly brownish. LEGS. Coxae dark with a coating of grey dust like that on the pleura. Remainder of legs black or very dark brown, only thinly dusted so not obscuring the shining cuticle. Coxae externally and femora ventrally covered with rather long, white hairs, on the femora longer than the greatest depth of the femora. Femora dorsally and tibia rather densely covered with short, white hairs. ABDOMEN. All tergites black and shining, but very inconspicuously dusted so not as glossy as completely undusted cuticle would be (compare gonocoxite). Tergite one more densely brown dusted so not shining, laterally reflexed margins of all tergites densely grey dusted obscuring the ground colour. All tergites with narrow to very narrow, sharply demarcated yellow apical margins, visible at least laterally even if dull yellow, often obsolete on disc and tapering away towards sternites. Sternites black and densely grey dusted obscuring the ground colour, apical yellow margins absent or very obscure. Tergites and sternites all covered with fairly long, erect yellow hairs, longest on sternites. GENITALIA. Conspicuously large and globular compared to most Protypusia gen. nov., but typical for Parusia gen. nov. Gonocoxites black to dark brown, shining, composed of two rounded hemispheres separated by a deep sulcus. Gonostylus sharply curved, apical three-fifths at right angles to basal twofifths. Epandrium shining black to dark brown, apico-lateral corners rounded. Both gonocoxite and epandrium covered in long whitish to pale yellow hairs, longest and densest on gonocoxite. Female Differs from the male in its more broadly separated eyes, about five times the diameter of the front ocellus, about one-fifth head width at narrowest point, hind ocellus separated from the eye margin by approximately one and a half times the diameter of that ocellus. A few hairs on the middle part of the frons laterally (sometimes very inconspicuous). Hairs on legs and abdomen shorter, conspicuously so on the abdomen. Apical yellow margins to tergites significantly broader and brighter, not or hardly narrowing medially, tergite eight not noticeably more shining than preceding tergites. Proximal part of genital fork flat (only two dissected specimens), spathulate, bluntly pointed. Common spermathecal duct attached to distal end of main sclerotized part of genital fork at an obtuse angle or at right angles so pointing towards tip of abdomen or dorsally. Remarks Very closely allied to Pru. loewi and occurring alongside it at the type locality although at a higher altitude. Differs mainly in the form of the epiphallus, non-genitalic features being very subtle. Based on the few specimens available, this species is smaller, darker, with less shining thoracic vittae and reduced yellow apical margins to the tergites. Females differ subtly in the form of the genital fork but may not always be reliably determinable. However, in the specimens dissected so far the form of the genital fork and attachment of the common spermathecal duct is consistent. Distribution Southeastern Spain., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 141-144, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958

Protypusia strymonas gen. et sp. nov. urn:lsid:zoobank.org:act: 5D2D1180-1764-4057-9044-52869C88F438 Fig. 56 Etymology Noun in apposition, named after the River Strymonas that flows through Lake Kerkini, Northern Greece. Type material Holotype GREECE • ♂; “ Kerkini, Ecotourism site 65 m, N41°8′15.6″ E23°13′1″, 19 June 2008, leg. G. Ramel pan trap”; NHMUK. Paratypes GREECE • 2 ♂♂, 2 ♀♀; “ Kerkini, Ecotourism site 65 m, N41°8′15.6″ E23°13′1″, 19 June 2008, leg. G. Ramel pan trap”; NHMUK • 1 ♂, 1 ♀; “same data”; PCDG • 1 ♀; “ 1–7August 2006, leg. G. Ramel Malaise ”; PCDG • 1 ♂; “ 15-21 August 2006, leg. G. Ramel Malaise ”; PCDG • 1 ♀; “ Lithotopossite, N41° 13740 E23° 21714, 20-26 June 2005, leg. G. Ramel Malaise ”; NHMUK • 1 ♀; “ Krousia Site 190 m, N41°11′32.4″ E23°3′59.5″, 8–14 August 2007, leg. G. Ramel Malaise ”; PCDG • 1 ♀; “ 11–17 July 2007, leg. G. Ramel Malaise ”; PCDG. Description MEASUREMENTS. Body length: 1.8–2.4 mm. Wing length: 1.8–2.2 mm. Male HEAD. Gena plus mouth margin very narrow, no wider than anterior ocellus, the shining yellowish mouth margin significantly broader than the linear, often hardly visible dusted gena. Gena broadening fairly abruptly just below antennae, running into long narrow frons, at antennal insertions no wider than scape, covered with grey to brown dust. Eyes narrowly separated by about half the diameter of the anterior ocellus or even less. Vertex between the eyes densely grey-brown dusted, similar to frons. Ocellar tubercle dark in ground colour, dark brown dusted, all ocelli contiguous with the eye margins or narrowly separated, especially anterior ocellus. Yellow hairs on ocellar tubercle about as long as width of vertex at its broadest, often sparse or even absent. Eye facets small and equal throughout. Occiput dark in ground colour, densely coated with grey-brown dust. Occiput covered with pale yellow to white hairs, upper ones just behind eyes relatively long, almost equalling hairs on ocellar tubercle, those on ventral side of head similar only a little longer. Antennae black with a brown or grey cast on scape and pedicel, postpedicel a little longer than scape and pedicel together, conspicuously contracted below in apical third, tip truncate (postpedicel rather variable in size and shape). Palps small but readily seen, pale greyish-white with a few inconspicuous pale hairs apically. Proboscis black, not exceptionally long, about equal to thorax including scutellum, hairless dorsally, the basoventral membrane pale-whitish. THORAX. Dark ground colour obscured by dense grey to brown dust, often with a golden or olive tinge, patterned with sharply defined dark brown paramedian vittae back to just rear of wing bases and antehumeral vittae, the latter broadly separated at thoracic suture. The median area in front of scutellum and posterior to the paramedian vittae with darker brown, poorly defined area; often a well-defined dark brown spot on supra-alar area just behind transverse suture (obscure in some). Hairs of mesonotum white to pale yellow, rather sparse, relatively short, arranged biserially along acrostichal and uniserially along dorsocentral lines, more general on lateral parts of mesonotum. Scutellum dark grey-brown, dusting similar to adjacent parts of mesonotum, more or less darker longitudinal mid-line, yellowish to white hairs very scant on disc longer around margin. Pleura greyer dusted than mesonotum, pronotum and posterior two-thirds of the anepisternum, with short yellow to white hairs. Anterior spiracle conspicuously white. WING. Membrane hyaline with the faintest brownish tinge, the veins brown, yellower basally. Crossvein r-m between one third and mid-point of discal cell, distinctly beyond m-cu. Anal lobe fairly well developed with evenly convex margin, about as broad as anal cell. HALTERE. Whitish, stem slightly infuscated especially at base, knob with a clear dark brown dorsal spot. LEGS. Dark brown, coxae variable with dull yellow apices, trochanters murky yellow to brown, hind ones often with blackish rim, femora narrowly yellow basally and apically, often obscure, tibia and base of basitarsi dull yellow tending to be infuscated dorsally, remainder of tarsi blackish. Coxae grey dusted like pleura, femora more thinly grey dusted. Coxae, femora and tibia clothed with short white hairs, longer on the coxae, fore and mid-femora posteriorly, hind femora antero-ventrally often also a little longer haired. ABDOMEN. Dark brown dusted with greyer dust laterally all tergites with narrow but clear yellow apical margins, broadest on middle tergites and laterally, extending right to edge of tergites. All tergites with variable white to pale yellow hairs shorter than the length of respective tergite especially on basal tergites, hairs a little longer laterally. Sternites similar to tergites but more evenly grey dusted, apical margins whiter, hairing short and sparse. GENITALIA. Large for Protypusia gen. nov., but a little smaller than in Pro. punctipennis, concolourous with abdomen, but more obviously grey dusted, tip of epandrium dull yellow-brown, less dusted so shinier, gonocoxite also with shiny yellow apices, much longer-haired than tergites, epandrium shorterhaired but still slightly longer-haired than tergites. Female Differs from the male in its broadly separated eyes, about one fifth head width, hind ocelli separated from eye margin by a one and a half times their diameter. Frons dusted as occiput, a little greyer, a darker brown line down middle from front ocellus to antennae, this dark area continued behind ocellar tubercle onto occiput. Frons with short sparse hairs antero-laterally. Postpedicel averages a little shorter. Hairing of thorax and legs shorter. Abdomen also short haired but not much different to male. Legs very similar to male, in some more extensively yellow. Remarks See Pro. punctipennis and Pro. kerkini gen. et sp. nov. Only eight specimens known from two samples at almost the same location in the village of Lithotopos to the south of Lake Kerkini, Northern Greece. Presumably more widely distributed but Usiini have a tendency towards endemism so could be very restricted. Distribution Greece (Lake Kerkini)., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 109-112, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958

Apolysis turkmenica (Paramonov, 1947) comb. nov. Parageron turkmenicus Paramonov, 1947: 218. Type material Holotype TURKMENISTAN • ♂; “ Askhabad, Transcaspia 23 April 1926 1 ♂ (leg. S.J. Paramonov)/ Oligodranes modestus Lw. S. Paramonov det. ♂ / Parageron turkmenicus sp. Nov ♂ S. Paramonov det./ Holotypus / Zool. Mus. Berlin”; ZMHB. Other material examined UZBEKISTAN • 1 ♂; “Samarkand, along river Zeravshan N39°38′ E67°04′ 22 May 1989 1 ♂ ”; NHMUK • 1 ♂; “ 2 km E, pasture N39°39′ E67°01′ 22 May 1989 1 ♂ (leg. Barták, coll. D.J. Greathead)”; NHMUK • 1 ♀; “Kisilkum, Fedchenko/Coll. H.Loew/ Typus / Oligodranes modestus cotypus Lw ♀ /КиЗилъкумъ [Kizilkum]/ Usia sp. Dr. E.O. Engel det/Zool. Mus. Berlin [this specimen is part of the type series of Oligodranes modestus Loew, 1873 (placed in Parageron by Evenhuis & Greathead (1999), Evenhuis & Greathead (2015) and in Protypusia in this review) but is certainly an Apolysis and very likely an example of A. turkmenicus]”; ZMHB. Remarks Unfortunately, the holotype now lacks antennae making a generic diagnosis difficult. However, it is a very small, slender-bodied species with holoptic eyes, a long, narrow discal cell and broadly developed anal lobe, all characters typical of Apolysis. The genitalia are very small and in the undissected specimen were retracted into the apical tergites rendering them invisible. Upon dissection, it became evident that the genitalia are of the Apolysis type, rotated 90°, gonostylus sub-bifurcate, and epiphallus simple. A particularly interesting feature is the apparent absence of the lateral ejaculatory apodeme, a condition found in other Apolysis but never in Usiini. Comparison with two further males from the Greathead collection (in NHMUK) and dissection of a probable female provide further evidence for the placing of this species in Apolysis. Distribution Turkmenistan, Uzbekistan., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 154-155, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Paramonov S. J. 1947. Uebersicht der mit der Gattung Usia Latr. (Bombyliidae, Diptera) naechstverwandten Gattungen. Eos 23: 207 - 220. Available from http: // hdl. handle. net / 10261 / 149924 [accessed 25 Jan. 2023].","Loew H. 1873 Beschreibungen europaischer Dipteren. Systematische Beschreibung der bekannten europaischen zweiflugeligen Insecten. Von Johann Wilhelm Meigen. Zehnter Theil oder vierter Supplementband. Beschreibungen europaischer Dipteren. Dritter Band. H. W. Schmidt, Halle.","Evenhuis N. L. & Greathead D. J. 1999. World Catalog of Bee Flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden.","Evenhuis N. L. & Greathead D. J. 2015. World Catalog of Bee Flies (Diptera: Bombyliidae) web site. Available from http: // hbs. bishopmuseum. org / bombcat / [accessed 25 Jan. 2023]."]}

Ectopusia additivaneura (Carles-Tolrá, 2009) gen. et comb. nov. Fig. 33 Parageron additivaneura Carles-Tolrá, 2009: 64. Etymology The specific name refers to the distinct additional crossvein present between veins R 2+3 and R 4 (‘ additivus ’ = ‘additional’ in Latin; ‘ neuron ’ = ‘vein’ in Greek). Type material Holotype SPAIN • ♂; “ Almería, 10-5-2004 Parque Natural Cabo de Gata –Níjar, pitfall, A. Aguirre leg./ Parageron additivaneura sp. n. ♂ Holotipo M. Carles-Tolrá det.”; PCMC-T. Paratype SPAIN • 1 ♀; “ Almería, 3-5-2004 Parque Natural Cabo de Gata –Níjar, pitfall, A.Aguirre leg./ Parageron additivaneura sp. n. ♀ Paratipo M. Carles-Tolrá det.”; PCMC-T. Redescription (from two specimens in spirit, so some details will not be comparable with descriptions from pinned material) MEASUREMENTS. Body length: 2.9–4.4 mm. Wing length: 3.4 mm. Male HEAD. Gena relatively narrow, at narrowest point about as broad as scape is long, broadening above into frons, entirely yellow in ground colour, a very narrow shinier border to the oral opening below. Frons converging dorsally to its narrowest point just in front of front ocellus were it is separated by four times the diameter of front ocellus. Frons yellowish-white with short adpressed white hairs anteriorly. Ocellar tubercle darker brownish, ocelli arranged in an equilateral triangle, the hind ocelli a little more than the diameter of that ocellus from eye margin, a few short, white hairs close to hind ocelli. All eye facets small. Occiput dark brown becoming blacker towards neck; covered with long white hairs longer than length of scape and pedicel combined, especially dense and ‘shaggy’ below. Antennae with scape and pedicel light brown a little darker than frons, third segment lighter, yellowish, especially apically. Scape quadrate, pedicel a little deeper than long, postpedicel, a little less than twice as long as scape and pedicel together, swollen basally with narrow, parallel-sided tip. Scape and pedicel with numerous short white hairs, third segment sparsely haired towards tip on dorsal side. Palps small but readily visible, slightly swollen apically, whitish-yellow with conspicuous white apical hairs about three quarters the length of the palp. Proboscis moderate, about twice frons length, brown, tapering but without swollen base, naked, the basoventral membrane pale brown. THORAX. Mesonotum blackish in ground-colour, yellowish post pronotal lobe, notopleuron and post alar calli; densely coated with grey dust, no darker or undusted vittae apparent. Mesonotum largely naked but white hairs on post pronotal lobe, anterior slope of mesonotum, notopleuron and a very few near scutellum. Scutellum as mesonotum but disc yellower, long white hairs present basally. Pleura yellowish-brown, a little darker on ventral parts of katepisternum and meron with long white hairs on katepisternum dorsally and anepisternum dorso-medially. WING. Additional crossvein present between veins R 2+3 and R 4. Membrane clear and veins whitish except where infuscated. Fork R 2+3 /R 4+5, base of R 4 and crossveins R 2+3 -R 4, r-m, m-m and m-cu infuscated brownish on vein and narrowly on to adjacent membrane. Crossvein r-m a little beyond middle of the discal cell, conspicuously beyond m-cu. Anal lobe well developed with conspicuously convex margin, obviously broader than anal cell. HALTERE. Knob whitish, stem slightly infuscated, especially at base. LEGS. Coxae slightly paler brown than pleura. Femora, tibia and tarsi yellowish-brown. Legs covered with white hairs, longer and more erect on the coxae and femora posteriorly, tibia hairs short and adpressed. ABDOMEN. Tergites brownish on basal two thirds, with fairly well demarcated yellow posterior margins on remainder. All tergites with pale hairs from a half to equalling the length of each tergite longer laterally. Sternites similar but paler and yellower, infuscated basally, hairing as on tergites. GENITALIA. Brown, paler apically, pale gonostyli, covered in fairly long white hairs as on tergites. Female As male except gena a little broader and frons wider, evenly narrowed from front to vertex, eyes smaller, lateral ocelli a little more distant from eye margin. Frons infuscated medially so less clear yellow than in male. Antennae a little shorter, not so abruptly contracted apically. Mesonotum with slightly paler paramedian vittae separated by a narrow, dark acrostichal stripe. Abdomen tapering to a point. Remarks This recently discovered species is clearly distinct from all other Usiini and it is difficult to know where its affinities may lie. The female genitalia suggest affinity with Parageron s. str., with rather weakly sclerotised genital fork and unpigmented vaginal plate. However, males of this genus are holoptic or nearly so with differentiated eye facets. The gonostyli also show some similarity to those seen in Parageron s. str., with a prominent internal ‘thumb’-like process, but the epiphallus really does not conform, being closest to Protypusia flavipalpis gen. et sp. nov., otherwise a quite different species. It seems most probable that Ectopusia gen. nov. is basal to Parageron s. lat. Distribution So far only known from the type locality in Almería, Southern Spain., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 32-35, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Carles-Tolra M. 2009. Two new bombyliid species from Spain (Diptera, Bombyliidae). Boletin de la Sociedad Entomologica Aragonesa 44: 63 - 66."]}

Protypusia raydahensis (El-Hawagry & Al Dhafer, 2016) gen. et comb. nov. Fig. 55 Parageron raydahensis El-Hawagry & Al Dhafer, 2016: 1307. Etymology After the locality “Garf Raydah” where the types were collected. Type material Holotype KINGDOM OF SAUDI ARABIA • ♂; “ Not seen but photographs studied. Asir.Abha. Raydah, 1811.884’ N, 4224.435’E, Alt. 2387 m asl, 8 June 2014, Sweep-net Leg. El-Hawagry”; KSMA. Paratypes KINGDOM OF SAUDI ARABIA • 1 ♂; “not seen but photographs of EFC specimens studied Asir. Abha. Raydah, 1811.884’ N, 4224.435’E, Alt. 2387 m asl, 7 June 2014, Sweep-net Leg. El-Hawagry”; EFC • 2 ♀♀; “ Asir. Abha. Raydah, 1811.884’ N, 4224.435’E, Alt. 2387 m asl, 7 June 2014, Malaise Leg. Al Dhafer, H; Fadl, H; Abdel Dayem, M; El Torkey, A; El Gharbawy, A”; KSMA • 1 ♀; “same data”; EFC. Other material examined Metatypes KINGDOM OF SAUDI ARABIA • 1 ♂; “ Asir. Abha. Raydah, 18°11.749’ N, 42°23.345’E, Alt. 1614 m asl, 5 September 2015, M.T.9 [Malaise Trap 9] Leg. Al Dhafer, H; Fadl, H; Abdeldayem, M; El Torkey, A; El Gharbawy, A; Soliman,A/ Parageron raydahensis El-Hawagry & Al Dhafer, 2016 ♂ ”; PCDG • 1 ♀; “same data. Parageron raydahensis El-Hawagry & Al Dhafer, 2016 ♀ ”; PCDG (donated by KSMA). Redescription MEASUREMENTS. Body length: 2.7–3.2 mm. Wing length: 3.6–4.0 mm. Male HEAD. Frons and gena densely silvery dusted entirely obscuring ground colour, longer erect hairs almost absent, can look more yellowish from some angles. Eye margin of frons sigmoid when viewed from directly above. Silvery dusted gena narrowing below antennae, at narrowest point narrower than the more shining, yellowish mouth margin, slightly protuberant. Occiput and ocellar tubercle dark in ground colour fairly densely covered with whitish dust, browner between ocelli, slightly thinner and more shining on elongate triangle behind ocelli. Ocellar tubercle with fine, pale yellow hairs about as long as width across hind ocelli, occiput with fine white, dorsally not over-topping ocellar tubercle, becoming longer ventrally but not particularly dense. All ocelli in contact with eye margin. Eyes confluent for slightly longer than the length of vertex (from photos about 1:1.1). Ommatidia conspicuously enlarged in the upper half to two fifths of the eyes, the transition to the smaller ones occupying the lower part distinct but not very abrupt. Antennae blackish, sparsely whitish dusted, scape approximately as long as broad, pedicel shorter, approximately one and a half times as broad as long. Postpedicel rather elongate, parallel-sided, roundly blunt-ended, with a small point dorsally immediately beyond the subapical sulcus, almost twice as long as scape and pedicel together and three times as long as deep. Sensilla in subapical sulcus whitish translucent so relatively conspicuous. Short, sparse whitish hairs on scape and pedicel, postpedicel with very short, sparse hairs dorsally, longer and more frequent before sub-apical sulcus. Palps small, slightly clavate, dark brown to black with a few short white hairs apically. Proboscis relatively long, approximately three to three and a half times the length of the head, longer than head, thorax and scutellum combined, dorsally entirely devoid of hairs, black, basoventral membrane brownish-yellow to dirty white. THORAX. Dark ground colour obscured by whitish-grey to brownish dust. Mesonotum with pronounced dark-brown to blackish, subshining paramedian and antehumeral vittae the latter widely divided at thoracic suture a very faint round dark spot between wing base and posterior antehumeral vittae. The grey acrostichal line is narrower than to about as wide as the paramedian vittae at their widest (about middle), anterior antehumeral vittae as broad or broader than the grey dusted area between paramedian and antehumeral vittae. Whole of mesonotum and scutellum covered with fine, whitish hairs, coarser and longer laterally. Paramedian and antehumeral vittae bare, acrostichal hairs bi- triserial, dorsocentrals in four to fine irregular lines. Scutellar hairs rather longer, especially laterally. Dusting of scutellum as mesonotum but thinner so shining cuticle visible centrally and in posterior view distinct paler grey dusting baso laterally. Pleura grey dusted as notopleuron, a little greyer and denser than on disc of mesonotum with similar hairs on pronotum, sparsely scattered across hind half of anepisternum and in antero-dorsal corner of katepisternum. WING. Membrane hyaline, faintly brown tinted, the veins brown, paler on subcosta and basally. Crossvein r-m within or at basal third of the discal cell just a little beyond m-cu.Anal lobe broad with conspicuously convex margin, clearly broader than anal cell. HALTERE. White to pale yellow, base of stem slightly infuscated, in some a clear brown infuscation on base of knob dorsally. In other examples with base of knob more or less infuscated with brown. LEGS. Largely black with a coating of thin white dust, denser, similar to pleura, on coxae. Femora with the tips narrowly clear yellowish, a little redder on hind femora. Hind tibia narrowly yellow at extreme base, other tibia more extensively yellow, especially mid tibia dorsally. Adpressed fine whitish hairs clothe legs, rather longer, hairs on four anterior femora posteroventrally, on hind pair anteroventrally and on coxae. Hind tibia with a fairly regular uniserial row of darker setae dorsally on apical half. ABDOMEN. Tergites dark brown to black, tending to be blacker basally, browner close to pale apical margins, thinly brownish-grey dusted, subshining, laterally and on reflexed margins much more densely yellow-grey dusted, denser than that on pleura and almost obscuring ground colour. Tergite one with dusting on reflexed lateral margins not strongly delimited, merging into dusting of disc which is a little more densely dusted than tergite two. Each tergite with sharply demarcated pale yellowish apical margins, mostly narrower than the thickness of a tibia. Sternites densely grey dusted as reflexed margins of tergites with pale apical margins paler and narrower than those on respective tergites. Tergites and sternites all covered with sparse, fine, silky whitish hairs, longest on lateral margins of tergites, tergite one hairless on disc except pale apical margin which has relatively short, adpressed hairs. Tergites two and three also bare basally, much of disc only sparsely haired. GENITALIA. Relatively large when compared to other holoptic species. Gonocoxites black to dark brown, apices reddish brown, yellowish grey dusted, subshining, paler tips shining, undusted, with whitish hairs similar to those on lateral margins of tergites. Epandrium rectangular when viewed laterally, blackish brown basally with broad yellow apical margin, especially the triangularly pointed apical corners, covered with short, adpressed whitish hairs similar to those on tibia. Gonostyli and epiphallic complex essentially identical to Pro. negevi gen. et comb. nov. as far as can be seen in undissected specimen. Female Differs from the male in its broadly separated eyes, the frons dark in ground colour, except anteriorly between antenna where brownish yellow, densely grey-brown dusted, any hairs minute and confined to anterior third and lateral margins, a small dark shining spot centrally. Hind ocelli separated from eye-margin by about twice the diameter of that ocellus. Frons about a quarter to two-fifths head width. Mesonotum darker markings can be less conspicuous, the antehumeral vittae very obscure, or as male. Hairing of thorax, legs and abdomen shorter, significantly so on the abdomen. In dried specimen apical sternite strongly convex, sharply ridged apically, fairly thinly yellow-grey dusted, subshining. Remarks In the key provided by Engel (1932) this taxon runs to Pro. inornata; and to Usia grisea Efflatoun with the key in Efflatoun (1945). It has not been possible to study male specimens of either species, but from Efflatoun’s (1945) descriptions, Pro. raydahensis gen. et comb. nov. is significantly smaller than Pro. inornata and larger than Usia grisea, and from both by more extensive pale bases to the four anterior tibia and more thinly dusted mesonotum with shining paramedian and antehumeral vittae. Also, it seems likely that Usia grisea in fact belongs in Apolysis based on its very small size and Efflatoun’s (1945: pl. 20 fig. 319) illustration of the antennae but is currently retained in unplaced Usia. Protypusia raydahensis gen. et comb. nov. is very closely allied to Pro. negevi gen. et comb. nov., the male and female genitalia being exceedingly similar. Protypusia negevi is a much paler, densely dovegrey dusted species with very inconspicuous, vaguely darker paramedian and antehumeral vittae. Distribution Kingdom of Saudi Arabia, mountains of the Asir province in the southwest of the country., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 105-109, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Engel E. O. 1932. Bombyliidae. In: Lindner E. (ed.) Die Fliegen der palaarktischen Region. Vol. 4. pt. 3. E. Schweizerbart, Stuttgart.","Efflatoun H. C. 1945. A monograph of Egyptian Diptera. Part VI. Family Bombyliidae. Section 1: Subfamily Bombyliidae Homoeophthalmae. Bulletin de la Societe Fouad Ier d'Entomologie 29: 1 - 483."]}

Protypusia kerkini gen. et sp. nov. urn:lsid:zoobank.org:act: 82A88DD4-C822-404F-8C71-B3DB47C9F344 Fig. 50 Etymology Noun in apposition, after the type locality, Kerkini, in Northern Greece. Type material Holotype GREECE • ♂; “ Kerkini, Midway site 750 m, N41°18′49.8″ E23°16′35.6″, 23-29 June 2008, leg. G. Ramel Malaise ”; NHMUK. Paratypes GREECE • 1 ♂; “ Kerkini, Lithotopos site, N41° 13740 E23° 21714, 20-26 June 2005, leg. G. Ramel Malaise ”; NHMUK • 1♀; “ Kerkini site, N41° 15181 E23° 19866, 20-26June 2005, leg. G. Ramel Malaise ”; NHMUK • 3 ♂♂, 3 ♀♀; “ Tree 4 330 m, N41°16′33.4 E23°12′49.8, 14-20 July 2008, leg. G. Ramel pan trap”; OUMNH • 1 ♂, 1 ♀; “ Krousia Mts. Site 190 m, N41°11′32″ E23°3′59″, 27 June-3 July 2007, leg. G. Ramel Malaise ”; PCDG • 2 ♂♂, 1 ♀; “ Krousia Site 190 m, N41°11′32.4″ E23°3′59.5″, 11-17 July 2007, leg. G. Ramel Malaise ”; PCDG • 3 ♂♂, 3 ♀♀ (pinned); “ Strymon F-Plain 55 m, N41°16’09” E23°19’39”, 21 May 2009, leg. G. Ramel Yellow Pan traps”; PCDG • 1 ♂, 2 ♀♀ (pinned); “same data”; NHMUK • 24 ♂♂, 10 ♀♀ (in spirit); “same data”; NHMUK • 1 ♂, 1 ♀; “ Roupel’s Gorge 60 m N41°17′32″ E23°19′35″, 7 June 2009, leg G. Ramel Yellow Pan trap ”; PCDG. Description MEASUREMENTS. Body length: 1.7–2.8 mm. Wing length: 1.7–3.0 mm. Male HEAD. Gena very narrow, linear, almost invisible, the shining yellow mouth margin significantly broader than gena, about as wide as width of front ocellus. Gena broadening abruptly just below antennae, running into long narrow frons, at antennal insertions hardly as wide as scape, covered with whitish-grey dust, a little darker and browner centrally. Eyes very narrowly separated by about half the diameter of anterior ocellus. Vertex between the eyes densely brown dusted, a little darker than the frons. Ocellar tubercle dark in ground colour, dark brown dusted, all ocelli contiguous with the eye. Pale hairs on ocellar tubercle about as long as width across hind ocelli, often sparse or even absent. White hairs on frons also short and inconspicuous. Eye facets small and equal throughout. Occiput dark in ground colour, densely coated with grey-brown dust, area behind ocelli a little darker. Occiput covered with pale yellow to white hairs, upper ones short, those on ventral side of head a little longer. Antennae black with a brown or grey cast on scape and pedicel, postpedicel conspicuously longer than scape and pedicel together, obviously contracted below in apical third, convex, swollen below in basal two thirds, tip truncate. Palps small but larger than in many related species, easily seen, pale whitish with a few inconspicuous pale hairs apically. Proboscis black, fairly long, about equal to head-thorax length (with scutellum), hairless dorsally, the basoventral membrane pale whitish. THORAX. Dark ground colour obscured by dense grey to brown dust, often with an olive tinge, patterned with sharply defined dark brown paramedian vittae back to just rear of wing bases and antehumeral vittae, the latter broadly separated at thoracic suture. Area in front of scutellum with a clear prescutellar vittae running forward almost to paramedian pair; supra-alar area just behind transverse suture lacking any defined spot. Hairs of mesonotum white to very pale yellow, relatively short, arranged along acrostichal and dorsocentral lines irregularly bi- to tri-serial more general on lateral parts of mesonotum. Scutellum dark grey-brown dusting similar to adjacent parts of mesonotum, yellowish to white hairs sparse on disc longer around margin. Pleura greyer dusted than mesonotum, pronotum and posterior two-thirds of the anepisternum, with short white hairs. WING. Membrane hyaline with the faintest brownish tinge, the veins brown, yellower basally. Crossvein r-m between one third and mid-point of discal cell, distinctly beyond m-cu. Anal lobe well developed with evenly convex margin, clearly broader than anal cell. HALTERE. Clear white knob lacking defined brown dorsal spot, stem slightly infuscated especially at base. LEGS. Femora and coxae brown to black with trochanters yellow, hind ones more or less darkened, base and tips of femora narrowly yellow, tibia and base of basitarsi yellow, apical tarsal segment black. Coxae grey dusted like pleura, femora also grey-dusted but more thinly so. Coxae, femora and tibia clothed with short white hairs, longer on the coxae, fore and mid-femora posteriorly, hind femora antero-ventrally. ABDOMEN. Dark brown dusted with greyer dust laterally with all tergites with broad, conspicuous yellow margins, broadest on basal tergites and laterally, extending right to edge of tergites. All tergites with white to pale yellow hairs mainly longer than the length of respective tergite, longest laterally, shorter and sparser on disc. Sternites similar to tergites but distinctly more grey-dusted, pale apical margins more obscure. GENITALIA. Large for the genus, but still not quite as large as in Pro. punctipennis, dusting similar to lateral part of abdomen, epandrium shining apically, yellow to brownish or blackish. Gonocoxite also shining and more brownish apically. Both epandrium and gonocoxite with long hairs as on tergites, or even longer on gonocoxites. Female Differs from the male in its broadly separated eyes, about one fifth head width, hind ocelli separated from eye margin by a little more than their diameter. Frons dusted as occiput, a little greyer, a darker brown line down middle from front ocellus to antennae, short sparse hairs antero-laterally. Postpedicel averages a little shorter. Hairing of thorax, legs and abdomen shorter, significantly so on the abdomen. Femora often much paler, almost as yellow as tibia. Remarks This species forms a distinct group with Pro. punctipennis and Pro. strymonas gen. et sp. nov., all of which are clearly very closely related. They are characterised by their relatively large male genitalia, approaching the condition seen in Parusia gen. nov. However, the epiphallic complex and female genitalia are very much of the same type as seen in more typical members of Protypusia gen. nov. These three species are confined to the eastern Mediterranean, and while Pro. punctipennis is widespread, the other two are currently only known from the vicinity of Lake Kerkini in Northern Greece, an example of the propensity for endemicity within the Usiini. Distribution Greece., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 90-92, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958

Protypusia dimonica (Zaitzev, 1996) gen. et comb. nov. Figs 13, 24, 29, 45 Parageron dimonicus Zaitzev, 1996: 693. Parageron dominicus, error for dimonicus Zaitzev, 1996 – Evenhuis & Greathead 1999 (acting as First Revisers). In his type description, Zaitzev (1996) used the name “ dominicus ” throughout except in the abstract where he used dimonicus. The spelling of dimonicus was selected as the correct original spelling because it derives from the town of Dimona where the types were collected. Etymology Named after the town of Dimona in the Israeli Negev close to the type location. Type material Holotype ISRAEL • ♀; “Loc. No. 18, C. Negev, N. ′Negarot 11 km SE, Mizpe Ramon 17 April 1994, Volkovitsh & Dolgovskaya leg./ Holotypus, Parageron dimonicus V. Zaitzev 96”; TAU. Paratype ISRAEL 1 ♀; “Loc. No. 18, Central Negev, N. ′Negarot 11 km SE, Mizpe Ramon 17 April 1994, V. Zaitzev leg./ Paratypus, Parageron dimonicus V. Zaitzev 96”; TAU. Other material examined ISRAEL • 1 ♀; “ Ein Mur, 19 April 1975 A. Freidberg / griseola ”; TAU • 1 ♀; “ Avdat, 11 April 1975 A. Freidberg ”; TAU • 2 ♀♀; “ 19 April 1975 ”; TAU • 1 ♀; “ Ein Mur, 31 March 1981, F. Kaplan ”; TAU • 1 ♂; “ Jerusalem, Mishot Rotem, 7 March [19]65, M. Weichselfish ”; TAU. MOROCCO • 9 ♂♂, 5 ♀♀; “ Ouarzazate, Tizi-n-Bachkoum 1500 m N30º49′06.0″ W07º15′33.4″ 16 April 2008, leg. J. Dils & J. Faes ”; PCJD • 6 ♀♀; “ Tafraout, Tafraout 1000 m N29º44′19.0″ W08º57′51.4″ 13 April 2008 leg. J. Dils & J. Faes ”; PCMB • 1 ♂; “ 3 May 1961, Maroc Sahar, Taidelt/[leg.] P. duMerle 4333/ Parageron griseus Param, V. Zaitzev det. 2002”; MNHN. Redescription MEASUREMENTS. Body length: 2.6–3 mm. Wing length: 1.9–3.2 mm. Male HEAD. Frons and mouth margin silvery grey, ground colour completely obscured, lacking longer erect hairs. Silvery dusted gena very narrow, almost linear below, much narrower than the rather protuberant, shiny black to brown mouth margin (in Israeli specimens dusted gena a little broader). Occiput and ocellar tubercle dark in ground colour densely and evenly covered with grey dust and outstanding white hairs, those on ocellar tubercle hardly longer than width of vertex across hind ocelli, longer in Israeli specimens. Hairs on occiput relatively short, in face view only just visible, becoming longer below onto jowls. All ocelli in contact with eye margin, making an equilateral triangle. Eyes confluent for slightly longer than the length of vertex (difficulty to assess because the eyes come together at very acute angles both above and below). Ommatidia conspicuously enlarged in the upper half to three fifths of the eyes, the transition to the smaller ones occupying the lower part distinct but not very abrupt. Antennae black, postpedicel mid-length, rather convex below, deepest beyond middle, with a small point dorsally immediately beyond the subapical sulcus, about one and a half times as long as scape and pedicel together. Antennae sparsely and very short-haired above, a few longer but inconspicuous ones immediately before the subapical sulcus. Palps minute and slender, slightly clavate, dark brown to blackish, the white apical setae shorter than the length of the palps. Proboscis moderately long, a little more than head-thorax length (including scutellum), naked dorsally, black except for the dirty-white basoventral membrane in some examples. THORAX. Dark ground colour obscured by grey dust, tending to be browner on disc in Moroccan specimens. Mesonotum with distinct darker paramedian vittae from front to just beyond wing bases (often darker anteriorly in Moroccan specimens) and more diffuse darker antehumeral vittae widely divided at thoracic suture (anterior one blackish in Moroccan specimens). Behind thoracic suture, and below the antehumeral vittae is a vague, roundish darker spot, barely visible in Israeli specimens. Whole of mesonotum and scutellum with variably long, white hair (longer in Israeli specimens), tending to form acrostichal and dorsocentral lines, more generally distributed laterally. Paramedian and antehumeral vittae mostly hairless. Scutellar hairs apically almost as long as scutellum and confined to periphery, naked on disc. Pleura concolourous with mesonotum, with similar hairs on pronotum and posterior twothirds of the anepisternum. WING. Membrane hyaline, at most very faintly brown tinged, the veins pale yellow-brown, yellower basally, subcosta yellow. Crossvein r-m very close to the base of the discal cell, barely beyond m-cu. Anal lobe broad with triangularly convex margin, obviously broader than anal cell. HALTERE. Pale yellow, base of stem slightly infuscated. LEGS. Coxae blackish with coating of grey dust same as pleura. Femora predominantly blackish or dark brown with a covering of grey dust, the tip yellow, more brownish on hind femora. Tibia and tarsi blackish or brown, base narrowly yellow. Coxae and front four femora furnished with moderately long white posterior hairs, hind femora with rather shorter anterior hairs, legs otherwise covered with short, adpressed white hairs. ABDOMEN. Tergites dark blackish-brown with dense grey dusting tending to be darker, subshining on disc of first two tergites, laterally. Each tergite with sharply demarcated pale yellow apical margins, sometimes broader on central tergites, tapering to a point on reflexed lateral margins. On average yellow apical margins narrower than width of hind basitarsus, often conspicuously so. Sternites similar to tergites. Tergites and sternites all covered with fairly long, erect white hairs, longest on lateral margins of tergites. GENITALIA. Small often partially contracted into apical tergites. Gonocoxites black, grey dusted with tips shinier and browner, covered in long white hairs. Epandrium basally black, grey dusted, with very broad yellow apical margin, covered in short, dense white hairs, erect medially, apically directed on yellow part. Cerci brown. Female Differs from the male in its broadly separated eyes, the frons dark in ground colour, densely grey dusted with short, white hairs anteriorly, a small dark undusted spot centrally. Frons about one third head width. Ocelli forming an equilateral triangle. Antennae tending to be shorter than in male. Eye facets equalsized throughout. Mesonotal dusting rather more yellowish, darker vittae tending to be less conspicuous. Hairing of thorax, legs and abdomen shorter, significantly so on the abdomen. Yellow apical margins on tergites tending to broaden towards tip. Vaginal plate smaller and less pigmented in Israeli specimens than in Moroccan specimens. Remarks Despite the wide distribution from Israel to Morocco, there is very little variation between populations. Moroccan specimens tend to have less clearly pale knees, males have longer vestiture and a rather browner mesonotum. There are minor differences in the genitalia of the few specimens dissected but no more than can often be found within populations. Using the key in Efflatoun (1945), this species will run to Pro. tewfiki (Efflatoun), but Efflatoun’s choice of characters in his key is bizarre and Pro. tewfiki can readily be eliminated on abdominal pattern. Based on Efflatoun’s descriptions, the Smithsonian Archives plates, and the photos from EFC, it is probably closest to Pro. deserticola (Efflatoun). The possibility that Pro. dimonica is a synonym of this species has to be considered. Unfortunately, I have been unable to examine the types of Pro. deserticola held in Cairo, but photographs suggest that it differs in colour of dusting and has more strongly marked paramedian and antehumeral vittae. The genitalia of both sexes of Pro. deserticola need to be studied to confirm its conspecificity or otherwise with Pro. dimonica. Distribution Israel, Morocco., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 72-75, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Zaitzev V. F. 1996. On the Bombyliidae (Diptera) of Israel. I. Entomologicheskoe Obozrenie 74 (4) [1995]: 902 - 912. [In Russian.]","Evenhuis N. L. & Greathead D. J. 1999. World Catalog of Bee Flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden.","Efflatoun H. C. 1945. A monograph of Egyptian Diptera. Part VI. Family Bombyliidae. Section 1: Subfamily Bombyliidae Homoeophthalmae. Bulletin de la Societe Fouad Ier d'Entomologie 29: 1 - 483."]}

Protypusia vagans (Becker, 1906) gen. et comb. nov. Figs 8, 58 Usia vagans Becker, 1906: 221. Etymology From Latin ‘ vagus ’ = ‘wandering’, ‘vague’. Type material Lectotype TUNISIA • ♂; “ Gafsa, Biró/ vagans Beck. ♂ det. Becker / Lectotypus /Zool. Mus. Berlin [leg. Lajos Biró, abdomen greased, head glued back on upside down]” ZMHB. Other material examined MOROCCO • 4 ♀♀; “Oujda, Plateau du Rekkam 100 km E Ain-Benimathar 1150 m N33°44.254 W2°59.095, 25 April 2010, Leg. Dils J. Faes J. ” PCJD. Redescription MEASUREMENTS. Body length. 2.2–2.6 mm. Wing length: 2.3–3.0 mm. Male HEAD. Gena and mouth margin narrow, about as broad as the length of the pedicel with pale grey-dusted gena barely wider than the tip of the palps, mouth margin shiny. Frons rather small, triangular and not tumid, silky white without outstanding hairs. Eyes meeting for about 9–10 facets, equal to or greater than the length of the vertex. Ocellar tubercle dark in ground colour, densely grey-brown dusted, all ocelli contiguous with the eye margins. White hairs on ocellar tubercle noticeably longer than the width of vertex at its broadest. Eye facets large in upper half, small in lower half, the division between them, just below the antennal insertion, not well marked. Occiput dark in ground colour, densely coated with grey-brown dust, covered with whitish hairs about half as long as those on ocellar tubercle, becoming longer and silky below. Antennae blackish with scape and pedicel more brownish, postpedicel about 1.4 times as long as scape and pedicel together, rather oval in shape with no more than a slight point dorsally immediately beyond the subapical sulcus. Palps small, subclavate, pale brown with several white hairs apically as long as palp. Proboscis moderate, about as long as thorax and scutellum, hairless dorsally, the basoventral membrane blackish-brown. THORAX. Dark ground colour obscured by brownish-grey dust, with vaguely darker paramedian vittae extending back two thirds the length of the mesonotum, also faint, obscurely defined, darker antehumeral vittae broken at thoracic suture. Hairs of mesonotum pale yellow-white, rather long, longer than ocellar hairs, acrostichals irregular, dorsocentrals irregularly uniserial, hairs on hind third and lateral parts of mesonotum more general, more dense on notopleuron, area above wing bare. Scutellum rather more yellowish brown dusted than mesonotum, hairs equal in length to than those on mesonotum, generally scattered dorsally but lateral margins bare. Pleura concolourous with mesonotum, pronotum and posterior two-thirds of the anepisternum, with long, white hairs. WING. Membrane somewhat milky with inconspicuous brown shades around fork of R 2+3 and R 4+5, fork of R 4 and R 5 and the crossveins r-m, m-m and m-cu the veins pale yellow basally, darker in rather more than apical half. Crossvein r-m a little beyond basal third of the discal cell, clearly beyond m-cu. Anal lobe very well developed with conspicuously convex margin, much broader than anal cell. HALTERE. Yellow, stem and base of knob clearly suffused with brown. LEGS. Coxae concolourous with pleura. Femora and tibia rather browner, not densely dusted, very tip of femora and narrow base of tibia yellow. Legs covered with white hairs, longer on the coxae, fore and mid-femora posteriorly, hind femora antero-ventrally and hind tibia dorsally. ABDOMEN. Greased but probably grey-brown dusted similar to thorax. Tergite one brownish laterally and along apical margin, remaining tergites also with narrow dull brownish posterior margins. All tergites with long white hairs on reflexed lateral margins, dorsally hairs shorter and largely confined to the posterior half of each tergite. Sternites similar to tergites. GENITALIA. Small even for holoptic Protypusia gen. nov., dark in ground colour, tip of gonocoxite yellower, densely covered in brown dust, covered in white hairs. Female Differs from male in having widely separated eyes, frons almost a third head width, densely olive-grey dusted with small median black spot. Laterally, close to eyes and level with anterior ocellus, is a uniserial row of relatively long, proclinate hairs, front half of frons with more generally distributed shorter hairs. Hairs on ocellar tubercle shorter, a little longer than distance across hind ocelli. Hairs of mesonotum, pleura and scutellum very much shorter and rather sparser, dusting with a definite yellow-brown colour. Legs generally with shorter hairs. Infuscation on wing veins can be rather obscure, most conspicuous on crossveins. Abdomen with apical yellow margins of tergites a little broader and more obvious, hairing of tergites and sternites significantly shorter than in male. Remarks The male and female genitalia suggest close affinities with Pro. dimonica and Pro. hyalipennis, the shape of the gonostyli especially being very similar in these three species. Protypusia hyalipennis especially is very similar, but the mesonotal pattern and hyaline wings readily distinguish it. Although both species occur in Morocco, Pro. hyalipennis is known north of the Atlas Mountains while Pro. vagans to the south. Protypusia dimonica also occurs on the Saharan side of the Atlas Mountains in Morocco, so could be sympatric with Pro. vagans, but is readily identified by its general blue-grey colour and entirely hyaline wings. Distribution Morocco, Tunisia., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 115-117, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Becker T. 1906. Usia Latr. Berliner Entomologische Zeitschrift 50 [1905]: 193 - 228."]}

Protypusia punctipennis (Loew, 1846) gen. et comb. nov. Fig. 54 Usia punctipennis Loew, 1846: 417. Etymology From Latin ‘ punctus ’ = ‘pierced’ and ‘ penna ’ = ‘feather’, ‘wing’; ‘spotted-wings’. Type material Lectotype (here designated) [TURKEY] • ♂; “[headless, front left and hind right legs missing; otherwise in good condition.] Small square blank label/ Usia punctipennis m./9691/ Coll. H. Loew / Type /Zool. Mus. Berlin”; ZMHB. Paralectotype UNKNOWN • 1 ♀; “[antennae missing; otherwise in good condition.] Small square blank label/ Typus / Zool. Mus. Berlin ”; ZMHB. Other material examined GREECE • 7 ♂♂, 13 ♀♀; “Fokida, Itea 0 m, N38°25′53″ E22°23′49″ 15 May 2001 ”; PCJD. TURKEY • 4 ♂♂, 3 ♀♀; “ Icel, Ќarakűtűk, 549 m, N37°8′28.7″ E34°50′20.1″ 3 June 2005 ”; PCJD • 7 ♂♂, 6 ♀♀; “ 663 m N37°8′27.6″ E34°50′23.6″ 5 June 2005 ”; PCJD • 8 ♂♂, 5 ♀♀; “ Calaman, 683 m N37°11′33.1″ E34°48′33.9″ 5 June 2005 (leg. J. Dils & J. Faes)”; PCJD. ISRAEL • 1 ♀; “ Tel Aviv beach, 10 June 1996 (leg. B. Merz & A. Freidberg)”; TAU • 1 ♀; “Merzliya, 28 August 1977, F. Kaplan ”; TAU • 2 ♂♂; “ 5 km E, No’omi, saline, 31°54′ N 35°30′ E, 16 March 2005, L. Friedman ”; TAU. CYPRUS • 1 ♂, 1 ♀; “Akamas Peninsular, Lara beach 0 m 28 April 2002 / 2t.28 dunes, meadow (leg. B. Merz, J. Deeming, M. Ebejer & P. Gatt)”; NMWC. Redescription MEASUREMENTS. Body length: 2.0–3.0 mm. Wing length: 1.9–3.6 mm. Male HEAD. Gena plus mouth margin very narrow, no wider than anterior ocellus, the shining yellow mouth margin significantly broader than linear, often hardly visible dusted gena. Gena broadening abruptly just below antennae, running into long narrow frons, at antennal insertions hardly as wide as scape is long, covered with whitish-grey or brown dust, often a little darker and browner centrally, frons paler dusted in Israeli specimens. Eyes clearly separated, by distinctly more than half diameter of anterior ocellus. Vertex between the eyes densely brown dusted, often a little darker than the frons, but paler and uniform in Israeli specimens. Ocellar tubercle dark in ground colour, dark brown dusted, hind ocelli contiguous with the eye, front ocellus narrowly separated. Pale hairs on ocellar tubercle rather longer than the width across hind ocelli. White hairs on anterior part of frons short and inconspicuous. Eye facets small and equal throughout. Occiput dark in ground colour, densely coated with brown to grey-brown dust. Occiput covered with pale yellow to white hairs, upper ones short, falling well short of tips of hairs on ocellar tubercle, those on ventral side of head a little longer but not dense. Antennae black with a brown or grey cast on scape and pedicel, postpedicel variable, longer than scape and pedicel together, obviously contracted below in apical third, convex, swollen below in basal two thirds, tip truncate. Palps relatively large, easily seen, pale yellow with a few very inconspicuous pale hairs apically. Proboscis black, rather short, about equal to thorax length (with scutellum), but longer in Israeli specimens where longer than head plus thorax, hairless dorsally, the basoventral membrane pale-whitish. THORAX. Dark ground colour obscured by dense grey to brown dust, often with an olive tinge, patterned with sharply defined dark brown paramedian vittae back to just rear of wing bases and antehumeral vittae, the latter broadly separated at thoracic suture. A dark prescutellar vittae running forward almost to paramedian pair, less clear but still present in some specimens; supra-alar area just behind transverse suture lacking any defined marking in many, but clear in some. Hairs of mesonotum white to very pale yellow, relatively short, arranged along acrostichal and dorsocentral lines irregularly bi- to tri-serial more general on lateral parts of mesonotum. Scutellum dark grey-brown dusted similar to adjacent parts of mesonotum, often with darker longitudinal mid-vittae, sparse yellowish to white hairs on disc longer around margin. Pleura greyer dusted than mesonotum, pronotum and posterior two-thirds of the anepisternum, with short white hairs. WING. Membrane apart from faint brownish tinge, with clear brown spots over junctions of R 2+3 and R 4+5, base of R 4, r-m, m-m and m-cu. Veins brown, subcosta yellower basally. Crossvein r-m approximately at mid-point of discal cell, or even beyond, well beyond m-cu. Anal lobe moderately developed with evenly convex margin, equal or a little broader than anal cell. HALTERE. Knob variable, from clear white (some Israeli specimens) to more or less dark brown dorsally (Greece and Turkey), stem slightly infuscated especially at base. LEGS. Femora and coxae brown to black, coxae with narrow yellow apical rim, femora sometimes yellowish basally, tips of femora yellow. Trochanters dark black to brown. Tibia and base of basitarsi yellow, apical tarsal segments black, hind tibia sometimes infuscated dorsally and apically. Coxae grey dusted like pleura; femora also grey-dusted but more thinly so. Coxae, femora and tibia clothed with short white hairs, longer on the coxae, fore and mid-femora posteriorly, hind femora antero-ventrally. ABDOMEN. Dark brown dusted with greyer dust laterally on reflexed margins, all tergites with conspicuous yellow apical margins, broadest on basal tergites and laterally, narrowing to edge of tergites (margins broader and brighter in Israeli specimens). All tergites with white to pale yellow hairs mainly longer then the length of respective tergite, longest laterally, shorter and sparser on disc. Sternites similar to tergites but distinctly more grey-dusted, pale apical margins more obscure. GENITALIA. Large for the genus, only some members of Parusia gen. nov. with proportionately larger male genitalia, dusting similar to lateral part of abdomen, epandrium shining apically, yellow to brownish or blackish. Gonocoxite also shining and more brownish apically. Both epandrium and gonocoxite with long hairs as on tergites, or even longer on gonocoxites. Female Differs from the male in its broadly separated eyes, about one fifth head width (a little wider in Israeli specimen), hind ocelli separated from eye margin by a little more than their diameter (more in Israeli specimen). Frons dusted as occiput, a darker brown line down middle from front ocellus to antennae, short sparse hairs antero-laterally. Postpedicel averages a little shorter. Hairing of thorax, legs and abdomen shorter. Femora often much paler, brown or sometimes almost as yellow as tibia. Wing spotting often more distinct and can include a spot at tip of anal cell. Haltere variable, knob with dark infuscation in Israeli specimen. Remarks Very closely allied to Pro. kerkini gen. et sp. nov. and Pro. strymonas gen. et sp. nov. Relatively widely distributed and so exhibiting some variability with more southern examples (Israel) being generally paler and with entirely white knob to haltere (2 males), or brown infuscated (1 female) and longer proboscis. No significant differences could be found in the epiphallic complex between Greek/Turkish and Israeli specimens. Distribution Cyprus, Greece, Israel, Turkey., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 102-105, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Loew H. 1846. Fragmente der Kenntniss der europaischen Arten einiger Dipteren-Gattungen. Linnaea Entomologica 1: 319 - 530. Available from https: // www. biodiversitylibrary. org / page / 46583212 [accessed 25 Jan. 2023]."]}

Protypusia zimini (Paramonov, 1947) gen. et comb. nov. Fig. 59 Parageron zimini Paramonov, 1947: 216. Etymology Named in memory of Leonid Sergeevich Zimin (1902–1970), who collected the types. Type material Lectotype (here designated) UZBEKISTAN • ♂; “ Tschangyr, northwest Bukhara, 13 June 1930 1 ♂ (leg. L.S. Zimin)/ Parageron zimini sp. nov. ♂ S. Paramonov det. Syntypus /Zool. Mus. Berlin.” ZMHB. Paralectotypes UZBEKISTAN • 2 ♀♀; “locality as lectotype 10 June 1930 ”; ZMHB • 1 ♂ [dissected], 1 ♀; “same locality 13 June 1930 ”; ZMHB • 1 ♀; “same data 17 June 1930 ”; ZMHB • ♂ ♀ in copula, 2 ♀♀; “same data 20 June 1930 ”; ZMHB. Other material examined IRAN • 1 ♂, 1 ♀; “ Ghazvin, road to Zereshk 1682 m, N36°24’128” E50°5’434” 28 June-7 July 2009, pan trap Leg. B. Gharali ”; PCDG. Redescription MEASUREMENTS. Body length; 3.9–5.1 mm. Wing length: 4.1–5.2 mm. Male HEAD. Frons and gena pale yellow in ground colour, the frons slightly tumescent covered with silky off-white dusting, lacking longer hairs. Gena moderately broad, at narrowest point about as broad as length of pedicel, more shining mouth margin a little narrower, together almost as broad as scape length. Occiput and ocellar tubercle dark in ground colour densely covered with grey dust and outstanding white hairs, ocelli contiguous with eyes. Eyes confluent for slightly longer than the length of vertex. Ommatidia conspicuously enlarged in the upper half of the eyes, sharply contrasting with the smaller ones occupying the lower half. Antennae blackish, tip of pedicel can be paler, postpedicel rather long with a small point dorsally immediately beyond the subapical sulcus. Palps small and slender, yellowish to brown, darker apically, the white apical setae shorter than the length of the palps. Proboscis very long, about equal to body length, naked dorsally, the basoventral membrane yellow. THORAX. Dark ground colour obscured by dove-grey dust, in some becoming more yellowish on posterior of mesonotum and scutellum, very faint darker paramedian vittae on anterior half or slightly more of mesonotum, antehumeral vittae may be indicated. Whole of mesonotum and scutellum evenly covered with moderately long, silky white hair, largely absent on darker vittae, on the front of the mesonotum hairs longer than tibia is thick at its mid-point. Pleura with similar hairs on pronotum, posterior twothirds of the anepisternum, and a few in upper middle of katepisternum. Metepimeron naked, can be yellow (Uzbekistan). WING. Membrane hyaline, the veins pale yellow, or becoming brown apically. Crossvein r-m from basal quarter to third of the discal cell, opposite or a little beyond m-cu. Anal lobe broad with conspicuously convex margin, broader than anal cell. HALTERE. Pale yellow, base of stem slightly infuscated. LEGS. Coxae dark with a dense coating of grey dust. Femora predominantly dark with a covering of grey dust, up to apical quarter clear yellow (Uzbekistan specimens) or just narrowly yellow (Iranian specimens). Front and mid-tibia clear yellow, hind tibia infuscated apically (Uzbekistan specimens) or infuscation more extensive (Iranian specimens). Tarsi black with the basitarsi more or less yellow basally in some. Legs covered with silky white hairs, rather variable, longer on Iranian male. ABDOMEN. Tergite one entirely yellow (Uzbekistan specimens) or with just apical margin yellow (Iranian specimens), remaining tergites black with dense grey dusting and sharply demarcated apical yellow margins (Uzbekistan specimens) or poorly defined pale margins (Iranian specimens), these becoming narrower rearwards. Sternites similar but yellow apical margins clearer. Tergites and sternites all covered with fairly long, erect white to very pale yellow, rather wavy hairs. GENITALIA. Relatively large when compared to other holoptic species. Gonocoxites black, grey dusted with broad yellow tips, covered in long whitish hairs. Epandrium basally black, grey dusted, with very broad yellow apical margin and narrow lateral margins, covered in short whitish hairs. Female Differs from the male in its broadly separated eyes, the frons pale yellow and hairless anteriorly, densely pale-grey dusted on hind half, laterally with 1–2 rows of tiny hairs. Legs yellow except for coxae basally (Uzbekistan specimens) or similar to the male, grey dusted, more or less infuscated tarsi largely blackish. Abdomen much more extensively yellow than in the male with less well defined blackish-brown bases to the tergites (except T1) and sternites (Uzbekistan specimens) or similar to male. Hairs on the thorax and abdomen a little shorter, most clearly on the tergites. Remarks Although twelve specimens were available, they originate from just two widely separated localities, 10 from the type locality and two from Iran. There is quite a degree of variation between these two locations notably in the extent of yellow on tergites and legs. Uzbekistan specimens have much yellower tergites and legs compared to the two specimens from Iran. However, in most features, including the structure of both male and female genitalia, they differ very little so I am as confident as I can be that they are conspecific. The illustrations provided by Zaitzev (1966) and ascribed to Pro. grisea (Paramonov) are certainly not Pro. grisea and appear very similar to this species or, perhaps the closely related Pro. negevi. The description, while brief, could fit either species, closer to Pro. negevi on size but more likely to be Pro. zimini on range. Without seeing the specimens it is not possible to be confident what species Zaitzev had before him. Distribution (? Georgia), Iran, Uzbekistan., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 119-121, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Paramonov S. J. 1947. Uebersicht der mit der Gattung Usia Latr. (Bombyliidae, Diptera) naechstverwandten Gattungen. Eos 23: 207 - 220. Available from http: // hdl. handle. net / 10261 / 149924 [accessed 25 Jan. 2023].","Zaitzev V. F. 1966. [Parasitic Flies of the Family Bombyliidae (Diptera) in the Fauna of Transcaucasia.] Nauka, Moscow & Leningrad. [In Russian.]"]}

Apolysis marginata (Brunetti, 1909) comb. nov. Usia marginata Brunetti, 1909: 228. Type material Holotype (not examined) INDIA • ♂; “Simla [= Shimla], Himachal Pradesh 16 May [19]09 leg. Dr. Annandale (Brunetti 1909)”; NZSI. Remarks Very likely belongs in Apolysis, non-type material probably of this taxon in the NHMUK was examined and belongs in Apolysis. Examination of the type to confirm this combination is desirable. Removed to unplaced species of Usiinae in Evenhuis (2015). Distribution India (Himachal Pradesh)., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 151-153, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Brunetti E. 1909. New Indian Leptidae and Bombyliidae with a note on Comastes osten Sacken, v. Heterostylum Macquart. Records of the Indian Museum 3: 211 - 230.","Evenhuis N. L. & Greathead D. J. 2015. World Catalog of Bee Flies (Diptera: Bombyliidae) web site. Available from http: // hbs. bishopmuseum. org / bombcat / [accessed 25 Jan. 2023].","Efflatoun H. C. 1945. A monograph of Egyptian Diptera. Part VI. Family Bombyliidae. Section 1: Subfamily Bombyliidae Homoeophthalmae. Bulletin de la Societe Fouad Ier d'Entomologie 29: 1 - 483."]}

Apolysis flavipes (Efflatoun, 1945) comb. nov. Usia flavipes Efflatoun, 1945: 212. Type material Holotype (not examined) EGYPT • ♀; “ Wadi El-Lega (South Sanai) June 1941 leg. Efflatoun (Efflatoun 1945)”; ESEC. The female holotype in ESEC is destroyed, photos show that only the pin, mount and labels exist. Other material examined Neotype (here designated) ISRAEL • ♀ (glued to small white card point, in perfect condition); “ Arava Valley, Iddan, side wadi below date orchard, - 640ft. el., malaise, 17-IV-1996, M.E. Irwin, 30°49’05” N 35°16’55” E”; TAU. Additional specimens (from Neotype locality) ISRAEL • 1 ♀; “ Arava Valley, Iddan, side wadi below date orchard, - 640ft. el., malaise, 12-IV-1996 ” TAU • 1 ♀; “same location 14-IV-1996 ”; TAU • 1 ♀; “same location 17-IV-1996 ”; TAU • 2 ♂♂; “same location 18/ 19-IV-1996 ”; TAU • 1 ♂, 2 ♀♀; “same location 22-IV-1996 ”; TAU • 1 ♂; “same location 25-IV-1996 ”; TAU • 2 ♀♀; “same location 28-IV-1996 ”; TAU • 2 ♀♀; “same location 30-IV-1996 ”; TAU. Remarks Efflatoun (1945) described this species from just one female specimen, now unfortunately destroyed, although not before a plate was completed of it. Searching for the type, photographs of the relevant drawers from the Egyptian Entomological Society collection were made and show just a pin, mount and labels standing over this name. I thus conclude the holotype is destroyed and propose a neotype for it to fix the name and help avoid confusion with other similar-looking species in this genus. Efflatoun’s (1945) original description plus the plate in the Smithsonian Institution Archives [Image no. SIA2012-7882] refer to a distinctive species, most particularly the contrasting ivory-white scape is a feature not seen in other species. The neotype and all other specimens in the series cited above share this unusual feature and fit very closely with Efflatoun’s description and the plate in the Smithsonian archives. There is a degree of variation in the series (as would be expected) that Efflatoun could not have appreciated from his single specimen. The legs vary in the degree of infuscation, in some front femora and tibia not quite clear yellow. Pleura often more extensively yellow than is suggested from the type description. Base of postpedicel externally often pale (as in Efflatoun’s type), but in some this is not apparent. Efflatoun’s holotype was collected in Wadi El-Lega (South Sinai). This is about 320 km south of the collecting locality of the neotype in the Negev. Although not close, the situation is very comparable, a wadi in rocky desert with sparse vegetation (from Google Maps) so a similar biotope. Distribution Egypt (Sinai), Israel (Negev)., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 150-151, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Efflatoun H. C. 1945. A monograph of Egyptian Diptera. Part VI. Family Bombyliidae. Section 1: Subfamily Bombyliidae Homoeophthalmae. Bulletin de la Societe Fouad Ier d'Entomologie 29: 1 - 483."]}

Apolysis melanderi nom. nov. Oligodranes bicolor Melander, 1946: 473. Type locality: USA (California) [Lectotype designated by Evenhuis & Greathead (1999: 667) in USNM]. [Preoccupied by Apolysis bicolor (Efflatoun, 1945)]. Distribution Nearctic: USA (California)., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on page 156, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958, {"references":["Melander A. L. 1946. Apolysis, Oligodranes and Empidideicus in America (Diptera, Bombyliidae). Annals of the Entomological Society of America 39: 451 - 495. https: // doi. org / 10.1093 / aesa / 39.3.451","Evenhuis N. L. & Greathead D. J. 1999. World Catalog of Bee Flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden.","Efflatoun H. C. 1945. A monograph of Egyptian Diptera. Part VI. Family Bombyliidae. Section 1: Subfamily Bombyliidae Homoeophthalmae. Bulletin de la Societe Fouad Ier d'Entomologie 29: 1 - 483."]}

Protypusia argentata gen. et sp. nov. urn:lsid:zoobank.org:act: 930472B2-293B-460D-8AA6-0318EBC9D4C9 Figs 25, 39 Etymology From Latin ‘ argentum ’ meaning ‘silver’, refers to the silver-grey dusting and white hairs covering much of the bodies of both sexes. Type material Holotype MOROCCO • ♂; “ 21 March 2006, Tiznit, Arbaa-sahel 320 m, N29°39′26.5″ W09°52′11.4″ Leg, Dils, J.- Faes, J.”; NHMUK. Paratypes MOROCCO • 1 ♀; “ 21 March 2006, Tiznit, Arbaa-sahel 320 m, N29°39′26.5″ W09°52′11.4″ ”; NHMUK • 1 ♂, 7 ♀♀; “ 21 March 2006, Tiznit, Arbaa-sahel 320 m, N29°39′26.5″ W09°52′11.4″ ”; PCJD • 1 ♀; “ 22 March 2006, Tiznit, Arbaa-sahel 330 m, N29°409′19.7″ W09°51′18.5″ ”; PCJD • 2 ♂♂, 3 ♀♀; “ 25 March 2009, Ouarzazate, Amerzgane 1350 m, N31°01′02.9″ W07°13′45.9″ ”; PCDG • 1 ♀; “ 16 March 2009, Tiznit 330 m, Arbaa sahel N29°39′26.4″ W09°52′11.5″ Leg, Dils, J.-Faes, J.”; PCJD. Description MEASUREMENTS. Body length: 4.0– 5.6 mm. Wing length: 4.1–6.0 mm. Male HEAD. Gena broad, wider than width of proboscis at base, broadening out above into frons, and somewhat inflated, all silvery-grey dusted except for a very narrow, blackish border to the oral opening level with proboscis. Frons covered with long, dark-brown to blackish hairs, the longest equalling the length of the postpedicel, these hairs, after a glabrous area laterally, continuing down to level with proboscis, leaving a narrow, glabrous gap before the silvery occiput hairs start. Holoptic, but eyes only touching for about three facets, the frons drawn out to a fine point. Ocellar tubercle black, subshining but with grey dusting apparent from some angles, a small silvery-grey triangle in front of anterior ocellus, the hind ocelli almost in contact with hind corners of eyes; ocelli form an equilateral triangle. Silvery hairs on ocellar tubercle about as long as scape and pedicel combined. Eye facets clearly enlarged in upper two thirds, about twice the size of lower facets. Occiput densely coated with silver-grey dust except for a triangular patch behind ocellar tubercle which is more thinly dusted. Either side of this triangular area, above neck, are darker brown dusted areas, the occipital callosities almost black. The whole occiput with a long, silvery vestiture, the hairs becoming longer below forming a ‘beard’ below the eye. Post occipital hairs dorsally not differentiated, not over-topping hairs on ocellar tubercle, from front extending above eyes hardly more than height of ocellar tubercle. Antennae blackish, grey dusted, especially on the scape, postpedicel significantly longer than scape and pedicel together, the sensilla in the apical sulcus contrastingly white. Scape and pedicel clothed with silvery hairs, the longest on the scape longer than its diameter; silvery hairs on postpedicel confined to dorsum and usually shorter, longest just before the sulcus. Palps short, claviform, black, thickly clothed with long silvery hairs. Proboscis long, about equal to the head and mesonotum combined, black, including basal membrane, laterally with short brown hairs along its base. THORAX. Mesonotum with blackish ground-colour variably obscured by pale silver-grey dust covering postpronotal lobe, notopleuron, above wing bases, and thinly in front of scutellum (smaller specimens from higher altitude) to densely covered with yellow-grey dust, more thinly on paramedian and antehumeral vittae (larger specimens from lower altitude). Hairs of mesonotum silvery, almost as long as antennae, relatively dense, especially anteriorly, sparser on paramedian vittae and close to scutellum. Scutellum varying from subshining blackish, dusted grey laterally, to densely yellow-grey dusted, with darker, more brownish disc, covered with long silvery hairs dorsally as long, peripherally longer than those on mesonotum. Pleura densely pale grey dusted, pronotum and anepisternum with long, silvery hairs, katepisternum with long silvery hairs ventrally and on anterior half. WING. Membrane hyaline, very faintly yellowish basally, the veins yellowish-brown, paler basally, the subcosta entirely yellow. Crossvein r-m between basal third and middle of the discal cell, clearly beyond m-cu, a little variable.Anal lobe well developed with conspicuously convex margin, broader than anal cell. HALTERE. Pale yellowish-white, base of stem infuscated. LEGS. Coxae concolourous with pleura and with the same long, silvery hairs. Femora and tibia silverygrey dusted, from some angles appearing darker on apices of tibia and tarsi. Femora covered with white hairs, long and dense on fore and mid-femora posteriorly, even longer but less dense on hind-femora. Tibia and tarsi covered with short, silver hairs, the tibia also with longer hairs in proximal halves, especially dorsally. ABDOMEN. Tergites subshining blackish on disc, more or less dusted silvery-grey apically and laterally, Extent of dusting varies from complete on apical segment to a narrow apical rim and on reflexed sites basally in smaller examples from higher altitude. Some specimens have very narrow, sharply defined yellow posterior margins, disappearing laterally. All tergites with long silvery hairs up to about twice as long as the length of each tergite. Sternites densely silver-grey dusted, pale apical margins distinct, becoming broader distally, hairs as long as those on tergites. GENITALIA. Small, similar to Pro. grata, somewhat ventrally deflected, dark ground colour completely obscured by silver-grey dust, slightly less so on tips of gonocoxite, cerci brown, gonostyli black. Hypopygium clothed with silvery hairs, longest on gonocoxite. Female Differs from the male in its broadly separated eyes, frons about one third head width, densely grey dusted, slightly less so on ocellar tubercle. Hairs on frons rather shorter, extending back to level of hind ocelli, the hairs extending down the gena silvery to yellowish-brown rather than black. Silvery post ocellar hairs a little shorter but still exhibits the ‘bearded’ effect below eyes. Hairs on mesonotum, tergites and legs a little shorter, especially dorsally. Mesonotum more densely grey dusted with paramedian and antehumeral vittae vaguely indicated in some specimens. Scutellum uniformly grey dusted. Tergite similar but with broader, more conspicuous apical yellow margins. Remarks Very similar to the sympatric Pro. grata, including the male and female genitalia, and the two species have been collected together at the same site in Morocco. Males are distinctive due to the coating of silvery dusting and largely very long silvery hairs. Females can be very similar to female Pro. grata, but are more uniformly grey dusted on mesonotum and abdomen, hardly showing the darker mesonotal vittae of female Pro. grata. Distribution Morocco., Published as part of Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., pp. 1-162 in European Journal of Taxonomy 863 (1) on pages 52-55, DOI: 10.5852/ejt.2023.863.2081, http://zenodo.org/record/7767958

Supplemental material, sj-docx-1-tej-10.1177_20417314221113746 for Comparison of bone formation mediated by bone morphogenetic protein delivered by nanoclay gels with clinical techniques (autograft and InductOs®) in an ovine bone model by Cameron Black, David Gibbs, Josephine McEwan, Janos Kanczler, Marta Peña Fernández, Gianluca Tozzi, Jonathan Dawson and Richard Oreffo in Journal of Tissue Engineering

Malignant pleural mesothelioma (MPM) is a highly lethal cancer of the lining of the chest cavity. To expand our understanding of MPM, we conducted a comprehensive integrated genomic study, including the most detailed analysis of BAP1 alterations to date. We identified histology-independent molecular prognostic subsets, and defined a novel genomic subtype with TP53 and SETDB1 mutations and extensive loss of heterozygosity. We also report strong expression of the immune-checkpoint gene VISTA in epithelioid MPM, strikingly higher than in other solid cancers, with implications for the immune response to MPM and for its immunotherapy. Our findings highlight new avenues for further investigation of MPM biology and novel therapeutic options. Significance: Through a comprehensive integrated genomic study of 74 MPMs, we provide a deeper understanding of histology-independent determinants of aggressive behavior, define a novel genomic subtype with TP53 and SETDB1 mutations and extensive loss of heterozygosity, and discovered strong expression of the immune-checkpoint gene VISTA in epithelioid MPM. See related commentary by Aggarwal and Albelda, p. 1508. This article is highlighted in the In This Issue feature, p. 1494

Additional file 1. ROSES reporting standards.

Additional file 2. Database search strings and bibliographic databases used in review.

Additional file 6. Guide to data extraction spreadsheet.

Additional file 12. Forest plots for effect sizes reported in experimental studies and variance distribution over meta-analysis models.

Additional file 7. Study validity assessment frameworks for experimental mesocosm and experimental field studies.

Additional file 4. Other citation requests and “test set” of highly relevant citations.

Additional file 10. Meta-analysis checklist (from [57]).

Additional file 13. R markdown file of the meta-analysis, showing forest plots with mean effect sizes for categorical moderating factors, results of three-level mixed effects models, and meta-regression plots for continuous moderating factors.

As part of the ‘immune landscape of cancer’, six immune subtypes were defined which describe a categorization of tumor-immune states. A number of phenotypic variables were found to associate with immune subtypes, such as nonsilent mutation rates, regulation of immunomodulator genes, and cytokine network structures. An ensemble classifier based on XGBoost is introduced with the goal of classifying tumor samples into one of six immune subtypes. Robust performance was accomplished through feature engineering; quartile-levels, binary gene-pair features, and gene-set-pair features were computed for each sample independently. The classifier is robust to software pipeline and normalization scheme, making it applicable to any expression data format from raw count data to TPMs since the classification is essentially based on simple binary gene-gene level comparisons within a given sample. The classifier is available as an R package or part of the CRI iAtlas portal.Code / Tool availabilitySource Code https://github.com/Gibbsdavidl/ImmuneSubtypeClassifierWeb App Tool https://www.cri-iatlas.org/

Despite great progress in Pipunculidae (Insecta: Diptera) systematics during the past decades, the Mediterrane-an fauna of big-headed flies remains largely unknown. Here, we present new faunistic and taxonomic data for 98 named species from Cyprus, Egypt, France, Greece, Italy, Malta, Morocco, Portugal, Spain, Tunisia, and Turkey, based on our own collecting efforts and museum specimens. Besides 56 first national records, the paper includes the description of Cephalops(Semicephalops) brachium Kehlmaier Withers sp. n. from France and Spain, and of TomosvaryellaositoKehlmaier, Gibbs Withers sp. n. and Tomosvaryella pugiunculus Kehlmaier Gibbs sp. n. from the Balear Islands (Spain). Furthermore, two new synonymies are proposed: Tomosvaryella lyneborgi (Coe, 1969) = Tomosvaryella cilitar-sis (Strobl, 1910); and Tomosvaryella glabrum (Adams, 1905) = Tomosvaryella pilosiventris (Becker, 1900). Lectotypes are designated for Tomosvaryella pilosiventris (Becker, 1900) and Tomosvaryella vicina (Becker, 1900). The following species need to be deleted from the Spanish checklist due to misidentifications: Cephalops subultimus Collin, 1956, Eu-dorylas montium (Becker, 1897), and Tomosvaryella nigronitida Collin, 1958.

Malignant pleural mesothelioma (MPM) is a highly lethal cancer of the lining of the chest cavity. To expand our understanding of MPM, we conducted a comprehensive integrated genomic study, including the most detailed analysis of BAP1 alterations to date. We identified histology-independent molecular prognostic subsets, and defined a novel genomic subtype with TP53 and SETDB1 mutations and extensive loss of heterozygosity. We also report strong expression of the immune-checkpoint gene VISTA in epithelioid MPM, strikingly higher than in other solid cancers, with implications for the immune response to MPM and for its immunotherapy. Our findings highlight new avenues for further investigation of MPM biology and novel therapeutic options. SIGNIFICANCE: Through a comprehensive integrated genomic study of 74 MPMs, we provide a deeper understanding of histology-independent determinants of aggressive behavior, define a novel genomic subtype with TP53 and SETDB1 mutations and extensive loss of heterozygosity, and discovered strong expression of the immune-checkpoint gene VISTA in epithelioid MPM.See related commentary by Aggarwal and Albelda, p. 1508.This article is highlighted in the In This Issue feature, p. 1494.

The Cancer Genome Atlas (TCGA) has catalyzed systematic characterization of diverse genomic alterations underlying human cancers. At this historic junction marking the completion of genomic characterization of over 11,000 tumors from 33 cancer types, we present our current understanding of the molecular processes governing oncogenesis. We illustrate our insights into cancer through synthesis of the findings of the TCGA PanCancer Atlas project on three facets of oncogenesis: (1) somatic driver mutations, germline pathogenic variants, and their interactions in the tumor; (2) the influence of the tumor genome and epigenome on transcriptome and proteome; and (3) the relationship between tumor and the microenvironment, including implications for drugs targeting driver events and immunotherapies. These results will anchor future characterization of rare and common tumor types, primary andrelapsed tumors, and cancers across ancestry groups and will guide the deployment of clinical genomic sequencing.

We performed an extensive immunogenomic analysis of more than 10,000 tumors comprising 33 diverse cancer types by utilizing data compiled by TCGA. Across cancer types, we identified six immune subtypes-wound healing, IFN-γ dominant, inflammatory, lymphocyte depleted, immunologically quiet, and TGF-β dominant-characterized by differences in macrophage or lymphocyte signatures, Th1:Th2 cell ratio, extent of intratumoral heterogeneity, aneuploidy, extent of neoantigen load, overall cell proliferation, expression of immunomodulatory genes, and prognosis. Specific driver mutations correlated with lower (CTNNB1, NRAS, or IDH1) or higher (BRAF, TP53, or CASP8) leukocyte levels across all cancers. Multiple control modalities of the intracellular and extracellular networks (transcription, microRNAs, copy number, and epigenetic processes) were involved in tumor-immune cell interactions, both across and within immune subtypes. Our immunogenomics pipeline to characterize these heterogeneous tumors and the resulting data are intended to serve as a resource for future targeted studies to further advance the field.

The Adaptation Design Tool of the Corals & Climate Adaptation Planning (CCAP) project was developed to help coral reef managers incorporate climate‐smart design into their plans, programs, and projects at any stage of planning and implementation. The CCAP Adaptation Design Tool (henceforth referred to as the Design Tool can be used to incorporate climate change adaptation into programs using existing planned actions as a starting point, and also to consider development of additional climate‐smart strategies as needed. Most managers will be working from existing plans responsive to previously established management objectives, with climate change already taken into account to varying degrees.

Recent studies, including genome-wide association studies, have identified several putative low-penetrance susceptibility loci for melanoma. We sought to determine their generalizability to genetic predisposition for multiple primary melanoma in the international population-based Genes, Environment, and Melanoma (GEM) Study. GEM is a case-control study of 1,206 incident cases of multiple primary melanoma and 2,469 incident first primary melanoma participants as the control group. We investigated the odds of developing multiple primary melanoma for 47 SNPs from 21 distinct genetic regions previously reported to be associated with melanoma. ORs and 95% confidence intervals were determined using logistic regression models adjusted for baseline features (age, sex, age by sex interaction, and study center). We investigated univariable models and built multivariable models to assess independent effects of SNPs. Eleven SNPs in 6 gene neighborhoods (TERT/CLPTM1L, TYRP1, MTAP, TYR, NCOA6, and MX2) and a PARP1 haplotype were associated with multiple primary melanoma. In a multivariable model that included only the most statistically significant findings from univariable modeling and adjusted for pigmentary phenotype, back nevi, and baseline features, we found TERT/CLPTM1L rs401681 (P = 0.004), TYRP1 rs2733832 (P = 0.006), MTAP rs1335510 (P = 0.0005), TYR rs10830253 (P = 0.003), and MX2 rs45430 (P = 0.008) to be significantly associated with multiple primary melanoma, while NCOA6 rs4911442 approached significance (P = 0.06). The GEM Study provides additional evidence for the relevance of these genetic regions to melanoma risk and estimates the magnitude of the observed genetic effect on development of subsequent primary melanoma.

Usia similis Paramonov, 1950 (Figure 16, Plate XIX) Usia similis Paramonov, 1950: 369. Usia aenea: Engel, 1932: 67 Textfig. 22, misidentification. Type material examined. Type locality Spain. The lectotype, designated by Zaitzev & Kandybina (1983), is in ZIS and it has not proved possible to visit or to borrow the specimen or obtain photographsduring this study. However, Paramonov���s description is quite adequate to be certain that the non-type material examined in the current study belongs to this taxon. Other material examined. Spain, Puerto, Santa Maria [C��diz July 1905], [leg.] Dusmet/ M.N.C.N., Madrid [♂♀] (MNCN); Meigen/ 926, 40/ Usia atrata, Spanian / U. aenea Rossi, det. Becker [♂ in Meigen coll. MNHN]; Punta, Paloma (pris Tarifa, Cadiz), 0.50m, 3 May 1970, [leg.] P. duMerle [1 pair in cop, 10 ♂ 9 ♀ in MNHN]. Etymology. from Latin similis meaning ���similar���, presumably to aeneoides, which Paramonov described in the same work. Diagnosis. A medium sized, robust blue-black species very similar to U. aeneoides Paramonov, endemic to Iberia. Although hairs on head, mesonotum short they are clearly evident, especially laterally, white to pale yellow. Oral margin narrow but wider than in U. aeneoides, occiput densely grey dusted, including depressed area behind vertex, frons relatively narrow, with longitudinal and diagonal grooves. Mesonotal cuticle with strong serrate punctures that give it a very rough appearance, few smooth shining areas away from paramedian vittae. Wings in most specimens with clear pattern of yellow base, brown mid-anterior patch and clear tip and trailing edge. Genitalia large and prominent, lateral horn-like projections of gonostylus usually readily seen and rule out all but U. aeneoides Paramonov. Furca with distinct step between ���arms��� and apical part. Redescription. Measurements. Body length. 4.4���4.8 mm. Wing length. 3.9���4.9 mm. Male. Head. Mouth-margin shining black, narrow but quite evident, about as broad as palps at widest point, obsolete above where it meets the dusting of the frons. Frons narrow (although a little wider than in U. aeneoides), a fifth to sixth head width, shining black, the frons longitudinally striated anteriorly between broad, dense white dust spots which occupy half the length of the frons. Between dust spots and ocellar area the cuticle is diagonally grooved. Frons narrowest at vertex where narrower than third antennal segment, widening slightly to mid-point where it widens more abruptly before resuming its original angle to the anterior of the frons. Lateral ocellus separated from eye margin by about the shortest diameter of that ocellus. Vestiture short, on ocellar area about as long as the distance between lateral ocelli (including ocelli), yellowish white. Occiput densely grey dusted including on triangular area behind vertex which is slightly less densely dusted. White hairs covering occiput similar to those on disc of mesonotum, a little shorter above, longer below. Antennae black, third antennal segment about 1.5 x longer than the scape and pedicel together, relatively elongate, almost bare dorsally, no more than a few inconspicuous very short setae. Proboscis black, hairless, about as long as thorax; palps short, dark brown, clavate with pale hairs, the longest conspicuously shorter than the length of the palps. Thorax. Mesonotum and scutellum shining black (blue-black in some lights) with strong surface sculpture of stellate hair insertions. Wrinkles tending to become more transverse laterally and on the bare paramedian lines. Pronotum, dorsum of post pronotal lobe, adjacent part of mesonotum, notopleuron and end of thoracic suture grey dusted. Scutellum shining with shallow irregular transverse wrinkles. Mesonotal hairing yellowish white, about long as those on ocellar tubercle on disc, longer marginally, acrostichals in 4���5 irregular lines, as wide as the paramedian lines, laterally uniformly distributed, most directed towards middle of mesonotum (pale hairs on disc easily lost). Scutellum uniformly short haired, longest hairs on the margin about half the length of the scutellum. Pleurae black with a covering of dense grey dust except narrow anterodorsal margin of anepisternum which is shining black and undusted (very narrow in one specimen). Pronotum and posterodorsal corner of anepisternum with short yellowish white hairs. Wing. Wing membrane faintly brownish yellow tinged distally. Basal to the humeral cross-vein veins and membrane yellow contrasting with the brown infuscated costal cell, first basal cell, basal half of R 1 (except extreme base which forms a narrow, clear ���window���) and R 2 + 3, vein m-cu and extreme base of discal cell. This forms a clear pattern, best appreciated with the naked eye, similar to that of U. aeneoides and U. aenea. Anal lobe slightly convex but clearly narrower than anal cell, r-m approximately at to a little before middle of discal cell. Haltere. Yellowish white, becoming slightly brown at base of stem. Legs. Entirely black to dark brown, the coxae grey dusted anteriorly and relatively long haired; femora with the longest pale posteroventral hairs barely longer than half the greatest depth of a femur, tibia and tarsi with a covering of minute yellow-brown setulae only. Abdomen. Shining black, with faint bluish reflections, cuticle with conspicuous hair insertions and transverse welts which give the tergites a rough texture, a little more dense on basal tergites. Tergites transversely impressed with base and apical margin reflexed to form an undulating, ridged profile. Disc of the tergites covered with very short, whitish hairs which do not or only just overlap each other. Laterally and on the down-curved margins (especially apically) the tergites are clothed with relatively longer, whitish hairs, as long as visible part of apical tergite (in normally set specimens). Sternites densely grey dusted, also with similar, whitish hairs along their posterior halves. Genitalia. Rather large, conspicuous, globular, as long as the four apical visible tergites, shining black with rough cuticle similar to that of mesonotum. Gonocoxite covered in short, dense, yellowish to golden hairs, including on the conspicuous ���horns��� either side. Viewed caudally the lobes of the gonocoxite are widely separated, between them a hairless, concavity, the cuticle reddish brown, with the gonostyli and tip of epiphallus usually easily visible. Gonostylus ���T���-shaped with an extra point externally, reddish brown, epiphallus with two flattened paddle-shaped lobes apically, reddish brown. Epandrium fairly long, evenly convex with short, yellowish white hairs. Female. As male except as follows: conspicuously broader oral margin which does not disappear above, almost as wide as length of the pedicel. Frons more widely separated, about a quarter head width, diverging evenly (almost) from vertex to front. Hairs on mesonotum and scutellum rather shorter, especially on scutellum. Anepisternum with shining anterior margin tends to be a little wider. Genitalia. Tip of abdomen coming to a rounded point; apical tergites much less densely punctate. Apical sternite transverse, tumescent either side, ridged and falling away steeply proximally, triple point apically. Furca pigmented, the arms long and broad, attached to the apical part at a distinct step, lacks any pigmented vaginal plate. Discussion. This very distinctive species is clearly a sibling species of U. aeneoides Paramonov which has closely similar male and female genitalia and much the same general appearance. Its relationship to other Usia species is less obvious, while it shares the same wing pattern with U. aenea (Rossi), there is little else in common with this species. Before Paramonov recognised this taxon in his 1950 revision, both U. similis and U. aeneoides were confused with U. aenea. Although Becker (1906 b) recognised specimens which fit the description of U. aeneoides, he seems to have had a female only so did not give it a name, referring to it as U. aenea var. However, he appears to have completely overlooked U. similis despite examining a male specimen in the Meigen collection in Paris which he named U. aenea. It is possibly this error that misled Engel (1932) into illustrating the genitalia of U. similis under U. aenea. Distribution. Endemic to Iberia where so far only known from Spain. Only 24 specimens seen during this study, 21 of them from a single date/locality, the other three from two dates. Known collection localities are along the Atlantic coast of Andalusia between Cadiz and Tarifa, most recently in 1970. Seems to be a rare and very local species., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 70-73, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Paramonov, S. J. (1950) Bestimmungstabelle der Usia - Arten der Welt (Bombyliidae, Diptera). Eos, 26, 341 - 78.","Engel, E. O. (1932) Bombyliidae. In: Lindner, E., (Ed.), Die Fliegen der palaarktischen Region. Vol. 4. pt. 3. E. Schweizerbart, Stuttgart, pp. 49 - 96.","Zaitzev, V. F. & Kandybina, M. N. (1983) Fam. Bombyliidae, In: Zaitzev, V. F., Kandybina, M. N., Nartshuk, E. P. & Rikhter, V. A. (Ed.), Catalog of the type specimens in the collection of the Zoological Institute of the Academy of Sciences of the USSR. Insecta (Diptera). Part 1. Families Acroceridae, Nemestrinidae, Bombyliidae. Nauka, Leningrad [= St. Petersburg]. pp. 12 - 44. [in Russian.]","Becker, T. (1906 b) Usia Latr. Berliner Entomologische Zeitschrift, 50 (1905), 193 - 228."]}

Usia florea (Fabricius, 1794) (Plate XV) Voluccella florea Fabricius, 1794: 412. Usia cuprea Macquart, 1834: 383. Usia gagathea Bigot, 1892: 374. Type material of V. florea examined. LECTOTYPE here designated; Algeria or Tunisia, florea / Volucella / ZMUC 0 0 503993 [1 ♀in ZMUC in poor condition with head, much of thorax, right wing, and all legs but mid-right missing]; PARALECTOTYPE; ZMUC 0 0 503994 [1 unsexed in ZMUC, only dorsum of thorax, scutellum and wings surviving]. Originally described from an unspecified number of specimens. Two syntypes found in ZMUC, both in poor condition. The specimen which has the abdomen largely intact, and so could be sexed, was selected as the lectotype. Type material of U. cuprea examined. LECTOTYPE here designated; Italy, Museum Paris, Sicile, leg. A. Lefebvre, Macquart/ TYPE / Usia cuprea [1 ♂ in MNHN in good condition]. Originally described from an unspecified number of specimens. A single male found in MNHN was selected as lectotype. Type material of Usia gagathea examined. HOLOTYPE; Algeria, Alger [Algiers], Usia gagatea [sic.] ♂, n.sp. Inedit, Quincy Octobre 1886. J. Bigot, Dr. Seriziat [leg.]/ U. florea Fab. Det. Becker / U. gagathea Ex coll. Bigot. [1 ♂ in OUMNH]. Other material examined. Algeria, North Algeria, Hamman Rhira, May 1911, (Rothsch. & Hart.)/ Algeria (Hon. L.W. Rothschild & Dr E.J.O. Hartert) 1914 - 57 1 ♂; Algeria, Algiers, May 1908 (Hon. W. Rothschild) 1908 - 242, 1♀ (BMNH); La Calle, Museum Paris, Alg��rie, Coll. H. Lucas 78 - 49 / 1201. [blue label]/ U. claripennis, H. Lucas det. [2 ♂]; 1211 / Usia claripennis, Macqt. [♀]; L���Alma, 7 May 1893, Museum Paris, Algerie, P. Lesne 6-97 [♂♀]; Museum Paris, Env. D���Alger, Bov�� 126 - 38 / 126, 38 [on reverse of round blue label]/ U. cuprea, H. Lucas det. [♀]; Museum Paris, Alg��rie, coll. H. Lucas 78 - 49 [♀]; Ouled Messelem, May 1893 / Museum Paris, Alg��rie, P. Lesne 6-97 [♀] (MNHN); France, Nyons, vi [♂] (MNHN). Etymology. florea from Latin meaning ���of flowers���; cuprea from Latin meaning ���copper���; gagathea from Greek ����������ε�� meaning coal as in coal black. Diagnosis. A fairly large black, species from northern Algeria. All shining black without any obvious coloured reflections, vestiture mid-length, not a very hairy-looking species. Occiput shining and undusted on upper part, thinly grey dusted below, hairs on ocellar tubercle shorter than distance across frons at vertex, frons in front of anterior ocellus essentially glabrous. Mesonotal vestiture mid length, longest acrostichal setulae about as wide as this line of hairs. Scutellar hairs much the same as median mesonotal hairs, the marginal ones usually about half mid-scutellar length in females (but these can be few or even missing in rubbed specimens), much shorter in males. Anepisternum mostly shining, a narrow band of dusting along posterior margin. Genitalia fairly large, epandrium conspicuously wider than long, very open apically, the aedeagus usually clearly visible, its hooked tip protruding beyond apex of gonocoxite; furca largely unpigmented, membranous, a small but distinct vaginal plate present. Redescription. Measurements. Body length. 6.5 ���9.0mm. Wing length. 6 ���8.7.0mm. Male. Head. Mouth-margin fairly narrow, narrower than the tip of the palps at its broadest, almost disappearing below, shining black. Frons rather matt and wrinkled, grey dusting laterally on anterior half, the dust spots separated by an undusted area a little more than one third of the frons width at this point. Frons narrowest level with anterior ocellus, conspicuously wider than width across lateral ocelli, lateral ocellus separated from eye margin by about the diameter of that ocellus up to twice its diameter. Vestiture not very long, on ocellar area shorter than the frons is wide at this point, whitish to yellow-brown. Occiput shining black on a fairly wide strip behind upper eye margin, thinly dusted centrally and below. Antennae black, third segment longer than the scape and pedicel together, all segments with very short setae above. Proboscis black, hairless, shorter than head and thorax together; palps short, black to brown, clavate with pale hairs, the longest shorter than the length of the palps. Thorax. Mesonotum and scutellum shining black except for a very narrow area of light dusting along suture between post pronotal lobe and mesonotum and above the wings root. Smooth except for hair insertions, bare paramedian lines extending back behind the mid point of the mesonotum, each about as broad as the midline which has 5���6 lines of hairs. Hind part of mesonotum slightly wrinkled close to scutellum, scutellum shallowly, transversely wrinkled. Mesonotum and scutellum covered with mid-length pale yellowish-white hair, that on the disc of the mesonotum conspicuously shorter than narrowest width of frons (i.e. longer than in U. accola but shorter than U. atrata), even shorter on hind part of mesonotum and scutellum. Pleurae black with a rather thin covering of grey dust except anepisternum, which is shining black, ventral part of katepisternum which is subshining. Posterior third of anepisternum and pronotum covered with rather longer yellowish white hairs. A few pale hairs on the middle of the katepisternum in some individuals. Wing. Wing membrane faintly yellow tinged becoming more yellow basally, the veins also yellow although conspicuously darker than the wing membrane, base of stem of R darker brown and base of costa blackish. Anal lobe as broad as anal cell, r-m before middle of discal cell. Haltere. Yellow, becoming brown at base of stem. Legs. Entirely black, the coxae grey dusted with pale yellow hairs externally; femora with rather long, pale hairs, those on the ventral surface of the hind tibia longer than depth of tibia but shorter than depth of femora; tibia and tarsi with a covering of minute brown setulae only. Abdomen. Shining black, the disc of the tergites covered with short, neat, whitish hairs. Laterally and on the down-curved margins the tergites are clothed with relatively longer, shaggier whitish yellow hairs (similar to U. maghrebensis Gibbs sp. nov.). Sternites also with similar long, paler whitish hairs. Genitalia. Very large, occupying about half the ventral surface of the abdomen, shining black or partially brown and covered in long, pale hairs similar to those on the lateral margins of the tergites, longest on the gonocoxite. In pinned specimens the very broad epandrium and the distinctive ���hooked��� tip of the epiphallus can usually be seen. The single French specimen differs slightly in having the lateral corners of the epandrium somewhat elongated and squared off in lateral view. Gonostyli also differ in being deeper basally. Female. As male except as follows: noticeably wider frons, the lateral ocellus separated from the eye margin by conspicuously more than the diameter of that ocellus. Oral margin a little boarder. Vestiture on ocellar area conspicuously shorter but the scutellum has longer marginal hairs up to half mid-scutellar length, sometimes inconspicuous. Abdomen conical tapering to a rounded point. Apical sternite characteristic, triangular with a longitudinal ridge, the tip with an apical point and lateral blunt angles. Discussion. The two remaining types of U. florea are in poor condition and one so damaged that it cannot be sexed. The best preserved specimen is a female but careful examination of the underside of the tip of the abdomen reveals the characteristic last sternite allowing confident identification with non-type material. The lectotype of U. cuprea Macquart is problematic because of the surprising collecting locality in Sicily. Usia are highly prone to endemicity, many with restricted ranges and only one, U. versicolor (Fabricius), is known to occur in both Algeria and Sicily. If U. florea really does occur in both Algeria and Sicily then it should also be found in the intervening countries of Tunisia and perhaps even Malta, as is the case for U. versicolor. Further, extreme eastern Algeria and northern Tunisia is occupied by another, apparently parapatric, member of the florea - group, U. accola. If any member of the florea -group were to occur in Sicily, then it is U. accola that would be expected. The provenance of the French specimen is even more worrying, labelled simply ���Nyons vi [June]��� with no year or collector indicated. That there should be a population of this species in south-central France, but with no records at all from the Mediterranean coast of France, mainland Italy, Spain or Morocco, seems highly unlikely. For both these specimens an error in labelling seems to be a far more likely possibility, or perhaps vagrancy or accidental importation. In his original description, Macquart (1834: 383) said that Mr Alexandre Lefebvre found several specimens [of U. cuprea] in Sicily and that the species is perhaps a variety of U. aenea Rossi (Macquart 1834: 383). It is clear from the descriptions in this work that Macquart understood U. aenea to be a species with a dark shade across the wing and that his U. cuprea lacked this marking. But it is very surprising that he thought it might be a variety of U. aenea and must have overlooked the strikingly different genitalia if he really had the type of U. cuprea before him. The ten specimens in the Bigot collection at OUMNH listed in Evenhuis & Greathead (1999) as syntypes show no evidence of being such. There are no Macquart labels on them, just Bigot���s locality labels. When Macquart described the taxon in 1834, Bigot was just 16 years old and had not yet begun to build the collection that, in the 1850 s, he sent to Macquart to describe. So it is extremely improbable that he should have Macquart types from this early period in his collection (A.C. Pont, pers. comm.). The genitalia of the holotype of Usia gagathea Bigot differ in some details from other specimens of this species. The data label gives merely ��� Alger ��� for collecting locality. Bigot often seems to have removed original labels, replacing them with his own. There is only one label on this specimen giving location, collector and name all in Bigot���s hand so this clearly is not the label applied by the original collector, if there was one. It is possible then that the specimen came from elsewhere in Algeria, not Algiers itself, so might be form a different population than most other specimens. However, the small morphological differences are more likely to be individual variation. Distribution. Algeria, (Italy, Sicily? Based solely on the Lectotype male of U. cuprea possibly a labelling error), (France, based on single specimen with limited data, reliability cannot be assessed). All specimens with date data are over 90 years old, none being seen from the inter-war period when numerous naturalists were still active in Algeria. The lack of recent records from Algeria is unsurprising, but there is reason to be concerned for the continued existence of this species. If it ever occurred naturally in Europe it must have been very local and rare and now probably extinct., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 58-61, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Macquart, P. J. M. (1834) Histoire Naturelle des Insectes. Dipteres. Ouvrage accompagne de Planches. Tome premier. N. E. Roret, Paris, 578 pp.","Evenhuis, N. L. & Greathead, D. J. (1999) World Catalog of Bee Flies (Diptera: Bombyliidae). Backhuys Publishers, Leiden, 1 - 756."]}

Usia accola Becker, 1906 (Figure 20, Plate XI) Usia accola Becker, 1906 a: 14 [1906 b: 214]. Type material examined. LECTOTYPE here designated; Tunisia, [locality indecipherable] 1896 / Tunis 40720 / accola Beck / Typus / Zool. Mus. Berlin. [♂ in ZMHB; excellent condition, head greasy]. Originally described from an unspecified number of specimens of both sexes in the Becker collection, now housed in ZMHB. However, only a single male syntype was located in ZMHB so this one becomes the lectotype. Other material examined. Tunisia, Tabarka, 13 May 1995, 1♂, meadow, stream leg. M.J. Ebejer (PCME); Cap Negro, 15 May 1995, ♀ leg. J.C. Deeming (NMWC); Museum Paris, Tunisie, Ain-dhaham, H. Gadeau de Kerville 1908, May, June, Usia florea [♂] (MNHN); Algeria, La Calle, W LeBoul, Usia florea, Museum Paris, Coll. E. S��guy 1919 [♂] (MNHN). Etymology. from Latin, meaning ���near, nearby, a neighbour���. Diagnosis. A large black, very short-haired species of northern Algeria and Tunisia. All shining black, the few available specimens lacking coloured reflections, all vestiture very short, more so than in sympatric species. Occiput shining and undusted behind upper eye margin, otherwise thinly grey dusted, hairs on ocellar tubercle shorter than distance between lateral ocelli, frons in front of anterior ocellus glabrous. Mesonotal vestiture very short and neat, very even in length, acrostichal setulae distinctly shorter than this line of hairs. Marginal scutellar hairs much less than half median length of scutellum. Anepisternum dusted on approximately posterior half. Genitalia small within the florea group, with epandrium longer than wide, apicolateral extensions square; furca with tip membranous. Redescription. Measurements. Body length. 6.4���6.6mm. Wing length. 6.0��� 6.2mm. Male. Head. Oral margin narrow, almost linear although vanishing below shining black. Frons shining black, obliquely wrinkled posterolaterally, narrowest a little in front of anterior ocellus, approximately one fifth head width, distinctly wider than width across lateral ocelli, lateral ocellus separated from eye margin by one to two times the diameter of that ocellus. Frons grey dusted laterally on anterior half, the dust spots separated by an undusted area rather more than one third of the frons width at this point. Hairing very short, on ocellar area shorter than or a little longer than the diameter of a lateral ocellus, brown to dark brown, frons in front of anterior ocellus glabrous. Occiput shining black above adjacent to the eyes and on the depressed area behind the ocellar tubercle, thinly grey dusted centrally and below. White hairs covering occiput similar to those on mesonotum but a little longer especially below. Ommatidia uniform in size across the eyes. Antennae black, third segment longer than the scape and pedicel together, relatively narrow apically, swollen below basally, all segments with very short setae above. Sensilla small, brown and inconspicuous. Proboscis black, hairless, about as long as head and thorax together; palps short, dark brown, clavate with pale hairs, the longest conspicuously shorter than the length of the palps. Thorax. Mesonotum and scutellum shining black except for an area of light dusting on anterior slope between paramedian lines and post pronotal lobe, upper margin of post pronotal lobe and along lower notopleural area and above the wings. Cuticle smoother than in related species, hair insertions not conspicuous, on disc separated by their diameter or a little more. Bare paramedian lines extending back to about the mid point of the mesonotum, each a little narrower than the midline which has 5���6 irregular lines of hairs, uniform width throughout its length. Hind part of mesonotum slightly wrinkled medially close to scutellum, scutellum rather deeply, transversely wrinkled. Pleurae black with a covering of grey dust except approximately anterior half of anepisternum and an irregular patch in the middle of the anepimeron which are shining black and undusted. Mesonotum and scutellum covered with very short whitish hairs, that on the disc of the mesonotum conspicuously shorter than the distance between the lateral ocelli, the majority of hairs overlapping by less than twice the distance between them. Posterior half of anepisternum and pronotum covered with rather longer whitish hairs. A few fine pale hairs on the middle of the katepisternum. Wing. Wing membrane conspicuously brownish yellow tinged anteriorly and basally, fading to a grey cast posteriorly and apically, the veins yellow, base of costa blackish. Anal lobe well developed with smoothly curved posterior margin, as broad as anal cell, crossvein r-m at or before middle of discal cell. Haltere. Yellow, becoming slightly brown at base of stem. Legs. Entirely black, the coxae grey dusted with fine white hairs on outer surface; femora shining, undusted with the longest pale ventral hairs barely longer than half the greatest width of a femur, tibia and tarsi with covering of minute dark setulae. Abdomen. Undusted black, lacking bronzy or bluish reflections, dulled by closely packed punctures that are joined by transverse furrows, the disc of the tergites covered with very short, whitish hairs barely overlapping each other. Laterally and on the down-curved margins the tergites are clothed with relatively longer, more yellowish hairs. Sternites also with similar long, pale yellowish hairs along their posterior halves. Genitalia. Prominent and conspicuous but smaller than in related species, shining black the gonocoxite covered in long, yellowish hairs similar to those on the lateral margins of the tergites, hairs on epandrium much shorter. In dried specimens the apex of the epandrium and lateral lobes of the gonocoxites are paler, in the lectotype yellowish. The characteristic shape of the gonostyli can often be seen in such specimens. Female. As male except as follows: wider frons, shorter third antennal segment, hairs on down-curved margins of tergites shorter and paler. Apical sternite longer than wide, apically angled and with a median ridge. Furca with tip unpigmented, membranous, arms short and almost straight. Discussion. This species is clearly very closely related to U. florea (Fabricius) and most of the few specimens seen have been misidentified as this species. However, the male genitalia are very different so they cannot be mistaken if the epandrium and aedeagus are examined. Probably even more closely related to the longer-haired U. atrata (Fabricius) and U. maghrebensis Gibbs, especially the latter which has similar male and female genitalia. U. florea and U. maghrebensis occur further west and are not known to be sympatric with U. accola but U. atrata can be found with this species in northern Tunisia. Distribution. North east Algeria and adjacent northern Tunisia. The five specimens seen come from a small area either side of the Algeria-Tunisia border, none very far inland. Would appear to be rare and very range restricted., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 48-50, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Becker, T. (1906 a) Die Ergebnisse meiner dipterologischen Fruhjahrsreise nach Algier und Tunis 1906 [part]. Zeitschrift fur Systematische Hymenopterologie und Dipterologie, 6, 1 - 16.","Becker, T. (1906 b) Usia Latr. Berliner Entomologische Zeitschrift, 50 (1905), 193 - 228."]}

Usia aeneoides Paramonov, 1950 (Plate XVIII) Usia aeneoides Paramonov, 1950: 369. Usia aenea Rossi var.: Becker, 1906 b: 214. Type material examined. Type locality Spain. The lectotype, designated by Zaitzev & Kandybina (1983), is in ZIS and it had not proved possible to visit or to borrow the specimen or obtain photographs. However, Paramonov���s description is quite adequate to be certain that the non-type material examined in this study belongs to this taxon. Other material examined. Spain, Andalucia, Cortija del Alcazar, 990���1200m, N 36 �� 55 ��� 15 ��� W 4 �� 5 ��� 18 ���, 9 June 2006, leg. D.J. Gibbs [10 ♂ 12 ♀] (PCDG); El Pardo [Madrid], Arias Encobet/ 11 June 1905 / M.N.C.N, Madrid [♂]; El Pardo [Madrid], Arias Encobet/ 1906 / M.N.C.N, Madrid [♀]; Torre del Vinagre, 0 6 July [19] 88 / Sierra de Cazorla, Jaen Espa��a, Col. C.M. Herrera/ Usia aenea, Rossi, A. Sanchez-Terr��n det./ M.N.C.N, Madrid [♀]; 4 June 1963, Province Madrid, El Chorro, Leg. T.W. Tolman [♂] (MNCN). Etymology. from Latin aeneus meaning ���bronzy��� + - oides meaning resembling. Diagnosis. A medium sized, compact, exceptionally short-haired species endemic to Iberia. An all black species, in natural light with a clear bluish hue, nearly all hairs on head, thorax and legs very short and black to dark brown, longer yellow hairs on abdomen laterally and gonocoxite. Occiput densely grey dusted, oral margin very narrow, frons relatively narrow, with longitudinal and diagonal grooves. Mesonotal cuticle with strong serrate punctures that give it a very rough appearance, few smooth shining areas away from paramedian vittae. Wings in most specimens with clear pattern of yellow base, brown mid-anterior patch and clear tip and trailing edge. Genitalia large and prominent, lateral horn-like projections of gonostylus usually readily seen and rule out all but U. similis Paramonov. Furca with distinct step between ���arms��� and apical part. Redescription. Measurements. Body length. 4.3 ���6.0mm. Wing length. 3.9���5.2mm. Male. Head. Oral margin very narrow, linear, disappearing below, shining black to brownish, narrower than tip of palps. Frons black, shining but conspicuously grooved diagonally in hind half, more shallowly so between dust spots, rather matt due to microsulpture. Anterolateral grey dust spots occupying most of width of frons, black, subshining middle strip narrower than each duster area. Eyes separated at their narrowest by about one sixth to one seventh head width, a little more than width across lateral ocelli inclusive, almost parallel in hind half, abruptly widening then continuing almost parallel to insertion of proboscis. Lateral ocellus separated from the eye margin by the diameter of that ocellus or slightly more. Ocellar tubercle shining, undusted, minute black setulae less than diameter of a lateral ocellus, frons in front of anterior ocellus glabrous. Occiput densely grey dusted, the black ground colour obscured including on most of the triangular depressed area behind ocellar tubercle. A fairly even covering of short black hairs across the dusted area, shorter and more even above, significantly longer close to lower oral margin. Ommatidia uniform in size across the eyes. Antennae black, third segment, almost longer than twice length of scape and pedicel together, parallel sided, sensilla inconspicuous, translucent brownish. All antennal segments with minute brown setulae above. Palps small, black, slightly browner basally, clavate tip wider than oral margin, the dark apical setae about half as long as the length of the palps. Proboscis shorter than headthorax length, black, dorsally hairless. Thorax. Mesonotum shining black with faint bluish hue under natural light, grey dusting on post-pronotal lobe and adjacent parts of mesonotum, notopleuron near suture and above wing root. Disc densely punctate, the individual punctures raised and stellate, often coalescing to create a rough texture to cuticle. On disc any areas of smooth shining cuticle small and usually cross-striated. Paramedian vittae usually traceable by their different surface sculpture, faintly transversely striate and hardly punctured, extending from anterior slope of thorax almost to wing insertions. Mesonotal vestiture very short, black setulae only readily visible on notopleuron and above wings, even shorter ones on anterior slope, disc essentially hairless but in some tiny dark setulae present barely longer than diameter of a puncture. Acrostichal punctures between the bare paramedian vittae in about 3���4 irregular rows. Scutellar punctuation and hairing as adjacent parts of mesonotum, looks naked but tiny black setulae often present. Pleurae uniformly grey dusted largely obscuring ground colour, except for the anterior, dorsal and posterior margins of anepisternum and a small median patch on anepimeron which are shining black. Short black setulae like those on notopleuron on posterior third of the anepisternum. Wing. Membrane with distinct brown markings extending from costa to vein M and faintly to Cu, distally to end of subcostal cell and just beyond r-m, contrasting with yellow base proximal to the humeral crossvein. Veins all dark brown except proximal to humeral crossvein where they are paler, yellowish. Crossvein r-m a little before the middle of discal cell, conspicuously beyond dm-cu. Anal lobe relatively narrow, trailing edge straighter proximally, convex medially, a little over half the width of anal cell. Haltere. Yellow, base of stem slightly brownish. Legs. Coxae black with a coating of grey dust like that on the pleurae. Remainder of legs black, undusted, shining below the setulae. Coxae externally with rather sparse short, black hairs, femora ventrally with short brown hairs, shorter than depth of tibia, hardly different from dorsal hairs. Tibia rather densely covered with short, recumbent, yellow-brown setulae. Abdomen. All tergites shining black, often with faint bluish reflections, slight dusting on down-curved margins, only obvious on first tergite. Each tergite densely punctured, the punctures tuberculate, often joined to neighbours by transverse ridges, each puncture with a tiny brown setae, hardly longer than width of a puncture. Apical rim of each tergite narrowly impunctate, smooth and shining. Sternites grey dusted, faintly punctate, with relatively long, erect pale yellow hairs. Genitalia. Large and globular, occupying much of underside of abdomen. Epandrium flaskshaped with rather triangular apicolateral blunt extensions, longer than wide, mostly shining black, the cuticle with simple punctures, evenly coated with short pale yellow hairs. Gonocoxite approximately square, shining black, orange-yellow apically where gonostyli articulate. Apicolaterally the gonocoxite has a conspicuous blunt ���thumb���- like process surmounted by a brush of golden hairs. Gonostyli ���T���-shaped with an inner, basal lobe, apex of epiphallus with two divergent, paddle-shaped lobes that can often be seen in pinned specimens. Female. As male except as follows: oral margin significantly broader, but still disappearing below. Frons noticeably wider, about one-sixth head width, lateral ocelli separated from eye margin by about twice their diameter, diverging more evenly from vertex. Diagonal grooves in front of anterior ocellus deeper and minute setulae on front of frons between grey dust spots more apparent. Abdomen tends to be more aeneous with coppery, gold and even red reflections (incandescent light). Genitalia. Tip of abdomen coming to a rounded point; apical tergites much less densely punctate. Apical sternite almost square, rather large and convex, apical notch very small, barely visible. Furca pigmented, the arms long and attached to the apical part at a distinct step, lacks any pigmented vaginal plate. Discussion. This very distinctive species is clearly a sibling species of U. similis Paramonov which has closely similar male and female genitalia and much the same general appearance. Its relationship to other Usia species is less obvious, while it shares the same wing pattern with U. aenea (Rossi), there is little else in common with this species. Distribution. Endemic to Iberia and so far only found in Spain from Madrid south to Andalucia. Specimens in collections are rather few so seems to be a local species but can occur in large numbers when found., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 68-70, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Paramonov, S. J. (1950) Bestimmungstabelle der Usia - Arten der Welt (Bombyliidae, Diptera). Eos, 26, 341 - 78.","Becker, T. (1906 b) Usia Latr. Berliner Entomologische Zeitschrift, 50 (1905), 193 - 228.","Zaitzev, V. F. & Kandybina, M. N. (1983) Fam. Bombyliidae, In: Zaitzev, V. F., Kandybina, M. N., Nartshuk, E. P. & Rikhter, V. A. (Ed.), Catalog of the type specimens in the collection of the Zoological Institute of the Academy of Sciences of the USSR. Insecta (Diptera). Part 1. Families Acroceridae, Nemestrinidae, Bombyliidae. Nauka, Leningrad [= St. Petersburg]. pp. 12 - 44. [in Russian.]"]}

Usia bicolor Macquart, 1855 (Figure 1, Plate III) Usia bicolor Macquart, 1855: 103 (83). Type material examined. HOLOTYPE: [Egypt] Usia bicolor ♀ [sic.] Macq./ U. bicolor Ex coll. Bigot. [seems to have been incorrectly sexed by Macquart. ♂ in OUMNH]. Other material examined. Israel, Tel Mareshna, 2km WNW Hebron, 357m, 17 March 1986, (leg. A. Shmida)/ at Ornithogalum tenuifolium (White) Liliaceae (OUMNH) ; Tel Aviv Country Club beach, 14 March 1995 ♂; Herzliyya, 11 March 1995 ♀; 14 March 1995 ♀ (leg. B. Merz); Shuneh Benjaminah, Coastal Plain, Palestine, 6 April 1953, O. Theodor [♂ 3 ♀]; dunes, Hederah, Coastal Plain, Palestine, 5 April 1953, O. Theodor [2 ♀]; Bet-Jibrin, Southern Palestine, 25 March 1954, O. Theodor [♂] (TAU) Herzliyya, N 32 �� 11 ��� E 34 �� 48 ���, 15 March 2008, 2♂ 5 ♀ (leg. A. Nir) (PCDG). Jordan, As Salt, 32 032N - 35 44 E, Al Balqa Gov., 13 April [20]02 [♀]; Aqraba, 32 43 N - 35 47 E, Irbid Gov., 25 March [20]02, on Linum pubescens Banks [♀] (BMNH ex Greathead coll.). Lebanon, Alma a Shaib, 3 April 1978, D. Gerling [♀] (TAU). Turkey, I��el, Gűzeloluk, N 36 �� 45 ���01.3��� E 34 ��07��� 47.3 ���, 1335 m, 26 May 2005, 3♂ 12 ♀; Hatay, Yunushani, Yayla Dagi, 600 m, 8 May 2000, 3♂ 2 ♀ (leg. J. Dils & J. Faes); Hatay, Belenpas E-side, 25 June 1992 1 ♂ (leg. J.P. Borrie); Hatay, 20 km N of Botash, 26 April 2000, 1♀ (leg. J.A.W. Lucas) (PCJD). Etymology. from Latin bi+color for ���two-coloured���, presumably referring to the bluish reflections on some specimens. Diagnosis. Variable in size, from small to quite large, one of the most frequently met with Usia in the eastern Mediterranean. An all black species, often with a faint bluish sheen and short, erect whitish vestiture. Occiput densely grey dusted, male eyes separated by distance across lateral ocelli, a series of distinct hairs bordering eye margins in front of anterior ocellus. Mesonotal vestiture short and neat, very even in length, Tibia similarly clothed with short, even hairs. Wings hayline with a brownish infuscation basally. Genitalia typical of the group differing only subtly from close congeners; apices of gonocoxite convergent, tips closely approximated or touching; furca with tip membranous. Redescription. Measurements. Body length. 2.5 ���9.0mm. Wing length. 2.5���6.5mm. Male. Head. Oral margin narrow, becoming narrower rearwards and disappearing below, shining black to brownish. Frons black, shining, conspicuously grey-dusted anterolaterally often distinctly longitudinally striated medially in front of depressed area in front of anterior ocellus. Eyes separated at their narrowest by about one fifth to one sixth head width, about equal to width across lateral ocelli. Lateral ocellus separated from eye margin by about half to two thirds diameter of that ocellus. Ocellar tubercle shining, undusted, eye margins in front of anterior ocellus diverging relatively evenly to rear of oral opening. Translucent white hairs on ocellar tubercle averaging width of frons, a few decidedly longer than width across lateral ocelli, rear of frons in front of anterior ocellus with distinct uniserial rows of proclinate setulae close to eye margins, about as long as scape, anterior part of frons with scattering of short setulae across most of width in places. Occiput densely grey dusted, black ground colour obscured except on triangular area behind ocellar tubercle, which is shining black. A fairly dense covering of pale yellowish hairs across dusted area, shorter, more even above, significantly longer close to lower oral margin. Ommatidia uniform in size. Antennae black, third segment, a little less than twice length of scape and pedicel together, narrow swollen below basally, concave medio-ventrally, sensilla inconspicuous, translucent brownish. All antennal segments with very short yellowish hairs above, longest just before apical sulcus. Palps small, black, apically, clavate tip a little narrower than oral margin, pale brown apical setae about half as long as length of palps. Proboscis shorter than head-thorax length, black, dorsally hairless. Thorax. Mesonotum shining black, sometimes with bluish reflections, grey dusting on anterior slope of mesonotum laterally, post-pronotal lobe, notopleuron and above wing root. Disc densely puncto-rugose, hairs issuing from confluent welts producing rough-textured, but still shining cuticle. Paramedian vittae usually just discernable by their different surface sculpture, more transversely wrinkled, extending from anterior slope of thorax almost to wing insertions. Mesonotum fairly densely clothed with fine whitish vestiture, on disc and posteriorly short, neat, erect to slightly reclinate, tending to proclinate close to scutellum, even in length, no hairs longer than those on upper occiput. Laterally hairs are longer and erect or inclined inwards (Figure 1). Acrostichal hairs between he bare paramedian vittae multiserial, entirely erect to slightly reclinate, widening posteriorly, dorsocentrals not distinguishable from the general vestiture. Scutellum shining as adjacent parts of mesonotum, hair length as on mesonotum or even shorter. Pleurae uniformly grey dusted largely obscuring ground colour, no areas of shining cuticle. Long white hairs, longer than any on mesonotum, on pronotum and upper and posterior half of anepisternum. Wing. Membrane brown tinged, this infuscation more intense in cells c, sc, base of r 1, and all but tip of first basal cell in most individuals, squama very pale yellow, veins all brown, darkest at base of C and Rs. Crossvein r-m at or a little before the middle of discal cell, conspicuously beyond dm-cu. Anal lobe moderately well developed, with evenly curved trailing edge, about two-thirds width of anal cell. Haltere. Yellow, base of stem slightly brownish. Legs. Coxae black with coating of grey dust as on pleurae. Remainder of legs black, undusted, shining below setulae. Coxae externally and femora ventrally covered with moderately long, white hairs, on mid and hind-femora a little more than greatest depth of tibia, shorter on front femora. Femora dorsally and tibia rather densely covered with short, recumbent, white hairs. Abdomen. All tergites shining aeneous black, often with bluish reflections, dusted on down-curved margins of first to third or fifth tergites. Each tergite densely puncto-rugose, surface sculpture of transverse welts each furnished with a hair, very narrow, smooth, shining apical margin. Disc uniformly and densely covered with very short, pale brownish, recumbent hairs, longer, whiter, more erect laterally. Sternites grey dusted, faintly punctate, with fairly long, erect white to very pale yellow hairs. Genitalia. Relatively small and simple, globular. Epandrium flaskshaped with apicolateral blunt extensions but lacking any median ���V���-notch, mostly shining black, cuticle punctate, evenly coated with pale yellow curved hairs. Gonocoxite about twice as broad as long, shining black basally, rufous brown apically, tips closely approximate or touching, haired as epandrium, gonostyli simple, curved with basal ���thumb���. Female. As male except as follows: oral margin decidedly broader, in lateral view visible to rear of oral opening. Frons a little wider, about one-fifth head width, lateral ocelli separated from eye margin by about their diameter or a little more. Dusting of frons more conspicuous and extensive, extending down between eye and oral margin. Mesonotal vestiture often slightly shorter, the paramedian vittae often obscure so acrostichal rows difficult to distinguish. Genitalia. Tip of abdomen coming to rounded point; apical sternite subrectangular, the basal part partially divided from main part, apical notch present but not deep, although often visible in undissected specimens. Main part of apical sternite with raised transverse ridge clothed with long whitish hairs. Furca arms very long, but tip poorly sclerotised, membranous so difficult to see, conspicuous, rectangular, dark-pigmented vaginal plate. Discussion. This common eastern Mediterranean species is very close to U. crinipes Becker and possibly conspecific, but the two are allopatric or parapatric and separated attitudinally. Also close to U. lata Loew, which shares the dusted occiput and epandrium lacking apical V-notch. Distribution. Egypt (where only known from the holotype collected by Bigot at Cairo (Macquart 1855) so perhaps rare but likely to be found in the Sinai), Iran, Israel (very common), Jordan, Lebanon, Turkey (coastal provinces from Mersin to Hatay). As far as can be ascertained from locality data, this is a lowland species, especially abundant on coastal dunes in Israel, but with occasional specimens up to 1335m in Turkey., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 26-29, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090

Usia putilla Becker, 1906 (Figure 12; Plate IX) Usia putilla Becker, 1906 b: 215, stat. rev. Type material examined. LECTOTYPE here designated; France, putilla, Lyon, Coll. Winthem / putilla det. Becker/ Type / Usia angustifrons B., Dr E.O. Engel det. [♂ in NMW]. PARALECTOTYPES; putilla, Lyon, Coll. Winthem / putilla det. Becker/ Type / Usia angustifrons Beck., Dr E.O. Engel det. [♂ in NMW]; 103 [on micropin]/ putilla, Lyon, Coll. Winthem / putilla det. Becker/ Type / Usia angustifrons B., Dr E.O. Engel det. [♀in NMW]; putilla, Lyon / putilla, Lyon, Coll. Winthem / putilla det. Becker/ Type / Usia angustifrons B., Dr E.O. Engel det. [♀ in NMW]. Originally described from an unspecified number of specimens of both sexes from Winthem���s collection, Vienna. Four syntypes were found in NMW, the male in best condition selected as lectotype. Other material examined. Spain, Cha��e (Segovia) 18.v��� 3.vii 2000, T. Malaise [♂] (leg. J.F. G��mez) (PCDG). France, Cavaliea [sic. = Cavalaire, Var], June [19]06, Museum Paris, ex coll. Hesse [♀ MNHN]. Etymology. from Latin putus, meaning pure or clear, + illa, a diminutive suffix; perhaps referring to its shininess and small size. Diagnosis. A small to very small shining black species, known only from a few specimens from France and Spain. Entirely black, vestiture fine, relatively sparse and silvery white, hairs erect or slightly inclined. Occiput shining, male frons narrow, eyes separated by less than distance across lateral ocelli, rear half of frons parallel, silvery dusting coalescing in front of anterior ocellus or, in females, dust spots separated. Acrostichal setulae proclinate in all specimens seen. Pleurae dusted apart from small shining dorsal and posterior margin on anepisternum in some individuals. Wings faintly yellow-grey tinged, distinctly yellow basally, veins yellow, becoming darker distally, anal lobe narrow but trailing edge clearly curved, more than half width of anal cell. Genitalia typical of group, differing only subtly from close congeners, most importantly in membranous apical margin of epandrium, which extends for more than half length of sclerite. Redescription. Measurements. Body length. 3.4���4.5mm. Wing length. 2.7���3.4mm. Male. Head. Mouth-margin very narrow, almost absent towards rear of oral opening, frons at narrowest point a little less than distance across lateral ocelli, equal to three times diameter of anterior ocellus, lateral ocellus separated from the eye margin by about its radius or less. Ocellar tubercle shining black, anterolateral silvery grey dust spots on frons extending back to narrowest part where they coalesce, a covering of fine pale setulae on anterior part of frons. Occiput shining black without any noticeable pollinosity except in the two vertical occipital tumescence but even here black ground colour not obscured. Hairing moderate length, on ocellar area about as long as width of vertex at narrowest point. Proboscis black, hairless, about as long as the head and thorax together; palps very short and thin, black to brown with short pale hairs apically; antennae black, third antennal segment longer than the scape and pedicel together, all segments shortly-haired above. Thorax. Mesonotum and scutellum shining black except for dorsum of post pronotal lobes, anteroventral corner of notopleuron and small areas above the wings. Shining between punctures, in some individuals these punctures rather stellate, bare paramedian lines relatively conspicuous and extending back to the mid point of the mesonotum, each a little broader than the acrostichal area which has 5���6 lines of proclinate hairs, most of hairs on disc of mesonotum also proclinate. Mesonotum covered with short white hairs, that on the disc no longer than distance between lateral ocelli, on scutellum about as long as vertex at level of lateral ocelli. Laterally hairing a little longer and denser, the hairs erect or inclined inwards. Pleurae black with a covering of grey dust sufficiently dense to obscure the black ground colour except for small shining patch along dorsal and posterior margin of anepisternum. Posterior half of anepisternum and pronotum with rather longer white hairs. Wing. Wing membrane clear, only faintly tinged yellowish-grey, becoming yellow basally, the veins also yellow basally becoming conspicuously darker brownish in the apical half. Anal lobe narrow, but with rounded margin, just a little narrower than anal cell (France) to a little over half width of anal cell (Spain, Figure 12), r-m a little before the middle of the discal cell. Haltere. Whitish yellow, slightly infuscated at base of stem. Legs. Entirely black, the coxae grey dusted as in pleurae; femora ventrally with mid-length, pale hairs, up to length of width of tibia, tibia and tarsi with a covering of short, white setulae, longer on tibia. Abdomen. Shining aeneous black, the disc of the tergites covered with short, whitish hairs issuing from transverse welts. Laterally and on the down-curved margins the tergites are clothed with rather longer, whitish hairs. Middle tergites with a shining, impunctate apical margin often with longitudinal grooves. Sternites grey dusted with whitish hairs confined to the posterior margin. Genitalia. Relatively large and conspicuous for this species group, few useful details visible in pinned specimens, shining black and covered in whitish hairs. Most useful character is the membranous apical ���V��� notch of the epandrium which extends for more than half of the sclerite. Female. As male except as follows: broader, though still very narrow, oral margin, and conspicuously wider frons, lateral ocellus separated the eye margin by about diameter of that ocellus. Posterior half of frons shining black, parallel sided, about four times diameter of anterior ocellus, as wide or wider than length of longest hairs on ocellar tubercle. Anterior half of frons widening, lateral margins each side covered in dense whitish dust obscuring ground colour, central two-thirds shining black or dark brown. Third antennal segment sometimes a little shorter. Anal lobe broader, margin more convex, as broad as anal cell. Abdomen conical tapering to rounded point. Discussion. This small shining black species is clearly very closely allied to U. manca Loew, and could possibly be the end of a cline of this species. If specimens come to light between north Italy and central south France that show intermediate characters then U. putilla Becker might need synonymising under U. manca. However, in the absence of such specimens the differences, though small, are sufficient to retain it as a full species. Engel (1932) treated this taxon as a subspecies of U. angustifrons Becker and he wrote that he could understand why Becker described it again under the name U. putilla. If Engel had the types of U. putilla in front of him then his decision is inexplicable, even without dissection these two species are abundantly distinct. Perhaps Engel did not have the U. putilla types available, in which case he might have worked from Becker���s description alone. Becker (1906 b) did list some useless features in his key (e.g., colour of wing membrane and position of vein r-m), but the width of the frons and nature of occiput, accurately described by Becker, should have been sufficient to convince Engel he was dealing with a distinct taxon. In his description Engel (1932) mentioned several characters (entirely dusted lower frons and dusted occiput) that do not fit Becker���s description of U. putilla. Distribution. A rare and little known species so far only known from the Rhone valley near Lyon, and the Mediterranean coast, southern France and a single specimen from central Spain., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 42-45, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Becker, T. (1906 a) Die Ergebnisse meiner dipterologischen Fruhjahrsreise nach Algier und Tunis 1906 [part]. Zeitschrift fur Systematische Hymenopterologie und Dipterologie, 6, 1 - 16.","Becker, T. (1906 b) Usia Latr. Berliner Entomologische Zeitschrift, 50 (1905), 193 - 228.","Engel, E. O. (1932) Bombyliidae. In: Lindner, E., (Ed.), Die Fliegen der palaarktischen Region. Vol. 4. pt. 3. E. Schweizerbart, Stuttgart, pp. 49 - 96."]}

Usia greatheadi Gibbs sp. nov. (Figure 6, Plate VI) Type material examined. HOLOTYPE; Turkey, 5 June 2005, I��el, Camalan, 663m, N 37 �� 11 ��� 33.1 ��� E 34 �� 48 ��� 33.9 ���, Leg. Dils J. Faes J. [♂ in NMWC]. PARATYPES; Greece, Lesvos, Ag. Georious, S of Mytilini/ 24 April 2001, [leg.] D.J. Greathead [6 ♂ 5 ♀ in BMNH]. Turkey, 5 June 2005, I��el, Camalan, 663m, N 37 �� 11 ��� 33.1 ��� E 34 �� 48 ��� 33.9 ���, Leg. Dils J. Faes J. [2 ♂ 6 ♀ in PCJD, 1 ♀ in NMWC]; 29 April 2001, Mulğa, Mumcular (Karaova), 200m, N 37 �� 4 ��� 9 ��� E 27 �� 40 ��� 54 ���, Leg. J. Dils-J. Faes [8 ♂ 6 ♀ in PCJD, 1 ♀ NMWC, 1 ♂ in OUMNH]; 22 April 2001, Mulğa, Reşadiye Yarimadasi, N 36 �� 45 ��� 51 ��� E 27 �� 45 ��� 59 ���, Leg. J. Dils-J. Faes [12 ♂ 6 ♀ in PCJD, 1 ♀ in NMWC]; 24 April 2001, Mulğa 50m, Yuvacik, N 36 �� 49 ��� 41 ��� E 28 �� 11 ��� 27 ���, Leg. J. Dils-J. Faes [4 ♂ 2 ♀ in PCDG, 1 ♀ in NMWC]; 6 June 2005, Adana, Pozanti, 656m, N 37 �� 23 ��� 51 ��� E 34 �� 51 ��� 20.3 ���, Leg. Dils J.��Faes J. [6 ♀ in PCJD, 1 ♀in NMWC, 1 ♀ in OUMNH]; Antalya Prov., Olympos, 25km S Kemer, 26 April 2000, 0m, leg. Merz & Senay [♂ in MHNG]; Cavusk��y (Andrasan), 28 April 2000, 50m, leg. Merz & Senay [1 ♂ 2 ♀ in MHNG]; Kursunlu Selalesi, 29 April 2000, 150m, leg. Merz & Senay [5 ♂ 1 ♀ in MHNG]; Phaselis, 10km S Kemer, 27 April 2000, 0m, leg. Merz & Senay [♂ in MHNG]; Mulğa, Sazak-Otmanlar, 1050m, N 37 ��0��� 13 ��� E 28 �� 55 ��� 6 ���, 11 June 2005, leg. D.J. Gibbs [2 ♂ 3 ♀ in PCDG]; Road 635 between G��ltarla and Calallar, N 36 �� 31 ��� 35 ��� E 30 ��00��� 39 ���, 850m, 10 May 2001, leg. D. Mann & Barclay [1 ♂ in OUMNH]; Rt. 685, 30km N Antalya, 400m, 5 May 2000, A. Freidberg, H. Ackerman, L. Friedman [2 ♀ in TAU]. Other material examined. Greece, Lesvos, 10 km S Mytilene, 100m N 39 ��0��� 51 ��� E 26 �� 35 ��� 26 ��� Poor phrygana simple landscape/ blue pantrap#4, 23��� 25 April 2004 (leg. Dahm/Greenwell/Lamborn/Messinger)/ Aegean University 0 0 0 6721 1 ♂; 0 0 0 6747, 1♂; 23 April 2004 10.45���11.15 (leg. M. Greenwell)/ Aegean University 0 0 14073 1 ♀; 9.1 km S Mytilene, 130m 39 ��02' 19 "N 26 �� 35 ' 58 "E, Phryganic habitat complex landscape/ blue pantrap#5, 11��� 13 May 2004 (leg. Dahm/Greenwell/Lamborn/Messinger)/ Aegean University 0 0 0 7005 1 ♂; white pantrap#2, 16��� 18 May 2004 (leg. Dahm/Lamborn/Messinger)/ Aegean University 0 0 0 4534, 1♀; 8.9 km SSE Mytilene, 60m, N 39 ��02��� 16 ��� E 26 �� 36 ��� 13 ���, Olive grove complex landscape/ white pantrap#2, 16��� 18 April 2004 (leg. Dahm/Greenwell/Lamborn/Messinger)/ Aegean University 0 0 0 2842 1 ♂. Etymology. Named in memory of David John Greathead who helped me considerably in getting started with this revision. Diagnosis. A relatively small blue-black species that appears to be fairly common in western Turkey and on Lesvos. An all black species, often with bluish and coppery reflections and short, erect whitish vestiture. Occiput densely grey dusted, including triangle behind ocellar tubercle, oral margin very narrow, in male almost invisible, frons narrow, male eyes separated by no more than distance across lateral ocelli, about one seventh head width. Anterolateral dust spots extending almost to level of anterior ocellus, in male coalescing. Mesonotal vestiture short and neat, very even in length. Wings brown tinged, most intense in first basal cell, both veins and wing yellower basally. Genitalia typical of the group differing only subtly from close congeners; epandrium with apical membranous ���V���. Description. Measurements. Body length. 3.5���5.7mm. Wing length. 2.9���4.5mm. Male. Head. Oral margin exceedingly narrow, barely discernable even close to insertion of proboscis, shining black above, yellow below. Frons black, rather matt due to rough surface sculpture, conspicuously grey-dusted, the dusting extending almost to anterior ocellus, the area in front of anterior ocellus dusted across whole width of frons (Figure 6). Eyes separated at their narrowest by about one seventh head width, no more than the width across lateral ocelli inclusive. Lateral ocellus separated from the eye margin by less than half the diameter of that ocellus. Ocellar tubercle shining, undusted, eye margins in front of anterior ocellus parallel initially before diverging relatively rapidly, then running parallel to insertion of proboscis before diverging rapidly to rear of oral opening. Translucent white hairs on ocellar tubercle averaging a little longer than width across lateral ocelli, rear of frons forward of anterior ocellus often with tiny, adpressed setulae close to eye margins, absent or invisible in some males, front of frons with a scattering of short setulae across most of width. Occiput grey dusted, the black ground colour just showing through, including on the triangular area behind ocellar tubercle. A fairly dense covering of translucent white hairs across dusted area, shorter and more even above, significantly longer close to lower oral margin. Ommatidia uniform in size across the eyes. Antennae black, third segment, a little less than twice length of scape and pedicel together, narrow swollen below basally, concave medio-ventrally, sensilla inconspicuous, white to translucent brownish. All antennal segments with very short yellowish hairs above, longest just before apical sulcus. Palps small, black, apically, clavate tip noticeably broader than the very narrow oral margin, the pale brown apical setae about two thirds as long as the length of the palps. Proboscis about as long as thorax and scutellum together, black, dorsally hairless. Thorax. Mesonotum shining black, often with bluish and coppery reflections, grey dusting on upper part of post-pronotal lobe, notopleuron narrowly and above wing root. Disc densely punctorugose, anterolaterally the individual punctures often discernable and separated by about their diameter. On disc surface sculpture shagreened and transversely wrinkled effacing the individual punctures. Paramedian vittae usually discernable by their different surface sculpture, more strongly transversely wrinkled, extending from anterior slope of thorax almost to wing insertions. Mesonotum fairly densely clothed with fine whitish vestiture, on the disc and posteriorly these are short, neat, erect to slightly proclinate, longer and orientated inwards laterally, most hairs little longer than those on upper occiput. Acrostichal hairs between the bare paramedian vittae multiserial, entirely erect to slightly proclinate, the front most more reclinate, widening posteriorly, dorsocentrals not distinguishable from the general vestiture. Scutellum shining as adjacent parts of mesonotum, hairing as mesonotum. Pleurae uniformly grey dusted largely obscuring ground colour, often thinner in upper front corner of anepisternum. White hairs, similar to those on mesonotum, on pronotum and upper and posterior corner of the anepisternum. Wing. Membrane brown tinged, this infuscation more intense in first basal cell in most individuals, the veins brown, but sc yellower and all veins proximal to basal cells yellow, membrane here also yellow tinged, squama very pale yellow. Crossvein r-m at or a little before the middle of discal cell, conspicuously beyond dm-cu. Anal lobe relatively narrow, trailing edge only slightly curved, about half to two-thirds width of anal cell. Haltere. Pale yellow, base of stem slightly brownish. Legs. Coxae black with a coating of grey dust like that on the pleurae. Remainder of legs black, undusted, shining below the setulae. Coxae externally and femora ventrally covered with moderately long, white hairs, on the mid and hind femora a little more than the greatest depth of the tibia, slightly shorter on front femora. Femora dorsally and tibia rather densely covered with short, recumbent, white hairs. Abdomen. All tergites shining aeneous black, often with bluish, greenish or coppery reflections, undusted. Each tergite densely puncto-rugose, the punctures seriate, shining apical margin with longitudinal groves. Disc uniformly and densely covered with very short, whitish, recumbent hairs, longer, and more erect laterally. Sternites grey dusted, faintly punctate, with fairly long, erect white to very pale yellow hairs. Genitalia. Relatively small and simple, globular. Epandrium flask-shaped with apicolateral blunt extensions and a distinct median ���V���-notch, in undissected specimens this can often be discerned as an apicomedian crease. Shining black, the cuticle punctate evenly coated with pale yellow hairs. Gonocoxite about twice as broad as long, shining black basally, rufous brown apically, the tips well separated, haired as epandrium, gonostylus simple, curved with basal ���thumb���. Female. As male except as follows: oral margin broader, although still narrower than most species. Frons appreciably wider, about one-sixth head width, lateral ocelli separated from eye margin by about their diameter. Dusting of frons extending up to level of anterior ocellus but not coalescing, centre of frons shining black. Lateral setulae on frons in front of anterior ocellus more numerous and conspicuous but still short. Mesonotal and scutellar vestiture often slightly shorter. Genitalia. Tip of abdomen coming to a rounded point; apical sternite rhomboidal, apical notch present but small, difficulty to see in undissected specimens. Main part of apical sternite domed and transversely wrinkled clothed with long whitish hairs. Furca typical of the group, tip sclerotised but variable in structure, pointed, blunt-ended or partially membranous, arms long and strongly curved, conspicuous triangular, dark-pigmented vaginal plate. Discussion. A distinct species, externally very similar to U. bicolor Macquart but with male genitalia closer to U. manca Loew. Probably not very closely related to either but clearly part of the lata species group. Distribution. Greece (Lesvos), western half of Turkey, from sea level to just over 1000m., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 33-36, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090

Usia notata Loew, 1873 (Figure 15, Plate XXI) Usia notata Loew, 1873: 201. Type material examined. LECTOTYPE here designated; Uzbekistan, Кизилъкумъ [= Kizil Kum]/ 9692 / Usia notata Lw / Coll. H. Loew/ Type / Zool. Mus. Berlin [♀ ZMHB]; PARALECTOTYPE; Kazakhstan, Узуната [= Uzunata]/ Coll. H. Loew/ Typus / Zool. Mus. Berlin [♀ ZMHB]. Originally described from an unspecified number of specimens. Two syntypes were found in ZMHB, both females, the specimen in best condition being selected as lectotype. Other material examined. Turkmenistan, Bairam-Ali, 28 April 1934, leg. N. Shibanov [♀] (ZMUM). Etymology. from Latin notatus, meaning ���marking���, probably referring to the brown spot in the middle of the wings. Diagnosis. A small to medium-sized Usia of central Asia. Mostly black with a faint greenish hue, immediately distinguished from all known species by the yellow anterior half of the frons, sharply demarcated from the shining black hind half. Occiput mostly shining, frons remarkably short and relatively wide, proboscis much shorter than other species, third antennal long. Mesonotal vestiture short, white, marginal scutellar hairs less than half midlength of scutellum. Knees narrowly but distinctly yellow, tergites all dark, sternites all pale. Wings clear but for a brownish infuscation over middle of first basal cell. Redescription. Measurements. Body length. 2.4 ���5.0mm. Wing length. 2.2���5.5mm. Male. Material not seen, but Paramonov (1950) stated ���males extremely similar to females���. Female. Head. Noticeably small and short, frons barely longer than width at antennae. Mouth-margin strikingly broad, clearly visible from directly above, convex, about as broad as length of pedicel; shining, undusted. Anterior half of frons clear yellow, the frons laterally with silvery dust spots leaving central thirds shining undusted. The yellow front of the frons sharply demarcated from the shining black hind half, ocellar tubercle and occiput, latter only dusted on occipital ���cushions���. Frons narrowest level with hind corners of eyes, broadening anteriorly, lateral ocellus separated from eye margin by about the diameter of that ocellus. Vestiture very short, on ocellar area no longer than distance between lateral ocelli; on black part of frons a group of short, forwardly curved white hairs occupy lateral quarters. Antennae relatively long, scape brown about a little longer than wide, pedicel blackish, brown ventrally, quadrate, third segment black, very narrowly brownish as base, elongate, narrow, longer than width of frons. Proboscis black, hairless, shorter than most species of Usia, about 1.5 ���2.0 times the length of the head; palps short, dark brown, clearly clavate with pale hairs, the longest almost as long as the palps. Thorax. Broad relative to head (as in U. lata Loew), mesonotum and scutellum shining black except for light dusting narrowly on notopleuron and above wing base. Surface conspicuously punctate, although punctures small and simple, separated by their diameter or a little more, with a tendency towards transverse wrinkles in hind part, scutellum decidedly transversely wrinkled. Bare paramedian lines inconspicuous, apparent only on anterior slope of mesonotum. Pleurae predominantly black with a dense covering of grey dust largely obscuring the ground colour; area around anterior spiracle yellow, hind half of anepimeron dull brown to yellow below dust. Mesonotum and scutellum covered with short, arcuate pale yellowish or white hairs, those on disc orientated towards head, those laterally a little longer and orientated towards disc of mesonotum, scutellar hairing similar. Anepisternum and pronotum with hairing similar to that on notopleuron, covering whole sclerite. Wing. Wing membrane clear, only faintly yellow tinged but for a distinct, diffuse brown spot in the first basal cell above the base of the discal cell, spreading to the fork of veins R 2 + 3 and R 4 + 5. Veins yellow basally, beyond R 2 + 3 ���R 4 + 5 fork R and M veins plus r-m are darker and browner. Anal lobe as broad as anal cell, margin convex; r-m just before the middle of the discal cell. Haltere. Yellow, becoming slightly infuscated at base of stem. Legs. Predominantly shining black, tips of femora and bases of tibia narrowly yellow-brown. Coxae grey dusted and haired as on anepisternum; femora with relatively long, pale hairs, the longest as long as the depth of the femora medially. Tibia and tarsi with a covering of minute pale setulae, a little longer and more erect on tibia, especially basally. Abdomen. Shining aeneous black, the disc of the tergites covered with very short, pale yellow hairs, appearing naked at low power. Laterally and on the down-curved margins the tergites are clothed with rather longer, whitish yellow hairs. Sternites mostly pale yellow with inconspicuous short pale hairs, last visible sternite darker brownish. Genitalia. Furca pigmented, apical part rather robustly constructed, arms long and strongly curved; apical sternite wider than long, brownish, darker than foregoing sternites. Discussion. A very distinct species with no obvious affinities with any other known species. The very short head and proboscis, long third antennal segment, very broad oral margin and pattern of yellow on head are not found in any other species. The female genitalia also do not suggest any close relationship to other species. Distribution. Only three specimens seen, one each from Kazakhstan, Turkmenistan, Uzbekistan. Records from Tajikistan are likely to be correct. The area is certainly very under-recorded, but even so the scarcity of specimens suggests that it is far from being a common species., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 76-77, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Loew, H. (1873) Systematische Beschreibung der bekannten europaischen zweiflugeligen Insecten. Von Johann Wilhelm Meigen. Zehnter Theil oder vierter Supplementband. Beschreibungen europaischer Dipteren. Dritter Band. H. W. Schmidt, Halle, viii + 320 pp.","Paramonov, S. J. (1950) Bestimmungstabelle der Usia - Arten der Welt (Bombyliidae, Diptera). Eos, 26, 341 - 78."]}

Usia angustifrons Becker, 1906 a (Figure 9, Plate XXII) Usia angustifrons Becker, 1906 a: 14 [1906 b: 212]. Type material of U. angustifrons examined. LECTOTYPE here designated; Tunisia, V [May], 52720 / angustifrons Beck / Typus / Zool. Mus. Berlin [♂] (ZMHB). PARALECTOTYPES; Tunisia, V [May], 52720 / angustifrons Beck / Typus / Zool. Mus. Berlin [1 ♂ 1 ♀] (ZMHB). Originally described from an unspecified number of specimens of both sexes. Three syntypes were found in ZMHB, the male in best condition was selected as lectotype. Other material examined. Tunisia, 10km E. of Kairoun, 5 May 1975, leg. J.W. Ismay 2 ♂ 3 ♀ (OUMNH). Etymology. From Latin angustus meaning ���narrow��� + frons meaning ���forehead���, ���brow��� or ���front���. Diagnosis. Small shining black species known only from Tunisia. Entirely black, sometimes with faint gold and green reflections, especially in females, vestiture fine, white, acrostichals erect or reclinate. Occiput shining along narrow strip just behind upper eye margin, otherwise thinly dusted. Male eyes separated by diameter of anterior ocellus, more closely approximate than any other Usia species, front half of frons entirely grey dusted obscuring ground colour. Female frons parallel in posterior half, abruptly diverging in anterior half, grey dust spots broad, central undusted strip narrower than an individual dust spot. Longest scutellar vestiture about half scutellar mid-length but variable, longest hairs can be few or missing through damage. Pleurae entirely grey dusted. Wings clear, anal lobe very narrow, a little more convex than in female but still narrower than anal cell in female. Redescription. Measurements. Body length. 6.5 ���9.0mm. Wing length. 6���7mm. Male. Head. Mouth-margin very narrow, almost absent, dark brown or black. Frons at narrowest point narrower than any other Usia species, no wider than the diameter of the anterior ocellus, lateral ocellus almost touching the eye margin. Ocellar tubercle shining black, frons at narrowest point dull but still black, anterior part of frons entirely pale grey dusted obscuring dark ground colour. Occiput black, shining, undusted on a narrow band behind upper eye margin, otherwise with a thin covering of pale dust that does not obscure ground colour, triangular area behind vertex more thinly dusted. Ommatidia all of uniform size. Hairing moderate length, on ocellar area about as long or a little longer than width of vertex level with lateral ocelli, frons glabrous. Antennae black, third segment longer than the scape and pedicel together, all segments with very short setae above, sensilla small, brown or pale. Proboscis black, hairless, about as long as the head and thorax together; palps very short and thin, black to brown with short pale hairs apically;. Thorax. Mesonotum and scutellum shining black except anterior slope of between paramedian lines post pronotal lobes, dorsum of post pronotal lobes, most of notopleuron and above wings which is grey dusted. Disc shining except for relatively simple punctures, many separated by areas of smooth cuticle greater than diameter of a puncture. Bare paramedian lines relatively conspicuous, extending back to beyond the mid point of the mesonotum, each about as broad as the midline which has 5���6 lines of hairs. Pleurae with a covering of grey dust sufficiently dense to obscure the black ground colour. Mesonotum and scutellum covered with mid-length pale yellowish white hair, that on the disc of the mesonotum about as long as the width of the vertex at the lateral ocelli. Pronotum and posterior edge of anepisternum with a scattering of white hairs. Wing. Wing membrane clear, becoming yellow basally and in costal cell, the veins also yellow basally becoming conspicuously darker brownish black in the apical half. Base of stem of R darker brown, base of C blackish. Anal lobe narrow, almost linear, conspicuously narrower than anal cell, r-m a little beyond the middle of the discal cell. Haltere. Yellow, becoming brown at base of stem. Legs. Entirely black, the coxae grey dusted as in pleurae; femora with rather short, pale hairs, the longest not exceeding the greatest depth of the tibia, tibia and tarsi with only a covering of short setulae, pale on tibia, browner on tarsi. Abdomen. Shining aeneous black, the disc of the tergites covered with short, whitish hairs, the hairs arising from transverse welts forming a rugose surface sculpture. Laterally and on the down-curved margins the tergites are clothed with rather longer, whitish hairs. Sternites grey dusted with whitish hairs confined to the posterior margin. Genitalia. Fairly small and partially enclosed in apical tergites, few useful details visible in pinned specimens, shining black, tip of gonocoxite often browner, covered in yellowish hairs, longest on the gonocoxite. Female. As male except as follows: broader, thought still linear, oral margin and the conspicuously wider frons, the lateral ocellus separated from the eye margin by about the diameter of that ocellus. Posterior half of frons shining black, parallel sided, about four times the diameter of the anterior ocellus and as wide or wider than the length of the longest hairs on the ocellar tubercle, a few tiny, proclinate hairs close to eyes forward of anterior ocellus. Anterior half of frons widening rather abruptly, more than lateral third each side covered in dense whitish dust obscuring ground colour, central shining black strip narrower than dusted areas. Third antennal segment sometimes shorter. Anal lobe broader, margin more convex, almost as broad as the anal cell. Abdomen conical tapering to a rounded point, apical sternite evenly convex, wider than long, apical emargination small and shallow, in dry specimen a square spot of pale dust discernable apicomedially. Discussion. The affinities of this small African species are not at all obvious. It has a pattern of dusting on the occiput similar to some of the U. florea group but is otherwise quite dissimilar. The narrow anal lobe and form of the gonostyli are reminiscent of some of the U. bicolor group but the epiphallus and female genitalia do not conform to that group. It does not seem to be closely related to any known species. Distribution. Only known from Tunisia. Specimens are few, so it appears to be rare and local., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 77-80, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Becker, T. (1906 a) Die Ergebnisse meiner dipterologischen Fruhjahrsreise nach Algier und Tunis 1906 [part]. Zeitschrift fur Systematische Hymenopterologie und Dipterologie, 6, 1 - 16.","Becker, T. (1906 b) Usia Latr. Berliner Entomologische Zeitschrift, 50 (1905), 193 - 228."]}

Key to genera of Usiini Becker 1. Disc of mesonotum largely obscured by dust and hairs on abdomen pale (white to yellow). If with shining mesonotal vittae (which, in U. loewi, occasionally coalesce such that disc is largely devoid of dust) abdomen with yellow apical margins, disc of tergites matt, laterally densely grey-dusted [all holoptic species and all but one with yellow apical margins to tergites belong here]................................................................................................ 2 - Either disc of mesonotum shining and undusted and tergites shining largely undusted or abdominal hairs black (if with yellow apical margins to tergites, U. transcaspica, tergites, except first, shining, undusted, even laterally)...................... 3 2. Anal lobe well developed, at least as wide as anal cell; entirely pale-haired species (white to yellow). Grey, brown or blackish dusted species, abdomen matt usually with pale apical margins to tergites. Furca variable, often heavily sclerotised........................................................................................... Parageron Paramonov - Anal lobe narrow, linear in ♂, almost linear in ♀, narrower than anal cell; with some dark brown hairs on mesonotum and scutellum (these can fade). Olive-grey and brown-dusted but rather thinly so subshining (somewhat variable). Epandrium with shining patches. Furca V-shaped folded over at distinct ���seam���. Genitalia (see Gibbs 2011 pl I).............................................................................................. Usia martini Fran��ois (see Gibbs 2011) 3. All hairs on head, thorax and abdomen clearly black............................................................................................. Usia subgenus Micrusia Evenhuis and Usia versicolor (Fabricius) (see Gibbs 2011) - At least hairs on down-curved lateral margins of tergites pale yellowish (only one species with dark vestiture, U. aeneoides, in which the dark mesonotal hairs are very short spicules)............................................. Usia Latreille, Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 7-8, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Gibbs, D. (2011) A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 1: Usia subgenus Micrusia (= blackhaired species) and Usia martini Francois. Zootaxa, 2960, 1 - 77."]}

Gibbs, David (2014): A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto. Zootaxa 3799 (1): 1-85, DOI: http://dx.doi.org/10.11646/zootaxa.3799.1.1

Usia annetteae Gibbs sp. nov. (Plate II) Type material examined. HOLOTYPE; Lebanon, 25km SE of Tripoli, Horsh Ehden Nature Reserve /, Alt. 1300���1600 m, 29 May��� 3 June 2001, J.C. Deeming [♂ NMWC]. PARATYPES; Israel, Mt. Hermon, 1600 m, Israel, 28 May 1980, leg. Kugler [♀ TAU]; 600 m, Hermon, 9 June 1975, F. Kaplan [♂ TAU]; 1800 m, Mt. Hermon, 12 June [19] 96, Merz & Freidberg leg. [1 ♂ 3 ♀ MHNG]; 1600 m, Mt. Hermon, 29 May 2000, leg. B. Merz [2 ♂ MHNG]; Mt. Hermon, Astra, 9 July 1987, A. Freidberg [♂ TAU]; Mt. Shalhavit, 1900 m, 9 July 1987, I. Nussbaum [2 ♂ 1 ♀ TAU]; Hermon Mt., 1750m, 8 July 1987, I. Naussbaum [♂ TAU]; Hermon, 1600 m, 23 May 1994, F. Kaplan & A. Freidberg [♂ TAU]; 1800 m, Har Hermon, 12 June 2003, T. Stern [♂ TAU]; 1800m, Har Hermon, 11 & 12 June 2003, A. Freidberg [1 ♂ 3 ♀ TAU]. Lebanon, 25km SE of Tripoli, Horsh Ehden Nature Reserve/ Alt. 1300���1600 m, 29 May��� 3 June 2001, J.C. Deeming [♂♀, NMWC]; Horsh Ehden NNR, Mixed forest 1500 m, 29 May��� 1 June 2001, [leg.] P. Gatt [2 ♂ PCME]. Etymology. Named in honour of Annette Greathead, who generously allowed me access to the literature and collection of the late David Greathead. Diagnosis. A relatively small shining black species but very variable in size, known only from Lebanon and the Golan Heights on the Israli/Syrian border (disputed territory. Entirely black, lacking metallic reflections, vestiture relatively sparse, fine and white so appearing a rather naked fly. Occiput shining, male eyes separated by the distance across lateral ocelli, mesonotal vestiture orientated inwardly or forward, but the anterior acrostichal reclinate. Wings brown-tinged with anal lobe a little narrower than anal cell. Genitalia typical of the group differing only subtly from close congeners. Description. Measurements. Body length. 3.0��� 6.4mm. Wing length. 2.5���5.1mm. Male. Head. Oral margin very narrow, almost linear, almost disappearing below, shining blackish to vaguely brown. Frons black, shining, with conspicuous linear silvery dust spots laterally on anterior half, depressed in front of anterior ocellus where slightly more rugose. Eyes separated at their narrowest by less than one sixth head width, equal to width across lateral ocelli. Lateral ocellus separated from eye margin by about diameter of that ocellus. Ocellar tubercle shining, undusted, eye margins in front of anterior ocellus diverging relatively evenly to rear of oral opening. Translucent white hairs on ocellar tubercle averaging width of frons, rear of frons in front of anterior ocellus with a few inconspicuous, pale, proclinate hairs laterally, anterior part of frons with denser hairing across most of width. Occiput shining black with grey dusting confined to occipital callosities, especially on external side, pale hairs, shorter and more even above. Ommatidia uniform in size across eyes. Antennae black, third segment, a little less than twice length of scape and pedicel together, narrow and parallel sided or slightly swollen basoventrally, sensilla inconspicuous, translucent pale brownish. All antennal segments with very short whitish hairs above, longest just before apical sulcus. Palps small, black, apically about as broad as oral margin at widest, pale apical setae about two-thirds length of palps. Proboscis shorter than head-thorax length, black, dorsally hairless. Thorax. Mesonotum shining black, grey dusting on dorsum of post pronotal lobe and above wing root. Disc relatively sparsely punctured, many areas with smooth shining cuticle, the punctures separated by distancesignificantly wider than a puncture, anterior slope of thorax and anterior part of paramedian vittae entirely impunctate, latter shallowly transversely wrinkled. Paramedian vittae extending from anterior slope of thorax to about half way to wing insertions. Mesonotum relatively sparsely clothed with pale whitish vestiture, in places very faintly yellow tinged. All hairs nearly perpendicular, those on disc and posteriorly slightly leaning forward. Laterally hairs longer, inclined towards middle of mesonotum. Acrostichal hairs between bare paramedian vittae in 4���5 irregular lines, posterior ones proclinate, anterior ones reclinate, dorsocentrals indistinguishable from the general vestiture. Scutellum shining as adjacent parts of mesonotum, erect-hairs on disc as dense as on rear of mesonotum. Pleurae grey dusted except for ventral half of katepisternum and all but lower hind corner of anepisternum, which are polished black. Fairly long white hairs on pronotum and posterior half of anepisternum. Wing. Membrane with brownish tinge often a bit more intense in middle of first basal cell, squama very pale yellow, veins yellow basally, browner in apical half. Crossvein r-m at or slightly before middle of discal cell, conspicuously beyond dm-cu. Anal lobe broad, with evenly curved trailing edge, a little narrower than anal cell. Haltere. Pale yellow, base of stem slightly brownish. Legs. Coxae black with coating of grey dust like that on pleurae. Remainder of legs black, undusted, shining below setulae. Coxae externally and femora ventrally covered with moderately long, white hairs, on femora a little more than greatest depth of tibia. Femora dorsally and tibia rather densely covered with short, recumbent, white hairs. Abdomen. All tergites shining black, undusted, each densely puncto-rugose, apical margin a little less densely punctate, instead more shining. Disc uniformly and densely covered with pale brownish, recumbent hairs, longer paler and more erect laterally. Sternites grey dusted basally, more shining apically, faintly punctate, with fairly long, erect white to very pale yellow hairs. Genitalia. Relatively small and simple, globular. Epandrium flask-shaped with apicolateral blunt extensions and median ���V���-notch extending half length of epandrium, shining black, cuticle punctate, evenly coated with pale yellow erect hairs. Gonocoxite about twice as broad as long, shining black, haired as epandrium, gonostylus simple, curved with basal ���thumb���. Female. As male except as follows: frons wider, about one-fifth head width, diverging a little more anteriorly, lateral ocelli separated from eye margin by twice their diameter. Dusting of frons more conspicuous and extensive. Oral margin a little wider, especially medially and with a greater tendency to be brownish. Occipital usually with some dusting above neck. Genitalia. Tip of abdomen coming to a rounded point; apical sternite almost square, with an apical notch, in undissected specimens it can be seen to be apically silvery dusted. Furca typical of the group, with pointed tip and with long arms, conspicuous dark-pigmented vaginal plate. Discussion. This species is very similar to the two European species U. manca Loew and U. putilla Becker, also small, shining black species with an undusted occiput. These three species have very similar male and female genitalia and are clearly sibling species. Indeed if it were not for the intercession of manca between them, then putilla and annetteae would probably be better considered conspecific at opposite ends of a cline. However, manca differs from both in having a linear anal lobe to the wing. So far there is no evidence that any of these species occur in either the Balkans or Turkey so annetteae is very widely separated from manca in Italy and putilla in France and Spain. Distribution. Northern Israel (Golan Heights), Lebanon from 600 to 1900m., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 24-26, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090

florea species group Included species: U. accola Becker, 1906 U. atrata (Fabricius, 1798) U. maghrebensis Gibbs sp. nov. U. cornigera Gibbs sp. nov. U. florea (Fabricius, 1794) U. vestita Macquart, 1846 Diagnosis. All are very similar in external appearance, averaging larger than members of the lata group and relitively robust. Integument black with a green or aeneous shine often apparent. All species with at least the upper part of the occiput shining, undusted. Anal lobe broad with strongly curved trailing edge. Although male genitalia are quite different between species, all have relatively small basal and lateral ejaculatory apodemes compared to the rest of the aedeagus. Female terminalia all have a heavily sclerotised apical sternite, longer than wide with relatively elongated apical processes, thus producing a deep cleft between them. Discussion.. This group encompasses six species that represent a radiation confined to the Maghreb, North Africa. All appear to be closely related, differing externally mainly in the length of the vestiture, from very short (U. accola) to exceptionally long (U. vestita). There is also some variation in the degree of brownish infuscation of the wings, two species being distinctly infuscated, one slightly, the remainder yellow tinged. However, the genitalia are abundantly distinct almost all parts showing remarkable modifications in one or more species. Female genitalia are also diverse in form, the furca particularly varying from entirely sclerotised and pigmented to almost entirely membranous and transparent., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 47-48, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Becker, T. (1906 a) Die Ergebnisse meiner dipterologischen Fruhjahrsreise nach Algier und Tunis 1906 [part]. Zeitschrift fur Systematische Hymenopterologie und Dipterologie, 6, 1 - 16."]}

Usia transcaspica Paramonov, 1950 (Plate X) Usia transcaspica Paramonov, 1950: 351. Type material examined. HOLOTYPE; Turkmenistan, Transcaspia mont, Kopet-dag Tshulli, 21���25 May 1913, A. Hohlbeck / 1104 / Usia transcaspica sp. nov. S. Paramonov det. ♀ / Holotypus / Zool. Mus. Berlin [♀ in ZMHB]. Other material examined. Turkey, Adiyaman, Nemrutdagi, 1500���2100m, 1 June 1985, ♂ (leg. H. v. Oorschot) (PCJD); Armenia, 7km N Armash, 2 July 1959, ♀ (leg. L. Zimina) (ZMUM); Iran, Ghazvin, 1463m, road to Abazar village, N 36 �� 17 ��� E 50 �� 10 ���, pan trap, 16���17 June 2009, Leg. Babak Gharali [♂♀] (PCDG). Etymology. named for the type locality east of the Caspian Sea. Diagnosis. Perhaps one of the most readily identified Usia species, superficially more reminiscent of some Parageron. A strikingly marked black and yellow species, Both sexes easily identified by the combination of shining black disc of mesonotum, densely grey dusted marginally, largely undusted, shining black tergites with broad yellow apical margins and legs yellow but for blackish apical tarsomeres. Similarly patterned species of Parageron always have densely dusted or at least matt second to terminal visible tergite and all sympatric species have densely grey dusted disc of mesonotum. Redescription. Measurements. Body length. 3.7 ���6.0mm. Wing length. 3.3���5.2mm. Male. Head. Oral margin narrow but quite prominent above oral margin, disappearing below in lateral view, shining brown. Frons blackish in ground colour, sometimes paler medially adjacent to eye margin, grey-dusted except for small rounded shining patch, on some extending forward of antennal bases. Eyes separated at their narrowest by between one sixth and one fifth head width, about equal to width across lateral ocelli inclusive. Lateral ocellus separated from the eye margin by about half to two thirds the diameter of that ocellus. Ocellar tubercle shining, thinly dusted, subshining, eye margins in front of anterior ocellus diverging slightly, then relatively abruptly before continuing parallel to insertion of proboscis, there diverging to rear of oral opening. Fine white hairs on ocellar tubercle short, barely longer than space between lateral ocelli, frons in front of anterior ocellus glabrous. Occiput black in ground, often browner below close to oral opening, densely grey dusted, except small triangular area behind ocellar tubercle which is thinly dusted. Upper part of occiput with fairly dense covering of fine white hairs with conspicuously procurved tips, the hairs thinning out and becoming shorter below, opposite to the usual condition in Usia. Ommatidia uniform in size across the eyes. Antennae black, sometimes yellower on scape and pedicel, third segment, a little less than twice length of scape and pedicel together, narrow swollen below basally, narrowing apically, sensilla white. All antennal segments with very short fine white setulae above. Palps relatively long, yellow, barely clavate tip a little wider than oral margin, short white hairs below, those on tip hardly longer than width of palp a widest point. Proboscis relatively long, approximately equal to headthorax length, black, dorsally hairless. Thorax. Mesonotum with a shining black patch on disc of variable extent, broadly grey dusted on anterior slope of mesonotum outside paramedian vittae, leaving a shining black stripe extending to the pronotum or almost so, post-pronotal lobe, notopleuron along thoracic suture, above wing root, prohumeral area and rear quarter to third of disc up to scutellum. Disc finely punctured, the hair insertions well separated by smooth shining cuticle. Paramedian vittae can be distinct, impunctate and finely transversely striated, reaching to wing insertions, or almost invisible. Mesonotum clothed with fairly long, fine silky white vestiture, erect to slightly reclinate, tending to proclinate close to scutellum, hairs as long as those on upper occiput. Laterally the hairs are longer and erect or inclined inwards. Acrostichal hairs between the bare paramedian vittae multiserial, entirely erect to slightly reclinate, widening slightly posteriorly, dorsocentrals not distinguishable from the general vestiture. Scutellum dusted as adjacent parts of mesonotum, hairing as mesonotum, cuticle smooth, unwrinkled. Pleurae uniformly grey dusted largely obscuring ground colour, except for a small area of shining cuticle on anepisternum anterodorsal in some individuals. Fine white hairs, like those on mesonotum, on pronotum and upper half of the anepisternum. Wing. Membrane clear, lacking any infuscation, veins all yellow, darkening slightly distally. Crossvein r-m at or a little before the middle of discal cell, conspicuously beyond dm-cu. Anal lobe well developed, with strongly curved trailing edge, wider than anal cell. Haltere. Yellow, base of stem slightly brownish. Legs. Coxae brown, becoming blacker basally, with a coating of grey dust like that on the pleurae. Remainder of legs yellow, undusted, shining below the setulae, except tip of basitarsus and apical tarsal segments which are dark brown to blackish. Coxae externally covered with moderately long, white hairs, especially apically. Femora, tibia and tarsi covered with short setulae, even those on femora shorter than width of tibia. Abdomen. All tergites shining black on disc with broad yellow apical margins that continue on to down-curved margin where it narrows slightly, tergite one thinly dusted laterally. Each tergite densely punctate, the punctures somewhat transverse forming a surface sculpture that gives the appearance of transverse striations. Disc uniformly and densely covered with fine, pale yellow, recumbent hairs, longer and more erect laterally. Sternites blackish brown with thin grey dusting, faintly punctuation almost invisible, with fairly long, recumbent pale yellow hairs. Genitalia. Relatively large and conspicuous, elongate. Epandrium yellow with apicolateral extensions bluntly pointed, the tips black, in dorsal view apical margin rectangularly emarginated, shining, the cuticle distinctly punctate evenly coated short pale yellow hairs. Gonocoxite about twice as broad as long, dusky basally, yellow in apical half, the tips curving towards each other but well separated, haired as epandrium, gonostyli slightly curved, truncate with basal ���thumb��� and an external ���step���. Female. As male except as follows: oral margin decidedly broader, more convex but still disappearing below. Frons a little wider, about one-fifth head width, a little wider than distance across lateral ocelli. Mesonotal vestiture often slightly shorter. One female from Armenia with black coxae, trochanter and all but tip of femora. Genitalia. Tip of abdomen coming to a rounded point; apical sternite rather rectangular, yellow, the basal part partially divided from the main part, no apical notch present but short median projection. Main part of apical sternite with raised basal boss, sclerite clothed with very short white hairs. Furca typical of the group, with rounded tip sometimes with short spine apically, arms long, conspicuous rectangular, dark-pigmented vaginal plate. Discussion. This very distinctive species is by far the most differentiated of the lata -group. Apart from the very obvious pattern of yellow, it has more extensive mesonotal dusting, longer occipital hairs above than below, smoother more finely punctured mesonotum and scutellum and broader anal lobe than in other species. In the latter two characters it approaches the condition seen in some Parageron with similar abdominal pattern. However, the male and female genitalia clearly place it in the lata -group and lack the diagnostic characters of Parageron so the similarities with Parageron are coincidental. Only five specimens have been seen which exhibit considerable variation in size, extent of mesonotal dusting and leg colour in the females. When much more material becomes available it will be worth looking to see if there is more than one species involved. However, amongst the four dissected specimens there no clear evidence of more than a single species. Distribution. Armenia, western Iran, eastern Turkey and Turkmenistan, poorly studied areas but does not appear to be very common., Published as part of Gibbs, David, 2014, A world revision of the bee fly tribe Usiini (Diptera, Bombyliidae) Part 2: Usia sensu stricto, pp. 1-85 in Zootaxa 3799 (1) on pages 45-47, DOI: 10.11646/zootaxa.3799.1.1, http://zenodo.org/record/286090, {"references":["Paramonov, S. J. (1950) Bestimmungstabelle der Usia - Arten der Welt (Bombyliidae, Diptera). Eos, 26, 341 - 78."]}