Parageron, Paramonov, 1929

Gibbs, David, 2023, A world review of the bee fly tribe Usiini (Diptera, Bombyliidae) - Part 3: Parageron Paramonov s. lat., European Journal of Taxonomy 863 (1), pp. 1-162 : 8-31

Key to the species of Parageron , Parusia and Protypusia

[♂ Protypusia emeljanovi , ♂ Protypusia modesta ( Loew, 1873) , ♂ Protypusia tewfiki ( Efflatoun, 1945) , ♂ Protypusia xizangensis (Yang & Tang, 1994) unknown]

1. Males ................................................................................................................................................. 2

– Females ........................................................................................................................................... 32

2. A discrete round or oval brown to shiny black spot on either side of the mesonotum on the thoracic suture ( Fig. 2 , pt. 1-TSS) (can be small and inconspicuous in some pale-yellow species, do not confuse with brown spot behind thoracic suture above wing base in some species) ....................... 3

– Without these discrete spots on thoracic suture, mesonotum either uniformly dusted or with longitudinal vittae of various forms, sometimes a vague brown spot behind the thoracic suture .... 6

3. Proboscis about 1–2× head length (excluding antennae); predominantly yellow, darkened at most on apical third of labium/labella, labrum swollen basally ( Fig. 1 ) (only slightly in pale yellow variant and specimens from Israel which probably belong here) [very variable and possibly more than one species] ................................................................................... ♂ Parageron lutescens ( Bezzi, 1925)

– Proboscis 2.5–4 × head length (excluding antennae); predominantly black, particularly labium/ labella, labrum dark up to the point where it starts to widen basally, basal swelling less striking, never wider than distance between outer edges of scape ( Fig. 2 ) ..................................................... 4

4. Eyes very narrowly separated, by less than the diameter of an adjacent facet, proboscis black, 2.5–3.5 × head length, not swollen basally. Thorax, including post pronotal lobe and scutellum, dark in ground colour under whitish dust; legs, including coxae, darkened with paler knees ............................................................................. ♂ Parageron erythraeus ( Greathead, 1967)

– Eyes confluent for a distance equal or more than length of ocellar triangle, proboscis black, 2–4 × head length, dark yellow and slightly swollen basally. Thorax yellow with darker disc beneath whitish dust, post pronotal lobe and scutellum yellow in ground colour; legs, including coxae, yellow ..... 5

5. In ventral view apex of epiphallus pickaxe-shaped, epandrium more elongate and parallel-sided ( Fig. 38a, e ). The limited material seen larger, frons almost bare, the longest hairs much shorter than pedicel, dorsal hairs on scape and pedicel longer, postpedicel lacking accessory points ( Fig. 3 ). Central Asia ....................................................... ♂ Parageron orientalis Paramonov, 1929 stat. rev.

– In ventral view apex of epiphallus simply rounded, without pointed lateral extensions, epandrium less elongate, trapezoidal, widest at base ( Fig. 37a, e ). Frons clearly pubescent, the longest hairs almost as long as the pedicel, dorsal hairs on scape and pedicel quite evident, some almost as long as pedicel, postpedicel usually bifid apically with a dorsally directed point just beyond the sulcus ( Fig. 4 ). Eastern Mediterranean (including Cyprus) .................... ♂ Parageron longilingua sp. nov.

6. Eyes touching (holoptic) ................................................................................................................... 7

– Eyes separated (dichoptic), sometimes only narrowly so ............................................................... 20

7. Frons and gena with conspicuous long hairs, some longer than the basal antennal segment (scape) ( Fig. 5 ) .............................................................................................................................................. 8

– Frons and gena practically bare, lacking hairs longer than the basal antennal segment (scape), if some hairs nearly as long as scape these confined to frons, gena below insertion of antennae bare ( Fig. 6 ) ............................................................................................................................................ 10

8. Silvery-grey species, tergites basally matt black contrasting with dove-grey apical half or more, margins narrowly yellow. Morocco..................................... ♂ Protypusia argentata gen. et sp. nov.

– Tergites matt black, dorsally no grey dusting divides the blackish colour from the clear yellow apical margins. Algeria, Morocco, Tunisia, Spain and France.................................................................... 9

9. Smaller species, long hairs on frons and gena erect, black to brown (sometimes pale brown), the longest hairs almost reach the tip of the antennae ( Fig. 5 ). Mesonotum matt black with three grey dusted stripes, acrostichal line short, dorsocentral lines almost reaching scutellum .......................... ......................................................................... ♂ Protypusia grata (Loew, 1859) gen. et comb. nov.

– Larger species, hairs on frons and gena tending to be more down-curved or wavy-tipped, silky white, falling well short of antennal length (although exceeding scape plus pedicel length) ( Fig. 7 ). Grey dusted mesonotal stripes bolder, acrostichal line extending beyond middle, dorsocentral lines coalescing in front of scutellum ........... ♂ Protypusia incisa ( Wiedemann, 1830) gen. et comb. nov.

10. Wing 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 ( Fig. 8 ). Morocco and Tunisia.......................................................... .................................................................... ♂ Protypusia vagans ( Becker, 1906) gen. et comb. nov.

– Wing clear ........................................................................................................................................11

11. Paramedian vittae blackish or brown and confluent on disc, any dusted acrostichal line only apparent anteriorly, fading out and becoming indistinct approximately at thoracic suture ( Figs 9 , 11–12 ). [Halteres with clear dark dorsal spot.] ............................................................................................ 12

– Mesonotum densely grey, brownish or olive dusted paramedian and antehumeral vittae either poorly defined, or if darker and distinct, dusted acrostichal line clear and merging with general dusting behind end of paramedian vittae ( Fig. 10 ). Halteres with knob pale or infuscated. [Scutellum grey dusted, concolourous with paler parts of mesonotum.] .................................................................. 14

12. Velvety black species, legs black, grey dusted dorsocentral lines barely reaching level of wing bases, tergites matt black without paler apical margins, scutellum matt black ( Fig. 9 ) ................................ ....................................................................... ♂ Protypusia ornata ( Engel, 1932) gen. et comb. nov.

– Dark brown species, knees obscurely paler reddish-brown, grey or brown dusted dorsocentral lines exceeding level of wing bases, continuing to the scutellum (sometimes only faintly) ( Figs 11–12 ), tergites dark brown or olive-grey dusted, sternites and often tergites with very narrow paler apical margins, scutellum may be dark brown dusted in part or entirely .................................................. 13

13. Palps tiny, brownish, slender with a few fine apical hairs. Dorsal surface of scape and pedicel with pubescence shorter than length of these segments. Mesonotum blackish or brown dusted with three paler greyish dusted lines, the dorsocentral ones continuing faintly to scutellum ( Fig. 11 ). Tergites with very narrow paler apical margins ............ ♂ Protypusia hyalipennis ( Séguy, 1941) gen. et comb. nov.

– Palps conspicuously larger, yellow, clavate with a fan of long apical hairs. Dorsal surface of scape and pedicel with pubescence longer than length of these segments. Mesonotum blackish brown dusted with two clear paler brown dorsocentral lines continuing strongly to scutellum ( Fig. 12 ). Tergites lacking paler apical margins ................................................................. ♂ Protypusia flavipalpis gen. et sp. nov.

14. Palps white. A very small dove-grey dusted species from Central Asia. [Reflexed lateral margin of tergite one predominantly yellow, reflexed margin of remaining tergites not contrasting with brown dusted disc, this possibly an artefact of the condition of the holotype ♂ examined? Epandrium dark in ground colour or only narrowly yellow.] Genitalia very different to the following species .......... ............................................................... ♂ Protypusia grisea ( Paramonov, 1947) gen. et comb. nov. N.B. From here on several species are included where specimens could not be examined and genitalia have not been studied. Characters for these are taken from type descriptions and photographs. Always double check by dissection and comparison with reference material.

– Palps black to brownish, at least partially (if palps very small can be impossible to see in poor specimens). If palps partly pale yellowish then larger species with wing length> 4 mm. Small to medium species from North Africa, Central Asia and the Middle East .......................................... 15 [♂ Protypusia xizangensis ( Yang & Yang, 1994) gen. et comb. nov. will probably run here but ♂ unknown] [♂ Protypusia modesta ( Loew, 1873) gen. et sp. nov. will probably run here but ♂ unknown]

15. Postpedicel relatively robust, ratio of depth:length 1:1.9–2.6. Ventral margin usually convex throughout (sometimes straight basally) curving dorsally evenly to tip in apical half so narrowly blunt-ended ( Fig. 13 ) ...................................................................................................................... 16

– Postpedicel elongate length:depth 1:2.4–3.1. Ventral margin straight or even slightly concave in apical half, abruptly curved up at tip so broadly blunt-ended ( Fig. 14 ) ......................................... 17

16. Mesonotum densely dove-grey dusted with paramedian and antehumeral vittae indistinct, slightly darker grey. Moroccan specimens have middle of mesonotum browner. Frons typical. Wing hyaline .................................................... ♂ Protypusia dimonica ( Zaitzev, 1996) gen. et comb. nov.

– Mesonotum brownish-grey to deep olive-buff dusted with paramedian and antehumeral vittae better defined, darker brown. Frons more prominent [by extrapolation from female distance from antennal insertion to eye greater than depth of postpedicel but no male specimen examined]. Wings with distinct light brownish tinge ............ ♂ Protypusia deserticola ( Efflatoun, 1945) gen. et comb. nov. [The unknown ♂ of Protypusia tewfiki ( Efflatoun, 1945) gen. et comb. nov. from Egypt will probably key here. Should be readily identified if it has same unique abdominal pattern seen in the ♀. See Fig. 57 ]

17. Mesonotum densely pale dove-grey dusted with poorly defined paramedian and antehumeral vittae where dusting a little thinner so only slightly subshining ( Fig. 10 ). Pale creamy apical margins to tergites about as wide as width of basitarsus. Haltere yellowish-white, unmarked ........................ 18

– Mesonotum more brownish-grey dusted with darker blackish-brown paramedian and antehumeral vittae much more obvious, thinly dusted and clearly shining. Pale margins to tergites often narrower than basitarsus. Haltere knob either pale or brown infuscated ....................................................... 19

18. Tibia clear yellow, or at least in basal quarter dorsally, hind pair with greatest tendency to be infuscated apically and dorsally, especially so in Iranian specimens. The slightly darker paramedian vittae distinctly wider than acrostichal line (at least in Iranian specimens). Slightly larger species wing length usually> 4 mm. Central Asia, Iran................................................................................. ............................................................... ♂ Protypusia zimini ( Paramonov, 1947) gen. et comb. nov. [♂ Protypusia emeljanovi ( Zaitzev, 1975) gen. et comb. nov. will probably run here but ♂ unknown]

– Legs black or dark brown with yellow colour of knees very narrow, extending down tibia less than median width of tibia. The slightly darker paramedian vittae distinctly narrower than acrostichal line. Slightly smaller species, wing length <4 mm. Israel.................................................................. .................................................................... ♂ Protypusia negevi ( Zaitzev, 1996) gen. et comb. nov.

19. Smaller species, body length <4 mm. Mesonotum relatively thinly grey-brown dusted, the blackishbrown paramedian vittae very thinly dusted and clearly shining. Paramedian vittae on disc as wide or wider than acrostichal line. Pale colour on tibia often more extensive, extending one quarter to one third down from base dorsally. Haltere often with distinct brown dorsal spot or at least brown infuscation at base of knob. Saudi Arabia.......................................................................................... ............................... ♂ Protypusia raydahensis (El-Hawagry & Al Dhafer, 2016) gen. et comb. nov.

– [From Plate ( Fig. 49 ) and original description, no specimen seen.] Larger species, body length> 4 mm. Mesonotum relatively more densely grey dusted, the blackish paramedian vittae narrower than the acrostichal line on the disc. Pale colour on tibia very narrow, confined to extreme base. Knob of haltere whitish, unmarked. Egypt and (? Senegal) ................................................................ ................................................................... ♂ Protypusia inornata ( Engel, 1932) gen. et comb. nov. [♂ Usia grisea Efflatoun, 1945 will probably run here if it belongs in Usiini . Haltere knob with brown spot, very small species from Egypt. See Fig. 68 .]

20. Hairs of frons continuing on to gena, below level of insertion of antennae at least a few hairs conspicuously longer than the basal two antennal segments (scape and pedicel) ( Fig. 5 ) ............. 21

– Hairs of frons not continuing down gena, stopping at about the level of insertion of antennae, any hairs below this conspicuously shorter than the basal two antennal segments (scape and pedicel) ( Fig. 6 ) ............................................................................................................................................ 22

21. Acrostichal line grey dusted similar to the dorsocentral lines, viewed from directly above, scutellum dark matt brown concolourous with darker parts of mesonotum. Canary Islands ............................. ........................................................................ ♂ Protypusia striata ( Báez, 1982) gen. et comb. nov.

– Acrostichal line brown dusted contrasting with the grey dusted dorsocentral lines, viewed from directly above, scutellum grey-brown dusted decidedly paler than the darker parts of mesonotum. Eastern Mediterranean ( Egypt, Israel) ...... ♂ Protypusia separata Gibbs & Theodor gen. et sp. nov.

22. At least tips of femora and bases of tibia (knees) yellow ............................................................... 23

– Legs entirely black .......................................................................................................................... 25

23. Wings with brownish spots at base of R 2+3, base of R 4, and over crossveins r-m, m-m and m-cu; eyes clearly separated, usually by more than the diameter of the front ocellus; genitalia larger ............... ............................................................. ♂ Protypusia punctipennis ( Loew, 1846) gen. et comb. nov.

– Wings clear, unmarked; genitalia smaller; eyes usually separated by no more than the diameter of front ocellus .................................................................................................................................... 24 N.B. The reliability of characters used to separate the following two species is uncertain so best to check epiphallic complex

24. Knob of halteres white; a narrow, defined, dark prescutellar vittae present; hairs on tergites 2–4 longer than respective tergite ................................................... ♂ Protypusia kerkini gen. et sp. nov.

– Knob of halteres with distinct dorsal brown spot; area of mesonotum immediately in front of scutellum with broad, diffuse darker patch; hairs on tergites 2–4 shorter than respective tergite ...... ............................................................................................. ♂ Protypusia strymonas gen. et sp. nov. N.B. In the following species external characters are subtle, difficult to interpret and could easily be lost through damage or wear. Close examination of epiphallus in situ or after dissection needed to confirm identity.

25. Epandrium with apico-lateral corners exerted (often visible in undissected insect), hypopygium very large relative to size of insect, as large as remainder of abdomen ( Fig. 15 ) [paramedian vittae narrower than median line, hairs on frons variable, on average a smaller insect] .............................. ........................................................................................................ ♂ Parusia faesae gen. et sp. nov.

– Epandrium with apico-lateral corners rounded, hypopygium smaller relative to size of insect, not as large as remainder of abdomen ( Fig. 16 ) [paramedian vittae often equal to or wider than median line, on average a larger insect] .............................................................................................................. 26

26. Hairs on front part of frons short and inconspicuous, shorter than scape and pedicel together ( Fig. 17 ) (if anal lobe very narrow check ♂ Usia martini François, 1969 see Gibbs 2011) ......................... 27

– At least some hairs on front part of frons longer than the scape and pedicel together ( Fig. 18 ) .... 29

27. Tip of epiphallus viewed laterally rather slender, only slightly expanded apically ( Fig. 65a ). [Paramedian and antehumeral vittae usually with no more than a faint sprinkling of brownish dust, the shining black integument of the mesonotum clearly evident] ...................................................... ............................................................................ ♂ Parusia loewi ( Becker, 1906) gen. et comb. nov.

27a. A much darker insect with the grey dusting of the mesonotum much reduced and thinner .............. .................................................................................................................. ♂ Parusia loewi dark form

27b. A paler insect looking much more like typical aurata ; grey dusting on mesonotum more extensive and denser such that shining paramedian and antehumeral vittae narrower than the grey dusted median and dorsocentral lines ................................................................. ♂ Parusia loewi pale form

– Tip of epiphallus viewed laterally conspicuously expanded apically, either T-shaped or fish-tailed with thick stem. Paramedian and antehumeral vittae dark brown dusted, completely or largely obscuring the shining black integument of the mesonotum, at most subshining ............................ 28

28. Eyes separated by at least twice the diameter of the front ocellus. Genitalia characteristic, particularly the shape of the epiphallus ( Fig. 67a–b ). Algeria, Tunisia, Corsica, Sardinia ....................................... ......................................................... ♂ Parusia taeniolata (Costa, 1883) stat. rev., gen. et comb. nov.

– Eyes separated by little more than the diameter of the front ocellus. Genitalia characteristic, particularly the shape of the epiphallus ( Fig. 66a–b ). Spain........................ ♂ Parusia propinqua gen. et sp. nov.

29. Inner margin of gonostylus at base strongly incised, more or less a semicircle ( Fig. 20 ). Frons narrower, eyes separated by about the depth (height) of the postpedicel. Hairs on front of frons clearly shorter than those on ocellar tubercle ( Fig. 19 ) .................................................................. 30

– Inner margin of gonostylus at base straighter, much less than a semicircle (but may be sharply bent in apical half) ( Fig. 22 ). Frons wider, eyes separated by almost twice the depth (height) of the postpedicel. Some hairs on front of frons as long as those on ocellar tubercle ( Fig. 21 ) ............... 31

30. Dark thoracic vittae relatively poorly marked, thinly dusted so only a little more shining than the rest of the thorax. Genitalia characteristic ( Fig. 60a–d ). Spain.......... ♂ Parusia almeria gen. et sp. nov.

– Dark thoracic vittae relatively well marked, hardly dusted so shining cuticle clearly visible. Genitalia characteristic ( Fig. 62a–e ). Morocco........................................... ♂ Parusia benoisti gen. et sp. nov.

31. Apical third of gonostylus bent in almost at a right angle ( Fig. 22 ). Epiphallus diagnostic ( Fig. 61a– b ) [genitalia smaller, yellow band on tergite one wider and paler yellow]. Morocco / Algeria........... ...................................................................... ♂ Parusia aurata ( Fabricius, 1794) gen. et comb. nov.

– External curve of gonostylus more even, only sharply bent at tip. Epiphallus diagnostic ( Fig. 63a–b ) [genitalia larger, yellow band on tergite one narrower and darker yellow]. Libya............................. ................................................................................................... ♂ Parusia cyrenaica gen. et sp. nov. FEMALES [ Usia grisea could possibly belong here, ♀ unknown]

32. Frons hairs continuing on to gena, some of the hairs below the antennal insertion longer than the scape ( Fig. 23 ) ................................................................................................................................ 33

– Any pubescence on frons not usually extending on to gena, any hairs below the antennal insertion shorter than the scape ( Fig. 24 ) ....................................................................................................... 37

33. Gena (dusted area not including more shiny mouth margin) at its narrowest part as wide as or wider than width of proboscis level with the tips of the palps ( Fig. 25 ) .................................................. 34

– Gena at its narrowest part narrower than width of proboscis level with the tips of the palps ( Fig. 26 ) .......................................................................................................................................... 36

34 Hairs on frons and gena silky white, somewhat shaggy and barely longer than scape and pedicel combined, continuing unbroken all the way to the ventral side of the head; hairs on dorsolateral surface of proboscis minute ................. ♀ Protypusia incisa ( Wiedemann, 1830) gen. et comb. nov.

– Hairs on frons and gena dark brown to almost white, some clearly longer than scape and pedicel combined, lower part of the gena hairless or almost so ( Fig. 23 ); hairs on dorsolateral surface of proboscis basally longer and conspicuous ...................................................................................... 35

35. Hairs on frons and gena blackish to brown; mesonotum with conspicuous, darker brown to blackish paramedian and antehumeral vittae; tergites dorsally dull black with sharply defined yellow apical margins ............................................................ ♀ Protypusia grata (Loew, 1859) gen. et comb. nov.

– Hairs on frons whitish, on gena tinged yellowish or white; mesonotum with paramedian and antehumeral vittae faint, just perceptibly darker grey than general pale grey dusting; abdomen pale grey dusted, each tergite more or less dull black basally with sharply defined yellow apical margins .............................................................................................. ♀ Protypusia argentata gen. et sp. nov.

36. Dark paramedian vittae no wider than median acrostichal line, antehumeral spot in front of thoracic suture narrower than grey dusted dorsocentral lines. Canary Islands ................................................ ........................................................................ ♀ Protypusia striata ( Báez, 1982) gen. et comb. nov.

– Dark paramedian vittae wider than median acrostichal line, antehumeral spot in front of thoracic suture wider than grey dusted dorsocentral lines. Eastern Mediterranean, Egypt.............................. ................................................................... ♀ Protypusia separata Gibbs & Theodor gen. et sp. nov.

37. Disc of mesonotum either yellow or the disc darkened under the pale dust and with a discrete, velvet-black to brown spot either side on the thoracic suture (do not confuse with less discrete spot sometimes present behind the thoracic suture above wing base) ................................................... 38

– Mesonotum either uniformly pale grey dusted, or with longitudinal darker vittae, lacking discrete velvety spots on the thoracic suture ................................................................................................ 41

38. Viewed dorsally, proboscis entirely black, no perceptible swelling basally. Scape whitish, contrasting with darker pedicel and blackish postpedicel. Eritrea..... ♀ Parageron erythraeus ( Greathead, 1967)

– Viewed dorsally, proboscis not entirely black, at least yellowish basally where distinctly, although often only slightly swollen. Scape yellow, hardly contrasting with pedicel and postpedicel, latter might be a shade darker ......................................................................................................................39 N.B. Some females of the next three species may not be reliably separable without associated males. Intergrades appear to occur and there could be unrecognised species here.

39. Proboscis short, 1–2 × head length (excluding antennae); proboscis almost entirely yellow, only darkened on apical quarter of labium/labella; base of labium often conspicuously swollen ............. ................................................................................................. ♀ Parageron lutescens ( Bezzi, 1925)

– Proboscis longer, 2.5–4 × head length (excluding antennae), mostly black, only yellow basally and never strongly swollen, from above narrower than width across basal antennal segments ............ 40

40. Larger, hairs on frons finer, shorter and more steeply inclined. Third antennal segment pointed, no obvious accessory point beyond sulcus (based on single specimen) ( Fig. 27 ). Iran, Central Asia .... ............................................................................ ♀ Parageron orientalis Paramonov, 1929 stat. rev,

– Smaller, hairs on frons a little longer and inclined closer to 45°, more bristle-like. Third antennal segment usually bifid apically, a more or less conspicuous point dorsally immediately beyond sulcus (can be missing and simply pointed as in Par. orientalis , best identified by associated males) ( Fig. 28 ). Eastern Mediterranean (including Cyprus) .................. ♀ Parageron longilingua sp. nov.

41. Tibia largely or entirely yellow ....................................................................................................... 42

– Tibia largely darkened, base and apex sometimes paler, sometimes basal half of front- and mid-tibia yellow ............................................................................................................................................. 46

42. Mesonotum pale grey dusted with slightly darker paramedian and antehumeral vittae only faintly indicated .......................................................................................................................................... 43

– Mesonotum yellow to greyish-brown dusted with clear dark brown paramedian and antehumeral vittae ............................................................................................................................................... 44

43. Scape dark, coxae with at least base dark, mesonotum and scutellum dark in ground colour ........... ............................................................... ♀ Protypusia zimini ( Paramonov, 1947) gen. et comb. nov.

– Scape yellow, front coxae yellow, mid and hind coxae dark, mesonotum laterally and scutellum wholly yellow in ground colour .......... ♀ Protypusia emeljanovi ( Zaitzev, 1975) gen. et comb. nov. [Based on type description, no specimens seen.]

44. Wings with brownish spots at base of R 2+3, base of R 4, and over crossveins r-m, m-m and m-cu ..... ............................................................. ♀ Protypusia punctipennis ( Loew, 1846) gen. et comb. nov.

– Wing clear, without brown spots over veins ................................................................................... 45

45. Halteres with white knob; mesonotum immediately in front of scutellum with well demarcated narrow dark prescutellar vittae ................................................ ♀ Protypusia kerkini gen. et sp. nov.

– Haltere knob with distinct dark brown spot dorsally; mesonotum immediately in front of scutellum with diffuse darker area ...................................................... ♀ Protypusia strymonas gen. et sp. nov.

46. Tergites 2–4 blackish with broad whitish-grey dusted median longitudinal stripe and whitish-grey lateral dusting visible in dorsal view. [All tergites with narrow pale apical margins, haltere white.] ( Fig. 57 ) Egypt........................................ ♀ Protypusia tewfiki ( Efflatoun, 1945) gen. et comb. nov.

– Tergites 2–4 uniformly coloured in dorsal view, either entirely dusted or entirely matt or velvety black other than yellow apical margins and occasionally grey dusting on recurved lateral margins of tergites can be visible from above .................................................................................................. 47

47. Mesonotum relatively densely pale grey dusted with no more than faintly darker paramedian and antehumeral vittae ( Fig. 29 ); if browner with evident mesonotal vittae then either wing infuscated on forks and cross-veins or palp pale yellowish ............................................................................. 48

– Mesonotum more thinly dusted grey or brown with contrastingly darker, more shining, brown or blackish paramedian and antehumeral vittae ( Fig. 30 ) ................................................................... 51 [If anal lobe very narrow check Usia martini François, 1969 see part 1 Gibbs 2011.]

48. Halteres with discrete dark-brownish infuscation on one side where stem meets knob ................. 49

– Halteres entirely white to pale yellow ............................................................................................ 50

49. Wings with faint but distinct brownish infuscation over cross-veins and forks; palps black; a browner dusted species ............................................ ♀ Protypusia vagans ( Becker, 1906) gen. et comb. nov.

– Wings clear, cross-veins and forks unmarked; palps pale yellowish; a greyer dusted species ........... ............................................................................................. ♀ Protypusia flavipalpis gen. et sp. nov.

N.B. The following four are all dove-grey species, often rather variable, and in two cases very little material is available. Not certainly separable on external characters; genitalia provide good characters but these do not divide dichotomously so read all four alternatives and check illustrations.

50. Tip of genital fork weakly sclerotised, although curved dorsally naturally, easily straightened and flattened. Vaginal plate large, patchily sclerotised, attached to genital fork by membranes only ( Fig. 45f–g ) ............................................ ♀ Protypusia dimonica ( Zaitzev, 1996) gen. et comb. nov.

– Tip of genital fork sclerotised such that it cannot be easily bent flat, but not strongly pigmented, curved back on itself and coming to a blunt point. Vaginal plate small and not pigmented, attached to genital fork by membrane only. Basal spermathecal duct swollen close to vaginal plate ( Fig. 51a– b ) ................................................................ ♀ Protypusia modesta ( Loew, 1873) gen. et comb. nov.

– Tip of genital fork strongly sclerotised and at least partly pigmented, strongly bent back on itself such that it cannot be flattened out, tip of genital fork ending bluntly, truncated. Arms of genital fork relatively short compared to the other three species. Vaginal plate small but strongly pigmented, attached to genital fork by membrane. Basal spermathecal duct not swollen close to vaginal plate ( Fig. 47e–f ) ........................................... ♀ Protypusia grisea ( Paramonov, 1947) gen. et comb. nov.

– Tip of genital fork strongly sclerotised and brown pigmented, strongly bent back on itself such that it cannot be flattened out, tip of genital fork ending bluntly or tapering to a point. Vaginal plate relatively large and strongly pigmented, attached to sclerotised parts of the genital fork. Basal spermathecal duct not swollen close to vaginal plate ( Fig. 52f–g ) ..................................................... .................................................................... ♀ Protypusia negevi ( Zaitzev, 1996) gen. et comb. nov.

51. Legs with contrasting paler knees. Abdomen with clear, if in some cases narrow pale yellow apical margins to the middle tergites ......................................................................................................... 52

– Legs entirely black, lacking paler knees. Tergites with or without pale apical margins ................ 55 N.B. From here on several species are included where specimens could not be examined and genitalia have not been studied. Characters for these are taken from type descriptions and photographs. Always double check by dissection and comparison with reference material.

52. Postpedicel elongate, parallel sided with broadly roundly blunt tip, often ventral margin of postpedicel in apical half concave, even if only slightly (similar to male Fig. 14 ). All tergites with clear pale brownish to yellow apical margins, even if narrow ........................................................................ 53

– Postpedicel short and deep, ventral margin convex and curving dorsally at tip to form a narrowly blunt tip (similar to male Fig. 13 ). In doubtful cases tergites with narrow, dull yellow apical margins from second to seventh tergites, broadest laterally, very obscure dorsally ..................................... 54 [ Usia grisea Efflatoun, 1945 , if it belongs in Protypusia , will probably run here. ♀ unknown.]

53. Larger species, body length> 4 mm. Tergites all with very broad yellow apical margins, often occupying a half or more of the visible length of respective tergite. Mesonotum greyer with paramedian and antehumeral vittae clearly visible by virtue of thinner dusting so more blackish than general dusting. Mesonotum anterior to scutellum lacking prescutellar vittae. Egypt and (? Senegal) ................................................................... ♀ Protypusia inornata ( Engel, 1932) gen. et comb. nov.

– Smaller species, body length <4 mm. Tergites with narrower pale apical margins, less than a half the length of the respective tergite. Mesonotum browner dusted, the paramedian and antehumeral vittae strongly marked dark brownish, relatively shiny. Mesonotum often with brown prescutellar vittae. Saudi Arabia.......... ♀ Protypusia raydahensis (El-Hawagry & Al Dhafer, 2016) gen. et comb. nov.

54. Postpedicel very robust, deep and strongly convex below. Frons more hairy, hairs on ocellar tubercle longer than distance between hind ocelli. Frons more prominent than typical, distance from base of antennae to eye greater than length of scape and pedicel combined [genital fork unknown]. Egypt.......................................................... ♀ Protypusia deserticola ( Efflatoun, 1945) gen. et comb. nov.

– Postpedicel less deep, but still with more or less convex ventral margin. Frons less hairy, hairs on ocellar tubercle barely longer then distance between posterior ocelli. [Genital fork only gently curved dorsally, weakly sclerotised apically, vaginal plate separated from main part of genital fork]. Morocco............................................... ♀ Protypusia hyalipennis ( Séguy, 1941) gen. et comb. nov.

55. Abdomen densely grey dusted, tergites 2–4 variably blackish basally, any very narrow pale apical margins confined to tergite one and apical tergites. Frons about a third head width, eye margins straight and diverging uniformly. Dusted gena clearly evident, together with more shining mouth margin approximately equal to the depth of the postpedicel .............................................................. ....................................................................... ♀ Protypusia ornata ( Engel, 1932) gen. et comb. nov.

– Abdomen matt black with grey dusting confined to reflexed lateral parts of tergites, all tergites with sharply demarcated yellow margins, even if narrow. Frons about a quarter head width, eye margins sinuate, more parallel sided near vertex. Dusted gena very narrow, together with more shining mouth margin obviously narrower than the depth of the postpedicel ........................................................ 56 N.B. The following species are not certainly separable if associated males are not available, even genitalia might not distinguish all specimens.

56. Basal spermathecal ducts thickened, about 2–3 × the diameter of the apical spermathecal ducts, if only 2× then much longer than ejection apparatus ( Fig. 61e–f ). Not Iberian ................................ 57

– Basal spermathecal ducts not thickened same diameter as the apical spermathecal ducts. Iberia..... ......................................................................................................................................................... 61

57. Basal spermathecal duct only slightly thickened, about 2× diameter of apical spermathecal duct. Proximal part of genital fork ‘waisted’ narrower at mid-point than at base and tip ( Fig. 61e–f ) ....... ...................................................................... ♀ Parusia aurata ( Fabricius, 1794) gen. et comb. nov.

– Basal spermathecal duct much thickened, about 3 × diameter of apical spermathecal duct ........... 58

58. Proximal part of genital fork in lateral view in line with or only shallowly angled relative to arms. ......................................................................................................................................................... 59

– Proximal part of genital fork in lateral view strongly angled relative to arms ............................... 60

59. Proximal part of genital fork simple, rod shaped (only one specimen). Basal spermathecal ducts longer than ejection apparatus and, if stretched out, exceeding tip of genital fork ( Fig. 63f ). Only known from type series from Al Jabal al Akhdar Mountains, Libya.... ♀ Parusia cyrenaica gen. et sp. nov.

– Proximal part of genital fork distally broad, converging proximally to a rounded tip. Basal spermathecal duct only a little longer than ejection apparatus and, if stretched out, not exceeding tip of genital fork ( Fig. 67e ). Tyrrhenian Islands and north Algeria and Tunisia.................................... ....................................................... ♀ Parusia taeniolata (Costa, 1883) stat. rev., gen. et comb. nov.

60. Proximal part of genital fork at an angle of less than 90° relative to arms. Proximal part of genital fork relatively narrow, tapering to a point. Arms relatively slender ( Fig. 62f–g ). Confined to northwest Morocco....................................................................................... ♀ Parusia benoisti gen. et sp. nov.

– Proximal part of genital fork at an angle of 90° or a little more relative to arms. Proximal part of genital fork broadest beyond middle, bluntly pointed. Arms relatively robust ( Fig. 64f–g ). Only known from northeast Morocco..................................................... ♀ Parusia faesae gen. et sp. nov.

61. Paramedian and antehumeral vittae shining, undusted or subshining, blackish, with a thin coating of dust ........................................................... ♀ Parusia loewi ( Becker, 1906) gen. et comb. nov. [part]

– Paramedian and antehumeral vittae dusted, not shining ................................................................. 62

62. Hairs on front part of frons short, much shorter than hairs on ocellar tubercle and shorter than length of scape and pedicel combined ( Fig. 31 ) ............................................................................................ ..................... ♀ Parusia loewi [part], ♀ Parusia propinqua gen. et sp. nov. (not reliably separated)

– Hairs on front part of frons relatively long, some almost as long as those on ocellar tubercle and as long as length of scape and pedicel combined ( Fig. 32 ) .............. ♀ Parusia almeria gen. et sp. nov.

Figures

Fig. 2. Parageron longilingua sp. nov., ♂.

Fig. 1. Parageron lutescens (Bezzi, 1925), ♂.

Fig. 38. Parageron orientalis Paramonov, 1929 stat. rev. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Epandrium lateral. g. Female genitalia ventral. h. Female sternite 8 ventral.

Fig. 3. Parageron orientalis Paramonov, 1929 stat. rev., ♂.

Fig. 37. Parageron longilingua sp. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Female genitalia ventral.

Fig. 4. Parageron longilingua sp. nov., ♂.

Fig. 5. Protypusia grata (Loew, 1859) comb. nov., ♂.

Fig. 6. Protypusia negevi (Zaitzev, 1996) gen. et comb. nov., ♂.

Fig. 7. Protypusia incisa (Wiedemann, 1830) gen. et comb. nov., ♂.

Fig. 8. Protypusia vagans (Becker, 1906) gen. et comb. nov., ♂.

Fig. 9. Protypusia ornata (Engel, 1932) gen. et comb. nov., ♂.

Fig. 11. Protypusia hyalipennis (Séguy, 1941) gen. et comb. nov., ♂.

Fig. 12. Protypusia flavipalpis gen. et sp. nov., ♂.

Fig. 10. Protypusia negevi (Zaitzev, 1996) gen. et comb. nov., ♂.

Fig. 13. Protypusia dimonica (Zaitzev, 1996) gen. et comb. nov., ♂.

Fig. 14. Protypusia negevi (Zaitzev, 1996) gen. et comb. nov., ♂.

Fig. 57. Protypusia tewfiki (Efflatoun, 1945) gen. et comb. nov., habitus, Smithsonian Institution Archives. Image SIA2012-7892.

Fig. 49. Protypusia inornata (Engel, 1932) gen. et comb. nov. a. Habitus male, Smithsonian Institution Archives Image SIA2012-7885-wing venation aberrant. b–d. Protypusia cf. inornata, Senegal specimen b. Female genitalia ventral. c. Female genitalia lateral. d. Female sternite 8 ventral.

Fig. 68. Usia grisea Efflatoun, 1945, habitus, Smithsonian Institution Archives. Image SIA2012-7883.

Fig. 15. Parusia faesae gen. et sp. nov., ♂.

Fig. 16. Parusia almeria gen. et sp. nov., ♂.

Fig. 17. Parusia loewi (Becker, 1906) gen. et comb. nov., ♂.

Fig. 18. Parusia aurata (Fabricius, 1794) gen. et comb. nov., ♂.

Fig. 65. Parusia loewi (Becker, 1906) gen. et sp. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Female genitalia ventral (Marbella). g. Female genitalia ventral (Salamanca).

Fig. 67. Parusia taeniolata (Costa, 1883) stat. rev., gen. et comb. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Epandrium dorsal. e. Female genitalia ventral.

Fig. 66. Parusia propinqua gen. et sp. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Epandrium dorsal. e. Female genitalia ventral.

Fig. 20. Parusia almeria gen. et sp. nov., ♂.

Fig. 19. Parusia benoisti gen. et sp. nov., ♂.

Fig. 22. Parusia aurata (Fabricius, 1794) gen. et comb. nov., ♂.

Fig. 21. Parusia aurata (Fabricius, 1794) gen. et comb. nov., ♂.

Fig. 60. Parusia almeria sp. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Epandrium dorsal. e. Female genitalia ventral.

Fig. 62. Parusia benoisti gen. et sp. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Female genitalia ventral. g. Female genitalia lateral. h. Female sternite 8 ventral.

Fig. 61. Parusia aurata (Fabricius, 1794) gen. et comb. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Epandrium dorsal. e. Female genitalia ventral. f. Female genitalia lateral.

Fig. 63. Parusia cyrenaica gen. et sp. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Female genitalia ventral.

Fig. 23. Protypusia grata (Loew, 1859) gen. et comb. nov., ♀.

Fig. 24. Protypusia dimonica (Zaitzev, 1996) gen. et comb. nov., ♀.

Fig. 25. Protypusia argentata gen. et sp. nov., ♀.

Fig. 26. Protypusia striata (Báez, 1982) gen. et comb. nov., ♀.

Fig. 27. Parageron orientalis Paramonov, 1929 stat. rev., ♀.

Fig. 28. Parageron longilingua sp. nov., ♀.

Fig. 29. Protypusia dimonica (Zaitzev, 1996) gen. et comb. nov., ♀.

Fig. 30. Protypusia. sp. cf inornata (Engel, 1932) gen. et comb. nov., ♀.

Fig. 45. Protypusia dimonica (Zaitzev, 1996) gen. et comb. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Female genitalia ventral. g. Female genitalia lateral.

Fig. 51. Protypusia modesta (Loew, 1873) gen. et comb. nov. a. Female genitalia ventral. b. Female genitalia lateral.

Fig. 47. Protypusia grisea (Paramonov, 1947) gen. et comb. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Epandrium dorsal. e. Female genitalia ventral. f. Female genitalia lateral.

Fig. 52. Protypusia negevi (Zaitzev, 1996) gen. etcomb. nov. a. Epiphallic complex ventral.b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Female genitalia ventral. g. Female genitalia lateral. h. Female sternite 8 ventral.

Fig. 64. Parusia faesae gen. et sp. nov. a. Epiphallic complex ventral. b. Epiphallic complex lateral. c. Gonocoxite ventral. d. Gonocoxite lateral. e. Epandrium dorsal. f. Female genitalia ventral. g. Female genitalia lateral.

Fig. 31. Parusia loewi (Becker, 1906) gen. et comb. nov., ♀.

Fig. 32. Parusia almeria gen. et sp. nov., ♀.